(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Bulletin"

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




ULLETIN 




Number 1 



ICAS-A78(1) 1-87 (1979) 



LIBRAR V 

AUG 7 1979 

NEW YUr\r\ 
BOTANICAL GARDE 
APRIL 1979 



Southern California Academy of Sciences 

Founded 6 November 1891, incorporated 17 May 1907 



OFFICERS 

Takashi Hoshizaki, President 

Fred G. Hochberg, Vice President 

Richard E, Pieper, Secretary 

Joseph E. Haring, Treasurer 

Jeffrey Chapman, Editor 

Gretchen Sibley, Assistant Editor 



1977-1979 

George Callison 
Robert A. Georges 
Takashi Hoshizaki 

Alan J. Mearns 
W. Walton Wright 



BOARD OF DIRECTORS 

1978-1980 

Kristin H. Berry 
Joseph E. Haring 
Mark M. Littler 
James G. Morin 
Donald J. Reish 



1979-1981 

John Baird 

Jules Crane 

F. G. Hochberg 

Richard E. Pieper 

Gloria Takahashi 



Membership is open to scholars in the fields of natural and social sciences, and to any perso 
interested in the advancement of science. Dues for membership, changes of address, and requests fc 
missing numbers lost in shipment should be addressed to: Southern California Academy of Science; 
the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 9000' 

Annual Members $ 12. C 

Life Members 150.C 

Fellows: Elected by the Board of Directors for meritorious services. 



The Bulletin is published three times each year by the Academy. Manuscripts for publication shoul 
be sent to the appropriate editor as explained in "Instructions for Authors" on the inside back cove 
of each number. AH other communications should be addressed to the Southern California Academ 
of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, Lc 
Angeles, California 90007. 



Date of this issue 23 July 1979 



Bull. Southern California Acad. Sci. 
78(1), 1979, pp. 1-19 

Systematics and Ecology of Benthic Phyllodocidae 

(Annelida: Polychaeta) off the 

Columbia River, U.S.A. 

Michael J. Kravitz and Howard R. Jones 

Abstract. — The benthic phyllodocid polychaete fauna of the continental shelf 
off the Columbia River, northern Oregon, and southern Washington, at depths 
of 11 to 97 m, is reported. Three species of Eteone and one species of Anaitides 
are newly described. Eteone (Mysta) barbata is newly reported in the Northeast 
Pacific Ocean. Ranges are extended for Eteone longa, Anaitides hartmanae and 
Anaitides longipes, the latter two species new to Oregon and Washington; E. 
longa is new to Oregon. New records are given for Eteone calif ornica and Eulalia 
levicornuta. The local distribution, including depths and sediment type(s), of each 
species is summarized. The distributions of all species in the genus Anaitides 
overlapped, while those of some species in Eteone were relatively segregated. 
The occurrence of E. longa following the dumping of dredged sediments from 
the Columbia River mouth is discussed. 

School of Oceanography, Oregon State University, Corvallis, Oregon 97331. 
Present Address (MJK): Virginia Institute of Marine Science, Department of 
Invertebrate Ecology, Gloucester Point, Virginia 23062. 



Phyllodocids are active, mainly benthic, polychaetes which are often brilliantly 
colored in life. The systematics and phylogenetic relationships within the family 
Phyllodocidae, as well as the geographical distributions of species in the polar 
basin and the Northwest Pacific Ocean, were treated by Ushakov (1972). Phyl- 
lodocid polychaetes have been reported from the Northeast Pacific Ocean off 
California by a number of workers, most notably Hartman (1936, 1968), and off 
Oregon and Washington by Carey (1972), Hartman and Reish (1950), Fauchald 
and Hancock (in press), Banse and Hobson (1974), and Richardson et al. (1977 — 
a Final Report to the U.S. Army Corps of Engineers). Of these, only Richardson 
et al. (1977) dealt with the area off the mouth of the Columbia River. Richardson 
et al. (1977), making use of species identifications in all the major phyla, defined 
the benthic invertebrate assemblages in a 435 square km area extending from 
northern Oregon to southern Washington (Fig. 1). Of the 38 polychaete families 
represented in this area, the family Phyllodocidae was the most speciose, con- 
taining 14 species. The present paper discusses the systematics and ecology of 
those species. Other polychaetes off the Columbia River will be treated in a 
forthcoming paper. 

Methods 

1,665 quantitative samples were taken at 119 stations off the Columbia River 
from December 1974 through June 1976 using a 0.1 m- Smith-Melntyre grab on 
board the R/V Cayuse. Locations of stations are shown in Fig. 1 and the cruises 
during which they were sampled are listed in Table 1 . Six replicate grab samples 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




i i 1 r 



• 


32 • 






K i "J? 


•«,»• K2 ° 




• * *2 6 K27 


• • 


22 «27 • • . 






K36« 


»K38 ."« 







Fig. 1. Station locations off the Columbia River. 



were obtained at each station during each cruise that the station was sampled. 
Two sets of six replicate grab samples were obtained at stations 14, 25 and 26 
during cruise A. Of each set of six replicate grab samples, five were sieved 
through a 1 mm mesh screen, the polychaetes retained on the screen preserved 
in 10% Formalin and identified, and one was used for sediment analysis. In rep- 
licate sets 16E, 17E, 24D, 29D, 47A, 217D, K20E and K38E, polychaetes were 
identified from four rather than five samples. In 7B, 7C, 7D, 7E, 7G, 8B, 8C, 8D, 
8E, 8G, 96A, 97A, 98A, 99A, 148B, 149B and 150B, polychaetes were identified 
from only three samples. Twenty-one grabs were taken at station 19 during cruise 
A, polychaetes identified from 20 of them. From cruise C onward, one grab of 
every replicate set was screened through a 0.5 mm screen and later divided into 
1 mm and 0.5 mm fractions. Though a number of phyllodocid polychaetes in the 
0.5 mm fractions were too small to be identified to the species level, none ap- 
peared to differ at the species level from those encountered in the 1 mm samples. 
Specimens in the 0.5 mm fractions are not reported herein. 

Study Area 

The major feature in the study area (Fig. 1) is the Columbia River, the largest 
river on the Pacific coast of North America. The Columbia River plume is delim- 



BENTHIC PHYLLODOCIDAE 

Table I. Sampling dates (cruises) of stations off the Columbia River. 

Cruises 1 



H 



Stations 



13, 25, 26 


X 




X 


X 


X 










11, 21 




X 


X 


X 


X 










1_5, 10, 14-16, 22 


X 




X 


X 


X 




X 






6-8, 12, 17, 23 




X 


X 


X 


X 




X 






18, 19 


X 


X 


X 


X 


X 




X 






20 




X 


X 


X 


X 


X 


X 






K7, Kll, K18, K22, K26, K31 










X 


X 


X 


X 


X 


28,31, 33 








X 


X 


X 


X 


X 


X 


24 




X 


X 


X 


X 


X 


X 


X 


X 


27 








X 


X 


X 




X 


X 


29 








X 


X 


X 


X 






K16 










X 


X 




X 


X 


32 








X 


X 










216-220 








X 












Kl, K5, K9, K14, K20, K27, K28, 




















K34, K36, K38, K40 










X 










9, 47-93, 95-102 


X 


















111-152 




X 
















94, 103 


X 


X 

















1 Cruises A, B, C, D, E, F, G, H, I are designated, respectively, as the following in Richardson et 
al., 1977: C7412B, C7501D, C7504B, C7506C, C7509E, C7510E, C7601A, C7604B, C7606A. 



ited by a 32.5% c salinity isopleth which may extend to a depth of about 40 m 
(Barnes et al., 1972). Salinities at 20 m or more below the surface vary little 
throughout the year, about 32% to 34% c (Duxbury, 1972). Bottom temperatures 
at these depths off the Columbia River probably do not differ appreciably from 
those reported off central Oregon (Adriana Huyer, pers. comm.), i.e. between 
6°C and 7°C to nearly 11°C (Huyer, 1977). Bottom currents off the Columbia 
River are discussed in Barnes et al. (1972). 

Stations directly off the Columbia River and south to 46°13'N, between 
124°08'W and 124°11.5'W, exhibited large seasonal variation in sediment com- 
position (Table 2A) in concordance with seasonal variation in river flow and wave 
conditions. Bottom sediments within this area contained the least mud (silt + 
clay) during the winter, when much of it is suspended in the water column by 
long-period surface waves, and the most mud during the summer or fall, when 
high river flow and moderate wave conditions favors deposition of silts and clays 
(Kulm et al., 1975). Sediment types of seasonally sampled stations within the 
above area changed from sand to muddy sand (silty or clayey sand) or sandy 
silt — or in one case, clayey silt. Stations outside this area (Table 2B) did not show 
large temporal variations in silt + clay content. Of these, the majority were sand. 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 2A. Sediments and Depths off the Columbia River: Stations within area characterized by 
large seasonal variation in silt + clay content. 1 





Range of 


] 


Percentage Silt 


+ Clay 








Winter 


Spring 


Summer 


Fall 






Cruise A 
and/or B 














Depths 2 


+ sometimes 




Cruise 


Cruise 


Sediment 


Station 


in meters 


G 


Cruise C 


D 


E 


Type(s) 3 


13 


18-20 


3.46 


14.91 


45.75 


39.28 


Sa, SSa 


14 


31-33 


0.99-1.39 


6.27 


5.16 


29.87 


Sa, SSa 


15 


42-46 


1.59-5.04 


21.00 


63.31 


23.26 


Sa, SSi 


16 


31-37 


8.59-37.06 


34.20 


82.94 


89.18 


Sa, SSa, CSi 


17 


31-33 


14.45-24.02 


26.68 


32.98 


78.15 


Sa, SSa, CSa, SSi 


22 


22-33 


10.82-14.01 


11.69 


20.77 


27.40 


Sa, SSa 


25 


15-18 


1.66-1.77 


3.54 


28.22 


38.22 


Sa, SSa 


26 


20-29 


1.34-1.65 


16.10 


48.09 


52.37 


Sa, SSa 


65 


49 


13.18 


— 


— 


— 


Sa, — 


70 


51 


2.48 


— 


— 


— 


Sa, — 


72 


26 


1.25 


— 


— 


— 


Sa, — 


74 


35 


1.42 


— 


— 


— 


Sa, — 


76 


37 


1.42 


— 


— 


— 


Sa, — 


77 


27 


1.16 


— 


— 


— 


Sa, — 


78 


22 


0.95 


— 


— 


— 


Sa, — 


79 


20 


1.21 


— 


— 


— 


Sa, — 


80 


20 


1.02 


— 


— 


— 


Sa, — 


81 


18 


5.35 


— 


— 


— 


Sa, — 


83 


18 


2.75 


— 


— 


— 


Sa, — 


86 


26 


1.16 


— 


— 


— 


Sa, — 


87 


40 


2.57 


— 


— 





Sa, — 


88 


44 


? 


— 








? 


89 


40 


16.14 


— 


— 





Sa, — 


92 


20 


5.70 


— 


— 





Sa, — 


93 


15 


2.86 


— 


— 


— 


Sa, — 


94 


16-17 


1.62-2.54 


— 


— 





Sa, — 


95 


47 


42.76 


— 


— 





SSa, — 


103 


24-31 


5.05-10.93 


— 


— 





Sa, — 


117 


31 


1.37 


— 


— 


— 


Sa, — 


118 


37 


1.63 


— 


— 





Sa, — 


119 


18 


3.99 


— 


— 





Sa, — 


126 


40 


17.34 


— 








Sa, — 


129 


18 


2.18 


— 


— 





Sa, — 


131 


15 


2.14 


— 


— 





Sa, — 


132 


15 


2.08 


— 


— 





Sa, — 


133 


15 


1.70 


— 








Sa, — 


134 


17 


1.97 


— 








Sa, — 


135 


22 


4.54 


— 








Sa, — 


136 


31 


16.20 


— 


— 





Sa, — 


137 


37 


28.82 


— 


— 


— 


SSa,— 



The area characterized by large seasonal variation in sediment composition is delineated in para- 
graph 2 under "Study Area." 

2 Depths are not corrected for tides or waves. 

3 Sa = sand, SSa = silty sand, CSa = clayey sand, SSi = sandy silt, CSi = clayey silt. Sediment 
types referred to in this paper are those of Shepard (1954). 



BENTHIC PHYLLODOCIDAE 5 

Table 2B. Sediments and Depths off the Columbia River: Stations outside of area characterized 
by large seasonal variation in silt + clay content. 1 





Range of Depths 2 


Percentage 




Station 


in meters 


Silt + Clay 


Sediment Type(s) 


1 


17-20 


1.24-2.84 


Sand 


2 


29-35 


1.21-1.73 


Sand 


3 


45-53 


1.68-3.16 


Sand 


4 


66-70 


3.13-6.23 


Sand 


5 


82-88 


14.49-28. 33 


Sand to Silty Sand 


6 


37-45 


19.53-26.90 


Sand to Silty Sand 


7 


55-66 


22.52-42.40 


Sand to Silty Sand 


8 


80-85 


26.54-31.13 


Silty Sand 


9 


81 


23.06 


Sand 


10 


15-17 


1.12-1.23 


Sand 


11 


11-13 


1.21-5.27 


Sand 


12 


15-16 


1.73-8.13 


Sand 


18 


40-46 


1.47-12.06 


Sand 


19 


29-31 


1.25-2.03 


Sand 


20 


22-24 


0.97-5.16 


Sand 


21 


17-20 


1.32-16.91 


Sand 


23 


27-31 


10.13-29.23 


Sand to Silty Sand 


24 


24-27 


1.42-1.76 


Sand 


27 


26-29 


0.76-1.58 


Sand 


28 


26-29 


1.35-1.97 


Sand 


29 


27-30 


0.92-1.78 


Sand 


31 


24-27 


1.47-1.85 


Sand 


32 


24 


1.22-1.76 


Sand 


33 


26-30 


1.27-1.90 


Sand 


47 


88-91 


17.01 


Sand 


48 


70 


5.02 


Sand 


49 


46 


2.21 


Sand 


50 


29 


2.47 


Sand 


55 


77 


17.27 


Sand 


56 


62 


7.14 


Sand 


58 


27 


1.29 


Sand 


61 


18 


1.90 


Sand 


62 


24 


1.34 


Sand 


66 


66 


21.11 


Sand 


68 


82 


27.32 


Silty Sand 


69 


68 


15.51 


Sand 


96 


67 


21.25 


Sand 


97 


97 


31.19 


Silty Sand 


98 


86 


27.26 


Silty Sand 


99 


68 


24.25 


Sand 


100 


48 


27.07 


Silty Sand 


101 


29 


14.81 


Sand 


102 


13 


2.61 


Sand 


111 


20 


1.63 


Sand 


113 


20 


1.15 


Sand 


114 


13 


1.46 


Sand 


124 


20 


1.37 


Sand 


125 


26 


1.11 


Sand 


141 


33 


18.38 


Sand 


146 


40 


60.48 


Sandy Silt 


147 


60 


33.34 


Silty Sand 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 2B. Continued. 





Range of Depths 2 


Percentage 




Station 


in meters 


Silt + Clay 


Sediment Type(s) 


148 


75 


33.42 


Silty Sand 


149 


84 


39.14 


Silty Sand 


150 


64 


9 


? 


151 


40 


12.64 


Sand 


152 


29 


11.83 


Sand 


216 


14 


1.21 


Sand 


217 


14 


1.06 


Sand 


218 


13 


1.26 


Sand 


219 


14 


1.09 


Sand 


220 


15 


0.95 


Sand 


Kl 


29 


1.28 


Sand 


K5 


22 


1.27 


Sand 


K7 


22-29 


1.19-1.69 


Sand 


K9 


22 


1.30 


Sand 


Kll 


22-27 


1.21-1.52 


Sand 


K14 


26 


1.08 


Sand 


K16 


25-27 


0.89-1.48 


Sand 


K18 


25-26 


0.88-1.27 


Sand 


K20 


24 


1.18 


Sand 


K22 


26-29 


0.91-1.37 


Sand 


K26 


24-27 


0.97-1.59 


Sand 


K27 


24 


1.45 


Sand 


K28 


29 


1.45 


Sand 


K31 


26-28 


0.89-2.20 


Sand 


K34 


27 


1.11 


Sand 


K36 


27 


1.00 


Sand 


K38 


27 


1.42 


Sand 


K40 


27 


1.56 


Sand 



1 The area characterized by large seasonal variation in sediment composition is delineated in para- 
graph 2 under "Study Area." 

2 Depths are not corrected for tides or waves. 



In general, the silt + clay content of sediments north of the river exceeded that 
of sediments south of the river. Stations south of the river in less than 40-50 m 
(excluding those within the area of large seasonal variation in sediment compo- 
sition) generally contained less than 5% silt + clay throughout the study period, 
the silt + clay fractions increasing further offshore. A detailed account of textural 
and mineralogical properties of sediments off the Columbia River is presented in 
Sternberg et al. (1977 — a Final Report to the U.S. Army Corps of Engineers). 
Depths sampled in this study ranged from 11 to 97 m (see Tables 2A and 2B). 

Species Accounts 

Records of collected specimens are listed under the appropriate species by 
station and cruise. For example, three specimens collected at station 10 during 
cruise A would appear as 10A (3). Holotypes have been deposited in the United 
States National Museum. Paratypes have been deposited in the United States 
National Museum, the Allan Hancock Foundation, Oregon State University, and 
the Virginia Institute of Marine Science. Representative specimen(s) of every 



BENTHIC PHYLLODOCIDAE 7 

species reported herein (except Paranaitis polynoides and Eulalia levicornuta) 
have been deposited in the Allan Hancock Foundation. 

Genus Eteone Savigny, 1818 

Eteone fauchaldi, new species 

(Fig. 2A-G) 

Records.— 2E (1); 3G (1); 6B (1), C (2), G (1); 12D (1); 13A (1); 14G (1); 16C 
(1), G (2); 17C (1); 18E (1); 19E (1, TYPE); 22C (1); 23E (1); 28F (1), H (1); 31H 
(1); 33E (1); 101A (1); 117B (1); 146B (1); K14E (1); K18E (1); K34E (2); K36E 
(1); K40E (1). 

Description. — Length to 62 mm; width 0.5 to 1.9 mm; number of segments to 
238. Prostomium long, trapezoidal with rounded anterior part having 2 pairs of 
slender, subulate antennae, the anterior pair slightly shorter (Fig. 2A). 2 widely 
set black eyes located slightly anterior to the small nuchal papilla at the posterior 
margin of prostomium. 

First segment about lH>-2!4 times as long as segment 2, provided laterally with 
2 pairs of tentacular cirri, similar in shape to the prostomial antennae. Ventral 
pair V/4-W2 times as long as dorsal pair. Segment 2 with small setigerous lobe. 
Setae on segment 2 absent in the largest specimens (39 mm and longer), well 
developed in a juvenile (2.5 mm), and poorly developed in other specimens. 

Dorsal cirri conspicuous, slightly asymmetrical, their posterior surfaces some- 
what concave. Ventral cirri broad. Both dorsal and ventral cirri surpass acicular 
lobes throughout the body; the ventral cirri are shorter than dorsal cirri. Presetal 
lip of neuropodia exceeds postsetal one. Large acicula reaches or surpasses pre- 
setal lip. Length-width ratio of dorsal cirri in anterior and median segments varies 
with size of animal (Figs. 2B, 2E and 2G) and increases posteriorly in all speci- 
mens. In a large specimen the dorsal cirri are rounded and broader than long 
anteriorly at segment 24 (Fig. 2B), as long as wide at about segment 60 (Fig. 2C), 
broadly conical by segment 90 (Fig. 2D), and somewhat longer posteriorly. An- 
terior dorsal cirri of holotype (22 mm) nearly as long as wide (Fig. 2E), gradually 
becoming longer than wide posteriorly (Fig. 2F). Anterior dorsal cirri of younger 
specimens may be longer than wide (Fig. 2G). Setae with shafts ending in 2 very 
unequal prongs, these with associated spines. Appendage finely denticulated on 
concave edge. 

Proboscis, when dissected, is tuberculate. Anal cirri long, tapering. 

Color is whitish in alcohol. 

It is our pleasure to name this species after Dr. Kristian Fauchald, who gave 
us so much of his time during our visits to the Allan Hancock Foundation. 

Remarks. — As noted in the above description, setae on segment 2 are absent 
in the largest specimens, well developed in a juvenile, and poorly developed in 
other specimens. Thus, the setae on this segment may be lost with growth. Elia- 
son {1962, fide Ushakov, 1972) observed a similar phenomenon in specimens of 
Eteone lactea Claparede from the Oresund. 

Systematic discussion. — Eteone fauchaldi is not likely to be confused with any 
other Pacific species but it bears considerable resemblance to E. heteropoda 
Hartman, found in the Northwest Atlantic Ocean and the Gulf of Mexico. A 
comparison of the two species reveals the anterior ends (including the tentacular 
cirri) and posterior ends to be similar but the structure of the dorsal cirri to differ. 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. A-G, Eteone fauchaldi new species. A, anterior end of a large specimen, dorsal view, 
x 100. B-D, from 39 mm specimen: B, parapodium from segment 24, anterior view, x 125; C, para- 
podium from segment 59, anterior view, x 125; D, parapodium from segment 90, anterior view, 
xl25. E-F, from holotype, approximately 22 mm: E, parapodium from segment 20, anterior view, 
x500; F, parapodium from segment 92, anterior view, x 125. G (10 mm specimen), parapodium from 
segment 18, anterior view, x500. 



BENTHIC PHYLLODOCIDAE 9 

In E. heteropoda the middle and posterior dorsal cirri are distinctly asymmetrical, 
but in E. fauchaldi the dorsal cirri are only slightly asymmetrical. 

Distribution. — Oregon to Washington in 15-50 m on sand, silty sand, and sandy 
silt (1). 

Eteone longa (Fabricius, 1780) 

Nereis longa Fabricius, 1780. 

Eteone longa Pettibone, 1954; Berkeley and Berkeley, 1948; Banse and Hobson, 
1974. 

Records.— ID (1), E (5), G (1); 2E (3), G (6); 3E (2); 4E (1); 10E (1), G (1); 19E 
(1), G (1); 20G (1); 24F (2), G (1), H (6), I (2); 28E (2), F (3), G (1), H (8), I (2); 
29E (3), F (1), G (1); 31E (3), F (5), G (2), H (3); 32E (1); 33E (2), F (2), H (1), 
I (2); K1E (2); K5E (1); K7E (2), F (2); K11E (2), F (1), G (1); K14E (2); K16E 
(1), H (1); K18E (1); K26E (2), F (2); K27E (2); K28E (2); K31E (1); K34E (1); 
K40E (1). 

Pettibone (1954) synonymized Eteone calif ornica Hartman, described from 
California, with E. longa. We consider the two as distinct species and have found 
both in the present study (see E. calif ornica below). E. longa has dorsal cirri 
which are longer than wide, bluntly conical, but E. calif ornica has broadly round- 
ed dorsal cirri that are as wide as long. Specimens from Arctic Alaska and Wash- 
ington identified as E. longa by Pettibone, 1954, were not examined by us, but 
the parapodium illustrated in that publication agrees with those of the present 
specimens. 

In the Northeast Pacific Ocean, E. longa is previously known from British 
Columbia and Washington. The present records extend the distribution southward 
to northern Oregon. 

Local distribution. — Off Oregon in 16-51 m on sand with less than 5% silt + 
clay. (One in 66 m on sand with 5.58% silt + clay.) 

Eteone calif ornica Hartman, 1936 
Eteone calif ornica Hartman, 1936, 1968. 

Records.— 2E (1); 3D (1), E (10); 4A (1), D (1), E (12), G (3); 5D (1), E (5), G 
(2); 6B (2), D (3), E (2), G (1); 7B (5), C (5), D (1), E (8), G (4); 8B (1), C (1), D 
(1), G (3); 9A (4); 13E (1); 16C (3); 18E (3); 19E (1); 20E (1); 28F (1); 48A (1); 
55A (2); 56A (1); 65A (14); 66A (1); 69A (3); 70A (4); 96A (4); 98A (1); 99A (4); 
100A (2); 147B (2); 148B (2); 149B (1); 150B (2). 

Eteone calif ornica differs from E. longa in having dorsal cirri as wide as long, 
broadly rounded, instead of longer than wide, bluntly conical. 

These records are the first obtained off the coast of Washington and Oregon, 
but the species has been recorded both to the north and south. 

Local distribution. — Oregon to Washington in 18-88 m on sand and silty sand. 

Eteone spilotus, new species 
(Fig. 3A-C) 

Records.— 5E (2), G(1);7B (1), C (6), D (2), E(ll), G (3); 8C(1), E (3), G (4); 
15A (1); 16A (7); 22A (2); 65A (24); 88A (32, TYPE); 89A (2); 95A (1); 96A (1); 
99A (1); 100A (8); 148B (2); 150B (1). 



10 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Description.— Length to over 26 mm; width 0.3 to 1 . 1 mm; number of segments 
to over 100. Prostomium slightly longer than wide, rounded anterior portion with 
4 subequal antennae (Fig. 3A). 2 black eyes near posterior margin; a small nuchal 
papilla at postmedian margin. 

Segment 1 about \Vi times as long as segment 2, with 2 pairs of subequal 
tentacular cirri longer than prostomial antennae. Segment 2 with setigerous lobes 
and setae well developed. 

Parapodia with small dorsal and ventral cirri (Figs. 3B and 3C). Symmetrical 
dorsal cirri inflated and subquadrangular, about as long as broad, somewhat lon- 
ger posteriorly. They reach or slightly surpass the acicular lobes throughout the 
body. In median and posterior segments neuropodial lips are subequal and elon- 
gated, ventral cirri reach to about middle of the acicular lobes (Figs. 3B and 3C). 
Setae with shafts ending in 2 equal prongs and numerous small spines. Appendage 
finely denticulated on concave edge. 

Proboscis smooth when everted. Anal cirri 2, thick, twice as long as broad. 

Color in alcohol pale, with distinct patches of black or brown pigment spots on 
dorsum of each segment — one patch at each side at base of parapodia, vaguely 
connected by a narrow transverse band; a smaller, medial patch anterior to trans- 
verse one on some segments. Pattern duplicated on ventrum. Dorsal cirri and 
some ventral cirri darkly pigmented. 

Etymology. — The specific name is derived from the Greek spilotos, meaning 
spotted, in reference to the distinct patches of pigment spots on the body. 

Systematic discussion. — Eteone spilotus differs from E. californica Hartman 
in body coloration and in having ventral cirri distinctly shorter than the acicular 
lobes. 

Distribution. — Oregon to Washington in 33-88 m on silty sand and sand with 
greater than 10% silt + clay. (One on sand with 1.59% silt + clay.) 

Eteone columbiensis, new species 
(Fig. 3D-G) 

Records.— HE (2); 21C (4), D (31, TYPE); 25D (1); 111B (8); 216D (2); 217D 
(2). 

Description. — Length about 34 mm; width 0.4 to 1 mm; number of segments 
to over 150. Anterior region of body dilated through 15 to 20 segments. Prosto- 
mium elongate and abruptly tapered such that its broad posterior portion com- 
prises about half the total length, sometimes less (Fig. 3D). Anterior margin of 
prostomium one third as wide as posterior margin, with 2 pairs of slender, sub- 
ulate, subequal antennae. 2 black eyes in posterior part of prostomium, a small 
nuchal papilla at the posterior margin. 

Segment 1 slightly longer than segment 2, with 2 pairs of tentacular cirri, dorsal 
pair shorter; both pairs slightly shorter than prostomial antennae. Segment 2 with 
setigerous lobes and setae well developed. 

Dorsal cirri are symmetrical, subrectangular and inflated, very small in anterior 
setigers (Fig. 3E). They are larger and surpass the acicular lobes from setiger 10- 
20 through the mid-region (Fig. 3F), becoming smaller posteriorly (Fig. 3G). Pre- 
and postsetal lips of neuropodia subequal. Ventral cirri small, same length as 
acicular lobes in anterior region of body. Proceeding posteriorly acicular lobes 
become narrow, surpassing ventral cirri (Figs. 3F and 3G). Setae in median and 



BENTHIC PHYLLODOCIDAE 



11 




Fig. 3. A-C, Eteone spilotus new species. A, anterior end, dorsal view, x 125. B-C, from a 20 
mm specimen: B, parapodium from segment 33, anterior view (diagram); C, parapodium from seg- 
ment 71, anterior view, x500. D-G, Eteone columbiensis new species. D, anterior end, dorsal view, 
x 100. E, parapodium from segment 7, anterior view, x500. F, a median parapodium, posterior view, 
x500. G, a posterior parapodium, posterior view, x500. 



12 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

posterior segments with shaft endings of equal length. Appendage finely dentic- 
ulated on concave edge. 

Proboscis smooth proximally; distal half with many small soft papillae. Anal 
cirri 2, thick, slightly clavate, twice as long as broad. 

Color in alcohol pale green, with 3 distinct patches of dark green or brown 
pigment spots on dorsum of each segment — one patch at each side at base of 
parapodia and one medial. Pattern duplicated on ventrum. Dorsal cirri occasion- 
ally pigmented. 

Etymology. — The specific name alludes to the Columbia River, in which region 
the present species was collected. 

Remarks. — Four gravid individuals were collected at station 21, two in April 
1975 (21C) and two in June 1975 (2 ID). The eggs are observed in the parapodial 
bases and give the egg-containing segments a light orange color. 

Systematic discussion. — Eteone columbiensis appears to be closely related to 
E. dilatae Hartman, but differs from it in the shape of the prostomium and the 
body coloration. The prostomium of E. dilatae tapers gradually from the posterior 
to anterior margin, while that of E. columbiensis is abruptly tapered. Eteone 
dilatae is pale greenish yellow, while E. columbiensis is pale green with three 
distinct patches of dark green or brown pigment spots on dorsum and ventrum 
of each segment. 

Distribution. — Within the mouth of the Columbia River to directly offshore in 
11-20 m on sand, and silty sand (1). 

Eteone sp. 
(Fig. 4A-B) 

Records.— AY) (1), E (1); 5G (1); 6E (1); 18E (1); 31E (1); K40E (1). 

These specimens, which belong to a single species, resemble Eteone spetsber- 
gensis Malmgren in the form of the dorsal cirri. Dorsal cirri are asymmetrical and 
their length-width ratio in median segments can vary between individuals (com- 
pare Figs. 4 A and 4B from two different specimens). Prostomium is about as long 
as wide. Setae on segment 2 are well developed. Ushakov (1972) and Pettibone 
(1954, p. 233) reported E. spetsbergensis as having setae poorly developed or 
absent on segment 2. Identification is uncertain at the present time. 

Local distribution. — Off Washington in 44 m on sand (1); off Oregon in 26-82 
m on sand. 

Eteone (Mysta) barbata (Malmgren, 1865) 

Mysta barbata Malmgren, 1865; Bergstrom, 1914. 
Eteone (Mysta) barbata Ushakov, 1972; Pettibone, 1954. 

Records.— 4E (1); 6C (1), D (1); 14D (1); 15D (1); 16G (1); 17B (1), D (2); 22D 
(1). 

Small individuals (about 9 mm) differ from large specimens in having a brown 
longitudinal stripe down each side of the dorsum. Also, the dorsal cirri of the 
small specimens have a greater length- width ratio, about 1 in posterior segments. 

The present records are the first in the Northeast Pacific Ocean. Previously 
known in the Pacific only on the Asiatic coast, to the sea of Japan. 



BENTHIC PHYLLODOCIDAE 



13 




A, 


0.12mm 


B, 


0.20mm 

i 


C, 


1.14mm 


D, 


0.57mm 

i 



Fig. 4. A-B, Eteone sp. A-B, median parapodia from two different specimens. C-D, Anaitides 
multipapillata new species. C (holotype), anterior end and proboscis, dorsal view. D. 40th parapo- 
dium. 



14 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Local distribution. — Northern Oregon to Washington in 26-66 m on sand (with 
greater than 5% silt + clay), clayey sand (1), and sandy silt (1). 

Genus Anai tides Czerniavsky, 1882 

Anaitides multipapillata, new species 

(Fig. 4C-D) 

Records.— 3D (1), G (1); 12E (2); 14C (1); 15D (1), G (1); 16D (1), G (1); 17B 
(1), D (1), E (1), G (1); 19D (1); 20D (1); 241 (1); 25E (1); 27H (1); 33G (1); 66A 
(1); 103B (1); 126B (1, TYPE); K31E (1); K40E (1). 

Description. — Length to 95 mm; width excluding parapodia to 5 mm; number 
of segments to 200. Cephalic lobe subcordate with 2 dark circular eyes, a post- 
median incision with a small occipital papilla, and a pair of small retractile nuchal 
processes at posterior margin of the cephalic lobe near base of first pair of ten- 
tacular cirri (Fig. 4C). Longest of four pairs of tentacular cirri drawn posteriorly 
reaches to segment 10-16. In preserved specimens, dorsum with dark iridescent 
transverse bars of olive green pigment, ventrum lighter, often whitish. Dorsal 
cirri usually olive green with a pale to white margin (Fig. 4D). Some preserved 
specimens are gray. 

Prostomium with 4 antennae and pigmented in two triangular, olive green 
patches (Fig. 4C). Distal part of proboscis tuberculate with tubercles arranged 
in six rows. Proximal portion of proboscis with 12 evenly spaced rows of 20-24 
papillae per row (Fig. 4C). Papillae large and turgid, often with red pigmented 

tips. Setae from third segment. Formula of anterior segments: I 1 + 0— + S-^r), 

with dorsal tentacular cirri on second segment the longest. Setae are homo- 
gomph compound spinigers, appendage finely denticulated along one edge, shaft 
distally spinous. 

Dorsal cirri large, subquadrangular, covering about 14 of the dorsum. Ventral 
cirri distally pointed, extending beyond setal lobes (Fig. 4D). Pygidium blunt. 
Anal cirri 2, cylindrical, tapering, about twice as long as width of pygidium. 

Etymology. — The specific name derives from the numerous papillae per row on 
the proboscis. 

Systematic discussion. — Anaitides multipapillata is distinguishable from A. 
groenlandica (Oersted) and A. mucosa (Oersted) by the absence of a distinct 
dorsal and ventral separation in the rows of proboscideal papillae and by the large 
size of these papillae. The papillae are more numerous than in A. mucosa. How- 
ever, large specimens of A. groenlandica may have about the same number of 
papillae per row as A. multipapillata. Identification of A. multipapillata requires 
examination of proboscis. 

Distribution. — South of Columbia River to directly off mouth in 15-66 m on 
sand, silty sand (1), clayey sand (1), sandy silt (2), and clayey silt (1). 

Anaitides groenlandica (Oersted, 1843) 

Phyllodoce (Anaitides) groenlandica Ushakov, 1972; Pettibone, 1954, 1963; 
Berkeley and Berkeley, 1948. 

Records.— 2D (1); 3E (2), G (1); 4D (3), E (3), G (3); 5D (2), E (1); 7B (1), D 
(4), E (1); 8B (2), C (1), D (5), E (6), G (2); 9A (4); 25E (2); 29D (1); 31E (1), G 



BENTHIC PHYLLODOCIDAE 15 

(1); 47A (2); 56A (1); 65A (6); 68A (1); 69A (2); 70A (7); 89A (1); 95A (5); 98A 
(1); 137B (1); 147B (3); 149B (1); 150B (1); K7E (1). 

The shape of the ventral cirri in our specimens varied from small to large 
animals. Small specimens had pointed ventral cirri as described for Anaitides 
mucosa (Oersted); large specimens had ventral cirri as described for A. groen- 
landica, i.e. oval with asymmetrical acuminate tip; intermediate sized specimens 
had ventral cirri intermediate in shape between A. groenlandica and A. mucosa. 
Anaitides groenlandica reportedly has a greater number of papillae per row on 
the proboscis than A. mucosa, 12-15 vs. 8-10 (Ushakov, 1972); 10-20 vs. 8-12 
(Pettibone, 1963); greater than 12 vs. not more than 10 (Banse and Hobson, 1974), 
and their color patterns differ (see respective descriptions in Ushakov, 1972). In 
our material, the larger specimens among those with groenlandica type ventral 
cirri had greater than 12 papillae per row on the proboscis and a color pattern as 
described for A. groenlandica, brown, with dark brown transverse stripes on 
dorsum; dorsal cirri brown with pale margins, thus fitting the definition of this 
species completely. All remaining specimens, those with groenlandica type ven- 
tral cirri, those with mucosa type ventral cirri, and those with intermediate 
shaped ventral cirri, had 12 papillae per row on the proboscis and the same color 
pattern described above. We therefore conclude that all our specimens belong to 
A. groenlandica; those with mucosa type ventral cirri are merely juvenile A. 
groenlandica. The largest specimen (6 mm wide) had about 25 papillae per row 
on the proboscis. As far as we know, no workers to date have reported greater 
than 20 papillae per row in specimens of A. groenlandica. 

Local distribution. — Oregon to Washington in 15-88 m on sand and silty sand. 

Anaitides hartmanae (Blake and Walton, 1977) 
Phyllodoce hartmanae Blake and Walton, 1977. 

Records.— 2G (1); 3G (1); 6E (1); 7C (2), D (4), E (1), G (2); 8C (1), G (1); 14A 
(1); 18B (1); 19E (1); 24H (3); 28E (1), G (1); 29G (1); 31G (1); 33D (1), E (1); 97A 
(1); 147B (1); K1E (1); K28E (1); K34E (1); K36E (1). 

The present species is herein referred to the genus Anaitides Czerniavsky, 
rather than Phyllodoce Savigny, because the papillae on the basal half of the 
proboscis are arranged in rows; see Fauchald (1977), p. 48. The rows of papillae 
are oblique (spirally arranged, according to Blake and Walton, 1977). 

Anaitides hartmanae is previously known only from California where it was 
described. The present records extend the distribution northward to Oregon and 
Washington. 

Local distribution. — Oregon to Washington in 26-97 m on sand and silty sand. 

Anaitides longipes (Kinberg, 1866) 

Phyllodoce (Anaitides) longipes Kinberg, 1866. 
Anaitides longipes Hartman, 1968. 

Records.— 3D (1); 8B (1); 15D (1); 18D (1). 

In the Pacific Ocean, Anaitides longipes is previously known from Chile and 
California. The present records extend the distribution northward to Oregon and 
Washington. 



16 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Local distribution.— Off Washington in 80 m on silty sand (1); off Oregon in 
44-53 m on sand and sandy silt. 

Genus Paranaitis Southern, 1914 
Paranaitis polynoides (Moore, 1909) 

Anaitis polynoides Moore, 1909. 

Paranaitis polynoides Hartman and Reish, 1950; Hartman, 1968; Ushakov, 1972. 

Phyllodoce (Paranaitis) polynoides Berkeley and Berkeley, 1948. 

Records.— 4E (1); 147B (1). 

Local distribution. — Off Washington in 60 m on silty sand (1); off Oregon in 66 
m on sand (1). 

Genus Eulalia Savigny, 1818 
Eulalia levicornuta Moore, 1909 

Eulalia levicornuta Moore, 1909; Hartman, 1968. 
Eulalia (Eulalia) levicornuta Banse and Hobson, 1968. 

Records.— 5C (1), D (1). 

These records are the first off Oregon, but the species has been recorded both 
to the north and south. 

Local distribution. — Off Oregon in 86-88 m on sand with greater than 15% 
silt + clay. (Note that both specimens were collected at the same station but 
during different cruises.) 

Genus Eumida Malmgren, 1865 
Eumida sanguine a (Oersted, 1843) 

Eulalia sanguinea Oersted, 1843. 

Eumida sanguinea Hartman and Reish, 1950; Hartman, 1968; Ushakov, 1972. 

Eulalia (Eumida) sanguinea Berkeley and Berkeley, 1948. 

Records.— 4E (1); 5C (1); 7D (1), E (1); 8B (2), D (1); 47A (1); 48A (2); 55A 
(2); 68A (2); 69A (2); 97A (1); 148B (1). 

Local distribution. — Oregon to Washington in 55-97 m on sand (with greater 
than 5% silt + clay) and silty sand. 

Distributional Patterns Within the Genera Eteone and Anaitides 

The local distribution, including depths and sediment type(s), for each phyl- 
lodocid species is given in the previous section. Seven species of Eteone occurred 
in the study area. Eteone columbiensis occurred only in shallow water (11-20 m) 
within and directly off the mouth of the Columbia River, in sand and silty sand. 
Eteone longa was restricted to sediments with less than 5% silt + clay and oc- 
curred only in 16-51 m off Oregon (except one specimen referred to under "local 
distribution" in the previous section), south of the area characterized by large 
seasonal variation in sediment composition. 1 Eteone spilotus occurred in sand 
with greater than 10% silt + clay and silty sand from Oregon to Washington 
(except one specimen referred to under "distribution" in the previous section). 



1 The area characterized by large seasonal variation in sediment composition is delineated (and 
discussed) in paragraph 2 under "Study Area." 



BENTHIC PHYLLODOCIDAE 17 

The distributions of the above three species did not overlap one another; that of 
E. colwnbiensis did not overlap any other species of Eteone. Eteone califomica, 
E. fauchaldi, E. barbata, and E. sp. each had a distribution which overlapped 
that of every species of Eteone except E. colwnbiensis. The first three occurred 
in sand and muddier (greater than 25% silt + clay) sediments from Oregon to 
Washington. Eteone sp. occurred off Washington and Oregon in sand. 

Each of the four species of Anaitides in the study area occurred in sand as well 
as muddier (greater than 25% silt + clay) sediments and their distributions over- 
lapped one another. Anaitides groenlandica occurred off Oregon, Washington, 
and directly off the Columbia River. Anaitides multipapillata occurred south of 
the Columbia River and directly off it. Anaitides hartmanae occurred off Wash- 
ington (21 specimens) and Oregon (18 specimens), but only one specimen (station 
97 A) was collected directly off the Columbia River, in deep water (97 m). In 
contrast to A. groenlandica and A. multipapillata, only one specimen of A. 
hartmanae (station 14A off Oregon) was found within the area characterized by 
large seasonal variation in sediment composition resulting from seasonal differ- 
ences in river flow and wave intensity. 1 Though the latter species occurred in 
sediments ranging from 1% silt + clay to 42% silt + clay, perhaps it is less tol- 
erant than the other two species to the temporal fluctuations in habitat within the 
above area. Only four specimens of A. longipes were collected, three off Oregon 
and one off Washington. 

Members of the family Phyllodocidae occurred commonly but in low numbers. 
Species were represented by only one or two individuals in the majority of rep- 
licate sets in which they occurred. For this reason, it is impossible to detect 
seasonality in the abundance of the phyllodocid species, even within the area 
characterized by large seasonal variation in sediment composition. 

One species which exhibited a temporal change in occurrence is Eteone longa. 
This species occurred commonly within and adjacent to the rectangular area 
delineated in Fig. 1 from cruise E (September 1975) to cruise I (June 1976), but 
only one specimen was collected prior to cruise E. (Considering E. longa -con- 
taining stations sampled both before cruise E and from cruise E onward, the 
species occurred in 1 of 33 replicate sets taken before cruise E, and in 32 of 39 
replicate sets taken between cruise E and I.) In July and August 1975, between 
cruises D and E, the U.S. Army Corps of Engineers dumped approximately 
460,000 m 3 of dredge material in the area delineated by the rectangle in Fig. 1 
(Richardson et al., 1977; this area is referred to as experimental site G in that 
publication). Prior to disposal, the median diameter of sediment in this area 
ranged from 2.70 to 2.89 4>. As the dredge material, derived from the mouth of 
the Columbia River, had a high percentage of 2.0-2.5 c/> size particles, the sedi- 
ment within the rectangular area following disposal was a coarser sand, the me- 
dian diameter of most stations between 2.1 and 2.4 (/>. Eteone longa appeared at 
this time (i.e. following disposal) outside the rectangular area, in sediments not 
affected by dumping, most of these finer grained (median diameter greater than 
2.7 (/>), as well as within the rectangular area, in finer sediments, and coarser ones 
(median diameter = 2.1-2.4 (/>) resulting from dumping. As E. longa did not show 
any preference for the coarser sediments, its sudden appearance in this general 
area is probably not due to the increase in sediment grain-size. Though it is 
tempting to postulate that E. longa was transported to the disposal vicinity from 



18 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



the mouth of the Columbia River during dredging and dumping operations, the 
species was never collected at the stations within or just outside the mouth. We 
are unable to explain the temporal disparity in the occurrence of E. longa. 

Acknowledgments 

Foremost, we thank Dr. Kristian Fauchald for his verification of several 
species, taxonomic advice, and critical review of the manuscript. Dr. Michael 
Richardson provided much information pertinent to the preparation of this paper, 
and has reviewed the manuscript. Dr. Andrew Carey provided the senior author 
with laboratory space and facilities. Dr. Keith Serafy offered many helpful sug- 
gestions in the preparation of the manuscript. Dr. Donald Boesch, Gary Gaston, 
and Gordon Bilyard critically reviewed the manuscript. Sam Mauldin did the 
illustrations for Anaitides multipapillata. Peter Smyth provided valuable sugges- 
tions relating to the art work. Discussions with William Colgate, Bruce Mundy, 
and Elizabeth Wilkins proved fruitful. This work was supported, in part, by U.S. 
Army Corps of Engineer Contracts DACW57-75-C-0137 and DACW57-76-C-0092. 

Literature Cited 

Banse, K., and K. D. Hobson. 1968. Benthic polychaetes from Puget Sound, with remarks on four 
other species. Proc. U.S. Nat. Mus., 125(3667): 1-53. 

. 1974. Benthic errantiate polychaetes from British Columbia and Washington. Fish. Res. Bd. 

Canada, Bull., 185:1-111. 

Barnes, C. A., A. C. Duxbury, and B. A. Morse. 1972. Circulation and selected properties of the 
Columbia River effluent at sea. Pp. 41-80 in The Columbia River estuary and adjacent ocean 
waters, bioenvironmental studies. (A. T. Pruter and D. L. Alverson, ed.), Univ. Washington 
Press, Seattle, xiii + 868 pp. 

Bergstrom, E. 1914. Zur Systematik des Polychaeten Familie der Phyllodociden. Zool. Bidr. Upp- 
sala, 3:37-224. 

Berkeley, E., and C. Berkeley. 1948. Annelida, Polychaeta errantia. Canadian Pacific Fauna, Fish. 
Res. Bd. Canada, 9b(l): 1-100. 

Blake, J. A., and C. P. Walton. 1977. New species and records of polychaeta from the Gulf of the 
Farallones, California. Pp. 307-321 in Essays on polychaetous annelids, in memory of Dr. 
Olga Hartman. (D. J. Reish and K. Fauchald, ed.), Allan Hancock Found., Univ. So. Cali- 
fornia, Los Angeles, 604 pp. 

Carey, A. G., Jr. 1972. Ecological observations on the benthic invertebrates from the central Oregon 
continental shelf. Pp. 422-443 in The Columbia River estuary and adjacent ocean waters, 
bioenvironmental studies. (A. T. Pruter and D. L. Alverson, ed.), Univ. Washington Press, 
Seattle, xiii + 868 pp. 

Duxbury, A. C. 1972. Variability of salinity and nutrients off the Columbia River mouth. Pp. 135- 
150 in The Columbia River estuary and adjacent ocean waters, bioenvironmental studies. (A. 
T. Pruter and D. L. Alverson, ed.), Univ. Washington Press, Seattle, xiii + 868 pp. 

Eliason, A. 1962. Weitere Untersuchungen fiber die Polychaetenfauna des Oresunds. Lunds Univ. 
Arsskr., N. F., Avd. 2, Bd. 58, No. 9:1-98. 

Fabricius, O. 1780. Fauna Groenlandica, systematice sistens, Animalia Groenlandica occidentalis 
hactenus indagata, quod nomen specificum, triviale, vernaculumque; synonyma auctorum plu- 
rium, descriptionem, locum, victum, generationem, mores, usum, capturamque singuli, pront 
detegendi occasio fuit, maximaque parti secundum proprias observationes, Hafniae, xvi + 
452 pp. 

Fauchald, K. 1977. The polychaete worms, definitions and keys to the orders, families and genera. 
Nat. Hist. Mus. Los Angeles Co., Science Series 28:1-190. 

, and D. A. Hancock. In press. Deep-water polychaetes from a transect off central Oregon. 

Allan Hancock Monogr. mar. biol. 

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. 



BENTHIC PHYLLODOCIDAE 19 

. 1968. Atlas of the errantiate polychaetous annelids from California. Allan Hancock Found., 

Univ. So. California, Los Angeles, 828 pp. 

, and D. J. Reish. 1950. The marine annelids of Oregon. Oreg. State Monogr. Stud. Zool., 

No. 6:1-64. 

Huyer, A. 1977. Seasonal variation in temperature, salinity, and density over the continental shelf 
off Oregon. Limnol. Oceanogr., 22:442-453. 

Kinberg, J. G. H. 1866. Annulata nova. Ofv. Svenska Vetensk. Akad. Forh., 22:239-258. 

Kulm, L. D., R. C. Roush, J. C. Harlett, R. H. Neudeck, D. M. Chambers, and E. J. Runge. 
1975. Oregon continental shelf sedimentation: interrelationships of facies distribution and sed- 
imentary processes. Jour. Geology, 83:145-175. 

Malmgren, A. J. 1865. Nordiska Hafs-Annulater. Ofv. Svenska Vetensk. Akad. Forh., 21:51-110. 

Moore, J. P. 1909. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of 
southern California in 1904. 1. Syllidae, Sphaerodoridae, Hesionidae and Phyllodocidae. Proc. 
Acad. Nat. Sci. Philadelphia, 61:321-351. 

Oersted, A. S. 1843. Annulatorum danicorum conspectus fasc. I. Maricolae. Copenhagen, 52 pp. 

Pettibone, M. H. 1954. Marine polychaete worms from Point Barrow, Alaska, with additional records 
from the North Atlantic and North Pacific. Proc. U.S. Nat. Mus., 103(3324):203-356. 

. 1963. Marine polychaete worms of the New England region. 1. Aphroditidae through Troch- 

ochaetidae. U.S. Nat. Mus., Bull., 227:1-356. 

Richardson, M. D., A. G. Carey, Jr., and W. A. Colgate. 1977. Aquatic disposal field investigations, 
Columbia River disposal site, Oregon; Appendix C: The effects of dredged material disposal 
on benthic assemblages. Technical Report D-77-30, U.S. Army Engineer Waterways Experi- 
ment Station, Vicksburg, Miss., 411 pp. 

Shepard, F. P. 1954. Nomenclature based on sand-silt-clay ratios. Jour. Sed. Petrology, 24:151-158. 

Sternberg, R. W., J. S. Creager, W. Glassley, and J. Johnson. 1977. Aquatic disposal field investi- 
gations, Columbia River disposal site, Oregon; Appendix A: Investigation of the hydraulic 
regime and physical nature of bottom sedimentation. Technical Report D-77-30, U.S. Army 
Engineer Waterways Experiment Station, Vicksburg, Miss., 332 pp. 

Ushakov, P. 1972. [Polychaeta 1. Polychaetes of the suborder Phyllodociformia of the Polar Basin 
and the northwestern part of the Pacific]. (In Russian.) Akad. Nauk SSSR. Zool. Inst. Fauna 
of the SSSR, n.s. 102, 271 pp. (translated 1974 by the Israel Program for Scientific Translation, 
Jerusalem). 

Accepted for publication January 2, 1978. 



Bull. Southern California Acad. Sci. 
78(1), 1979, pp. 20-31 

Recognition of Quaternary Wave Formed Marine Terraces on 

Santa Catalina Island 

William F. Samaras 1 and Joseph A. Gellura 2 

Abstract. — Four previously unconsidered and undescribed terrace locations 
have been identified as Quaternary wave cut, emergent, marine terraces. The 
presumption has persisted that there apparently are no identifiable, marine ter- 
races on Santa Catalina Island. This is, in large part, due to the absence of 
corroborative Pleistocene marine fossil assemblages and the extensive erosion of 
suspected terrace locations. However, there does exist an abundance of water 
rounded pebbles and cobbles. These are in relatively distinct concentrations in 
at least four locations, especially along the western margins of the island. Uti- 
lizing the methodology developed by Dodkins and Folk in 1970, the authors 
determined that the Maximum Projection Sphericities (i//p) of the pebbles and 
cobbles were generated by wave not fluvial action, thereby establishing them as 
artifacts from wave cut benches. 

1 Research Associate, Museum of Natural History of Los Angeles County, 900 
Exposition Blvd., Los Angeles, CA 90007, and Carson High School, 22328 So. 
Main St., Carson, CA 90745. 2 Earth Science Instructor, Cherry Hill West High 
School, Chapel Ave., Cherry Hill, New Jersey, 08002. 



The paucity of identifiable marine terraces on Santa Catalina Island has re- 
mained an enigma, albeit on a relatively local level, since Smith (1847) first ob- 
served that the most striking difference between Santa Catalina and the slopes 
of San Pedro Hill (Palos Verdes Peninsula, Los Angeles County) and of San 
Clemente Island is the marked absence of marine terraces. Lawson (1893) con- 
cluded that there is no trace of an elevated wave cut terrace, sea cliff, or strand 
line of any kind observable on the island. Smith (1897) later observed that the 
island yielded extremely poor Pleistocene subtidal and intertidal marine inver- 
tebrate faunal assemblages on its ill defined emergent terraces. 

The insular neighbors of Santa Catalina are San Clemente, San Nicolas, Santa 
Barbara, Anacapa, Santa Cruz, Santa Rosa and San Miguel Islands (Fig. 1). All 
have more or less well defined, wave formed, marine terraces. This indicates a 
period of extensive subsidence and subsequent emergence from the Late Pliocene 
through the Pleistocene. Therefore, Santa Catalina Island should not be an ex- 
ception to this pan-insular pattern of subsidence, emergence and subsequent ter- 
racing so characteristic of the islands as well as the coastal margin of Southern 
California. 

Structural Considerations 

Santa Catalina Island is a melange composed of rocks of the Catalina Schist 
(it is understood, although not absolutely proven, that this complex belongs to 
the Franciscan Formation) of Jurassic-Cretaceous age, capped by Tertiary vol- 
canics. 



1ARINE TERRACES ON SANTA CATALINA ISLAND 



21 



|; ^^r^^g, jk 




N 


i^-^^V 


\ 




6tf, S.l.e. Peok • 0„ *\. 


A 


1 


v • A ^ %■ 


6> 


II 


V.-.r; ■■ - V\> 




1 




^****^ii«i^.ujj 








Pacific 


^v7 \ y ^\_ 








1^ \ ^/°^ : ^s 










Ocean 


|t V : ° T 




TS-6^ 


\ *jJj[ilitile.Ha.bt,. ' •■'/ ■'. BlockJock Ml -i 




TS-5- 


—-^ttPV a**SvT' <? ''' * «Mmm "\- 




TS-8- 


_— ^P M &£*? ^**"- 20 '° ^h. 

jabsT"*^^' /^-^Oniobo Ml. £*** -. V 

r\\ Mmm Ml .Bonn.oq . / j, V ' '-2 




TS-4- 










•^TSS»A ^**t •'SK.. 1 ..\ 




% 


I^Pedro ^^ 


*"• • • • •*v«*« M « a L'"'"""-~r^ : ' /** i ~ ; vp 


?* Avolon 


^--V 


\i. ' *' Mmm ■ ^JLTT^f •' .' >...■» -A 






TS^-T-^^^^^ / 


f% \ 


* \ 


SollaVtrds >^ ,'■!% 
Pi. ^*fcv... 


~ jB&pl 


X: 







Fig. 1. Locations of wave cut marine terraces on Santa Catalina Island identified from Pleistocene 
'ave abraded pebbles and cobbles. The inset map indicates the geographic relationship of Santa 
!atalina Island (g) to the mainland and to the other channel islands: a. San Miguel; b. Santa Rosa; 
. Santa Cruz; d. Anacapa; e. San Nicolas; f. Santa Barbara; and, h. San Clemente. The map symbols 
idicate: TS is Terrace Site; Mmm is Middle Miocene marine; Tv is Tertiary volcanics; KJf is Catalina 
chist; • is Pre-Quaternary sedimentary deposits; ■ is a survey and sampling site; ▼ is previously 

efined terrace site; is the generalized separation between (Tv) capping volcanics and (KJf) 

xposed metamorphic rocks. 



The rocks of the metamorphic basement range from glaucophane and chlorite- 
ctinolite rich schist to hornblende-garnet schists. The capping volcanics consist 
i rhyolite, dacite, andesite and some basalt. Consequently, the highly dissected, 
Tonounced vertical relief of the island, with its lack of definable marine terraces, 
aay be due to differential patterns of weathering controlled by the diverse char- 
cter of its lithology. 

Former Indian inhabitants extensively used the relatively flat areas as habitation 
ites and middens so that invertebrate fossil material has either been obscured or 
nidentifiably incorporated, with midden deposits. A few areas of Santa Catalina 
sland geomorphically have the appearance of probable marine terraces. These 
Dcations are strewn with pebbles and cobbles that have been abraded by some 
orm of water action, either marine or fluvial. 

Pre-Quaternary Insular Submergence 

Evidence of Pre-Quaternary (Tertiary) periods of submergence exist An ex- 
tosure of poorly consolidated, unfossiliferous limestone is exposed in Cotton- 



22 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. The slide depressed, basin like marine terrace (site TS-7) behind Salta Verde Point on the 
west side of Santa Catalina Island. The Palisades (light colored cliffs) and the east end of the Island 
are in the background. 



wood Canyon (Fig. 1). This limestone unit unconformably overlies Catalina Schist 
and is overlain by Upper Miocene andesite. Two small deposits of fossiliferous 
limestone are located on the flanks of Mt. Banning (439 m a.s.l.) and Bald Peak 
(427 m a.s.l.) on the south side of Bullrush Canyon (Fig. 1). These limestones 
overlie earlier Miocene volcanics. They are intercalated by andesitic tuffs and 
lapilli and overlain by andesitic lavas of Upper Miocene age. A tuffaceous diat- 
omite deposit, extending from sea level at Big Fisherman Cove to about 150 m 
a.s.l. at Empire Quarry (Fig. 1), is intercalated by beds of lapilli and capped by 
Upper Miocene andesitic volcanics. 

Defined Terraces 

Apparent terraces were observed and described by Smith (1847) from the upper 
sections of Avalon Canyon (Fig. 1). Rounded pebbles that he found on the surface 
of apparently terraced areas in the vicinity of Little Harbor were interpreted as 



Table 1. Computed Percents of Certainty for wave formed vs. river formed pebbles and cobbles. 



Wave Formed 
Pebbles/Cobbles (Beach) 


River Formed 
Pebbles/Cobbles (Fluvial) 


4>p % Certainty 


<//p % Certainty 


0.60 95% 
up to 0.65 76% 


over 0.65 76% 
0.70 95% 



MARINE TERRACES ON SANTA CATALINA ISLAND 



23 









Fig. 3. General terrace deposited pebble shapes and measurements relative to Maximum Projec- 
tion Sphercities (i//p). a) andesite cobble approaching prolateness from site TS-5, L = 138.4 mm, I = 
81.8 mm, S = 53 mm, with a ijjp of 0.62; b) oblate andesite cobble from site TS-6, L = 105.3 mm, I = 

88.3 mm, S = 52 mm with a t//p of 0.66; c) oblate dacite pebble from site TS-4, L = 86 mm, I = 77 
mm, S = 30 mm with a t//p of 0.51; d) oblate andesite cobble from site TS-5, L = 104.7 mm, I = 68 
mm, S = 64.4 mm with a \\ip of 0.83; e) oblate andesite cobble from site TS-6, L = 11 1.6 mm, I = 

78.4 mm, S = 39.2 mm, with a i//p of 0.56; f) oblate dacite pebble from site TS-4, L = 71.4 mm, I = 
64.4 mm, S = 25.7 mm, with a t/>p of 0.52. 



having been abraded and subsequently deposited by ancient fluvial action as were 
pebbles later found in association with angular lithoclasts from terraces in Avalon 
Canyon. Smith (1933) identified a few surfaces he believed to be terraces, but 
conjectured their origins on the basis of notched and beveled salients and apparent 
strand lines. Bailey (1941) and Slosson (1973) identified these previously described 
areas as Quaternary terraces. 

Methodology 

Exploratory surveys over most of the island identified water rounded pebbles 
and cobbles from the flanks of Mt. Banning, along promontories such as Salta 
Verde Point and Indian Head Rock and on canyon slopes. The only location 
where Quaternary fossils (primarily molluscan) were found in association with 
water rounded pebbles was on the north slope above the mouth of Cottonwood 
Canyon (site TS-5). They were exposed along the margin of a landslide displaced 
block of blue schist. 



24 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Tables 2-6. Computed Maximum Projection Sphericities for pebble suites analyzed. In all the 
Tables (2-6) of computed Maximum Projection Sphericities, the pebble size classification is based on 
the L or Long Axis (Figure 6). In the column expressing Percent of Certainty (%), the letter F 
indicates Fluvial (river) origin while the letter W indicates Wave (beach) origin. Control Site (TS-8): 
Cottonwood Canyon Beach. 

Pebble Size (mm) 8-16 16-32 32-64 64-128 128-256 

Number of Pebbles 2 29 46 30 3 

Mean xj,p 0.76 0.62 0.55 0.54 0.56 

% Certainty 95% F 76% W 95% W 95% W 95% W 

Mean i//p for suite 0.572 

a 0.116 

% Certainty for suite 95% Wave-Formed 

Pebble Lithology: 1 10 andesite pebbles and cobbles 



The pebbles and cobbles observed and collected were primarily andesite. Al- 
though andesite is not always the predominant rock exposed, it is much more 
resistant to weathering than the metamorphics. The landslide modified, terraced 
area behind Salta Verde Point (Figs. 1 and 2) provided the exception to the 
predominance of andesite pebbles and cobbles. The pebbles and cobbles at this 
site are a green dacite porphyry (hornblende-quartz-diorite of Bailey, 1941). This 
is the most abundant rock type comprising the pebbles and cobbles on the con- 
temporary beaches in the immediate area of Salta Verde Point. 

The overall shape of the water abraded rocks tends to be more oblate rather 
than prolate (Fig. 3). Random pebble suites were collected from four possible 
terrace locations: 1) the top of the ridged terrace on the south side of Cottonwood 
Canyon (Site TS-4); 2) the remnant terrace on the slide block behind Indian Head 
Rock (Site TS-5); 3) Rancho Escondido terrace on the south side of Little Harbor 
(Site TS-6); and 4) Salta Verde Point terrace (Site TS-7) as in Fig. 1. A total of 
370 pebbles and cobbles were collected in random samplings from these locations. 
The number of rocks taken from each site was determined by abundance and 
availability, not by size or shape preference. 

Standardization (Table 2) was accomplished by collecting 110 structurally iso- 



Table 3. Site TS-4, South side of Cottonwood Canyon. 
Pebble Size (mm) 8-16 16-32 32-64 64-128 128-256 



Number of Pebbles 


8 


31 


46 


22 


Mean \jip 


0.778 


0.629 


0.593 


0.603 


% Certainty 


95% F 


76% W 


95% W 


95% W 



Mean »//p for suite 0.619 

or 0.115 

% Certainty for suite greater than 76% Wave Formed 

Pebble Lithology: 
71 andesite 18 quartz 8 scoria 
3 tuff 5 schist 2 metasandstone 



MARINE TERRACES ON SANTA CATALINA ISLAND 25 

Table 4. Site (TS-5), Behind Indian Head Rock. 

Pebble Size (mm) 8-16 16-32 32-64 64-128 128-256 

Number of Pebbles 26 32 43 29 6 

Mean i//p 0.691 0.687 0.526 0.573 0.583 

% Certainty 76% F 76% F 95% W 95% W 95% W 

Mean i/»p for suite 0.612 

o- 0.141 

% Certainty for suite 90-95% Wave Formed 

Pebble Lithology: 
118andesite 12 quartz 
4 schist 2 metasandstone 



tropic andesite pebbles and cobbles from a Holocene beach at the mouth of 
Cottonwood Canyon, Site TS-8 (Fig. 1). Shaping on this beach is controlled by 
moderate to heavy wave energy. The beach is approximately fifty meters wide 
with a foreshore slope averaging 10°. The beach faces west on the seaward side 
of the island. It grades from sand subtidally, coarse sand and gravel intertidal, to 
pebbles and cobbles in the supratidal. The backbeach grades into fluviatile sands 
deposited by Cottonwood Creek (Fig. 5). 

The shape development study conducted by Dodkins and Folk (1970) showed 
that beach pebbles have a much lower sphericity than river pebbles. They stated 
that, "the maximum projection sphericity averages 0.68 for rivers, 0.64 for low- 
wave-energy beaches, and 0.58 for high- wave-energy beaches." The authors uti- 
lized the method developed by Dodkins and Folk (1970) to identify the origin of 
the water rounded pebbles and cobbles found on the island. Because of the lack 
of fossils, reliance was placed on the determination of the mean sphericities of 
available gravels and scattered concentrations of water rounded pebbles. The 
long, intermediate and short axis of each pebble from each pebble suite was 
calculated (Fig. 6) and their individual Maximum Projection Sphericities (ipp) 
computed using the formula developed by Folk (1955): 



S 2 



LI 

Table 5. Site (TS-6), Rancho Escondido Terrace. 



Pebble Size (mm) 8-16 16-32 32-64 64-128 128-256 

Number of Pebbles 2 17 16 9 

Mean i/»p 0.64 0.66 0.58 0.44 

% Certainty 76% W 76% F 95% W 95% W 

Mean i//p for suite 0.587 

a 0.148 

% Certainty for suite 95% Wave Formed 

Pebble Lithology: 
14 andesite 14 schist 
12 quartz 4 metasandstone 



26 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 4. In situ, wave formed conglomerates exposed in a low road cut at marine terrace site TS- 
4 on the south side of Cottonwood Canyon. 



where S is the short axis; L the long axis; and I is the intermediate axis. The 
Maximum Projection Sphericity is a comparative relationship between the max- 
imum projection area of a pebble (LI) and the maximum projection area of a 
sphere of equal volume. The Mean Maximum Projection Sphericity for each 
pebble suite is computed by adding the individual ijjp together then dividing the 
sum by the total number of pebbles measured. The Percent of Certainty for each 
pebble size classification and each pebble suite is computed in Table 1 . 

The Percent of Certainty is a function of probability percentages. If the i|/p 
mean is under 0.65, there is a 76% probability that the pebble suite was wave 
formed (beach); if the mean ifjp is less than 0.60, there is a 95% certainty that the 



Table 6. Site TS-7, Salta Verde Terrace. 



Pebble Size (mm) 



8-16 



16-32 



32-64 



64-128 



128-256 



Number of Pebbles 
Mean i//p 
% Certainty 



15 


51 


14 


3 


0.709 


0.667 


0.582 


0.580 


95% F 


95% F 


95% W 


95% W 



Mean \jjp for suite 0.657 

cr 0.100 

% Certainty for suite 76% Fuvial 

Pebble Lithology: 
73 dacite porphyry 10 andesite 



MARINE TERRACES ON SANTA CATALINA ISLAND 



27 




Fig. 5. Profile of the control beach (Site TS-8 and Table 2) at the mouth of Cottonwood Canyon, 
a. subtidal zone, medium sized sand grain bottom; b. intertidal zone, grading from coarse sand to 
gravel; c. splash and supratidal zone, pebbles and cobbles; and, d. fluviatile sand in creek bed behind 
berm. 



pebble suite originated on a beach. Conversely, if the mean i//p is greater than 
0.65 there is a 76% probability that the pebble suite is from a river. The break 
point between fluvial and wave formed is from i//p 0.65 to 0.66. This is due to the 
difference between the mechanics of fluvial and beach abrasion and independent 
of lithology as long as the rocks are structurally isotropic (Dodkins and Folk, 
1970). 

Pebble size classification was adapted from Dodkins and Folk (1970). The anal- 
ysis on Santa Catalina Island necessitated the addition of a smaller pebble cate- 
gory, 8-16 millimeters. The addition was made to accommodate the abundance 
of small pebbles available in the areas surveyed and to provide the analysis with 
as random a selection, free from size bias, as possible. Of the 370 pebbles and 
cobbles collected, 336 were structurally isotropic (213 andesite, 42 quartz, 73 
dacite prophyry and 8 scoria). The remaining 34 rocks (23 schist, 8 metasandstone 
and 3 tuff) generally lacked flow structure or planar features. 

Terrace Site Description 

Terrace site TS-4 (Table 3) is on the south side of Cottonwood Canyon (Fig. 
1), at latitude 33°10' 18" north, longitude 1 18°27'42" west, on the west side of Santa 
Catalina Island. One hundred and seven pebbles and cobbles were collected from 
exposed gravels near a low road cut (Fig. 4). 

Site TS-5 (Table 4) is the remnant of a terrace situated on a slide block within 
a landslide complex located along the seaward flank of Cottonwood Canyon, 
behind Indian Head Rock. Although the terrace surface is physically lower in 
elevation (45 meters a.s.l.) than the adjacent TS-6 site (74 meters a.s.l.) it was 
obviously continuous prior to the sliding. It is located at latitude 33°10'36" north, 
and longitude 118°28'42" west, on the south side of Little Harbor. One hundred 
and thirty six pebbles and cobbles were collected from a small gravel exposure 
near the seaward margin of the slide block. 

Site TS-6 (Table 5) is the most distinctly terraced location surveyed and sam- 
pled. Its relatively broad, flat surface stretches inland past Escondido Ranch. 
There are no in situ gravels exposed on this terrace location. Rather, the pebbles 
are strewn about, generally becoming more numerous downslope near the sea- 
ward margin. Terrace TS-6 is located between Cottonwood Canyon to the south- 
east and Little Harbor on the northwest, at latitude 33°22'18" north, longitude 



28 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 6. Projection of the coordinates for computing the Maximum Projection Sphericity of a water 
rounded rock. L represents the maximum projected length, I the intermediate length (width) and S 
the short axis or thickness of the rock. 



118°28'42" west. This location is referred to as Rancho Escondido Terrace. Forty 
four pebbles were collected from this site. 

Site TS-7 (Table 6) is a slide modified, basin shaped terrace behind Salta Verde 
Point (Figs. 1 and 2). It is at latitude 33°19'12" north and longitude 118°25'12" 
west on the west side of Santa Catalina Island. Eighty three pebbles were col- 
lected from the surface of the terrace. The present terrace lies at an average 
elevation of 45 meters a.s.l. 



Discussion 

In three of the five sites sampled, including the control site (TS-8), the smallest 
pebble classification, 8-16 mm, produced a high mean i|/p indicative of fluvially 
formed rocks. On the other hand, in all of the sites sampled the large pebble size 



MARINE TERRACES ON SANTA CATALINA ISLAND 



29 




« 





Fig. 7. Anomalous cobble modifications. Fragments d, e, g, i, 1, and m are spalled off pieces of 
cobbles found in Indian middens on Santa Catalina Island. These were used as choppers, scrapers, 
gravers, etc. (Santa Catalina Island Museum Anthropological Collection). Spalls c and h are from 
Salta Verde Point terrace (TS-7) and j is a quartzite spall from Ripper's Cove, a' is a spall chipped 
by the authors from a prolate cobble from the Salta Verde Site (TS-7). The spalls illustrated are com- 
posed of dacite porphyry and quartzite. 



classifications, especially 128-256 mm, produced characteristically low i//p. These 
low mean sphericity values are indicative of wave formed rocks. This apparent 
bimodal condition had been found on high wave energy, gravel beaches by Dod- 
kins and Folk (1970). Under these conditions there is a tendency for smaller 
pebbles to be tossed about randomly, to be abraded more uniformly and to fall 
into the interstitial spaces between larger rocks where they are rolled back and 
forth by wave action rather than slide. In the case of sandy beaches, it was found 
that pebbles of all sizes have uniformly low sphericities. 

This condition of wave force discrimination in pebble shaping, producing wide 
range of Maximum Projection Sphericities, is evident in all of the sites sampled. 
We conclude, therefore, that similar physical conditions existed during the Qua- 
ternary period for at least 3 of the 4 sites sampled (sites TS-4, TS-5, and TS-6), 
and, in all probability, for the TS-7 site. The Cottonwood Canyon Beach control 
site is typical of the aforementioned physical parameters to pebble type devel- 
opment, namely, high wave energy (waves averaging over 0.6 m in height) and 
a gravel beach. These parameters have produced a low mean Maximum Projec- 
tion Sphericity for the site of 0.572, giving it at least a 959f certainty that the 
pebbles are wave formed. 



30 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Interpretation of Apparently Non-Supportive Data From Salta Verde 

The mean ijjp for the Salta Verde Point (TS-7) site (Figs. 1 and 2) in the two 
smaller pebble groupings (Table 6) indicate a fluvial origin. Conversely, the mean 
\\sp for the two larger pebble categories indicate a wave/beach controlled origin. 
This enigma is compounded by the fact that the TS-7 site is the most obviously 
terraced location of the four sampled and analyzed. Extensive field observations 
at this location turned up a peculiar repetition of broken cobbles above 128 mm 
size, rendering them useless for accurate measurement. Only three found in the 
128-256 mm range were not broken (Fig. 7). Contrarily, only a very few pebbles 
in the 64-128 mm range were found broken and none in the smaller ranges. The 
cobbles were also broken in an unusual fashion; rather than being broken in half 
through their short axis, the well rounded ends along their long axis were spalled 
off as by a deliberate, well directed blow (Fig. 7). In searching through the Santa 
Catalina Island Museum's Indian artifact collection, we found similar spalled off 
pieces identified as crude choppers and scrapers (Fig. 7). Also an abundance of 
midden sites in the Salta Verde area indicates that it was used extensively as a 
habitation area by the former Indian population. 

Therefore, the relatively high mean i//p of 0.656 for this particular terrace may 
well be directly related to the toolmaking needs of the Island's original inhabit- 
ants. The paucity of large pebbles, along with the broken condition of those 
found, seems to indicate that the Pleistocene deposited gravels served as readily 
available raw material for paleolithic toolmakers well into Holocene times. Con- 
sequently, the larger pebbles and cobbles with significantly lower sphericities, 
well into the wave formed category as indicated in Table 6, were apparently either 
removed from the area long ago or were rendered useless for interpretive data by 
being broken. This is also indicated by the average of the mean i//p for each size 
range of 0.634 rather than by the mean i//p for the entire suite. Given the evidence 
on hand, albeit much of it circumstantial, it is the authors' opinion that the Salta 
Verde Point Terrace was also formed by wave action during a period of submer- 
gence and subsequent emergence. 

Conclusion 

The mean i//p for the four suspected terrace sites sampled is 0.618, with a 
standard deviation (cr) of 0.028. This figure coincides almost exactly with the 
mean i//p for ten known ancient beach sites on the Atlantic and Gulf coasts sam- 
pled by Dodkins and Folk (1970) with a mean ifjp of 0.61. This coupled with the 
fact that the surrounding islands and coastal margin all show conclusive signs of 
Quaternary subsidence and emergence in the form of wave formed marine ter- 
races, supplies strong evidence that Santa Catalina Island is no exception. 

Acknowledgments 

We wish to extend our gratitude to Douglas Propst, superintendent, Santa 
Catalina Island Conservancy, for his personal interest and direction, and to the 
Santa Catalina Island Conservancy for providing grants-in-aid and facilities. We 
especially wish to thank Robert O'Guin, former Ranger Santa Catalina Island 
Conservancy, for unselfishly sharing his time and extensive personal knowledge 
of the island. 



MARINE TERRACES ON SANTA CATALINA ISLAND 31 

We wish to thank the Santa Catalina Island Company, Patricia Moore and the 
Santa Catalina Island Museum for generously providing materials and space. We 
gratefully acknowledge the help and dedication of Mary Bertsch. 

We wish to extend our appreciation to the many interested people of the city 
of Avalon for sharing with us the information about their island, and especially 
to Stanley Beus of Northern Arizona University and Martin Reiter of Los Angeles 
Harbor College for their valued criticisms and support and to Patricia Brown and 
Theresa Price for the editing and preparation of this manuscript. 

We acknowledge Carl Hubbs of Scripps Institute of Technology for asking the 
questions that led to the research for this paper. 

Literature Cited 

Bailey, Edgar H. 1941. Mineralogy, petrology, and geology of Santa Catalina Island, California. 

Unpubl. Dis., Stanford University, pp. 173-185. 
Dodkins, James E., and Robert L. Folk. 1970. Shape development on Tahiti-Nui. Jour. Sed. Petr., 

40(4): 1167-1203. 
Folk, Robert L. 1955. Student operator error in determination of roundness, sphericity and grain 

size. Jour. Sed. Petr., 25:297-301. 
Hobbs, William H. 1912. Earth features and their meaning. MacMillan Co., New York, pp. 256- 

257. 
Lawson, A. C. 1893. Pliocene diastrophism of the coast of Southern California. Univ. California 

Bull. Geology, 1:115-160. 
Slosson, James E. 1973. Geologic map of Santa Catalina Island, modified from original by E. H. 

Bailey, 1940, USGS. 
Smith, W. S. T. 1897. Geology of Santa Catalina Island. Proc. California Acad. Sci., 1:1-71. 

. 1933. Marine terraces on Santa Catalina Island. Amer. Jour. Sci., ser. 5, v. 25, 123 pp. 

Woodring, W. P. et al. 1946. Geology and paleontology of Palos Verdes Hills, California: U.S.G.S., 

Professional paper 207, 135 pp. 

Accepted for publication November 27, 1978. 



Bull. Southern California Acad. Sci. 
78(1), 1979, pp. 32-39 

Species Diversity and Distributional Relationships of 

Pseudoscorpions from Slash Pine (Pinus elliottii Eng.) 

in Florida (Arachnida: Pseudoscorpionida) 

Vincent Brach 

Abstract. — Nine species of pseudoscorpions associated with living and dead 
slash pine, Pinus elliottii Eng., in south central Florida were studied with respect 
to distribution and species diversity. Trees offering the greatest diversity of bark 
microhabitats have the greatest species diversity. Most of this diversity is in the 
fauna of the flake bark, which is the commonest and least patchily distributed 
bark microhabitat. However, some species may be unable to utilize this micro- 
habitat unless other types of bark microhabitats are available simultaneously. 

Archbold Biological Station, Rt. 2, Box 180, Lake Placid, Florida 33852. Pres- 
ent address: Department of Biology, Texas Eastern University, Tyler, Texas 
75701. Reprint address: Mr. Fred Lohrer, Archbold Biological Station, Rt. 2, Box 
180, Lake Placid, Florida 33852. 



Despite the fact that pseudoscorpions are widely distributed and often abundant 
in habitats containing crevices, the ecology of pseudoscorpions is described by 
Muchmore (1973) as being "sadly neglected." Hoff and Bolsterli (1956), Ressl 
and Beier (1958), Hoff (1959, 1961, 1963a) and Nelson (1975) have presented the 
results of systematic collections of pseudoscorpions from broad geographic areas 
together with some habitat information, the studies of Ressl and Beier (1958) and 
Hoff (1959) being the most detailed. A few species of pseudoscorpions have 
received detailed treatment with respect to particular ecological parameters such 
as desiccation resistance [Kensler, 1967 — Neobisium maritimum Leach; Cloudse- 
ly-Thompson, 1956 — Withius subruber (Simon) and Dactylochelifer latreillei 
(Leach)]. Faunal studies of restricted habitats in which pseudoscorpions occur 
as minor components have yielded some additional ecological information (e.g. 
Morton, 1954; Graves and Graves, 1969; Richards, 1971). Only the studies of 
Hoff and Clawson (1952), Chamberlin and Malcolm (1960) and the review of 
Weygoldt (1969) deal entirely with the distributional relationships of pseudoscor- 
pions with respect to a particular habitat (rodents' nests, caves and the sand 
islands near Beaufort, North Carolina respectively). However, no studies have 
been made of the distributional relationships and species diversity of the pseudo- 
scorpion fauna of a particular microhabitat. 

Among the most important microhabitats in which pseudoscorpions may be 
found are the subcortical or underbark spaces of trees. Several families of pseudo- 
scorpions (Cheliferidae, Cheiridiidae, Chernetidae and Atemnidae) are very com- 
monly encountered in such situations and several genera (e.g. Parachelifer, Den- 
drochernes) are almost exclusively associated with the loose or started bark of 
a variety of tree species, usually conifers. The types of bark found on trees are 
quite variable and offer widely varying conditions of crevice size, humidity and 
permanence which not only change from one tree species to another but also 



PSEUDOSCORPIONS FROM SLASH PINE 33 

with the age and state of health of a particular tree. Thus, a particular tree may 
show a microsuccession of arthropod communities during its existence (Blackman 
and Stage, 1924; Savely, 1939) and at different times may be expected to support 
different associations of pseudoscorpions as well. 

The purpose of this study was to investigate the species diversity and distri- 
butional relationships of the pseudoscorpions found on living and lightning struck 
slash pines, Pinus elliottii Eng., in south central Florida. 

Materials and Methods 

This study was conducted at the Archbold Biological Station, Highlands Coun- 
ty, Florida during October-April 1975-1976. The topography and vegetation types 
characteristic of the Station have been described elsewhere (Woolfenden, 1969). 
Pseudoscorpions were collected from 25 mature slash pines ranging from 30-80 
cm diameter at breast height growing in slash pine-turkey oak and scrubby flat- 
woods habitats. The method of sampling was to conduct a l /i hr timed search of 
the entire trunk from a height of 1.5 m to the ground. Pseudoscorpions were 
collected with an aspirator or a brush as they were exposed. Since the bark of 
living trees was found to contain pseudoscorpions only beneath exfoliating flakes, 
live bark was not removed from these, but the started bark on dead trees was 
removed from the trunk after the remaining flake bark had been searched. Almost 
all of the trees selected could be thoroughly sampled in the allotted time; com- 
parison with three untimed trial searches lasting over 1 hr suggested that at least 
90% of the pseudoscorpions present are collected within the first Vi hr. 

Lightning strikes on P. elliottii are a major cause of tree mortality at the Arch- 
bold Station. Thirteen of the 25 trees sampled had been killed by lightning (the 
passage of the bolt is visible as a spiral channel down the trunk) and exhibited 
a regular sequence of decay stages. The study of Baker ( 1973) on lightning caused 
mortality in the closely related loblolly pine, P. taeda L., at the Tall Timbers 
Research Station in Leon County, Florida, and my own observations of lightning 
killed slash pines at the Archbold Station suggested the following classification 
of decay stages: 

Stage 1. Live, healthy trees. Flake and live bark present, foliage dark green. Few 
or no pitch tubes from wood boring beetles visible. 

Stage 2. Freshly hit trees (approximately 1 mo). Foliage partly or all brown, 
beginning to fall off. No loss of bark, but pitch tubes from wood boring 
beetles evident. Lightning streaks clearly evident on trunk. 

Stage 3. Early decortication. Pitch tubes no longer evident. Bark not loose but 
removable with a knife. Flake bark reduced. Upper branches often miss- 
ing; no foliage. 

Stage 4. Intermediate. Tree reduced to trunk and a few stubs of larger branches. 
Extensive decortication on parts of trunk, especially above 2 m. Basal 
bark loose and easily removed in large strips with fingers. The wood 
and undersurface of the bark often moist, especially on the north side, 
and may support large termite colonies. Wood may be punky and "red- 
rotten" in places. 

Stage 5. Advanced. Decortication almost complete, with only a small region of 
easily removed bark fragments remaining at the trunk base. Wood be- 



34 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

neath the bark largely dry and "white-rotten." Sections of sapwood 
may be gone, producing a "necked" effect. The trunk often can be 
pushed over with slight effort. 

Of the 25 trees examined, 14 were in stages 3-5 with six in stage 3, seven in 
stage 4, and one in stage 5. A single tree was found in stage 2 but could not be 
quantitatively sampled because of its location on the landscaped Station grounds. 
However, a brief examination showed that this tree had not decayed long enough 
for its bark to differ significantly from that of a live tree. The remaining 11 trees 
were all stage 1 (had not been struck by lightning). 

Representative specimens of each species of pseudoscorpion collected were 
deposited in the Archbold Station collection. All species determinations were 
made by William B. Muchmore. 

Results 

Nine species of pseudoscorpions were collected at least once during the course 
of this study. These are: 

Family Tridenchthoniidae 

Verrucaditha spinosa (Banks) 
Family Cheiridiidae 

Cheiridium sp. 
Family Sternophoridae 

Gary ops depressa Banks 

Idiogaryops paludis (Chamberlin) 
Family Chernetidae 

Parachernes latus (Banks) 

Americhernes longimanus Muchmore 

Dendrochernes morosus (Banks) 
Family Atemnidae 

Paratemnus elongatus (Banks) 
Family Cheliferidae 

Parachelifer superbus Hoff 

The occurrence of each species with respect to the decay stage of the tree, 
number of specimens per collection, and type of bark from which collected is 
summarized in Table 1. 

Discussion 

Following a lethal lightning strike, a mature pine tree follows a roughly pre- 
dictable sequence of decay stages which spans a period of several years, ending 
when the tree falls over and becomes a log (Baker, 1973). During this time, its 
subcortical microhabitats undergo profound changes with respect to humidity and 
the number and types of available crevices. The trunk of a mature slash pine is 
typically covered with close fitting patches of tightly appressed bark which pro- 
vide little shelter where they join the trunk. However, the laminated, exfoliating 
exterior of the live bark provides a distinctive and unique microhabitat for those 
species which are extremely flat and are able to tolerate dryness. At the Archbold 
Station, the foremost of these species are the sternophorid pseudoscorpions G. 



PSEUDOSCORPIONS FROM SLASH PINE 



35 



Table 1. Summary of pseudoscorpion collections from 25 slash pines, Pinus elliottii, at the Arch- 
bold Biological Station, Highlands County, Florida during October-April 1975-1976. 



















# per 






Stage of Tree 






Total 


Coll. Site 






Start- 












# of 


# 


Flake 


ed Sap- 












Species 


1 


2 3 


4 


5 


Colls. 


Coll. 


Bark 


Bark wood 


V. spinosa 


— 


— 1 (7)* 


2(14) 


1 (6) 


4 


27 





27 


Cheirid. sp. 


2(7) 


— — 


— 


— 


2 


7 


7 





G. depressa 


7(19) 


— 5 (38) 


3 (30) 


1 (7) 


16 


194 


194 





I. paludis 


2(8) 


— 1 (3) 


1 (3) 


— 


4 


14 


14 





P. latus 


3(4) 


— 3(7) 


— 


— 


6 


11 


11 





A. longimanus 


— 


— 3(8) 


4(25) 


1 (8) 


8 


37 


11 


27 


D. morosus 


2(3) 


1 (3) 


2(6) 


— 


5 


12 


2 


3 


P. elongatus 


6( + ) 


— — 


4( + ) 


1 (5) 


15 


>200 


( + ) 


3 


P. superbus 


— 


— 3(7) 


3(8) 


1 (4) 


11 


19 


9 


9 1 



* The number of collections is followed by the number of specimens in each collection. 
+ Visually estimated at >20 per collection (explanation in text). 



depressa and /. paludis. P. latus and Cheiridium sp., while much less common, 
were also found exclusively in this microhabitat. G. depressa was usually abun- 
dant when it occurred at all, but typically each patch of flake bark (approximately 
50 x 50 mm) sheltered a single specimen. Hoff (1963b) found that 13 out of 15 
collections of G. depressa at the Archbold Station were from the exfoliating bark 
of standing slash pines, with some collections containing between 25 and 50 
specimens. Since the two collections not made from standing slash pines were 
small but still slash pine associated (stump bark and a rotten fallen limb), Hoff 
(1963b) concluded that G. depressa is restricted to slash pines. However, while 
a living slash pine may support a very large population of G. depressa, as soon 
as the tree dies and stops replacing its bark the population begins to deteriorate, 
a few individuals perhaps remaining as long as there are patches of flake bark 
left. 

/. paludis is a less common species which Hoff (1963b) found to be less re- 
stricted to slash pine than G. depressa, since he also collected it from beneath 
the bark of several species of hardwoods. Although Hoff (1963b) recorded /. 
paludis and G. depressa in one collection at the Archbold Station, I never found 
these two species together on the same tree. 

The tridenchthoniid pseudoscorpion V. spinosa is said to migrate from the soil 
to the subcortical spaces of dead trees during the summer over some parts of its 
range in southeastern North America (Weygoldt, 1969). This species was always 
found beneath the started bark of dead slash pines in stages 3-5, usually on those 
trees which were quite moist and rotten beneath the bark. V. spinosa was col- 
lected only four times from slash pine during this study but was found to be 
relatively abundant under the bark of fallen logs and in the moist litter beneath. 

The genus Parachelifer is interesting in that all of its species are predominately 
subcortical in occurrence and several species are restricted to particular species 
of trees. Hoff (1959) states that P. persi/nilis (Banks) and P. scabriculus (Simon) 
are associated with yellow pine (Pinus ponderosa Laws) and occasionally with 



36 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

other trees, while Hoff and Bolsterli (1956) report that P. longipalpus Hoff occurs 
"beneath the bark of broad-leafed deciduous trees' 1 in the Mississippi Valley. 
Parachelifer archboldi Hoff, which is sympatric with P. superbus in Florida but 
was not collected in this study, is apparently restricted to the subcortical spaces 
of live oaks {Quercus virginiana Mill) (Hoff, 1964). P. superbus often occurred 
beneath well started bark on dead slash pines at the Archbold Station but was 
most often encountered under the bark of trees which had begun to dry out and 
were approaching stage 5 of the decay sequence. 

The chernetid pseudoscorpions D. morosus and A. longimanus showed very 
different microhabitat preferences. D. morosus had the least restricted micro- 
habitat of all of the pseudoscorpions studied, occurring beneath the started and 
flake bark of living and dead slash pines in approximately equal numbers. This 
species was also collected from the punky sapwood of a decaying sand pine, 
Pinus clausa Sarg., in sand pine scrub at the Archbold Station and may occur on 
other trees as well. Nelson (1975) records Michigan specimens "under the bark 
of dead trees" and Hoff (1959) found that the related D. crassus Hoff is appar- 
ently associated with yellow pine logs. The wide distribution of D. morosus and 
perhaps other congeners may be due to the fact that these pseudoscorpions are 
commonly phoretic on beetles and other insects (Muchmore, 1971) and have been 
taken at light traps at the Archbold Station, presumably having been carried there 
by insects (Frost, 1966). On the other hand, at the Archbold Station, A. longi- 
manus is largely restricted to the moist subcortical spaces of stage 3-5 slash 
pines, only eight out of 37 specimens having been taken from beneath flake bark. 
Muchmore (1976) also records this species from light traps at the Archbold Sta- 
tion, suggesting phoretic dispersal. 

The two collections totalling seven specimens of Cheiridium sp. taken from 
beneath the flake bark of living slash pines may represent accidental associations, 
since the bark from which these specimens were taken was almost level with the 
forest floor. However, this or a very similar species is relatively common under 
the breast height flake bark of slash pines in suburban Coral Gables, Dade County, 
Florida, where G. depressa is rare or absent (my unpublished observations). The 
extremely small size of Cheiridium pseudoscorpions may make them suitable as 
prey items for G. depressa, which did not occur on the trees from which the 
specimens of Cheiridium sp. were collected. 

P. elongatus is in several ways a most atypical pseudoscorpion. It is always 
found in aggregations around semipermanent communal silken molting nests and 
exhibits a marked degree of social behavior involving cooperative prey capture 
and nest construction (Brach, 1978). The dense aggregations of this species made 
quantitative collecting difficult, since the members of a colony would scatter 
when uncovered and all could not be collected. I therefore visually estimated the 
sizes of the colonies. In most cases, the number of animals per colony was 
estimated at about 20. Over 200 specimens were collected during the course of 
this study, supporting Hoff s statement (Hoff, 1964) that P. elongatus is "one of 
the common Florida pseudoscorpions/ ' P. elongatus occurs under started and 
flake bark but shows a distinct preference for flake bark and is usually found 
within 0.25 m of the ground, often below the level of the litter. Since flake bark 
on the lowermost portions of the trunk is always the last to disappear from a dead 



PSEUDOSCORPIONS FROM SLASH PINE 37 

tree, P. elongatus, like G. depressa, can be expected to occur on slash pines in 
any stage of decay as long as at least some flake bark remains. 

The presence of a pseudoscorpion fauna on slash pine can be expected when- 
ever bark is still on the trunk. Only a single pseudoscorpion (P. superbus) was 
collected from the dry punky heart wood of a late stage 5 tree, and this animal 
may have been dislodged from its subcortical resting place during the course of 
collecting. Hence, it may be safely said that the pseudoscorpion fauna of slash 
pine in Florida is essentially a bark fauna. The pseudoscorpion fauna of slash 
pine bark can be divided into three categories based on microhabitat preference. 
G. depressa, I. paludis, P. latus, Cheiridium sp. and P. elongatus are more or 
less restricted to the dry flake bark, the last two species being further restricted 
to the lowermost portions of the trunk. P. superbus and D. morosus were found 
beneath flake and dry subcortical bark in approximately equal numbers; appar- 
ently humidity, rather than bark type per se, influences microhabitat selection by 
these species. V. spinosa and A. longimanus were largely or entirely restricted 
to the moist subcortical spaces of dead trees. Since the nine specimens of A. 
longimanus taken from beneath flake bark were all from dead trees, it is possible 
that A. longimanus, which is more heavily sclerotized than V. spinosa, may 
occasionally forage in low humidity microhabitats, returning periodically to the 
moist subcortical spaces to rehydrate. In the laboratory, I found that A. longi- 
manus desiccates rapidly in dry culture jars and must be provided with a moisture 
source, such as a cotton plugged vial of wet sand, for long term maintenance. 

The diversity of pseudoscorpions inhabiting slash pines at the Archbold Station 
probably is most strongly influenced by the relative permanence of their respec- 
tive microhabitats, the distribution of these microhabitats in time and space, and 
the history of each tree with respect to colonization. Clearly, the least patchily 
distributed microhabitat in time and space is that of flake bark, and therefore one 
would expect this microhabitat to show the largest number of species. Lightning 
struck trees constitute a small fraction of the total number of slash pines in an 
area at any given time. Since a dead tree has a "life span" of only a few years 
at best (Baker, 1973), during which its subcortical environments are changing 
continuously, it would be expected that pseudoscorpions restricted to the tran- 
sient, relatively rare microhabitat of the moist subcortical spaces of dead trees 
would be relatively few in species and patchy in occurrence, the latter phenom- 
enon probably reflecting the chances of colonization and extinction. V. spinosa 
was collected only four times, although 14 of the 24 trees examined had apparently 
suitable moist subcortical spaces. By contrast, G. depressa and P. elongatus 
were found on 16 and 15 of the 25 trees examined, respectively. This proportion 
might have been increased for both species had the number of live trees examined 
been proportionate to their abundance. Pseudoscorpions such as A. longimanus 
and P. superbus, which made use of more than one microhabitat, showed an 
intermediate abundance. 

The data of this study did not permit critical analysis of seasonal differences 
or of intraspecific competition or predation, any of which might be expected to 
have a significant effect on abundance and diversity. However, both Hoff (1963b) 
and I noted that G. depressa and /. paludis rarely occur together on the same 
tree. In the laboratory, P. elongatus readily used G. depressa as prey; in nature, 
this could be expected to produce some of the observed habitat partitioning of 



38 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

flake bark. However, it is likely that the most significant factor determining 
species diversity is the microhabitat diversity of the tree. The most diverse 
pseudoscorpion fauna observed on a single tree was six species, taken from the 
single stage 5 tree examined. This tree was somewhat unusual for a tree of its 
class in that it had patches of flake bark, moist and dry subcortical spaces, and 
a region of dry punky heartwood; it therefore possessed the greatest number of 
potential microhabitats of any of the trees examined. When the flake bark micro- 
habitat alone is considered, the greatest number of species collected from one 
tree is four (five trees), although eight of the nine species studied occurred in this 
microhabitat. Only P. elongatus occurred on all five of these trees. Thus, it 
appears that most of the diversity of the slash pine pseudoscorpion fauna occurs 
in the commonest and most permanent of the available microhabitats, even 
though two of the species (A. longimanus and P. superbus) are apparently unable 
to utilize this microhabitat unless started bark is also present, A. longimanus 
preferring the moist, and P. superbus the dry, subcortical spaces. 

Conclusions 

Although my data are limited with respect to the number of trees sampled, it is 
apparent that the greatest overall species diversity occurs on slash pines offering 
the greatest microhabitat diversity. The specific microhabitat showing the greatest 
species diversity is that offtake bark, which was present in varying amounts on 
all of the trees examined. The two commonest species, G. depressa and P. 
elongatus, are found exclusively beneath flake bark but differ greatly in behavior 
and distribution on the trunk, the former being a solitary species which ranges 
over the entire tree and the latter being a subsocial species which largely restricts 
its activities to the lowermost portions of the trunk. Two species, A. longimanus 
and P. superbus, appear unable to utilize the flake bark microhabitat unless 
started bark is also present; thus, they are not found on live trees. A single 
species, V. spinosa, is entirely restricted to the relatively rare and transient moist 
subcortical spaces and is one of the least commonly encountered species. All of 
the remaining species occur sporadically on slash pine but may either not be 
restricted to this tree species (e.g. /. paludis, D. morosus) or may not be able to 
maintain populations in the presence of the diffuse competition or predation pro- 
duced by the activities of the commoner, more closely slash pine adapted species 
such as P. elongatus and G. depressa. The maintenance of slash pine populations 
of these "fugitive species' ' at the Archbold Station and elsewhere may depend 
upon reservoir populations on other tree species and upon effective dispersal 
systems, such as phoresy. 

Acknowledgments 

I thank James N. Layne and the Archbold Biological Station staff for their 
generous support while I was in residence at the Station during 1975-1976 and 
William B. Muchmore for species determinations and much helpful advice. 

Literature Cited 

Baker, W. W. 1973. Longevity of lightning-struck trees and notes on wildlife use. Proc. Ann. Tall 
Timbers Fire Ecology Conf., 13:497-504. 



PSEUDOSCORPIONS FROM SLASH PINE 39 



Blackman, M. W., and S. S. Stage. 1924. On succession of insects living in the bark and wood of 

dying, dead, and decaying hickory. N.Y. State College of Forestry Tech. Bull., 17:1-269. 
Brach, V. 1978. Social behavior in the pseudoscorpion Paratemmus elongatus (Banks) (Pseudo- 

scorpionida: Atemnidae). Insectes Sociaux, 25:3-12. 
Chamberlin, J. C, and D. R. Malcolm. 1960. The occurrence of false scorpions in caves with special 

reference to cavernicolous adaptation and to cave species in the North American fauna. Amer. 

Midi. Nat., 64:105-115. 
Cloudsely-Thompson, J. L. 1956. Notes on Arachnida, 28. Biological observations and records. Ent. 

Mon. Mag., 92:193. 
Frost, S. W. 1966. Additions to Florida insects taken in light traps. Fla. Entomol., 49:243-251. 
Graves, R. C, and A. C. Graves. 1969. Pseudoscorpions and spiders from moss, fungi, rhododendron 

leaf litter and other microcommunities in the Highlands area of Western North Carolina. Ann. 

Ent. Soc. Amer., 62:267-269. 
Hoff, C. C. 1959. The ecology and distribution of the pseudoscorpions of North-Central New Mexico. 

Univ. N. Mexico Publ. Biol., 8:1-68. 

. 1961. Pseudoscorpions from Colorado. Bull. Amer. Mus. Nat. Hist., 122:409-464. 

. 1963a. Pseudoscorpions from the Black Hills of South Dakota. Amer. Mus. Novitates, 

2134:1-10. 
. 1963b. Sternophorid pseudoscorpions, chiefly from Florida. Amer. Mus. Novitates, 2150: 

1-14. 
. 1964. Atemnid and cheliferid pseudoscorpions, chiefly from Florida. Amer. Mus. Novitates, 

2198:1-43. 
, and D. L. Clawson. 1952. Pseudoscorpions from rodent nests. Amer. Mus. Novitates, 

1585:1-38. 
, and J. E. Bolsterli. 1956. Pseudoscorpions of the Mississippi River drainage basin area. 

Trans. Amer. Micros. Soc, 75:155-179. 
Kensler, C. B. 1967. Desiccation resistance of intertidal crevice species as a factor in their zonation. 

J. Anim. Ecol., 36:391-405. 
Morton, J. E. 1954. The crevice faunas of the upper intertidal zone at Wembury. J. Mar. Biol. Assn. 

U.K., 33:187-224. 
Muchmore, W. B. 1971. Phoresy by North and Central American pseudoscorpions. Proc. Rochester 

Acad. Sci., 12:79-97. 
. 1973. Ecology of pseudoscorpions — a review. Proc. Soil Microcommunities Conf. (Syra- 
cuse), 1:121-127. 
. 1976. Pseudoscorpions from Florida and the Caribbean area. 5. Americhernes, a new genus 

based upon Chelifer oblongus Say (Chernetidae). Fla. Entomol., 59:151-163. 
Nelson, S. 1975. A systematic study of Michigan pseudoscorpionida (Arachnida). Amer. Midi. Nat., 

93:257-301. 
Ressl, F., and M. Beier. 1958. Zur Okologie, Biologie and phanologie der heimischen Pseudoskor- 

pione. Zool. Jahr. (Syst.), 86:1-26. 
Richards, A. M. 1971. An ecological study of the cavernicolous fauna of the Nullarbor plain, southern 

Australia. J. Zool. (London), 164:1-60. 
Savely, H. E. 1939. Ecological relations of certain animals in dead pine and oak logs. Ecol. Monog., 

9:321-385. 
Weygoldt, P. 1969. The biology of pseudoscorpions. Harvard Univ. Press, Cambridge, Mass., 

145 pp. 
Woolfenden, G. E. 1969. Breeding-bird censuses of five habitats at Archbold Biological Station. 

Audubon Field Notes, 23:732-738. 

Accepted for publication January 4, 1979. 



Bull. Southern California Acad. Sci. 
78(1), 1979, pp. 40-45 

Summer Mean Diurnal Water and Air Temperature Distributions 
in the Cape Mendocino area, California 

Gerald Hannes 

Abstract. — This study is concerned with the spatial analysis of the mean air 
and water temperature distributions recorded at sampling sites located throughout 
the Cape Mendocino area of northern California. Employing a three hour sam- 
pling interval, it was found that the basic northerly air flow in summer was quite 
important to the resulting distributions of air temperatures computed for each 
individual site. Furthermore, elevation and distance from the coastline were also 
quite important to the resulting air temperature distributions. However, the mean 
water temperatures examined were slightly modified, if any, by the diurnal heating 
cycle. 

Department of Geography, California State University, Fullerton, California 
92634. 



Atmospheric temperatures in coastal environments are modified through heat 
exchanges with the marine environment (Hasse, 1971; Roll, 1965). For example, 
modifications can be brought about by the movement of marine air associated 
with a basic sea breeze flow (Hannes, 1977). Thus, the diurnal temperature range 
could be reduced by such factors as the cooler air temperatures associated with 
marine air or by the presence of a cloud cover. 

This study is concerned with the spatial analysis of the mean air and water 
temperature distributions throughout the Cape Mendocino area of northern Cal- 
ifornia. It deals with a number of sampling sites and the data employed extend 
from July 8, 1970, through August 17, 1970. 

This paper has several objectives: 1) to study the spatial distribution of the 
mean diurnal atmospheric temperature distributions obtained from a coastal north- 
south instrument transect; 2) to study the mean diurnal air temperature distri- 
butions gathered at weather stations in an east-west transect along a ridge; 3) to 
assess, if possible, the effects of elevation on the air temperature distributions; 
and, 4) to analyze the mean diurnal water temperature distribution and their 
possible relationships to the near coastal air temperature distributions. 

Data Source and Study Area 

The Cape Mendocino area of northern California was chosen because of the 
possible interactions of the cold upwelled waters with the atmosphere in the area 
(Reid et al., 1958), as well as the investigation of the possible effects of the general 
northerly air flow (Robinson et al., 1965) on air temperatures. With cold water 
offshore during the summer months, maximum land water temperature contrasts 
would occur. These thermal contrasts could then lead to a potentially strong sea 
breeze flow. But with a strong, persistent northerly summer air flow over the 
Cape Mendocino area, the basic land sea breeze circulation may be modified. 
This situation might lead to differences in air temperature distributions between 



WATER AND AIR TEMPERATURE DISTRIBUTIONS 



41 



4100 



. CAPE MENDOCINO AREA, 
NORTHERN CALIFORNIA 



45' ' 



D 15' - L 



BLUNT'S REEF, 
LIGHT SHIP 



^ 



• CITY 

■ STATION 




124 u 00 

I 



RINIDAD HEAD 
(EL. 180 FT.) 



41°00' 



# ARCATA 



• FERNDALE 
I NAVY 
(EL. 390 FT.) 



BUNKER HILL 
(EL. 2401 FT.) 



30' 



MOUNT PIERCE 
(EL. 3188 FT.) 



WALKER-GEORGE 
(EL. 1300 FT.) 



MATHEWS- MATTOLE 
(EL. 75 FT.) 



WINDY NIP B 
(EL. 2400 FT.) 



124°30' 



15' 



SML Carlo Lab CSUF 



I24°00' 



Fig. 1. Study Area 



42 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

upstream and downstream stations. Thus, the resultant mean air temperature 
distributions should indicate the interaction of the two previously mentioned mod- 
ifiers of air temperature (Hannes, 1974; Lahey, 1974). 

A majority of the data used in this study was gathered by standard weather 
instruments on loan from the National Center for Atmospheric Research and the 
University of Illinois. The Blunt's Reef Light Ship data were obtained on micro- 
film from the National Climatic Data Center, Asheville, North Carolina. 

Air temperatures were sampled at ten locations, while water temperatures were 
sampled at two locations (see Fig. 1). The variables studied were averaged by 
hours. In total, eight samples were obtained per day. Of the ten stations recording 
air temperatures, five stations are classified as low level coastal stations: Navy, 
Ocean House, Mathews-Mattole, Blunt's Reef, and Trinidad Head. One station, 
Walker-George, is intermediate in geographical location from the coast as well 
as in elevation. The other remaining four stations are classified as ridge top lo- 
cations: Windy Nip, Mount Pierce, Bunker Hill, and Mazzepa. The coastal lo- 
cations nearly make a north-south transect, while the ridge top sites are aligned 
in a basic east-west transect (see Fig. 1). Progressing inland from the coast, the 
ridge top sampling sites of Mazzepa, Bunker Hill, and Mount Pierce are found 
at increasing elevations respectively. 

The atmospheric temperature data were gathered on hygrothermographs. Each 
hygrothermograph had a seven day recorder. Since the Blunt's Reef Light Ship 
data were taken only every three hours, the continuous weekly hygrothermograph 
data were reduced to match the sampling interval of the Blunt's Reef Light Ship 
data. 

Water temperatures were gathered by the bucket method at Blunt's Reef and 
with a Barnes Infrared Radiation thermometer at Trinidad Head (see Fig. 1 for 
location). Once again, the Blunt's Reef data were sampled every three hours; 
therefore, the Trinidad Head water temperatures had to be extracted at three 
hour intervals to correspond to the Blunt's Reef sampling interval. 

Results 

The mean diurnal air temperatures and water temperatures are presented in 
Figure 2. These hourly values were obtained by averaging all values available for 
each individual hour sampled. As might be expected, geographical position with 
respect to the coast seems to have an important effect on the different air tem- 
perature distributions. For example, the minimum temperature at Navy, Maz- 
zepa, Bunker Hill, and Mount Pierce indicate a general increase in magnitude 
from the coastal low level site to the higher, more interior ridge top site of Mount 
Pierce. 

Air temperatures: coastal. — The coastal stations of Ocean House, Navy, and 
Trinidad Head have the lowest diurnal temperature ranges. However, the air 
temperatures of Blunt's Reef exhibit the smallest diurnal range. This small tem- 
perature range is probably due to the station's marine location. On the other 
hand, the Mathews-Mattole site has the greatest mean diurnal range. 

The Mathews-Mattole site is the southernmost low level coastal site (see Fig. 
1). With respect to the various intermediate ridge systems, this location is possibly 
very important to the resultant higher afternoon air temperature. If the northerly 
marine air flows over the various ridges and is forced to mix turbulently with air 



WATER AND AIR TEMPERATURE DISTRIBUTIONS 



43 



AIR TEMPERATURE COASTAL AIR TEMPERATURE INTERMEDIATE AIR TEMPERATURE: RIDGE TOP WATER TEMPERATURE 




-I I I I I I I I I I I I I L 





HOURLY MEAN TEMPERATURES JULY 8 to AUGUST 17, 1970 
Fig. 2. Water and Air Temperatures 



from the inversion layer aloft, then the marine air could be modified. This to- 
pographically induced mixing could cause the resultant high afternoon tempera- 
tures. Atmospheric wave activity is also suggested as a possible mechanism to 
mix the air turbulently over the various ridges. The possible existence of atmo- 
spheric waves in the Cape Mendocino area was shown by Hannes, 1972. 

Air temperatures: intermediate. — The only intermediate sampling station, 
Walker-George, has a temperature distribution similar to that of the Mathews- 
Mattole and Windy Nip (Fig. 2). Walker-George records its highest average tem- 
perature at 1600. The occurrence of the high temperature at 1600 is in general 
agreement with the high temperatures at the two other southerly stations, i.e., 
Mathews- Mattole and Windy Nip. Comparing the high at 1600 to the temperatures 
recorded at the ridge top locations, Walker-George's high is equal to or greater 
than those at Bunker Hill, Mazzepa, and Mount Pierce (see Fig. 2). Furthermore, 
Walker-George's early morning temperature declines in magnitude from 0100 to 
0700. This decrease is similar to the temperature changes attained at Ocean 
House, Mathews-Mattole, and Navy sites. However, the ridge stations do not 
exhibit major decreases during these morning hours. Thus, the greater diurnal 
range and the shape of the distributional curve is akin to the distributional curves 
at Mathews-Mattole and Windy Nip, and not to the ones located on the more 
northern ridge system (i.e., Mazzepa, Bunker Hill, and Mount Pierce). This is 
likely due to topographically induced mixing over the ridges (Hannes, 1972). 

Air temperatures: ridge top. — The last category of the air temperature distri- 



44 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

butions examined is that of the ridge top sites (Fig. 2). The shapes of the four 
distributional curves examined are somewhat different. The station with the low- 
est elevation, Mazzepa, has the smallest average hourly temperatures of all the 
ridge top sites. Further inland from the coast, and at higher elevations, Bunker 
Hill and Mount Pierce's maximum and minimum temperatures are greater than 
the more coastal site of Mazzepa, suggesting the decreasing effects of the marine 
environment and possible diurnal variations in the height of the inversion base. 
The exact nature of the marine inversion in this study area is unknown due to 
the lack of continous upper air data; however, from limited data sampled at 
Areata, Hannes (1972) found the morning inversion base approximately at 900 
meters. The marine inversion on the west coast of the United States also exhibits 
diurnal height fluctuations (Meitin and Stuart, 1977; Schroeder et al., 1967). Thus, 
the diurnal changes in the height of the inversion base could logically affect the 
air temperatures recorded at the ridge top stations. With a low inversion base, 
the ridge top stations could be under the influence of warm, dry air rather than 
cool, moist marine air. 

Windy Nip, the southernmost of all the stations (Fig. 1) has the highest average 
temperatures at all hours examined. These high values are probably due to the 
modification of the northerly air flow passing over the transverse ridges. In ad- 
dition to the high temperature values, there is another distinction between Windy 
Nip and the other ridge top sites. This difference is apparent during the early 
morning hours (0100 and 0400). At those hours, Windy Nip experiences a decline 
in air temperature while the stations to the north are nearly constant and in the 
case of Bunker Hill, there is a temperature rise. Thus, each individual site along 
the northern ridge experiences some form of heat importation at those hours 
(0100 and 0400); otherwise, under normal conditions, air temperatures should 
decline. Possibly local fluctuations in inversion height could affect the night tem- 
peratures at Bunker Hill. 

Furthermore, Windy Nip displays its highest average temperature at 1600 while 
the other ridge top sites experience their respective maximums at 1300. This fact 
suggests that there is a modification of the marine air downstream at Windy Nip, 
while this air is modified to a lesser extent along the more northerly ridge com- 
plex. Thus, the high temperature recorded at Windy Nip at 1600 is similar to the 
respective highs at Mathews-Mattole and Walker-George. 

Water temperatures. — The average hourly water temperature distributions at 
both Trinidad Head and Blunt' s Reef show very little variation throughout the 
diurnal cycle (Fig. 2). Trinidad Head's water temperature is diurnally slightly 
greater in magnitude than the water temperature at Blunt' s Reef. The major 
difference between the two distributional curves occurs in the late morning and 
early afternoon hours (0700 to 1600). The water temperatures recorded at Trinidad 
Head are decreasing while Blunt' s Reef water temperatures are increasing slight- 
ly. The reason for this difference is not fully understood at this time. Possibly 
the difference is due to various forms of ocean and air interactions occuring along 
the coast (Lahey, 1977). But in general, the water temperatures recorded at both 
locations are quite cool during the summer period examined. The cool water 
temperatures found off Cape Mendocino more than likely cause the small diurnal 
range in air temperatures at the low level sites. 



WATER AND AIR TEMPERATURE DISTRIBUTIONS 45 

Summary 

In a coastal area such as Cape Mendocino, the direction of general air flow 
along with local topography are very important to the resultant air temperatures 
experienced at a specific location. Ocean House, of all the land stations, is closest 
to the ocean; yet air temperatures at Navy were lower during the average diurnal 
cycle. Ocean House which is located on the leeward side of Cape Ridge probably 
experiences more modified air than Navy, which occupies a windward location. 

However, in regard to the east-west trending ridge top sites of Mazzepa, Bunker 
Hill, and Mount Pierce, location with respect to distance from the marine envi- 
ronment as well as elevation are suggested as the causes of their appropriate 
temperature distributions. Elevation is very important to the resultant diurnal air 
temperature distribution because of nearness to the base of the inversion layer. 
Since the nocturnal inversion layer is not stationary throughout the night, its 
fluctuations could directly influence the air temperatures at various ridge top sites 
and not the low level sites. It is possible for a ridge top site to be in an inversion 
layer during a portion of the night and not at any other hour. This might account 
for some of the early morning temperature rises (0400 at Bunker Hill). Also 
'breaking' atmospheric waves and turbulent mixing of the inversion layer air 
downward could account for heating and the early morning temperature rise. 

Concerning the water temperatures, it was found that they are slightly modified 
by the diurnal heating cycle. In fact, Trinidad Head experiences a slight temper- 
ature decline in the late morning and early afternoon hours (1000 to 1600). The 
cause of this decrease is not fully understood but could be due to different forms 
of up welling operating near each of the sampling sites. 

Literature Cited 

Hannes, G. 1972. An analysis of atmospheric temperature, pressure, winds, and sea surface tem- 
peratures, summer 1970 at Cape Mendocino, California. Unpublished Ph.D. Dissertation, Univ. 

Illinois, 289 pp. 

. 1974. Factor analysis of coastal air and water temperatures. J. Appl. Meteor., 13:3-7. 

. 1977. Surface air flow in coastal southern California. Bull. Southern California Acad. Sci., 

76:21-31. 
Hasse, L. 1971. The sea surface temperature deviations and the heat flow at the sea-air interface. 

Boundary Layer Meteor., 1:368-379. 
Lahey, J. 1974. On the relationship between sea tides, atmospheric pressure, winds, weather systems 

and upwelling along the northern California coast. Tethys, 6{ 1— 2):409— 418. 
. 1977. The relationship of sea tidal stirring along the Pacific coast of the United States to 

decreasing sea temperatures and coastal wind field intensification. Yearbook, Assoc. Pacific 

Coast Geog., 39:117. 
Meitin, R., and D. Stuart. 1977. The structure of the marine inversion in northwest Oregon during 

26-30 August 1973. Mon. Weather Rev., 105:748-761. 
Reid, J. et al. 1958. Studies of the California current system. California Coop. Fish. Invest. Rpts. 

July, 1956-Jan., 1958:27-56. 
Robinson, L., D. Eberly, and H. Cramer. 1965. Meteorology and atmospheric diffusion in the vicinity 

of the Humbolt Bay power plant. Pacific Gas and Electric Co., San Francisco, 335 pp. 
Roll, H. 1965. The physics of marine air. Academic Press, New York, 426 pp. 
Schroeder, M., M. Fosberg, O. Cramer, and C. O'Dell. 1967. Marine air invasion of the Pacific 

coast: a problem analysis. Bull. Amer. Meteor. Soc, 48:802-808. 

Accepted for publication January 4, 1979. 



Bull. Southern California Acad. Sci. 
78(1), 1979, pp. 46-55 

Niche Separation Within a Population of Freshwater Fishes in an 
Irrigation Drain Near the Salton Sea, California 

Allan A. Schoenherr 

Abstract. — The fish fauna of the King Street canal is a mixture of native and 
introduced species. Inflow from a thermal well at 42°C and irrigation runoff at 
22°C were responsible for a thermal gradient that, in addition to differences in 
flow, held fishes in remarkably pure species populations. On 19 March 1977, 
Cyprinodon macularius was found most abundantly in water 10 cm deep at 39°C. 
Gambusia affinis occurred in flowing water 25 cm deep at 32°C. Only Poecilia 
sphenops inhabited a cool water outflow 18 cm deep at 22°C. Downflow, Poecilia 
latipinna was taken most commonly in slow moving water up to 50 cm deep at 
26°C, and Notropis lutrensis occurred in riffles up to 25 cm deep at similar tem- 
peratures. Flooding during late summer 1977 and subsequent reconstruction of 
the canal obliterated most of the habitat diversity. All five species survived, albeit 
seriously reduced in number, and the species sorting that was previously observed 
also was no longer in evidence. Thermal differences remained, and a pond was 
constructed that impounded hot water. Later, on 17 July 1978, the pond included 
Cyprinodon macularius, Gambusia affinis, Poecilia latipinna and a new intro- 
duction, Tilapia zilli. Downstream, Cyprinodon macularius, Poecilia sphenops, 
and Notropis lutrensis were taken in flowing water. 

Division of Life Sciences, Fullerton College, Fullerton, California 92634. 



Introduction 



An interesting assemblage of freshwater fishes inhabits waterways and canals 
draining into the Salton Sea in Riverside and Imperial Counties, California (Table 
1). Most of these drains carry irrigation runoff, and represent permanent aquatic 
habitats. The fish fauna of these waterways is a mixture of aquarium species, 
escaped bait fishes, introduced game fishes, introduced "weed eaters," and one 
native form, a subspecies of Cyprinodon macularius, the desert pupfish. 

On the northwest side of the Salton Sea, in the King Street canal, a thermal 
outflow was responsible for marked diversity in habitats not apparent in other 
drains (canals). In most of these canals fish species are found in mixed assem- 
blages. Fish distributions in the King Street canal seemed to reveal preferences 
for distinct habitats, each occurring in virtually pure species populations. Flood- 
ing and subsequent reconstruction of the canal appeared to obliterate the distinct 
habitats and thus provided an opportunity to study the influence of habitat di- 
versity on fish distribution. 

This study documents nearly complete niche separation among introduced 
freshwater fishes in an "unnatural habitat," illustrating rapid selective forces in 
action, and it describes the effect of habitat alteration on niche stability. The 
study also describes habitat and potential threats to the desert pupfish, Cyprin- 
odon macularius, the only native fish in the area. 



NICHE SEPARATION OF FRESHWATER FISHES 



47 



Table 1. Fishes known to inhabit irrigation canals and waterways in the vicinity of the Salton Sea, 
Riverside and Imperial Counties, California. 



Species 



Reference 



Dorosomidae 



Cyprinidae 



Dorosoma petenense 

Carassius auratus 
Cyprinus carpio 
Notropis lutrensis 
Notemigonus chrysoleucus 

Cyprinodontidae 

Cyprinodon macularius 

Poeciliidae 

Gambusia affinis 
Poe cilia latipinna 
Poecilia mexicana 
Poecilia sphenops 
Poeciliopsis gracilis 
Xiphophorus helleri 
Xiphophorus variatus 

Centrarchidae 

Micropterus salmoides 
Lepomis cyanellus 



Ictaluridae 



Sciaenidae 



Cichlidae 



Gobiidae 



Ictalurus nebulosus 
Ictalurus punctatus 

Bairdiella icistia 
Cyonoscion nobilis 

Tilapia mossambica 
Tilapia zilli 

Gillichthyes mirabilis 



Black, pers. comm. 

Personal observation 
Mearns, 1975 
Personal observation 
Soltz, pers. comm. 

Personal observation 

Personal observation 

Personal observation 

Personal observation 

Personal observation 

Mearns, 1975 

Mearns, 1975 

St. Amant and Sharp, 1971 

Soltz, pers. comm. 
Personal observation 

Personal observation 
Soltz, pers. comm. 

Soltz, pers. comm. 
Personal observation 



St. Amant, pers. comm. 
Personal observation 



Personal observation 



Materials and Methods 

The canal that parallels King Street lies 7 km north of the Riverside Imperial 
County line on the northwest side of the Salton Sea. Observations and collections 
of fishes were made on 3 March 1977, 21 May 1977, 18 March 1978, and 17 July 
1978. The area provided a unique opportunity to study factors that influence fish 
distribution because depth varied only slightly (5 to 50 cm) and there was present 
a striking thermal gradient in addition to variations in turbulence and velocity. 
Flow was generally shallow and linear. Riffles alternated with laminar flow and 
undercut banks were present in association with meanders. A hot artesian well 
flowed into the canal. At its source the water was 46°C. Approximately 50 m 
downflow where the water cooled to 39°C fishes began to appear. From that point 



48 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



/■ 
HOT WATER 
SOURCE 

SHALLOW 
WASH 



RAPID 
CURRENT 



COLD WATER 
SOURCE 

22° 




N 



Fig. 1 . Diagram of upper portion of King Street canal indicating differences in stream flow and 
temperature (3 March 1977). Fishes occur in nearly pure species groups as follows: Cyprinodon 
maculariiis, shallow wash; Gambusia affinis, flowing water at 32°C; Poecilia sphenops, cold water 
source; Poecilia latipinna, undercut banks; and Notropis lutrensis, riffles. Temperatures in °C. 



to the mouth of the canal fishes were collected using aim nylon seine with ca. 
3 mm bar measure. Some specimens of each species were preserved in 10% 
formalin to insure correct identification. In order not to damage living fishes, 
counts of individuals were not made, and most fishes were released at their 
collection sites after identification. Temperatures at each collection site were 
recorded. Dissolved oxygen was determined using a Beckman Fieldlab Oxygen 



NICHE SEPARATION OF FRESHWATER FISHES 49 

Table 2. Habitat preferences for fishes inhabiting the King Street canal, Riverside County, Cali- 
fornia on 3 March 1977. 





Water 












Temp. 


Diss. 0, 


Depth 




Physical 


Species 


(°C) 


(mg/1) 


(cm) 


Flow 


habitat 


Cyprinodon macularius 


39 


6.8 


5-10 


slow 


shallow wash 


Gambusia affinis 


32 


7.0 


25 


moderate 


stream margin 


Poecilia sphenops 


22 


8.5 


18 


rapid 


stream source 


Poccilia latipinna 


26 


8.5 


50 


slow 


undercut banks 


Notropis lutrensis 


26 


8.7 


8-15 


rapid 


riffles 



Analyzer, and other chemical aspects of water quality were determined using a 
Bausch and Lomb Minispec 20 spectrophotometer. 

Results 

On 3 March 1977 collections of fishes revealed nearly complete species sep- 
aration along various parts of the canal (Fig. 1, Table 2). Only desert pupfish, 
Cyprinodon macularius , were found in the shallowest, hottest water. They were 
abundant in water up to 10 cm deep at 39°C. An adjacent area of faster moving 
water at 32°C and 25 cm deep was inhabited only by schools of mosquitofish, 
Gambusia affinis. A few meters downflow, cold water at 22°C bubbled from a 
submerged pipe. At that point a small pool, 18 cm deep, contained in abundance 
only one species offish. These were mollies of the "shortfin" group that have 
been identified by their dentition in accordance with Hubbs (1961) as Poecilia 
sphenops, although in coloration they resemble a variety of Poecilia mexicana 
known in the aquarium trade as "liberty mollies" (Miller, pers. comm.). Mature 
males and females of this variety have a distinctive orange band in the dorsal fin 
and carry no striking colors on the caudal fin. In the Johnson Street canal on the 
eastern side of the Salton Sea "typical" Poecilia mexicana have a metallic blue 
caudal fin with a distal orange band. These mollies are found in a similar habitat 
along with the porthole fish, Poeciliopsis gracilis. The assemblage in the Johnson 
Street canal has been described by Mearns (1975). 

Water mixed slowly as it flowed in the King Street canal, the southern half of 
the stream remaining noticeably cooler for approximately 100 m. Water in this 
channel varied from 1 to 4 m in width and never was more than 50 cm deep. It 
meandered numerous times. Slow moving water up to 50 cm associated with 
undercut banks alternated with broad, fast moving riffles ca. 8 to 15 cm deep. 
Temperature varied from 24° to 27°C. Sailfin mollies, Poecilia latipinna, inhabited 
slow moving portions of the run, and red shiners, Notropis lutrensis were found 
in the riffles. Species overlapped very little in these habitats, although some 
Poecilia sphenops occurred in faster moving water with Notropis in upper por- 
tions of the run where the water temperature was 27°C. Approximately 2 km 
downflow the stream broadened to a floodplain no more than 10 cm deep and up 
to 5 m wide. Cyprinodon abounded in this area among roots of cattails (Typha), 
and rushes {J uncus). 

On 21 May 1977 the area was revisited. Distribution offish species had changed 
little. Water temperatures varied slightly from the determinations of 3 March. 



50 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. View east of upper portion of King Street canal (3 March 1977). Cold water source to 
extreme right center and undercut bank to left of center. Shallow wash to lower right. 



The thermal water measured ca. 2° cooler. The cold source was the same. Dis- 
solved oxygen measurements ranged from 7.2 mg/1 at 37°C to 8.5 mg/1 at 22°C. 
Other measures of water quality such as total dissolved solids at l.l%o showed 
little variability between the stations. 

On 18 March 1978, one year later, the area was visited in order to assess the 
influence of flooding on the previously observed niche separations. Heavy rains 
occurred during late summer 1977 and winter 1978. The area barely was recog- 
nizable. Discussion with nearby residents revealed that flooding had obliterated 
the stream banks, allowing water to flow over 3 m deep and 15 m wide. Virtually 
all rooted vegetation was removed. The following February, the channel was 
rebuilt. Bulldozers reconstructed canal walls, the bottom was scraped, and por- 
tions were stabilized with rocks held in place by wire mesh and iron posts. Figures 
2 and 3 depict the canal before and after reconstruction. 

Reconstruction greatly reduced habitat diversity and sorting of fish species was 
poorly defined (Table 3). The hot water and cold water inflow pipes were in their 
former positions and a temperature gradient remained. Above the cold water 
inflow a small earthen dam had been constructed, apparently to impound the hot 
water. This impoundment was a full meter in depth at the center and apparently 
was being used by local residents for bathing, washing clothes, and swimming. 



NICHE SEPARATION OF FRESHWATER FISHES 



51 




Fig. 3. View east of King Street canal following reconstruction (18 March 1978). 



Water temperature in the upper 3 cm was 36°C and dissolved oxygen was 7.2 
mg/1. At the upper, shallow end of the pond numerous young Cyprinodon were 
collected and released. A small number of Poecilia latipinna and Gambusia af- 
finis also were collected in this shallow area although they were more common 
in deeper portions of the pond. 

Downstream ca. 2 km the water course broadened as before, although distinct 
high banks had been constructed. No fishes were collected here, and there was 
no rooted vegetation. At the mouth of the channel, where it opened into the 
Salton Sea, Poecilia latipinna was abundant. It also abounded along the edge of 
the Salton Sea, in areas where the substrate was an odoriferous, black organic 
ooze. 

On 17 July 1978 another visit to the canal revealed that a portion of the dam 
that impounded the water had broken down and the pond was only 60 cm deep. 
No collections were made, but visual observations were enhanced by very clear 
water. Apparently Tilapia zilli had been introduced since the last visit. Two adults 
were observed swimming in the pond. The surface temperature was 40°C. In a 
shaded shallow of approximately one square meter surface, in water 32°C and no 
deeper than 3 cm, were swarming a mixture of Cyprinodon macularius, Poecilia 
latipinna, Gambusia affinis, and juvenile Tilapia zilli- Backs of the fishes fre- 
quently protruded from the water giving the area the appearance of boiling. 

Collections of fishes from other canals in the Salton Sea area seemed to confirm 



52 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 3. Habitat preferences for fishes inhabiting the King Street canal, Riverside County, Cali- 
fornia on 18 March 1978. 



Species 


Water 

Temp. 

(°C) 


Diss. O, 
(mg/1) 


Depth 
(cm) 


Flow 


Physical 
habitat 


Cyprinodon macularius 


36 

27 


7.2 
8.3 


2-100 
10 


slow 
rapid 


pond margin 
riffles 


Gambusia affinis 


36 


7.2 


2-100 


slow 


pond center 


Poecilia latipinna 


36 

28 


7.2 
7.8 


2-100 
50 


slow 
slow 


pond center 
stream margin 


Poecilia sphenops 


27 


8.3 


10 


rapid 


riffles 


Notropis lutrensis 


26 


8.3 


10 


rapid 


riffles 



the preferences exhibited by the population in the King Street canal. Some dif- 
ferences, however, are worth noting. 

In the Garfield Street canal Cyprinodon macularius were collected as recently 
as 7 December 1975 in water 50 cm deep among roots of cattails. That area is 
now inhabited by Tilapia zilli which were introduced to the Salton Sea area in 
1971 for the purpose of weed eradication (St. Amant, pers. comm.). Tilapia are 
aggressive, feeding on detritus as well as aquatic vegetation. They probably also 
eat fish eggs. My collections seem to indicate that where pupfish formerly abound- 
ed, Tilapia are now common. Tilapia apparently were not present in the King 
Street drain until recently. Observations of relative abundance of the two species 
will be continued to determine if Tilapia appears to replace Cyprinodon. 

Desert pupfish formerly were abundant in shoreline pools at the edge of the 
Salton Sea (Barlow, 1958a, b; Walker et al., 1961), and even were able to dive 
into the reducing, malodorous mud without apparent harm (Lowe et al., 1967). 
During rainy months certain pools along the north shore still are inhabited by 
pupfish (Black, pers. comm.), but such habitat adjacent to the mouth of the King 
Street canal seemed to be inhabited solely by swarms of sailfin mollies. It likewise 
seemed true near the mouth of the Avenue 81 canal where seine hauls yielded 
only Poecilia latipinna. Perhaps something about newly inundated land is hostile 
to desert pupfish. The Salton Sea is rising rapidly. Surface level has risen ca. 60 
cm since September 1975 (Skjold, pers. comm.). Perhaps nesting is upset when 
the water level rises too rapidly. Perhaps there is too much organic material in 
the substrate or water. Perhaps peculiar combinations or lethal concentrations of 
mineral ions are responsible. Whatever the case, it appears that the natural range 
of Cyprinodon macularius in the area has shifted such that freshwater drains and 
water courses leading to the Salton Sea are now the primary habitat. 

Discussion and Conclusions 

The sorts of preferences indicated by this study are notable in view of the wide 
range of tolerances exhibited by these fish species. The hottest water inhabited 
by fishes in the King Street canal, 40°C, is either within, or not far above the 
known upper limit of these species as follows: Notropis lutrensis, 39.5°C (Brues, 
1928); Poecilia sphenops, 35.6°C (Miller, 1949); Gambusia affinis, 39.5°C (Hubbs, 



NICHE SEPARATION OF FRESHWATER FISHES 53 

1959); and Cyprinodon macularius, 44.6°C (Barlow, 1958a, b; Lowe and Heath, 
1969). Poecilia latipinna was recorded in the pond at 36°C and Tilapia zilli at 
40°C. Of course, temperature stratification occurs and fishes may not necessarily 
occur at the temperature where the reading was taken. Nevertheless, when sev- 
eral species occur together, and the habitat is suitably diverse it is notable that 
they disperse themselves according to definable preferences. 

Temperature tolerances notwithstanding, the oxygen supply of natural waters 
is probably the most frequent single factor influencing the life of desert fishes 
(Deacon and Minckley, 1975). In the King Street canal, however, all water is 
flowing and approaches saturation for dissolved oxygen. Prior to construction of 
the pond, depth varied only slightly, ranging from 5 to 50 cm, therefore, temper- 
ature and physical characteristics such as turbulence and velocity are most likely 
the significant parameters responsible for the observed sorting of species. Com- 
paring the King Street canal to other canals in the area, it leads one to the 
conclusion that temperature is a very important factor. The pronounced thermal 
gradient, from 46°C to 22°C superimposed upon differences in flow and slight 
differences in depth was responsible for a nearly pure sorting of species based on 
habitat preferences. Considering that all but one of the species is introduced it 
further indicates that selective forces act rapidly. 

The native pupfish, Cyprinodon macularius, formerly inhabited marshes, 
sloughs, and backwaters of the Colorado, Gila, and Sonoyta Rivers (Hubbs and 
Miller, 1941; Miller, 1961). That habitat in the Salton Sea area is most closely 
approximated by warm, shallow, slow moving waters, particularly in association 
with rooted aquatic vegetation or algal mats. Desert pupfishes also inhabit saline 
pools along the edge of the Salton Sea, at least during the rainy season, but these 
habitats now seem mainly to contain sailfin mollies, perhaps due to unstable 
conditions associated with the rapidly rising water, ca. 60 cm since September, 
1975. In the King Street canal Cyprinodon was most abundant in a shallow wash 
at 39°C. It also was common ca. 2 km downflow in shallow water among roots 
of cattails and sedges. 

"Liberty mollies," identified on the basis of dentition as Poecilia sphenops, 
appeared to inhabit only the King Street canal, and were found most abundantly 
near the outflow of water at 22°C. In some of the other canals this habitat is 
occupied by the closely related shortfin molly, Poecilia mexicana, and/or Poe- 
ci Hops is gracilis, neither of which was collected in the King Street canal. Red 
shiners, Notropis lutrensis were the only fishes collected in riffles of the King 
Street canal although they also were taken in small numbers with Poecilia sphe- 
nops in fast moving water other than riffles. By comparison, it is interesting to 
note that my collections of Notropis from the Gila and Verde Rivers in Arizona 
seemed to indicate that they preferred slow moving water deeper than 20 cm. 

The most common fishes of the canal system were sailfin mollies, Poecilia 
latipinna. In the King Street canal they preferred slow moving water in associ- 
ation with undercut banks. They reached their greatest abundance in brackish 
water near the mouth of the canal, and in shallows at the edge of the Salton Sea 
where the substrate had become an odoriferous, black, anaerobic gel. 

Mosquitofish, Gambusia affinis, occurred in slow moving waters at 32°C in the 
King Street canal. In other canals they were found in slow moving water with 
Poecilia latipinna, but advanced farther into vegetation, and not as far into the 



54 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

current as the mollies. Also, they seemed to avoid shallow or weedy areas in- 
habited by desert pupfishes. 

The impact of flooding or other forms of habitat destruction on niche prefer- 
ences is marked. In the Avenue 81 canal Notropis lutrensis used to be common 
along with Poecilia latipinna. Since the flooding of September 1976, however, it 
appears that Notropis may have been eliminated. After three pronounced floods, 
Cvprinodon macularius , the native species, and Poecilia latipinna occur there 
as before. Regarding Poecilia latipinna, it is interesting to note that it apparently 
was the only species extirpated from the Salt River near Mesa, Arizona by the 
flood of 1970 (Minckley, 1973). Apparently the ability of sailfin mollies and desert 
pupfish to tolerate saline waters of the Salton Sea enables them to reinhabit these 
canals after flooding. Perhaps the absence of such a refuge in the Salt River led 
to the demise of Poecilia latipinna in the Salt River. 

In the King Street canal, flooding and subsequent reconstruction dramatically 
reduced the diversity of habitat and the striking examples of niche separation 
observed in 1977 were obliterated. All original species were still present as re- 
cently as 17 July 1978 and some examples of species sorting were still observable, 
but they were far less obvious. Essentially two habitats remained, cool flowing 
water and impounded hot water. The former was inhabited by Notropis lutrensis, 
Poecilia sphenops, and a few Cvprinodon macularius. The impounded hot water 
apparently is used for washing clothes and human bathing. Recently Tilapia zilli 
was introduced here. Poecilia latipinna, Gambusia affinis, and Cyprinodon mac- 
ularius were found there with the Tilapia. The ability of this population of desert 
pupfishes to tolerate pollutants from human bathing and washing clothes as well 
as encroachment by Tilapia zilli remains to be seen. 

Literature Cited 

Barlow, G. W. 1958a. Daily movements of desert pupfish, Cyprinodon macularius, in shore pools 

of the Salton Sea, California. Ecology, 39:580-587. 

. 1958b. High salinity mortality of desert pupfish, Cyprinodon macularius. Copeia, 231-232. 

Brues, C. T. 1928. Studies on the fauna of hot springs in the western United States and the biology 

of thermophilous animals. Proc. Amer. Acad. Arts Sci., 63:139-228. 
Deacon, J. E., and W. L. Minckley. 1973. Desert Fishes. Pp. 385-488 in Desert Biology, Vol. II 

(G. W. Brown, Jr. ed.), Academic Press, xv 4- 601 pp. 
Hubbs, C. 1959. High incidence of vertebral deformities in two natural populations of fishes inhab- 
iting warm springs. Ecology, 40:154-155. 
. 1961. Isolating mechanisms in the speciation of fishes. Pp. 5-23 in Vertebrate Speciation 

(W. F. Blair, ed.), Univ. Texas Press, xvi + 642 pp. 
Hubbs, C. L., and R. R. Miller. 1941. Studies of the fishes of the order Cyprinodontes. XVII. Genera 

and species of the Colorado River system. Occ. Pap. Mus. Zool. Univ. Mich., 433:1-9. 
Lowe, C. H., and W. G. Heath. 1969. Behavioral and physiological responses to temperature in the 

desert pupfish, Cyprinodon macularius. Physiol. Zool., 42:53-59. 
, D. S. Hinds, and E. A. Halpern. 1967. Experimental catastrophic selection and tolerances 

to low oxygen concentration in native Arizona freshwater fishes. Ecology, 48:1013-1017. 
Mearns, A. J. 1975. Poeciliopsis gracilis (Heckel), a newly introduced poeciliid fish in California. 

Calif. Fish and Game, 61:251-253. 
Miller, R. R. 1949. Hot springs and fish life. Aquarium J., 20:286-288. 
. 1961. Man and the changing fish fauna of the American Southwest. Pap. Mich. Acad. Sci. 

Arts Lett., 46:365-404. 
Minckley, W. L. 1973. Fishes of Arizona. Sims Printing Co., Phoenix, xv + 293 pp. 



NICHE SEPARATION OF FRESHWATER FISHES 55 



St. Amant, J. A., and I. Sharp. 1971. Addition of Xiphophorus variatus (Meek) to the California fish 
fauna. Calif. Fish and Game, 57:128. 

Walker, B. W., R. R. Whitney, and G. W. Barlow. 1961. The fishes of the Salton Sea. Pp. 77-91 in 
The ecology of the Salton Sea, California, in relation to the sportfishery (B. W. Walker, ed.), 
State of California Department of Fish and Game, Fish Bulletin No. 1 13, 204 pp. 

Accepted for publication February 28, 1979. 



Bull. Southern California Acad. Sci. 
78(1), 1979, pp. 56-61 



Research Notes 

Two Asiatic Gobiid Fishes, Tridentiger trigonocephalus and 
Acanthogobius flavimanus, in Southern California 



While SCUBA diving in Los Angeles Harbor I collected mature adults and 
eggs of one Asiatic goby, Tridentiger trigonocephalus (Gill) and photographed 
another, Acanthogobius flavimanus (Temminck and Schlegel). Subsequent col- 
lections indicated that both are established in Los Angeles-Long Beach Harbors 
and vicinity, and that the latter species may be rapidly extending its range in 
southern California. 

The chameleon goby, T. trigonocephalus (Fig. 1), reaching a length of approx- 
imately 90 mm TL, is larger than most native Californian gobies, while the yel- 
lowfin goby, A. flavimanus, which grows to approximately 240 mm TL, is larger 
than all Californian gobies (Miller and Lea, 1976). The latter species seems to be 
highly adaptable to marine and fresh waters and able to extend its range rapidly 
(Brittan et al., 1970; Kukowski, 1972). 

Tridentiger trigonocephalus 

Tridentiger trigonocephalus occurs in marine and brackish waters from Hong 
Kong to the Amur River Basin, USSR, and the Japanese Islands (Fowler, 1961; 
Tomiyama, 1936), on hard and soft substrata (Carl L. Hubbs, Scripps Institution 
of Oceanography, pers. comrn.; Dotu, 1957, 1958). This goby has been reported 
from Los Angeles Harbor (Hubbs and Miller, 1965; Miller and Lea, 1976), and 
is established in San Francisco Bay (Ruth, 1964; Brittan et al., 1970) and Sydney 
Harbor, Australia (Hoese, 1973; Friese, 1973). 

On 1 June 1960, Jim Wright of Marineland of the Pacific observed two, but 
could only collect one goby at Fish Harbor in Los Angeles Harbor (John H. 
Prescott, New England Aquarium, pers. comm.). The single specimen (Fig. 1) 
was identified by Carl L. Hubbs as T. bifasciatus Steindachner, a synonym of T. 
trigonocephalus (Tomiyama, 1936; Koumans, 1940), and was the first capture of 
this goby in American waters. It is a 70.4 mm SL male captured on a rock jetty 
at a depth of 3 m. The specimen is deposited in the fish collection of Scripps 
Institution of Oceanography (SIO 77-337). This specimen is the basis of the report 
by Hubbs and Miller (1965) of this goby being in Los Angeles Harbor. 

On 24 May 1977, I collected three females, 49.2 to 57.2 mm SL, and twelve 
males, 56.7 to 77.3 mm SL, chameleon gobies in Los Angeles Harbor on the 
rocky "rip-rap" under Pier 228, Slips D and E, in depths of 0.3 to 3 m. Most 
specimens were collected from discarded bottles littering the bottom. The surface 
water temperature was 18°C and salinity was 34.3% f at the surface and at 3 m. 
Visibility was 1 to 2 m. These specimens are deposited in the fish collection of 
the Natural History Museum of Los Angeles County (LACM 36989-1). 

Gonadal inspections of all specimens revealed well developed eggs and testes. 
Gut examinations of all specimens revealed, in order of decreasing frequency: 
crustaceans, including gammarid and caprellid amphipods, copepods, isopods, 



RESEARCH NOTES 57 




Jf % - 



Fig. 1. The first Thdentiger trigonocephalus, captured in Los Angeles Harbor. Standard length 
is 70.4 mm. 



and ostracods; polychaete worms; gastropods; and fish eggs, as evidenced by one 
gut containing 478 eggs. 

On 22 September 1977, I collected two males, 44 and 78 mm SL, and a 66 mm 
SL female T. trigonocephalus from the same location. The larger male and the 
female, together with a clutch of 300 eggs, were collected from an aluminum beer 
can. 

Acanthogobius flavimanus 

Acanthogobius flavimanus occurs naturally in Japan, Korea, and China (Fow- 
ler, 1961; Tomiyama, 1936) in marine, brackish, and river waters (Okada, 1955). 
It has been introduced into Sydney Harbor, Australia (Hoese, 1973; Friese, 1973). 
This goby was first found in California in 1963 in the San Joaquin River and the 
Stockton Deepwater Channel (Brittan et al., 1963). Since then it has been reported 
from the Sacramento Delta, the San Francisco Bay area, and Bolinas Lagoon 
(Brittan et al., 1970), Elkhorn Slough (Kukowski, 1972), and Tomales Bay and 
Estero Americano (Miller and Lea, 1976). It has not heretofore been reported 
from southern California. 

An underwater photograph (Fig. 2) taken in Los Angeles Harbor under Pier 
228 on 22 September 1977, revealed the presence of A. flavimanus in the rocky 
"rip-rap." No specimens were captured. On 29 March 1978, biologists from Ma- 
rine Biological Consultants, Inc., Costa Mesa, California, took a 156 mm SL male 
(Fig. 3) in a gill net {W2 in. bar mesh) at a depth of 18 m in the back harbor, 
Channel 3, of Long Beach Harbor (John L. Wintersteen, pers. comm.). This 
specimen is deposited in the Natural History Museum of Los Angeles County 
(LACM 37346-1). The testes of this individual were not well developed. Stomach 



58 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. Underwater photograph of Acanthogobius flavimanus, taken in Los Angeles Harbor. 
Standard length is estimated at 100 mm. 



contents included a crab, ostracods, amphipods, copepods, and a large amount 
of unidentifiable digested material. 

On 15 April 1978, two specimens of A. flavimanus, 24 and 30 mm SL, were 
collected in a beach seine over the silty substratum of Upper Newport Bay, about 
37 km southeast of Los Angeles Harbor, by Michael H. Horn, of California State 
University at Fullerton, and Larry G. Allen, of the University of Southern Cal- 
ifornia (pers. comm.). They subsequently collected another specimen, 48 mm SL 
on 4 May 1978, and a fourth specimen, 57 mm SL, on 14 May 1978, from the 
same locality. These gobies are being retained in the fish collection of California 
State University at Fullerton (uncataloged). Stomach contents of these four go- 
bies included ostracods, copepods, and amphipods. 

On 26 June 1978, Eric H. Knaggs of the California Department of Fish and 
Game (pers. comm.) beach seined four yellowfin gobies, 66 to 90 mm SL, in the 
sandy mud bottom of the San Gabriel River, between the Westminster Avenue 
and Seventh Street bridges (Long Beach, Los Angeles County) about 13 km east 
of Los Angeles Harbor. These gobies are deposited in the Natural History Mu- 
seum of Los Angeles County (LACM 37711-1). On 31 July 1978 he collected two 
more specimens, 71 and 84 mm SL, from the Long Beach Swimming Lagoon, 
near the mouth of the Los Angeles River. These latter specimens were sacrificed 
for other studies. The salinity and temperature of the localities of these two 
collections are unknown, but both areas are influenced by marine and fresh 
waters. 



RESEARCH NOTES 59 




Fig. 3. Acanthogobius flavimanus , collected in Long Beach Harbor. Standard length is 156 mm. 

Discussion 

The occurrence of numerous mature and spawning adults is evidence that an 
established population of T. trigonocephalus exists in Los Angeles Harbor. The 
presence of small, probably locally spawned, A. flavimanus can be taken as 
evidence that it is established in Newport Bay. An observation in Los Angeles 
Harbor and a collection of yellowfin gobies in Long Beach Harbor waters and 
the San Gabriel River are strong evidence that this goby is established and may 
be expanding its range in southern California. 

Because of the nature of international shipping, there is little doubt that the 
Los Angeles-Long Beach Harbors are the locations of introductions of both 
gobies into southern California. There is no regular international shipping into 
Newport Bay or the San Gabriel River. The fact that these gobies only occur 
extralimitally in widespread Pacific port cities and vicinities would also seem to 
implicate shipping. 

The actual method of introduction is unknown, but suggested mechanisms gen- 
erally involve the transport of eggs on fouling organisms growing on the hulls, or 
in the seawater systems of ships (Brittan et al., 1963; Hubbs and Prescott in 
Hubbs and Miller, 1965; Dawson, 1973; Hoese, 1973). Another mechanism, that 
of egg transport on oysters, would not be applicable to Los Angeles-Long Beach 
Harbors, as no oysters have been introduced there. 

Dawson (1973) estimated that minimum ship transit time from Yokohama, Ja- 
pan to San Francisco was eight days in 1972. The average incubation time of eggs 
of T. trigonocephalus is 8.5 days at 20°C (Dotu, 1958), while that of A. flavimanus 
is about 28 days at 13°C (Dotu and Mito, 1955). Therefore, it is possible that eggs 
of both gobies could be laid on fouling organisms of ships hulls in the Orient, and 
be transported to the west coast of the United States in time for hatching. Sea- 
water temperatures along likely transpacific shipping routes are sufficient for egg 
development. 

Dawson (1973) convincingly related the recent (since the mid-1950's) increase 
in exotic introductions of Indo-West Pacific organisms into the Americas to the 
increased speeds of modern ships. John E. Fitch (California Department of Fish 
and Game, pers. comm.), suggested another mechanism of introduction of exotic 
marine organisms might occur when occupants of aquaria, found on many large 
ships, were dumped into foreign harbors. 

The problems associated with the introductions of exotic species into a habitat 



60 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

were discussed by Lachner et al. (1970). Such introductions, be they deliberate 
or accidental, are no longer unusual, and most ecologists would consider them 
unwise. 

It appears that these two Asiatic gobies have become a permanent part of 
southern California's ichthyofauna. One, T. trigonocephalus , seems to be re- 
stricted (in southern California) to Los Angeles, and probably Long Beach Har- 
bors, while A. flavimanus is extending its range down coast from Los Angeles 
Harbor, a situation reminiscent of the "explosive spread' 1 reported by Brittan et 
al. (1970) in the San Francisco Bay-Delta region. 

Virtually nothing is known about the effect of these gobies on native species. 
If A. flavimanus continues to expand into streams, lagoons, and bays down coast, 
there is a possibility that some populations of tidewater gobies, Eucyclogobius 
newberryi might be eliminated (Moyle, 1976), and staghorn sculpin, Leptocottus 
armatus might be displaced (Brittan et al., 1970). The longjaw mudsucker goby, 
Gillichthys mirabilis is collected in back bays and sloughs for use as bait in local 
lakes, the Salton Sea, Lake Mead, and the Colorado River (Turner and Sexsmith, 
1964). If A. flavimanus is collected with bait species, it could be transported to 
other marine and freshwater habitats, where it could become established. 

I wish to thank Carl L. Hubbs, John H. Prescott, John L. Wintersteen, Michael 
H. Horn, Larry G. Allen, and Eric H. Knaggs for making goby specimens and 
various data available to me. 

Herbert W. Frey and John E. Fitch read the manuscript and offered valuable 
suggestions. Robert L. Lavenberg, Lillian J. Dempster, Leslie Harris, Kenneth 
F. Mais, and Doyle A. Hanan assisted in various aspects. This work was accom- 
plished under Project DJ-F-27-D7, Federal Aid to Sport Fish Restoration, U.S. 
Fish and Wildlife Service. 

Literature Cited 

Brittan, M. R., A. B. Albrecht, and J. B. Hopkirk. 1963. An oriental goby collected in the San 

Joaquin River Delta near Stockton, California. Calif. Fish and Game, 49:302-304. 
, J. D. Hopkirk, J. D. Conners, and M. Martin. 1970. Explosive spread of the oriental 

goby Acanthogobius flavimanus in the San Francisco Bay-Delta region of California. Proc. 

Calif. Acad. Sci., 38:207-214. 
Dawson, C. E. 1973. Occurrence of an exotic eleotrid fish in Panama with discussion of probable 

origin and mode of introduction. Copeia, 1973:141-144. 
Dotu, Y. 1957. The bionomics and life history of the goby, Triaenopogon barbatus (Giinther) in the 

innermost part of Ariake Sound. Sci. Bull. Fac. Agri., Kyushu Univ., 16:261-274. 
. 1958. The bionomics and life history of two gobioid fishes, Tridentiger undicericus Tomi- 

yama and Tridentiger trigonocephalus (Gill) in the innermost part of Ariake Sound. Sci. Bull. 

Fac. Agri., Kyushu Univ., 16:343-358. 
, and S. Mito. 1955. On the breeding-habits, larvae and young of a goby, Acanthogobius 

flavimanus (Temminck et Schlegel). Japanese J., 4:153-161. 
Fowler, H. W. 1961. A synopsis of the fishes of China. Part IX. The gobioid fishes. Quart. J. Taiwan 

Mus., 14:203-250. 
Friese, J. E. 1973. Another Japanese goby in Australian waters: what next? Koolewong. Proc. Roy. 

Zool. Soc. New South Wales, 2:5-7. 
Hoese, D. F. 1973. The introduction of the gobiid fishes Acanthogobius flavimanus and Tridentiger 

trigonocephalus into Australia. Koolewong. Proc. Roy. Zool. Soc. New South Wales, 2:3-5. 
Hubbs, C. L., and R. R. Miller. 1965. Studies of cyprinodont fishes. XXII. Variation in Lucania 

parva, its establishment in western United States, and description of a new species from an 

interior basin in Coahuila, Mexico. Misc. Publ. Mus. Zool. Univ. Mich., 127:1-111. 



RESEARCH NOTES 61 

Koumans, F. P. 1940. On a collection of fishes from East Java. Z. Mededeelinger, 22:257-264. 

Kukowski, G. E. 1972. Southern range extension for the yellowfin goby, Acanthogobius flavimanus 
(Temminck and Schlegel). Calif. Fish and Game, 58:326-327. 

Lachner, E. A., C. R. Robins, and W. R. Courtenay, Jr. 1970. Exotic fishes and other aquatic 
organisms introduced into North America. Smithsonian Contrib. Zool., 59:1-29. 

Miller, D. J., and R. N. Lea. 1976. Guide to the coastal marine fishes of California. Calif. Fish and 
Game, Fish Bull., 157:1-249. (Revision, including addendum publ. by Div. Agric. Sci., Univ. 
Calif., Richmond.) 

Moyle, P. B. 1976. Inland fishes of California. U. Calif. Press. Berkeley, 405 pp. 

Okada, Y. 1955. Fishes of Japan. Maruzen Co., Ltd., Tokyo, 434 + 28 pp. 

Ruth, F. S. 1964. Habitat check list of the vertebrates of the San Francisco Bay Region California. 
Mimeo-type Corp., Walnut Creek, 16 pp. 

Tomiyama, I. 1936. Gobiidae of Japan. Japanese J. Zool., 7:37-112. 

Turner, C. H., and U. C. Sexsmith. 1964. Marine baits of California. Calif. Fish and Game, Sac- 
ramento, 71 pp. 

Accepted for publication August 16, 1978. 

Peter L. Haaker, California Dept. of Fish and Game, 350 Golden Shore, Long 
Beach, California 90802. 



Bull. Southern California Acad. Sci. 
78(1), 1979, pp. 61-67 



The Velvet Whalefish, Barbourisia rufa, added to California's 

Marine Fauna, with Notes on Otoliths of Whalefishes and 

Possible Related Genera 



California's north coast trawl fishermen normally sort their catches at sea and 
deliver the most desirable species (rockfishes, Sebastes spp. and Sebastolobus 
spp.; sablefish, Anoplopoma fimbria; lingcod, Ophiodon elongatus; and several 
kinds of pleuronectid flatfish) in a fresh state to shoreside markets where most 
are filleted, packaged, and frozen. If, when sorting his catch, a fisherman finds 
what he believes is a rare or unusual fish, mollusk, crustacean, or other organism 
he often will leave it at the market with a request that it be placed in their freezer 
and saved for the California Department of Fish and Game (DFG). Similarly, if 
while processing a load, plant personnel encounter an "odd-ball" (some animal 
of unusual color, size, or condition; or an organism that is tagged or one that 
cannot be identified by those present at the time; etc.) that specimen too will be 
saved. 

Unless forgotten or overlooked, this material is then turned over to the first 
DFG employee to stop at the plant during routine surveillance or sampling op- 
erations, and at that time he is informed (usually verbally) of the name of the boat 
and/or skipper who caught the oddity. It is then necessary for the DFG employee 
to contact the vessel skipper during his next trip to port, and obtain complete 
collecting data for the specimen or specimens in question. 

Unfortunately, as in the present case, an occasional item is saved for which 
collecting data cannot be obtained, even after making dozens of inquiries over a 
lengthy period of time. In these instances, it is necessary to make assumptions 



62 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

based upon available evidence, which if actual facts were known, might not hold 
up. The whalefish in question, 330 mm standard length (SL) and weighing 456 g, 
was picked up by DFG biologist Lawrence Quirollo at Eureka Fisheries in Cres- 
cent City on 18 June 1974, and represented the first record of Barbourisia rufa 
from the eastern Pacific. It was thought to have been in Eureka Fisheries' freezer 
for about one month before it was picked up, but even though an enlarged pho- 
tograph of the fish was circulated among all the then employed plant personnel, 
no one could remember having seen the fish previously, nor did they know who 
might have placed it in the freezer. The photograph also was shown to all fish- 
ermen who had made deliveries to Eureka Fisheries during the first six months 
of 1974, with similar negative results. It was assumed then, that the fish had been 
saved by some former employee of Eureka Fisheries, but several individuals who 
fitted this concept could not be located. 

An examination of fish landing receipts for several months prior to June 1974 
indicated that all catches delivered to Eureka Fisheries at Crescent City during 
that period had been made off California, most in the vicinity of Redding Rock. 
No records were found of deliveries by vessels that had been fishing off Oregon, 
so it must be assumed that this whalefish had been caught off California. 

Previously, based upon fewer than 20 specimens, Barbourisia rufa had been 
reported from a number of localities throughout the world: Gulf of Mexico (type 
locality), Indian Ocean near Madagascar, South Africa, Kurile Islands, Florida, 
Japan, and the south Pacific near New Caledonia (Trunov, 1968; Penrith, 1969; 
Fourmanoir, 1970a, 1970b; Maruyama and Ono, 1973). John R. Paxton, the Aus- 
tralian Museum, has informed me (personal comm.), however, that he has been 
able to locate published and unpublished capture records for 43 specimens of B. 
rufa, with eight from the Hawaiian area representing the closest previous captures 
to our shores. His records extended into the first few months of 1976. Only two 
of the 43 individuals were larger than the Californian whalefish: one from off 
South Africa at 345 mm SL, and one from lat. 22°14'S long. 02°56'E at 333 mm 
SL. Meristic data on the Californian Barbourisia, as determined from X-rays are: 
dorsal rays 22, anal rays 17, caudal rays 5 + 19 + 4 (dorsal rudimentary, prin- 
cipal, and ventral rudimentary), and vertebrae 44. The otoliths (sagittae) of this 
fish were removed and examined for evidence of age, and although numerous 
alternating opaque and hyaline zones could be discerned, these could not be 
interpreted as being annuli. The finely spinulose scales, which are unique to 
Barbourisia rufa, give a velvety texture to the skin which eased the task of 
coining a vernacular. 

Among recent publications, Rosen and Patterson (1969:457), after considerable 
discussion, state that they "feel confident that the correct position of the ceto- 
mimoids is within the Beryciformes. ,, They mention that Rondeletia bicolor lacks 
a swimbladder, as does Scopelogadus beani, but go no further in drawing con- 
clusions regarding affinities of these two genera (families). Earlier (p. 373), Rosen 
and Patterson, in discussing family Neoscopelidae, point out that in Scopelengys, 
one of the three genera assigned to the family, "the arrangement of the adductor 
mandibulae was found to be characteristic of all members of Berycoidei and of 
the single melamphaid [Melamphaes bispinosus] examined." 

Gosline (1971:134-136) divides the order Cetomimiformes into three groups of 
families depending upon placement of pelvic and dorsal fins. He states that Ce- 



RESEARCH NOTES 



63 




Fig. 1. Right sagittae from known whalefish genera (otolith length by height and fish standard 
length also noted): a) Barbourisia rufa 5.0 x 1.1 mm (330 mm SL); b) Gyrinomimus sp. 2.0 x 1.5 
mm (310 mm SL); c) Cetomimus sp. 1.4 x 1.2 mm (159 mm SL); d) Rondeletici loricata 1.2 x 1.9 
mm (83 mm SL); e) Cetostomus regani 1.3 x 1.3 mm (213 mm SL); Ditropichthys sroreri 1.4 x 
1.6 mm (SL unk.). 



64 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. Left sagittae of fishes belonging to suborder Stephanoberycoidei (otolith lengths and fish 
standard lengths in mm are noted): a) Stephanoberyx monae (2.4, 99.5); b) Scopelogadus bispinosus 
(2.3, 80); c) Gibberichthys latifrons (1.5, 95); d) Melamphaes lugubris (3.7, 41). Outline of sulcus has 
been indicated with broken, inked-in line. 



tomimus (Cetomimidae) "has been shown (Parr 1929) to have strong osteological 
affinities with myctophiform fishes," and includes it in his second group of fam- 
ilies (pelvic fins jugular or absent; dorsal fin posterior). In discussing his third 
group (pelvics subabdominal) he concludes that, "aside from Barbourisia, there 
seems no reason to think that all of the members of the present group are not 
beryciform derivatives, possible [sic] related to the Melamphaidae and Stepha- 
noberycidae." 

Finally, Nelson (1973:190) places the three whalefish families in the suborder 
Cetomimoidei (Order Beryciformes), following the lead of Rosen and Patterson 
(1969). He does not discuss their possible ancestry. 

Since whalefish otoliths previously had not been discussed, illustrated nor com- 
pared, and since I had material from all extant genera in the three whalefish 
families (Cetomimidae, Rondeletiidae and Barbourisiidae), it seemed worthwhile 
to determine if they might be useful in whalefish taxonomy or in discerning fa- 
milial relationships. 



RESEARCH NOTES 



65 




Fig. 3. Left sagittae of fish genera belonging to family Neoscopelidae (otolith lengths and fish 
standard lengths in mm are noted): a) Solivomer arenidens (1.5, 76); b) Neoscopelus sp. (8.0, 150); 
c) Scopelengys tristis (3.2, 172); d) Neoscopelus sp. (1.7, 37). Outline of sulcus has been indicated 
with broken, inked-in line. 



In using otoliths (sagittae) as a taxonomic character, the most important feature 
for determining family relationships (and ancestry) is the sulcus (groove on the 
inner face): its position, configuration and differentiation. Although, in the Ce- 
tomimidae the sulcus is slightly excavated, it appears to be a slightly raised "blip" 
in the center of the otolith; it opens to the margin anterodorsally via a presulcal 
trough. Ostium (anterior part of sulcus) and cauda (posterior part) are undiffer- 
entiated. In Gyrinomimus, Cetomimus and Ditropichthys (Figs, lb, c and f), the 
otolith has smoothly rounded margins, and appears trilobate in outline. The sulcus 
is encircled by a low, smooth ridge except at the point of entry for the presulcal 
trough. In Cetostomus (Fig. le), the otolith margin is roughened, and the sulcus 
is rimmed by a high, relatively sharp ridge, somewhat foliose in appearance. 
Otolith length in all four genera is equal to or slightly greater than otolith height. 

In Rondeletia (Fig. Id, Rondeletiidae), the sulcus is slightly excavated, circular 
in outline, and surrounded by a slightly raised rim. In contrast with cetomimid 
otoliths, the sagitta of Rondeletia is much higher than long, and the sulcus lies 
just inside the anterior margin. A very short presulcal trough leads to the margin, 
and as with cetomimid otoliths, ostium and cauda cannot be distinguished. 

In Barbourisia (Fig. la, Barbourisiidae) the sulcus is elongate and deeply ex- 
cavated. It is surrounded by a high, broadly rounded rim, and as in the other two 



66 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

whalefish families, a presulcal trough runs diagonally upwards to the anterodorsal 
margin of the otolith. The otolith is more than four times longer than high. 

Since otoliths were on hand from all families placed in the order Beryciformes 
by Nelson (1973) except Korsogasteridae, and from all families placed in suborder 
Myctophoidei by Greenwood, Rosen, Weitzman and Myers (1966), these were 
examined to see if they could shed any light on the problem of whalefish rela- 
tionships. 

Based only upon otolith characters, the suborder Cetomimoidei of Nelson 
(1973:190) should be more closely allied to his suborder Stephanoberycoidei (p. 
186) than Berycoidei (p. 187). Sagittae of all five melamphaid genera (Stepha- 
noberycoidei) have a centrally located, round or oval sulcus that is surrounded by 
a low rim, and there is a presulcal trough leading to the anterodorsal margin 
(Weiler, 1968). The sagittae of Scopelogadus spp. (Fig. 2b) show greater simi- 
larity to whalefish otoliths than do sagittae of Melamphaes spp. (Fig. 2d) which 
are typical of the other three melamphaid genera. Otoliths of Stephanoberyx 
monae (Fig. 2a) have sulcal characters that are intermediate between those found 
in melamphaid genera and Gibberichthys latifrons (Gibberichthyidae: 
Stephanoberycoidei, Fig. 2c). 

Only Scopelengys (Neoscopelidae, Fig. 3c) among myctophoid families has 
otoliths with sulcal characters similar to those found in whalefishes and melam- 
phaids. Sulcal characters of Neoscopelus (Fig. 3b and 3d) and Solivomer (Fig. 
3a), the other two genera in the Neoscopelidae, show closer relationships to the 
Myctophidae {Neoscopelus) and Berycidae {Solivomer) than they do to the whale- 
fishes. Nafpaktitis (1977) reviewed previous research on the Neoscopelidae, but 
still needed is an in-depth comparison of the three neoscopelid genera to deter- 
mine if the family as presently constituted accurately reflects relationships or is 
in fact a heterogeneous assemblage of oceanic misfits. 

Acknowledgments 

Special thanks go to Lawrence Quirollo, California Department of Fish and 
Game (DFG), Eureka, for picking up the specimen at the waterfront, for seeing 
that it was taken care of properly until he could send it to me, and for the many 
hours he subsequently put in trying to obtain capture data. Several individuals 
contributed otoliths that were critical to this study: Robert Lavenberg, Natural 
History Museum of Los Angeles County; Basil Nafpaktitis, University of South- 
ern California; and Bruce Robison, University of California, Santa Barbara. John 
Paxton, The Australian Museum, and Richard Rosenblatt, Scripps Institution of 
Oceanography furnished information on material in collections at their institutions 
and elsewhere. Lillian Dempster and W. I. Follett, California Academy of Sci- 
ences supplied some much needed references and other information. Jack W. 
Schott, DFG, Long Beach took the excellent otolith photos, and Micaela Wolfe, 
DFG, Long Beach, typed the manuscript. 

Literature Cited 

Fourmanoir, P. 1970a. Notes ichtyologiques (I). Cah. O.R.S.T.O.M., ser. Oceanogr., 8(2):19-34. 

. 1970b. Notes ichtyologiques (II). Cah. O.R.S.T.O.M., ser. Oceanogr., 8(3):35-46. 

Gosline, W. A. 1971. Functional morphology and classification of teleostean fishes. Honolulu, Univ. 
Press of Hawaii. 208 pp. 



RESEARCH NOTES 67 



Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. 1966. Phyletic studies of 

teleostean fishes, with a provisional classification of living forms. Amer. Mus. Nat. Hist., Bull., 

131(4):339-456. 
Maruyama, K., and K. Ono. 1973. On the second record of Barbourisia rufa from Japan. Jap. J. 

Ichthyol., 20(4):242-244 (Japanese text, English summary). 
Nafpaktitis, Basil G. 1977. Fishes of the western north Atlantic. Family Neoscopelidae. Sears Found. 

Mar. Res., Mem., 1(7): 1-12. 
Nelson, J. S. 1973. Fishes of the world. New York, John Wiley and Sons. 416 pp. 
Parr, A. E. 1929. A contribution to the osteology and classification of the orders Iniomi and Xeno- 

beryces. Bingham Oceanogr. Coll., Occ. Pap., 2:1-45. 
Penrith, M. J. 1969. New records of deep-water fishes from south west Africa. Cimbebasia, ser. A, 

l(3):59-75. 
Rosen, D. E., and C. Patterson. 1969. The structure and relationships of the paracanthopterygian 

fishes. Amer. Mus. Nat. Hist., Bull., 141(3):357-474. 
Trunov, I. A. 1968. The whalefish (Barbourisia rufa, Fam. Barbourisiidae) and the frilled shark 

(Chlamydoselachus anguineus, Fam. Chlamydoselachidae) in southwest African coastal 

waters. Voprosy Ikhtiologii, Acad. Sci., USSR, 8(1): 135-137. (English-language edition, Scrip- 

ta Technica Inc.) 
Weiler, W. 1968. Die Otolithen der bathypelagischen Familie Melamphaidae und ihre systematische 

Bedeutung. Senckenbergiana biol., 49(3/4):223-230. 

Accepted for publication October 8, 1978. 

John E. Fitch, California Department of Fish and Game, Long Beach, California 
90802. 



Bull. Southern California Acad. Sci. 
78(1), 1979, pp. 67-69 



The Larva of Onthophagus medorensis Brown with Notes on its 
Biology (Coleoptera: Scarabaeidae) 



Onthophagus medorensis Brown (Brown, 1929) is a small, fairly common green 
to blue dung beetle which occurs from Kansas south to southern Texas, east to 
Louisiana and Arkansas and west to the edge of the shortgrass prairie (Howden 
and Cartwright, 1963). According to Edmonds and Halffter (1978), the larva has 
not been described. From data accompanying previous collections (Howden and 
Cartwright, 1963), it would appear that O. medorensis is partial to shaded, sandy 
areas and occurs on rotting plant material in addition to a variety of dung types. 
Brown (1926, 1928) indicates that most specimens have been taking during the 
early summer. 

During the course of this study, two series containing four adult specimens 
each of O. medorensis were collected. Both series showed the metallic green 
phase of coloration described in Howden and Cartwright (1963). The first collec- 
tion was made from human dung found near a construction site in mixed pine- 
oak woodland five miles south of Shreveport, Caddo Parish, Louisiana on 5 May 
1977; these beetles were saved alive for rearing of the larvae. The other collection 
was made on 29 October 1977 from rabbit dung found in a mixed pine-oak wood 
on the Texas Eastern University campus, Smith County, Texas. Other dung 



68 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



2.0 mm 






Fig. 1. Third-stage larva of Onthophagus medorensis Brown: a. head capsule; b. mandibles; c. 
epipharynx; d. arrangement of setae on venter of last abdominal segment (diagrammatic). 



beetles taken with these collections were two O. oklahomensis Brown (Louisiana 
collection) and one Canthon sp. (Texas collection). 

The specimens of O. medorensis from the Louisiana collection were placed in 
a quart rearing jar filled to within 25 mm of the top with moist sand taken from 
the collection site and covered with a layer of fresh human dung. On 1 June 1977, 
the jar was emptied and the adult beetles and brood cells retrieved. Four sub- 
spherical brood cells measuring approximately 17 mm in diameter were found; 
each contained a third instar larva. All four larvae were fixed and preserved in 
hot Ward's insect larva preservative prior to dissection and description. 

The following description employs the terminology of Ritcher (1966). 

O. medorensis Brown, third-stage larva (description based on four specimens; 
Fig. 1): Maximum width of head capsule 1.68-1.71 mm; epicranial stem deeply 
forked basally on frons with a prominent median hump between the fork branch- 
es; maxillary stridulatory area with 6-11 short, conical teeth (x = 8.5); gibbosity 



RESEARCH NOTES 69 

on dorsum of third abdominal segment with two irregular to subquadrate patches 
of 55-68 weakly curved setae; venter of last abdominal segment with two sub- 
circular patches of 55-65 caudally directed setae; teeth of epipharyngeal phoba 
relatively long, coarse and very closely set; anterior epitorma very weakly scler- 
otized and variable in appearance; macrosensillae not evident; mandibular cusps 
prominent and heavily sclerotized; molar area of right mandible with a conspic- 
uous sclerotized projection at apex. 

The apparent absence of macrosensillae in the larvae of O. medorensis is note- 
worthy in that Ritcher (1966) states that the presence of a pair of macrosensillae 
constitutes one of the diagnostic features of larvae of this genus, although it was 
not used as a character in the most recent key to the scarabaeine larvae (Edmonds 
and Halffter, 1978). However, since larvae are known from only a small fraction 
of the 1500+ described species of Onthophagus, it is likely that this and perhaps 
other apparently diagnostic features will be found to have exceptions as the larvae 
of other species are described. 

Acknowledgments 
I thank James R. Green for aid in the identification of the specimens. 

Literature Cited 

Brown, W. J. 1926. Notes on the Oklahoma species of Onthophagus. Proc. Okla. Acad. Sci., 5:99- 

101. 
. 1928. An annotated list of the coprophagous Scarabaeidae known to occur in Oklahoma. 

Proc. Okla. Acad. Sci., 7:24-28. 

. 1929. Studies in the Scarabaeidae, III. Canadian Entomol., 61:204-214. 

Edmonds, W. D., and G. Halffter. 1978. Taxonomic review of immature dung beetles of the subfamily 

Scarabaeinae (Coleoptera: Scarabaeidae). Syst. Entomol., 3:307-331. 
Howden, H. F., and O. L. Cartwright. 1963. Scarab beetles of the genus Onthophagus Latreille 

north of Mexico (Coleoptera: Scarabaeidae). Proc. U.S. Nat. Mus., 1 14(3467): 1-135. 
Ritcher, P. O. 1966. White grubs and their allies; a study of North American scarabaeoid larvae. 

Oregon St. Monog. Entomol., 4:1-219. 

Accepted for publication January 2, 1979. 

Vincent Brach, 325 W. First St., Tyler, Texas 75701. 



INDEX TO VOLUME 74 

Acari, Gammaridacarus brevisternalis, 5-9 

Acarina, Hannemania, 15-19; new species, 161-163 

Amphibia, New species, 160-161 

Amphipod, Orchestoidea cornicidata infestation by Gammaridacarus brevister- 
nalis, 5-9; Grandidierella japonica, 104-108 

Anelosimus eximius, Biology, 37-41 

Aphrodita sonorae, n. sp., 75-80 

Applegate, Shelton P.: A new species of Paleocene Chimaeroid from California, 
27-30 

Araneae, Anelosimus eximius, 37—41 

Axiothella rubrocincta with Harmothoe cf. lunulata, 42-43 

Bachia, 113-121 

Baker, Robert J., see Greenbaum, I. F. 

Barmeyer, Robert A.: Predation on the isopod crustacean Porcellio scaber by 

the Theridiid spider Steatoda grossa, 30-36 
Beach, Douglas W., see Leatherwood, S. 
Brach, Vincent: The Biology of the social spider Anelosimus eximius (Araneae: 

Theridiidae), 37-41 
Brach, Vincent: Development of the whipscorpion Schizomus floridanus, with 

notes on behavior and laboratory culture, 97-100 
Brach, Vincent: A case of active brood defense in the thornbug, Umbonia cras- 

sicornis (Homoptera: Membracidae), 163-164 
Brusca, Gary J., see Flores, M. 
Bryothinusa, Study and key, 109-112 

California, Chigger infestation of tree frog, 15-19; crabs, 100-104; 
Eschrichtius robustus, 45-46; Grandidierella japonica, 104-109; 
Hannemania, 15-19; Micrarionta opuntia, 94-96; Neotrombicula superci- 
liaris, n. sp., 161-163; New terrestrial snail, 94-96; 
Paleocene chimaeroid, 27-30; two new species of Polychaetes, 75-80 

Canidae, Calcanea, 143-155 

Carlson, David C: Taxonomic characters of the genus Ochodaeus Serville with 
descriptions of two new species in the O. pectoralis LeConte species com- 
plex (Coleoptera: Scarabaeidae), 49-65 

Ceratias holboelli, Parasite on, 1-5 

Chapman, John W. and Julie A. Dorman: Diagnosis, systematics, and notes Gran- 
didierella japonica (Amphipoda: Gammaridea) and its introduction to the 
Pacific Coast of the United States, 104-108 

Chile, New frog species, 160-161 

Chimaeroid, Paleocene from Calif., 27-30 

Coelenterata, Bryothinusa, 109-112; Hydrichthys pietschi, n. sp., 1-5; Ocho- 
daeus pectoralis, 49-65 

Coleoptera, two new species, 49-65; Keys to Bryothinusa, 109-112 

Collins, Charles T., see Johnson, W. D. 

Crabs, Records and range in California, 100-104 



INDEX TO VOLUME 74 71 

Ctenophore, Pleurobrachia bachei, 10-15 
Cuckoo owlet, Metabolism, 44-45 

Dorman, Julie A., see Chapman, J. W. 

Dyer, William G.: Parasitism as an indicator of food sources in a cave adapted 
salamander habitat, 72-75 

Egoscue, Harold J.: Population dynamics of the kit fox in western Utah, 122-127 
Eschrichtius robustus, Born outside calving lagoons, 45-46 

Finstad, William O. and Donald R. Nelson: Circadian activity rhythm in the Horn 
Shark, Heterodontus francisci: effect of light intensity, 20-26 

Fish, Chimaeroid, 27-30; Heterodontus francisci, 20-26; Parasite on Ceratias 
holboelli, 1-5; Salmo gairdneri, 80-86 

Flores, Marilyn and Gary J. Brusca: Observations on two species of hyperiid 
amphipods associated with the ctenophore Pleurobrachia bachei, 10-15 

Frogs, Tree: Infestation by chiggers, 15-19 

Frog, New species, 160-161 

Gammaridacarus brevisternalis infests amphipod, 5-9 

Gastropoda, Micrarionta opuntia, new species, 94-96; Monadenia churchi, 93- 
94 

Grandidierella japonica , 1 04- 1 08 

Gray whales born outside calving lagoons, 45-46 

Greenbaum, Ira F., Robert J. Baker, and Don E. Wilson: Evolutionary impli- 
cations of the Karyotypes of the stenodermine genera Ardops, Ariteus, Phyl- 
lops, and Ectophylla, 156-159 

Haig, Janet and Mary K. Wicksten: First records and range extensions of crabs 

in California waters, 100-104 
Hanne mania infestation of tree frogs, 15-19 
Harmothoe cf. lunulata in tube of Axiothella rubrocincta, 42-43 
Harty, Francis M. and Howard J. Stains: Least and long-tailed weasels (Mustela 

nivalis and M. frenata) collected in Grundy County, Illinois, 44 
Hawk, owl, Metabolism, 44-45 

Heterodontus francisci, Circadian activity rhythm, 20-26 
Homoptera, Defense in Umbonia crassicornis, 163-164 
Hulsea heterochroma and H. vestita ssp. callicarpha hybridization, 86-92 
Hydrichthys pietchi, n. sp., 1-5 
Hyperiid amphipods in symbiosis with Pleurobrachia bachei, 10-15 

Illinois, Mustela nivalis and M. frenata occurrence, 44 

Isopod, Porcellio scaber preyed upon by Steatoda grossa, 30-36 

Johnson, Wayne D. and Charles T. Collins: Notes on the metabolism of the 
Cuckoo Owlet and Hawk Owl, 44-45 

Karyotypes of Ardops, Ariteus, Phyllops, and Ectophylla, 156-159 



7 2 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Kit fox, Population study, 122-127 

Kudenov, Jerry D.: The occurrence of the polynoid Harmothoe cf. lunulata from 

the tube of the maldanid Axiothella rubrocincta (Polychaeta), 42-43 
Kudenov, Jerry D.: Two new species of errant polychaetes from the Gulf of 

California, Mexico, 75-80 

Lamellibranch, Morphological and physiological studies, 66-72 

Leatherwood, Stephen and Douglas W. Beach: A California gray whale calf (Es- 

chrichtius robust us) born outside the calving lagoons, 45-46 
Legner, E. F., see Moore, I. 
Loomis, Richard G., see Welbourn, W. C, Jr. 
Loomis, Richard B. and Hans Megens: A new species of Neotrombicula (Acar- 

ina: Trombiculidae) from southern California, 161-163 
Lynch, John D.: A new Chilean frog of the extra-andean assemblage of Telma- 

tobius (Amphibia: Leptodactylidae), 160-161 

Malmgrenia hartmanae, n. sp., 75-80 

Martin, W. E.: Hydrichthys pietschi, n. sp. (Coelenterata) parasitic on the fish, 

Ceratias holboelli, 1-5 
Megens, Hans, see Loomis, R. B. 
Mexico, Two new species of polychaetes, 75-80 
Micrarionta opuntia, n. sp., 94-96 
Monadenia churchi Hanna and Smith, affinities, 93-94 
Moore, Ian and E. F. Legner: A study of Bryothinusa (Coleoptera: Staphylini- 

dae), comparing a tabular and a dichotomous key to the species, 109-112 
Morton, Martin L.: Seasonal cycles of body weights and lipids in belding ground 

squirrels, 128-143 
Murphy, Judith A., see Paparo, A. A. F. 
Mustela nivalis and M.frenata found in Illinois, 44 
Mytilus edulis, Morphological and physiological studies, 66-72 

Nelson, Donald R., see Finstad, W. G. 

Neotrombicula superciliaris, n. sp., 161-163 

New taxa: Aphrodita sonorae, 75-80; Hyrichthys pietschi, 1-5; Ischyodus zins- 
meisteri, 27-30; Malmgrenia hartmanae, 75-80; Neotrombicula superciliar- 
is, 161-163; Ochodaeus howdeni, 49-65; O. ritcheri, 49-65; Telmatobius 
grandisonae, 160-161 

Ochodaeus howdeni, O. ritcheri, n. sp., 49-65 
Ochodaeus Serville, Taxonomic characters, 49-65 
Orchestoidea corniculata, Infestation on, 5-9 

Paleocene Chimaeroid, 27-30 

Paparo, Anthony A. F. and Judith A. Murphy: A study of the morphological and 

physiological changes in the lamellibranch, Mytilus edulis, after 6-OH-DOPA 

administration, 66-72 
Parasitism and a salamander, 72-75 



INDEX TO VOLUME 74 73 

Payne, Thomas R.: Study on the development of the prior residence effect in 

rainbow trout (Salmo gairdneri), 80-86 
Plants: Huslea heterochroma and H. vestita ssp. 86-92 
Pleurobrachia bachei in symbiosis with hyperiid amphipods, 10-15 
Polychaetes, two new species, 75-80; Axiothella rubrocincta, 42-43 
Porcellio scaber, 30-36 
Presch, William: The evolution of limb reduction in the teiid lizard genus Bachia, 

113-121 
Prior residence, effect on development of rainbow trout, 80-86 

Roth, Barry: On the affinities of Monadenia churchi Hanna and Smith (Gastro- 
poda: Stylommatophora), 93-94 

Roth, Barry: Description of a new terrestrial snail from San Nicolas Island, Cal- 
ifornia (Gastropoda: Stylommatophora), 94-96 

Salamander habitat: Parasitism indicates food sources, 72-75 

Salmo gairdneri, Prior residence effect on development, 80-86 

Schizomus floridanus, Development and behavior, 97-100 

Scurlock, Donna: Infestation on the sandy beach amphipod Orchestoidea cor- 

niculata by Gammaridacarus brevisternalis (Acari: Laelaptidae), 5-9 
Shark, horn, 20-26 

Squirrels, belding ground, Seasonal cycles of body weight, 128-143 
Spider, social; Biology of, 37-41 
Stains, Howard J., see Harty, F. M. 

Stains, Howard J.: Calcanea of members of the Canidae, 143-155 
Steatoda grossa preys on Porcellio scaber, 30-36 
Stenodermine, Evolution and karyotypes, 156-159 
Symbiosis with hyperiid amphipods and Pleurobrachia bachei, 10-15 
Systematics, Diagnosis and notes on Grandidiella japonic a, 104-108 

Teiid lizard, Evolution of limb reduction, 113-121 
Telmatobius grandisonae, n. sp. 160-161 
Theridiid spider, 30-36 
Thornbug, Brood defense, 163-164 

Umbonia crassicornis, Brood defense, 163-164 
Utah, Population study of kit fox, 122-127 

Welbourn, W. Calvin Jr. and Richard B. Loomis: Hannemania (Acarina: Trom- 
biculidae) and their anuran hosts at Fortynine Palms Oasis, Joshua Tree 
National Monument, California, 15-19 

Whipscorpion, Schizomus floridanus, Development and behavior, 97-100 

Wicksten, Mary K., see Haig, J. 

Wilken, Dieter H.: Natural hybridization between Hulsea heterochroma and H. 
vestita ssp. callicarpha, 86-92 

Wilson, Don E., see Greenbaum, I. F. 



INDEX TO VOLUME 75 

Acarina, Two new species, 278-284 

Acipenser, Hermaphroditism, 119-126 

Africa, Lanternfish, 138-152 

Agladphamus juvenalis, 16-19 

Ahlstrom, Elbert H., H. Geoffrey Moser, and Michael J. OToole: Development 
and distribution of larvae and early juveniles of the commercial lanternfish, 
Lampanyctodes hectoris (Gunther), off the west coast of southern Africa 
with a discussion of phylogenetic relationships of the genus, 138-152 

Alepocephalid fish, 153-158 

Allen, Larry G., see Horn, M. H. 

Amphistichus argenteus, cranial osteology, 29-38 

Anderson, M. Eric, see McCosker, J. E. 

Arenivaga apacha nest structure and micro-climate, 273-277 

Argis levior (Rathbun) from California, 56 

Arndt, Rudolph G., see Rohde, F. C. 

Atz, James W., and C. L. Smith: Hermaphroditism and gonadal teratoma-like 
growths in sturgeon {Acipenser), 119-126 

Baskin, Jonathan N., see Feldmeth, C. R. 

Bats, Seasonal changes in body weight, 258-266 

Beierle, John W., Chris Degnen, Joseph J. Beierle, Donald R. Patten, and Floyd 

E. Durham: An analysis of the fluid contents in the postanal sac of the gray 

whale, Eschrichtius robustus, 5-10 
Beierle, John W., see Durham, Floyd E., 1-5 
Beierle, Joseph J., see Beierle, J. W. 
Bomolochus cuneatus, Fraser, 22-28 
Brasil, Polychaetes, 16-22 
Brattstrom, Bayard H.: A Pleistocene herpetofauna from Smith Creek Cave, 

Nevada, 283-284 
Bryozoa, Spathipora mazatlanica, 38-42 

Calcanea of Mustelinae, 237-248 

California, Copepods, 22-28; Decapod, 56; Fish from archaeological site, 13 1 — 
1 37; Lamprey, 60-67; Mellivorinae, melinae, mephitinae, and lutrinae, 249- 
257; Parasite on mudcrab, 55-56; Pit sculpin, 111-118; Spawning Grunion, 
198-203; Survey of fishes in bays and estuaries, 159-170 

Chamberlain, Dilworth W., see Matsudo, H. 

Chiroptera, Seasonal changes in Myotis thysanodes and M. lucifugus, 258-266 

Ciliary activity control, 49-54 

Cockroach, Nest structure and micro-climate, 273-277 

Cohen, Allen C, and Jacqueline L. Cohen; Nest structure and micro-climate of 
the desert cockroach, Arenivaga apacha (Polyphagidae, Dictyoptera), 273- 
277 

Cohen, Jacqueline L., see Cohen, A. C. 

Coloration in fishes, 183-198 

Copepods, New species, 22-28 



INDEX TO VOLUME 75 75 

Coryphaenoides carapinus, Life history, 203-211 

Cottus pitensis, Feeding ecology, 111-118 

Crustacea, Portunion conformis Muscatine, parasite in mudcrab, 55-56 

Ctenostomata, Spathipora mazatlanica, n. sp., 38-42 

Cymatogaster, aggregata Gibbons, 22-28 

Cyprinodon nazas, C. alvarezi, C. meeki, C. macrolepis, four n. sp., 68-75 

Degnen, Chris, see Beierle, J. W. 

Delmarva Peninsula, U. S., Lampreys, 99-111 

Dioplosyllis broadi, n. sp., 19-22 

Dipodomys ordii, 225-237 

Durham, Floyd E., and John W. Beierle: Investigations on the postanal sac of 

the gray whale Eschrichtius robustus, 1-5 
Durham, Floyd E., see Beierle, J. W. 

Ecology of Pit sculpin, 111-118 

Elbert, Stephen A., see Resh, V. H. 

Embiotocid fishes, Orientation, 170-183 

Enhydra lutris, Systematics and status, 267-270 

Entosphenus hubbsi, n. sp., 60-67 

Ergasilus lizae Kr0yer, 22-28 

Eschrichtius robustus. Postanal sac, 1-5; fluid contents of sac, 5-10 

Ethological data in Goodeid fishes, 84-99 

Fauchald, Kristian: Some Nephtydae (Polychaeta) from Ubatuba, Brasil, 16-19 

Fauchald, Kristian, see Mueller, G. J. 

Feldmeth, C. Robert and Jonathan N. Baskin: Thermal and Respiratory studies 
with reference to temperature and oxygen tolerance for the unarmored stick- 
leback Gasterosteus aculeatus williamsoni Hubbs, 127-131 

Fish, Alepocephalid, 153-158; Archaeological site, 131-137; Amphistichus ar- 
genteus, 29-38; Dover Sole, 270-273; Embiotocid, 170-183; Goodeid, 84- 
98; Grunion, 198-203; Hypsoblennius Gill, 183-198; Lampreys, 99-111; Lan- 
ternfish, 138-153; Notropis atherinoides, 76-84; Rattail, 203-211; Sculpin, 
111-118, Stickleback, 127-131; Sturgeon, 119-126; Survey offish in Califor- 
nia bays and estuaries, 159-170 

Fitzsimons, John Michael: Ethological isolating mechanisms in goodeid fishes of 
the genus Xenotica (Cyprinodontiformes, Osteichthyes), 84-99 

Follett, W. I.: Fish remains from an archaeological site at Rancho Carrillo on the 
Silver Strand, San Diego County, California, 131-137 

Gasterosteus aculeatus williamsoni Hubbs, Thermal and respiratory studies, 127— 

131 
Gaudin, Anthony J., see Morris, Steven L. 
Goodeid fishes, Ethological isolating mechanisms, 84-99 
Gregg, W. O. and W. B. Miller: Two new species of Helminthoglypta (Mollusca: 

Pulmonata) from San Diego County, California, 10-15 
Gray whale, Fluid contents of postanal sac, 5-10 
Gray whale, Postanal sac, 1-5 



76 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Haedrich, Richard L. and Pamela T. Polloni: A contribution to the life history of 
a small Rattail Fish, Coryphaenoides carapinus, 203-211 

Hanan, Doyle A.: A new species of cyclopoid copepod, parasitic on shiner 
surfperch, Cymatogaster aggregata Gibbons, in Anaheim Bay and Hunting- 
ton Harbor, California, with notes on Bomolochus cuneatus Fraser and Er- 
gasilus lizae Kr0yer, 22-28 

Helminthoglypta edwardsi, n. sp., 10-15 

Helminthoglypta walton, n. sp., 10-15 

Hemigrapsus oregonensis, 55-56 

Hendricks, Fred S., see Schmidly, D. J. 

Holobomolochus embiotocae, n. sp., 22-28 

Horn, Michael H.: In honor of Carl L. Hubbs, 57-59 

Horn, Michael H. and Larry G. Allen: Numbers of species and faunal resem- 
blance of marine fishes in California bays and estuaries, 159-170 

Hubbs, Carl L., 57-59 

Hypsoblennius Gill, Coloration, 183-198 

Isaacs, John D.: Reproductive products in marine food webs, 220-233 
Isopod, 55-56 

Jennings, Daryl E., see Resh, V. H. 

Kangaroo rat, Systematics of Dipodomys ordii, 225-237 
Key to the Cyprinodon exitnius complex, 68-75 
Kott, Edward, see Vladykov, V. A. 
Krumholz, Louis A., see Resh, V. H. 

Lampanyctodes hectoris (Giinther) development and distribution, Southwest Af- 
rica, 138-152 

Lampetra lamottenii, 99-111 

Lamprey, n. sp., 59-67; life history, 99-111 

Land snails, 10-19 

Lanternfish, 138-152 

Lapota, David: Incidence of the entoniscid parasite, Portunion conformis Mus- 
catine (Crustacea: Isopoda) in the mudcrab Hemigrapsus oregonensis from 
San Diego County, California, 55-56 

Leuresthes sardina, Tides, waves and spawning, 198-203 

Li, Hiram W. and Peter B. Moyle: Feeding ecology of the pit sculpin, Cottus 
pitensis, Ash Creek, California, 111-118 

Lizards, secondary palate in microteiid, 281-283 

Loomis, Richard B.: Two new species of the genus Pseudoschoengastia (Aca- 
rina, Trombiculidae) from Mexico, 278-281 

Losey, George S., Jr.: The significance of coloration in fishes of the genus Hyp- 
soblennius Gill, 183-198 

Marine food webs, 220-233 

Matsudo, Hitoshi and Dilworth W. Chamberlain: Notes on mycoplasma-like or- 



INDEX TO VOLUME 75 77 

ganisms in skin tumors of Dover Sole, Microstomus pacificus, taken near 

sewage outfalls, 270-273 
McCosker, John E. and M. Eric Anderson: Aquarium maintenance of mesopelagic 

animals: a progress report, 211-219 
Mesopelagic animals, Aquarium maintenance, 211-219 
Mexico, Bryozoan, 38-42; Pseudoschoengastia bisetosa and P. smithi, 278-281; 

Pupfish, 68-75 
Microstomus pacificus, 270-273 
Miller, Robert Rush: Four new pupfishes of the genus Cyprinodon, from Mexico, 

with a key to the C. eximius complex, 68-75 
Miller, W. B., see Gregg, W. O. 
Mollusca, Two new species, 10-15 
Morris, Steven L. and Anthony J. Gaudin: The cranial osteology of Amphistichus 

argenteus (Pisces: Embiotocidae), 29-38 
Moser, H. Geoffrey, see Ahlstrom, E. H. 
Moyle, Peter B., see Li, H. W. 
Mueller, George J. and Kristian Fauchald: A new species of Dioplosyllis (Poly- 

chaeta: Syllidae) from California, 19-22 
Muench, Kevin A., see Thomson, D. A. 
Murphy, Judith A., see Paparo, A. A. 
Mussel, Mytilus edulis, 49-54 
Mustelidae, Calcanea, 237-248, 249-257 
My otis thysanodes, and M. lucifugus, Seasonal changes in body weight, 258-266 

Nephtys acrochaeta, N. magellanica, N . squamata, 16-19 

Nevada, pleistocene herpetofauna, 283-284 

New taxa: Cyprinodon alvarezi, C. macropelis, C. meeki, C. naza, 68-75; Di- 
plosyllis broadi, 19-22; Entosphenus hubbsi, 60-67; Helminthoglypta ed- 
wardsi, H. walton, 10-15; Holobomolochus embiotocae, 22-28; Pseudos- 
choengastia bisetosa, P. smithi, 278-281; Spathipora mazatlanica, 38-42 

Notropis atherinoides, Vertebral variation, 76-84 

OTarrell, Michael J. and Eugene H. Studier: Seasonal changes in wing loading, 
body composition, and organ weights in My otis thysanodes and M. lucifugus 
(Chiroptera: Vespertilionidae), 258-266 

Ohio River, fish, 76-84 

Okkelbergia aepyptera, Life history, 99-111 

O'Toole, Michael, see Ahlstrom, E. H. 

Otter, sea, Systematics and status, 267-270 

Paparo, Anthony A. and Judith A. Murphy: The coordinated role of the cerebral 

and visceral ganglia in ciliary beating, 49-54 
Patten, Donald R., see Beierle, J. W. 
Petromyzonidae, n. sp., 60-67 
Photostylus pycnopterus. New data, 153-158 
Pisces, Amphistichus argenteus, cranial osteology, 29-38; Freshwater lampreys, 

99-111; Goodeid, behavior, 84-99 



78 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Polloni, Pamela R., see Haedrich, R. L. 

Polychaeta, Dioplosyllis broadi, n. sp. 19-22; from Brasil, 16-19; Egg capsule 

formation, 285-286 
Poly dor a ligni, Egg capsule formation, 285-286 
Polyphagidae, Nest structure and micro-climate, 273-277 
Portunion conformis Muscatine, 55-56 
Postanal sac of gray whale, 1-5, 5-10 
Presch, William: Secondary palate formation in microteiid lizards (Teiidae: Lac- 

ertilia), 281-283 
Pseudoschoengastia bisetosa, P. smithi, n. sp. 278-281 
Pupfish, 68-75 

Reish, Donald J., see Rice, S. A. 

Resh, Vincent H., David S. White, Stephen A. Elbert, Daryl E. Jennings, and 

Louis A. Krumholz: Vertebral variation in the emerald shiner Notropis ath- 

erinoides from the Ohio River: an apparent contradiction to "Jordan's Rule,' 1 

76-84 
Rice, Stanley A. and Donald J. Reish: Egg capsule formation in the polychaete 

Polydora ligni: confirmation of an hypothesis, 285-286 
Roest, Aryan I.: Systematics and the status of sea otters, Enhydra lutris, 267- 

270 
Rohde, Fred C, Rudolf G. Arndt, and Johnson C. W. Wang: Life history of the 

freshwater lampreys, Okkelbergia aepyptera and Lampetra lamottenii 

(Pisces: Petromyzonidae), on the Delmarva Peninsula (East Coast, United 

States), 99-111 

Schmidly, David J. and Fred S. Hendricks: Systematics of the southern races of 

Ord's Kangaroo Rat, Dipodomys ordii, 225-237 
Sculpin, pit, Feeding ecology, 111-118 
Shrimp, 56 

Smith, C. L., see Atz, J. W. 
Snake, Tantilla semicincta, 42-48 
Soule, Dorothy F., see Soule, John D. 
Soule, John D. and Dorothy F. Soule: Spathipora mazatlanica, a new species of 

burrowing bryozoa (Ctenostomata) from Mazatlan, Sinaloa, Mexico, 38-42 
South American colubrid snake variation, 42-48 
Spathipora mazatlanica, 38-42 
Stains, Howard J.: Calcanea of members of the Mustelidae. Part I, Mustelinae, 

237-248 
Stains, Howard J.: Calcanea of members of the Mustelidae. Part II, Mellivorinae, 

Melinae, Mephitinae, and Lutrinae, 249-257 
Stephens, John S., Jr., see Terry, C. B. 
Stickleback, 127-131 
Studier, Eugene H., see O'Farrel, M. J. 
Sturgeon, Hermaphroditism, 119-126 
Systematics, Dipodomys ordii, 225-237; Enhydra lutris, 267-270 



INDEX TO VOLUME 75 79 

Tantilla semicincta (Dumeril, Bibron, and Dumeril) variation, 42-48 

Teiidae, Secondary palate formation, 281-283 

Terry, Catherine B. and John S. Stephens, Jr.: A study of the orientation of 

selected embiotocid fishes to depth and shifting seasonal vertical temperature 

gradients, 170-183 
Thomson, Donald A. and Kevin A. Muench: Influence of tides and waves on the 

spawning behavior of the Gulf of California grunion, Leuresthes sardina 

(Jenkins and Evermann), 198-203 
Tumors, skin, with mycoplasma-like organisms, 270-273 

United States, Eastern lamprey, 99-111 

Vladykov, Vadim D. and Edward Kott: A new nonparasitic species of lamprey of 
the genus Entosphenus Gill, 1862, (Petromyzonidae) from south Central Cal- 
ifornia, 60-67 

Wang, Johnson, C. S., see Rohde, F. C. 

White, David S., see Resh, V. C. 

Wicksten, Mary K.: First record of Argis levior (Rathbun) from California (De- 
capoda: Crangonidae), 56 

Wilson, Larry David: Variation in the South American Colubrid snake Tantilla 
semicincta (Dumeril, Bibron, and Dumeril), with comments on pattern di- 
morphism, 42-48 

Wisner, Robert L.: New data on the rare alepocephalid fish Photostylus pycnop- 
terus, 153-158 



INDEX TO VOLUME 76 

Acari, Ceuthothrombium cavaticum, 135-137 

Acarina, n. sp., 69-72 

Age, Effects on predation, 173-185 

Air, Surface flow in coastal California, 21-31 

Algae, Morphology of Sorella, 5-15 

Appy, Ralph and Murray D. Dailey: A new species of Rhinebothrium (Cestoda: 
Tetraphyllidea) and redescription of three rhinebothriate species from the 
round stingray, Urolophus halleri Cooper in southern California, 116-127 

Autolytus (Regulatus) penetrans, n. sp., 42-48 

Aves: Sphyrapicus varius, 38-41 

Behavior of Pacific angel shark, 193-201 

Behavior of Elasmobranchs, 202-204 

Bellemin, Jeanne M. and Glenn R. Stewart: Diagnostic characters and color con- 
vergence of the garter snakes Thamnophis elegans terrestris and Thamnophis 
couchii atratus along the central California Coast, 73-84 

Bender, Kristen E., see Collins, C. T. 

Bennett, Stephen G., see Loomis, R. B. 

Brach, Vincent: Larvae of Onthophagus p. polyphemi Hubbard and Onthopha- 
gus tuberculifrons Harold (Coleoptera: Scarabaeidae) 66-68 

Brattstrom, Bayard H., see Hudson, D. M. 

Browning, M. Ralph; Interbreeding members of the Sphyrapicus varius group 
(Aves: Picidae) in Oregon 38-41 

Brusca, Richard C: Range extensions and new host records of cymothoid isopods 
(Isopoda: Cymothoidae) in the eastern Pacific Ocean, 128-131 

California, Amastigus acutus, n. sp. and gen., 57-60; Caprellidae survey, 146- 
167; characters and colors of garter snakes, 73-84; Pleistocene herpetofauna 
from Newport Beach, 16-20; Rhynchospio microcera, n. sp., 1-4; surface 
air flow, 21-31; Syllidae, n. sp., 42-48 

Cave crickets, Parasite of, 135-137 

Cerithidea calif ornica (Haldeman), Salinity and paleoecology, 60-63 

Cestoda, Five new morphotypes of Phillobothrium delphini, 99-110; redescrip- 
tion of three rhinebothriate species, 116-127; Rhinebothrium ditesticulum, 
n. sp., 116-127 

Ceuthothrombium cavaticum, biology, 135-137 

Coleoptera, Larvae of two scarab beetles, 66-68 

Collins, Charles T. and Kristen E. Bender: The natal pterylosis of the house 
finch, 209-211 

Crustacea, Family Caprellidae, 146-167 

Cunningham-Paparo, Kathleen, see Paparo, A. A. 

Cymatoceras pecosense, n. sp., 133-135 

Dailey, Murray D., see Testa, J. 
Dailey, Murray D., see Appy, R. 
DeLong, Nikki, see Guthrie, D. A. 



INDEX TO VOLUME 76 81 

DOPA decarboxylase inhibitors and chemical sympathectomy, 32-37 
Dorsey, John H.: A new species of Rhynchospio (Polychaeta: Spionidae) from 
San Clemente Island, California 1-4 

Eutrombicula bitarsala, n. sp., 69-72 

Fishes, Behavior of elasmobranchs, 202-204 
Funk, Richard S., see Tucker, J. K. 

Geology, Cedros island, 91-98 
Geomyidae, Carotid arteries in three genera, 63-66 

Guthrie, Daniel A. and Nikki DeLong: Carotid arteries in the rodent genera 
Pappogeomys, Geomys, and Thomomys (family Geomyidae), 63-66 

Hannes, Gerald: Surface air flow in coastal southern California, 21-31 

Herpetofauna from Newport Pleistocene deposits, 16-20 

Hipponoe gaudichaudi, Brooding behavior and protandry, 85-90 

House finch, Pterylosis, 209-211 

Hudson, Dennis M. and Bayard H. Brattstrom: A small herpetofauna from the 

Late Pleistocene of Newport Beach Mesa, Orange County, California, 16- 

20 
Hydrocoral, with polychaete, 42-48 

Isopoda, Range and hosts of cymothoid, 128-131 

Kilmer, Frank H.: Reconnaissance geology of Cedros Island, Baja California, 

Mexico, 91-98 
Knudtson, Bruce K. and Joseph R. Stimers: Notes on the behavior of elasmo- 

branch fishes exposed to magnetic fields, 202-204 
Kudenov, Jerry D.: Brooding behavior and protandry in Hipponoe gaudichaudi 

(Polychaeta: Amphinomidae), 85-90 

Lepidoptera, Hormone activity, 186-192 

Loomis, Richard B. and Stephen G. Bennett: A new species of Eutrombicula 

(Acarina: Trombiculidae) from lizards of Nevado de Colima, Jalisco, Mexico, 

69-72 

Martin, Donald M.: A survey of the family Caprellidae (Crustacea, Amphipoda) 
from selected sites along the northern California coast, 146-167 

McCranie, James R., see Wilson, L. D. 

Metridium senile, 168-172 

Mexico, Cedros island geology, 91-98; Eutrombicula bitarsala, n. sp., 69-72 

Mullen, David A., the striped dolphin, Stenella coeruleoalba, in the Gulf of 
California, 131-132 

Murphy, Judith, see Paparo, A. A. 

Mussel, Mytilus edulis, Ciliary activity, 111-115 

Mytilus edulis, Ciliary activity, 111-115 

Nautilida, Cymatoceros pecosense, n. sp., 133-135 



82 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Nelson, Donald R., see Standard, E. A. 

New taxa: Amastigos acutus, n. gen., and n. sp., 57-60; Autolytus (Regulatus) 
penetrans, 42-48; Cymatogeras pecosense, 133-135; Eutrombicula bitar- 
sala, 69-72; Rhinebothrium ditesticulum, 116-127; Rynchospio microlera, 
1-4 

Onthophagus p. polyphemi Hubbard and O. tubercnlifrons Harold, larvae, 66- 

68 
Oregon, Interbreeding of Sphyrapicus varius, 38-41 

Pacific Ocean, Isopod range and hosts, 128-131 

Paleoecology, Salinity and Cerithidea calif ornica (Haldeman), 60-63 

Paparo, Anthony A., Kathleen Cunningham-Paparo, and Judith Murphy: The 
effect of endogenous 5-HT on K ion enhancement of ciliary activity in the 
mussel Mytilus edulis, 111-115 

Paparo, Anthony A., Kathleen Cunningham-Paparo, and Judith Murphy: DOPA 
decarboxylase activities and potassium stimulation of lateral cilia on the gill 
of Mytilus edulis I. A response to DOPA decarboxylase inhibitors and chem- 
ical sympathectomy, 32-37 

Paparo, Anthony A., Richard E. Sparks, Judith A. Murphy, and Kathleen J. 
Cunningham-Paparo: The effect of potassium ions on the rate of ciliary ac- 
tivity in Sphaerium transversum I. A different response in small and large 
clam preparations, 139-145 

Patton, Donald R. and William F. Samaras: Unseasonable occurrences of gray 
whales, 205-209 

Paramyelois transit ella. Effects of hormone, 186-192 

Piltz, Fred: A new genus and species of polychaete (Family Capitellidae) from 
southern California, 57-60 

Phyllobothrium delphini, Five new morphotypes, 99-110 

Pleistocene, Herpetofauna from Newport, 16-20 

Polychaeta, Amastigos acutus, n. gen., n. sp., 57-60; Autolytus (regulatus) pen- 
etrans, n. sp., 43-48; Hipponoe gaudichaudi, 85-90; Rhynchospio micro- 
cera, 1-4 

Ponasterone A. effect on Paramyelois tranistella (Walker), 186-192 

Porras, Louis, see Wilson, L. D. 

Potassium ions and ciliary activity, 139-185 

Pterylosis of the house finch, 209-211 

Purcell, Jennifer F.: The diet of large and small individuals of the sea anemone 
Metridium senile, 168-172 

Rhinebothrium ditesticulum, n. sp., 116-127 
Rhynchospio microcera, n. sp., 1-4 

Samaras, William F., see Patton, D. R. 

Scott, David B. and Timothy L. Cass: Response of Cerithidea californica (Halde- 
man) to lowered salinities and its paleoecological implications, 60-63 
Sea anemone, Diet of Metridium senile, 168-172 
Serpentes, From tropical America, 49-56 
Snakes, garter, Characters and colors, 73-84 



INDEX TO VOLUME 76 83 

Sorella, Morphology, 5-15 

Sparks, Richard E., see Paparo, A. A. 

Sphaerium transversum, Ciliary activity, 139-145 

Sphyrapicus varius, Interbreeding, 38-41 

Squatina calif arnica, Behavior, 193-201 

Standora, Edward A. and Donald R. Nelson: A telemetric study of the behavior 

of free-swimming Pacific angel sharks, Squatina californica, 193-201 
Stenella coeruleoalba in the Gulf of California, 131-132 
Stewart, Glenn R., see Bellemin, J. M. 
Stewart, Joan A.: Morphology of Sorella in natural habitats and under laboratory 

conditions, 5-15 
Stimers, Joseph R., see Knudtson, B. K. 

Tantilla, Distribution and taxnonmic notes, 49-56 

Testa, Jacqueline and Murray D. Dailey: Five new morphotypes of Phylloboth- 
rium delphini (Cestoda: Tetraphyllidea), their relationship to existing mor- 
photypes, and their zoogeography, 99-110 

Texas, New species of Cymatoceros, 133-135 

Thamnophis elegans terrestris and T. couchii atratus, diagnostic characters and 
colors, 73-84 

Tropical America, Distribution and taxonomic notes on Tantilla, 49-56 

Tucker, John K. and Richard S. Funk: A new species of Cymatoceras (Nautilida: 
Cymatoceratidae) from west Texas, 133-135 

Urolophus hailed, Infected with species of Cestoda, 116-127 

Webb, James P., Jr., Pierre Robaux, and Glenn D. Campbell: Notes on the 
biology of Ceuthothrombium cavaticum (Acari: Trombidiidae), a parasite of 
cave crickets (Rhaphidophoridae: Ceuthophilus), 135-137 

Wells, Harrington: Selective pressure on predator and prey when an age class 
becomes a refuge from predation, 173-185 

Whales, gray, Unseasonable occurrences, 205-208 

Wilson, Larry David, James R. McCranie, and Louis Porras: Taxonomic notes 
on Tantilla (Serpentes: Colubridae) from tropical America, 49-56 

Woodwick Keith H., see Wright, J. D. 

Wright, John D. and Keith H. Woodwick: A new species of Autolytus (Poly- 
chaeta: Syllidae) commensal on a California hydrocoral, 42-48 

Yokoyama, Victoria Y.: Extraction of a phytoecdysone, ponasterone A, from 
Podocarpns gracilior Pilger, and its effect on Paramyelois transitella (Walk- 
er) (Lepidoptera: Pyralidae), 186-192 



INDEX TO VOLUME 77 

Alga, Southern California intertidal algae transect, 1-13; two brown algae new to 

California, 28-35 
Amblyosyllis speciosa Izuka, Notes on, 22-27 
Armstrong, John W. and Peter A. Jumars: Branchiate Dorvilleidae (Polychaeta) 

from the north Pacific, 133-138 

Benefits from Los Angeles City Government, 71-81 

Brach, Vincent: Notes on the biology of Lithurgus gibbosus Smith in Florida 

(Hymenoptera: Megachilidae) , 144-147 
Branchiate Dorvilleidae from the north Pacific, 133-138 
Bufo boreas and B. canorus, Sympatry and hybridization, 52-55 

California: Amblyosyllis speciosa Izuka, 22-27 '; Benefits from Los Angeles City 
Government, 71-81; Dodecaseta or aria, n. gen. and sp., 40-43; First report 
of Heteropodarke heteromorpha Hartmann-Schroder 1962, 82-84; First rec- 
ord of Plesionika, 84-87; Intertidal algal transect, 1-13; Mohave Desert 
annuals, 95-108; Sea otter survival, 109-115; Syllides reishi, n. sp., 22-27; 
two brown algae new to state, 28-35 

Calloplax Thiele in the Americas, 56-64 

Chagas' trypanosome carried by western conenose, 130-132 

Chemsack, John A. and E. G. Linsley: Additions to the Cerambycidae of the 
Revilla Gigedo Islands (Coleoptera), 138-142 

Cerambycidae, Additions from Revilla Gigedo Islands, 138-142 

Chiton, n. sp., in Galapagos, 36-39 

Chlorogalum pomeridianum (D.C.) Kunth, Pollination ecology, 124-129 

Coleoptera, Cerambycidae of Revilla Gigedo Islands, 138-142 

Collins, Charles R.: The natal pterylosis of hummingbirds, 14-21 

Collisella edmitchelli (Lipps), Taxonomic affinities, 65-70 

Cutleria cylindrica Okamura, New to California, 28-35 

Dawson, E. Yale, Resurvey of Southern California algal transects, 1-13 

Dodecaseta oraria, n. gen., and n. sp., 40-43 

Dorsey, John H.: A new species of Syllides (Polychaeta: Syllidae) with notes on 

Amblyosyllis speciosa Izuka from San Clemente Island, California, 22-27 
Dorsey, John H.: A first report of Heteropodarke heteromorpha Hartmann 

Schroder, 1962 (Polychaeta: Hesionidae) from California, 82-84 
Dorvilleidae, Branchiate forms, 133-138 

Ferreira, Antonio J.: A new species of chiton (Neoloricata: Ischnochitonidae 

from the Galapagos Islands, 36-39 
Ferreira, Antonio J.: The genus Calloplax Thiele, 1909 (Mollusca: Polyplaco 

phora) in the Americas, 56-64 
Florida, Biology of Lithurgus gibbosus Smith, 144-147 
Friedsen, Richard D., see Matson, J. O. 

Galapagos, Chiton, n. sp., 36-39 

Gastropoda, Collisella edmitchelli (Lipps), Taxonomic affinities, 65-70 



INDEX TO VOLUME 77 85 

Harrington-Coulombe, Linda J., see Leatherwood, S. 

Heacox, Albert E. and Paul C. Schroeder: First report of brooding in Sy Hides 

japonica Imajima (Syllidae: Polychaeta), 142-144 
Hemiptera, Western conenose, carrier of Chagas' trypanosome, 130-132 
Heteropodarke heteromorpha Hartmann-Schroder, First report in California, 82- 

84 
Hollenberg, George J.: Physiological Notes VIII. Two brown algae (Phaeophyta) 

new to California, 28-35 
Hubbs, Carl L., see Leatherwood, S. 
Hui, Clifford A. and Sam Ridgway: Survivorship patterns in captive killer whales 

(Orcinus orca), 45-51 
Hummingbirds, Natal pterylosis, 14-21 
Hybridization in Bufo boreas and B. canorus, 52-55 
Hymenoptera, Biology of Lithurgus gibbosus Smith, 144-147 

Irrigation, Effects on desert annuals, 95-108 
Ischnochiton macleani, n. sp., 36-39 

Johnson, Hyrum B., Frank C. Vasek, and Terry Yonkers: Residual effects of 

summer irrigation on Mojave Desert annuals, 95-108 
Jumars, Peter A., see Armstrong, J. W. 

Killer whales, Survivorship in captivity, 45-51 

Leatherwood, Stephen, Linda J. Harrington-Coulombe and Carl L. Hubbs: Relict 
survival of the sea otter in central California and evidence of its recent re- 
dispersal south of Point Conception, 109-115 

Liliaceae, Pollination ecology of Chlorogalum pomeridianum (D.C.) Kunth, 124— 
129 

Lindberg, David R.: On the taxonomic affinities of Collisella edmitchelli (Lipps) 
(Gastropoda: Acmaeidae) a late Pleistocene limpet from San Nicolas Island, 
California, 65-70 

Linsley, E. G., see Chemsak, J. A. 

Lithurgus gibbosus Smith, Biology, 144-147 

Los Angeles City government benefits, 71-81 

Matantia, From California and Western Mexico, 84-87 

Matson, John O. and Richard D. Friesen: The subspecific status of Onychomys 

torridus clams Hollister 1913 (Rodentia: Cricetidae), 116-123 
McCammon, James A. and Janet K. Stull: A new genus and species of Capitel- 

lidae (Polychaeta) from California, 40-43 
Mollusca, Calloplax Thiele in the Americas, 56-64 
Morton, Martin L. and Kenneth N. Sokolski: Sympatry in Bufo boreas and B. 

canorus and additional evidence of natural hybridization, 52-55 
Myriactula rivulariae (Suhr in Areschoug) J. Feldmann, New in California, 28- 

35 

Natantia, Plesionika from California and western Mexico, 84-87 



86 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Neoloricata, Ischnochiton macleani, n. sp., 36-39 

New taxa: Ischnochiton macleani, 36-39; Syllides reishi, 22-27; Dodecaseta 
or aria, n. gen. and n. sp., 40-43 

Onychomys torridus clams Hollister 1913, 116-123 
Oral History, Comparative study, 89-94 
Orcinus ore a, Survivorship, 45-51 

Pacific, north, Branchiate Dorvilleidae, 133-138 

Teter left because. . .' oral history, 89-94 

Phaeophyta, two brown algae new to California, 28-35 

Plesionika, First record in California and western Mexico, Plesionika beebei and 

P. mexicana, 84-87 
Pollination ecology of Chlorogalum pomeridianum (D.C.) Kunth, 124-129 
Polychaeta, Amblyosyllis speciosa lzuka, 22-27; Dodecaseta oraria, n. gen. and 

n. sp., 40-43; Heteropodarke heteromorpha Hartmann-Schroder 1962, 82- 

84; Syllides reishi, n. sp., 22-27; Branchiate Dorvilleidae, 133-138; Syllides 

japonica Imajima, 142-144 
Pterylosis of hummingbirds, 14-21 

Revilla Gigedo Island, Additional Cerambycidae, 138-142 

Ridgway, Sam H., see Hui, C. A. 

Rodentia, Status of Onychomys torridus clarus Hollister, 1913, 116-123 

Schroeder, Paul C, see Heacox, A. E. 
Sea Otter, Survival and redispersal, 109-115 
Sokolski, Kenneth N., see Morton, M. L. 

Stockhouse, Robert E., II and Harrington Wells: Pollination ecology of Chloro- 
galum pomeridianum (D.C.) Kunth, (Liliaceae), 124-129 
Stull, Janet K., see McCammon, J. A. 
Syllides japonica Imajima, Brooding, 142-144 
Sympatry in Bufo boreas and B. canorus, 52-55 

Taxonomic affinities of Collisella edmitchelli (Lipps), 65-70 

Thorn, Ronald M. and Thomas B. Widdowson: A resurvey of E. Yale Dawson's 
42 intertidal algal transects on the southern California mainland after 15 
years, 1-13 

Thomas, Henry B.: Some estimates of the distribution of benefits from Los An- 
geles city government activity, 71-81 

Thym, Nancy: 'Peter left because . . .' A comparative study of family oral his- 
tories, 89-94 

Triatoma p. protracta. Carrier of Chagas' trypanosome, 130-132 

Vasek, Frank C, see Johnson, H. B. 

Wells, Harrington, see Stockhouse, R. E., II 
Western conenose and Chagas 1 trypanosome, 130-132 
Whales, killer, 45-51 



INDEX TO VOLUME 77 87 

Wicksten, Mary K.: The species of Plesionika from California and Western Mex- 
ico (Natantia: Pandalidae), 84-87 

Widdowson, Thomas B., see Thorn, R. M. 

Wood, Fae, D., see Wood, S. F. 

Wood, Sherwin F. and Fae D. Wood: Additional notes on the Western Conenose, 
Triatoma p. protracta (Insecta: Hemiptera: Reduviidae), as a carrier of Cha- 
gas' Trypanosome, 130-132 

Yonker, Terry, see Johnson, H. B. 



INSTRUCTIONS FOR AUTHORS 

e BULLETIN is published three times each year (April, August, and November) and includes articles in English 
any field of science with an emphasis on the southern California area. Manuscripts submitted for publication 
juld contain results of original research, embrace sound principles of scientific investigation, and present data 
a clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide 
contributors. Consult also recent issues of the BULLETIN. Authors should strive for directness and lucidity, 
lieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous 
erence of pronouns, and logically placed modifiers. 

MANUSCRIPT PREPARATION 

t is strongly recommended that, before submitting a paper, the author ask qualified persons to review it. The 
hor is requested to submit at least two additional copies with the original, on 8V2 x 11 opaque, nonerasable 
:>er, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. 
otnotes should be avoided. Manuscripts which do not conform to the style of the BULLETIN will be returned to 
author. 

Vn abstract summarizing in concise terms the methods, findings, and implications discussed in the paper must 
•ompany a feature article. Abstract should not exceed 100 words. 

V feature article comprises approximately five to thirty typewritten pages. Papers should usually be divided into 
following sections: abstract, introduction, methods, results, discussion and conclusions, acknowledgments, and 
rature cited. Avoid using more than two levels of subheadings. 

\ research note is usually one to six typewritten pages and rarely utilizes subheadings. Consult a recent issue 
the BULLETIN for the format of notes. Abstracts are not used for notes. 

Abbreviations: Use of abbreviations and symbols can be determined by inspection of a recent issue of the 
ULETIN. Omit periods after standard abbreviations: 1.2 mm, 2 km, 30 cm, but Figs. 1-2. Use numerals before 
ts of measurements: 5 ml, but nine spines (10 or numbers above, such as 13 spines). The metric system of 
lights and measurements should be used wherever possible. 

i axonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International 
de of Botanical Nomenclature (Lawjouw et al.. 1956), the International Code of Nomenclature of Bacteria and 
|uses (Buchanan et al.. 1958), and the International Code of Zoological Nomenclature (Stoll. et al.. 1961). 
;;cial attention should be given to the description of new taxa, designation of holotype, etc. Reference to new 
la in titles and abstract should be avoided. 

'he literature cited: Entries for books and articles should take these forms. 
HcWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. 

Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm.. 54: 452-458. • 
irattstrom, B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), 
'Jniv. California Press, xii + 635 pp. 

Tables and figures (line drawings, graphs, or black and white photographs) should not repeat data contained in 
I text. The author must provide numbers and short legends for tables and figures and place reference to each 
• hem in the text. Legends should be typed on a separate sheet of paper and placed at the end of the manuscript. 
I strations and lettering thereon should be of sufficient size and clarity to permit reduction to standard page size; 
» inarily they should be no more than twice the size of intended reduction and should not exceed 8V2 by 1 1 inches 
r ize. Photographs must be printed on glossy paper. Submit one photoduplicated copy of each illustration. All 
I Orations accompanying Research Notes will be reduced to one column width. All half-tone illustrations will have 
iit screen (grey) backgrounds. Special handling such as dropout half-tones, special screens, etc., must be requested 
tnd will be charged to authors. 

cover illustration pertaining to an article in the issue or one of general scientific interest will be printed on the 
er of each issue. Such illustrations along with a brief caption should be sent to the Editor for review. 

PROCEDURE 

11 manuscripts should be submitted to the Editor, Jeffrey I. Chapman, School of Public Administration, Uni- 
»ity of Southern California, Los Angeles, California 90007. Evaluation of a paper submitted to the BULLETIN 

1 ins with a critical reading by the Editor; several referees also check the paper for scientific content, originality, 
clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent 
he author at which time he or she may be requested to rework portions of the paper considering these rec- 

i|nendations. The paper then is resubmitted and may be re-evaluated before final acceptance. 

roof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He or she 
uld promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations, legends, 
bibliographical references. He or she marks corrections on the galley (copy editing and proof procedures in 
c Manual) and promptly returns both galley and manuscript to the Editor. Manuscripts and original illustrations 
not be returned unless requested at this time. All changes in galley proof attributable to the author (misspellings, 
nsistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with 
printer, not the Editor. 



CONTENTS 

Systematics and Ecology of Benthic Phyllodocidae (Annelida: Polychaeta) 
off the Columbia River, U.S.A. By Michael J. Kravitz and Howard 
R. Jones 1 

Recognition of Quaternary Wave Formed Marine Terraces on Santa 

Catalina Island. By William F. Samaras and Joseph A. Gellura 20 

Species Diversity and Distributional Relationships of Pseudoscorpions 
from Slash Pine (Pinus elliottii Eng.) in Florida (Arachnida: Pseudo- 
scorpionida). By Vincent Brach 32 

Summer Mean Diurnal Water and Air Temperature Distributions in the 

Cape Mendocino area, California. By Gerald Hannes 40 

Niche Separation Within a Population of Freshwater Fishes in an Irriga- 
tion Drain Near the Salton Sea, California. By Allan A. Schoen- 
herr 46 

Research Notes 

Two Asiatic Gobiid Fishes, Tridentiger trigonocephalus and Acanthogobius flavimanus, in 

Southern California. By Peter L. Haaker 56 

The Velvet Whalefish, Barbourisia rufa, added to California's Marine Fauna, with 

Notes on Otoliths of Whalefishes and Possible Related Genera. By John E. 

Fitch 61 

The Larva of Onthophagus medorensis Brown with Notes on its Biology (Coleoptera: 

Scarabaeidae) . By Vincent Brach 67 

Index: 1975 through 1978 70 



COVER: Anaitides multipapillata, a new species of Polychaeta off the Columbia River, U.S.A. 



■£ 



IOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




ULLETIN 



r>lume 78 



Number 2 




E:AS-A78(2) 89-150 (1979) 



LIBRARY 

DEC 2 8 1979 

Nc/\l YOKK 



AUGUST 1979 



-~T 



Southern California Academy of Sciences 

Founded 6 November 1891, incorporated 17 May 1907 



OFFICERS 

Takashi Hoshizaki, President 

Fred G. Hochberg, Vice President 

Richard E. Pieper, Secretary 

Joseph E. Haring, Treasurer 

Jeffrey Chapman, Editor 

Gretchen Sibley, Assistant Editor 



1977-1979 

George Callison 
Robert A. Georges 
Takashi Hoshizaki 

Alan J. Mearns 
W. Walton Wright 



BOARD OF DIRECTORS 

1978-1980 

Kristin H. Berry 
Joseph E. Haring 
Mark M. Littler 
James G. Morin 
Donald J. Reish 



1979-1981 

John Baird 

Jules Crane 

Fred G. Hochberg 

Richard E. Pieper 

Gloria Takahashi 



Membership is open to scholars in the fields of natural and social sciences, and to any persi 
interested in the advancement of science. Dues for membership, changes of address, and requests 
missing numbers lost in shipment should be addressed to: Southern California Academy of Scienc 
the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 9000 



ersi 
ncej 



Annual Members $ 12. ' 

Life Members 150. 

Fellows: Elected by the Board of Directors for meritorious services. 



The Bulletin is published three times each year by the Academy. Manuscripts for publication shoi 
be sent to the appropriate editor as explained in "Instructions for Authors" on the inside back cov 
of each number. All other communications should be addressed to the Southern California Acader 
of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, L 
Angeles, California 90007. 

Date of this issue 5 December 1979 



Bull. Southern California Acad. Sci. 
78(2), 1979, pp. 89-106 

Osprey Distribution, Abundance, and Status in Western North 

America: III. The Baja California and 

Gulf of California Population 

Charles J. Henny 1 and Daniel W. Anderson 2 

Abstract. — An estimated 810 ± 55 pairs (minimum estimate) of ospreys {Pan- 
dion haliaetus) were nesting in the study area during our survey (24 March-1 
April 1977). Approximately 174 pairs nested along the Pacific side of Baja Cali- 
fornia, 255 pairs along the gulf side, 187 pairs on the Midriff Islands, and 194 
pairs in coastal Sonora and Sinaloa. Most nested on cliffs adjacent to the sea 
(59%); some nested on cactus in flat terrain (26%). Seven per cent nested on the 
ground, three percent nested in mangroves and other trees in the southern portion 
of the study area, and four percent nested on man-made structures. The extreme 
northwestern Baja California population that was extirpated early in this century 
has not recovered. However, several populations immediately to the south along 
the Pacific Coast now appear stationary. Pesticide residues in osprey eggs from 
Mexico were among the lowest reported for the species in North America. 

1 Patuxent Wildlife Research Center, U.S. Fish and Wildlife Service, 480 S. 
W. Airport Road, Corvallis, Oregon 97330. 2 Division Wildlife and Fisheries Bi- 
ology, University of California, Davis, California 95616. 



Fifty years ago the osprey was reported as a common coastal resident along 
both the Pacific and Gulf sides of Baja California and practically all the adjacent 
islands (Grinnell 1928). More recently, Friedmann et al. (1950:62) described the 
species as a "common resident from Baja California to Sinaloa and the Tres 
Marias Islands, and probably elsewhere on the Pacific Coast/' Kenyon (1947) 
surveyed the northwest coast of Baja California in 1946 and concluded that the 
osprey population had declined markedly in the last 30 to 40 years except possibly 
at Scammon's Lagoon. Jehl (1977) updated Kenyon's work on several north- 
western sites (through 1972), and concluded that the population had stabilized in 
the last 25 years. The historical distribution and abundance of ospreys in the 
remainder of Baja California and the Gulf of California is poorly understood. 

The present study was designed to determine the 1977 distribution and abun- 
dance of ospreys in coastal Baja California, its adjacent islands and the coast of 
mainland Mexico as far south as Mazatlan (Fig. 1). Studies using the osprey as 
an indicator species of pollutant contamination have been conducted on nearly 
every nesting population in the United States (see review, Henny, 1977). These 
osprey studies played an important role in documenting problems associated with 
DDT use in the northeast (Ames and Mersereau, 1964; Ames, 1966) and else- 
where, and in documenting the recovery in productivity that followed the reduced 
use and the eventual ban of the product in the United States. We believe that the 
osprey can play a similar indicator species role in coastal Mexico. Future prob- 
lems in the surveyed region may or may not be pollutant oriented, but an increase 
in human activity in the region is almost certain (Anderson et al., 1976). We 



90 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 





1,6° 114° 112° H0° 108° 






TIJUANA C\ _ 1 _ _ _ 

% c.r.j.y/ -^ UNITED S T'A T E S 
















32° 


Ni 7 T M ' "-*%. 

v I ^tei^ ^" 1 

v '^**V " ^~ - 














- 


32° 


















~'\ 






y ^***fc.- "~~~ ' 




















\, I \ /' ■ ' 










1 % 1 1 
•fc. ■:% I \ 






30° 


_[ \ ' I \ \ 

ll~ % #—[————_( ' 

PUERTO SAN TA°<\ ^< p tn Bom*/t nc V I "" 










CATARINA X- ^k \ % ' 






30° 




^i- FK\ ^.Guardian \ S N R A J 










V -%\Q''^ &-' „ 1 










\ VNl il ? 5 r 










\ Uc r ib £m : 










\.\ Sil </\""' \ 










San %' . , ' \ ^ • «?: ' 

Jj SANTO KJ J Lorenzo V, ( 
__) Si DOMINGOjS 4 «t 












28° 














IHatividod* t ^?f —4 l_ \v . '. 








L p ?^™<^7c^r\^ ^— ~^^5E — f- \ 1 








Punta Eugenia *y *»T' Logoon \ GUAruAS p> -. 
<&-'. SANTA ROSAL/a\ 1 * L 




28° 




>0 ^^vj- Sanlgnacio \jJSan Marcos "M^ \ 








"t 7 l\ L °°?Sv * |.S<w mez ^ ii< Ti; V 








1 o . ' V^H v Kk 1 yvJ 








/ Punta Abreojos fc o> $,TN Ildefonso I Vj^ 1 








-^ X. V ■*>%. 1 ^Ni / 






26° 


J £}^ Punta Santo DomingciNAi; 1 2-' ' 


( 






1 V \ Coronodo ffi i 








^t— U-Q&£J^Q__gCorrneri M \__ 

tip 1 / Donionte J tr — : "V 


) 






1 V X^onserro" LrOPOLOBAMPO \ \ 




26° 










a / \ *Coraimo ^s, *v ' 








m ' \TAUBIBICHE \ ^3* lP 

° #1^1 '"~Smto Cruz ^%i ■ I / ; 














24° 


MugaolenoBo,-^ >^j ; yEspi'ntu Punta BaraditoTL O "\ 

L____^ Santa Morganta* \. V \ » ^jy\ V 








— ^_l_____ ^-v ^-ifJN, ^Cw-'o'"* 


>*. ~ 


.•"v 






50 50 100 150 XL 1 






24° 




KILOMETERS \ a 


MAZATL AN 


0<< 4 






•°CABO SAN LUCAS 

1 i I 


>5 




116° 114° 112° 110° 108° 


IC 


6° 



Fig. 1. The Baja California and Gulf of California study area for ospreys. 



collected data in 1977 to form a basis for future comparisons. In addition to 
collecting the baseline data, it is our purpose to describe areas that we believe 
are critical habitat for ospreys. 

Methods 

We located occupied osprey nests from a fixed-wing aircraft (Cessna 210) flying 
60-100 m above the shorelines and inland up to 2 km. A pilot and two observers 
were present in the plane, and 80 h of flying time were logged. In areas where 
large cactus or mangrove forests existed adjacent to the shoreline, we flew tran- 
sects at increasing distances landward from the shore looking for osprey nests 



OSPREY POPULATION BAJA CALIFORNIA AND GULF 



91 



Table 1. Number of occupied osprey nests (nesting pairs) seen from the air and the ground. 



Location 



Air 



Ground 
n 9 



Both 
m 



Tot. Est. 

N 



Visibility 
Rate 
N/n, 



Los Angeles Bay (rock cliffs) 20 35 20 35.00 1.75 

Midriff Islands (rock cliffs) 
Kino Estero (cactus) 
Islands <26° N Lat. 

(rock cliffs) 
Scammon's Lagoon 

(small island ground nests) 
Pooled (except Scammon's Lagoon) 95 130 80 154.38 1.63 



51 


63 


41 


78.37 


1.54 


7 


9 


6 


10.50 


1.50 


17 


23 


13 


30.08 


1.77 


26 


23 


22 


27.18 


1.05 



until no more were observed. Generally, we did not search for nesting pairs more 
than 2 km landward from the shoreline. We made a single pass to census rocky 
cliffs adjacent to the shoreline or flat terrain with no cacti. It is possible that some 
birds nesting considerable distances from the coastline were missed during the 
survey. From fairly extensive surveys on the ground over the past eight years by 
DWA, we do not believe that any major osprey nesting concentrations were 
located outside our study area. A possible exception is near Puertecitos (northeast 
Baja California) which is discussed later. 

Nests were located by both air and ground observers. We located nests from 
the air during the period 24 March to 1 April. We classified them as occupied if 
an adult was present on or in the immediate vicinity of the nest, or if either young 
or eggs were in the nest. Occupied nests were missed if a bird was not at or near 
a nest when surveyed, the nest was abandoned before the area was surveyed, 
the nest was initiated after the area was surveyed, or we failed to see the nest. 
The ground studies, conducted by individuals involved with other studies, were 
generally made by boat, although one area was surveyed on foot and from a 
pickup truck. The ground studies were conducted within a few days to 2 weeks 
of the interval for the aerial survey. 

The ground studies made it possible to compare numbers of occupied nests at 
the time of census in various areas seen from air, ground, and both air and ground. 
Comparing data from both counts allowed us to obtain a visibility rate for ad- 
justing aerial counts to the total nesting population by use of a modification of 
the Petersen Estimator (see Henny et al., 1978a, b; Burnham and Henny, in 
prep.). We sampled a finite population of size N (N unknown) by use of two 
methods. The data were then recorded so that we knew the number of elements 
n l5 observed by method 1 (aerial survey), the number of elements n 2 , observed 
by method 2 (ground survey), and m, the number of elements observed by both 
methods. Then, 

nin 2 



N 



m 



is a reasonably good estimator of N. In this approach we assumed statistical 
independence of n x and n 2 . 

In sampling osprey nests, it is also necessary to assume N is not changing 
during the time between the ground and air survey. The ground and aerial counts 



92 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 2. A comparison by region and latitude of occupied and unoccupied osprey nests observed 
during the 1977 aerial survey (24 March-1 April). 



West Baja, 
L.C. 



East Baja, 
L.C. 



Midriff 
Islands 



Mainland 
Mexico 



Totals 



Latitude 


Occ. 


Unocc. 


Occ. 


Unocc. 


Occ. 


Unocc. 


Occ. 


Unocc. 


%Occ. 


>27°40' 


78 


75 


67 


53 


115 


103 


54 


38 


54 


26°00' to 27°40' 


27 


12 


47 


42 


— 


— 


— 


— 


58 


<26°00' 


4 


14 


43 


72 


— 


— 


43 


40 


42 


All Combined 


109 


101 


157 


167 


115 


103 


97 


78 


52 



The difference is not significant among locations, x 2 = 11.31, 8 d.f. (.20 < P > .10). 

were made within 2 weeks of each other; therefore, it is doubtful that significant 
changes had taken place. Then N/nj is a reasonably good estimator of the aerial 
visibility rate. The aerial count is multiplied by the aerial visibility rate to obtain 
the population estimates. Separate visibility rates were initially estimated for the 
nests in cacti, the nests on cliffs, and the ground nests on small islands. For this 
study, the nests on cliffs and cacti have been pooled because of their similar 
visibility (Table 1). Aerial visibility rates computed from locations with ground 
surveys were also utilized for similar areas, even though no ground survey was 
made. A comparison of the maximum number of nests seen from both air and 
ground with the estimated number of nests (N) shows that approximately 6% of 



Table 3. Distribution and abundance of nesting osprey s on the Pacific side of Baja California in 
1977. 



Type of nest substrate 



Location 



Cliff 



Cactus Ground Other 



Max. 
Observed 



Total 
Estimate 



Northwest Baja, L.C. 
Puerto Santa Catarina 

to Morro Santo Domingo 
Scammon's Lagoon and 

vicinity 
Natividad Island 
Cedros Island (west side) 
San Benitos Islands 

Subtotal 

Southwest Baja, L.C. 
Punta Eugenia to 

Punta Abreojos 
San Ignacio Lagoon 
Punta Santo Domingo to 

Cabo San Lucas 

Subtotal 
Grand Total 



17. 9 b 



22.8 
19.6 
26 



1.6 



12 



19.5 



— 


27.3 


22.8 


41 


50.1 


— 


— 


— 


14 


22.8 


— 


— 


— 


12 


19.6 


— 


— 


— 


23-29 (26) 


26 


1.6 


27.3 


22.8 


105 


138.0 






1.6 


1 


1.6 


— 


27.3 


— 


26 


27.3 


4.9 


— 


1.6 


4 


6.5 


4.9 


27.3 


3.2 


31 


35.4 


6.5 


54.6 


26.0 


136 


173.4 



b Decimal values included to eliminate rounding errors in total estimates. 
c Not surveyed in 1977, data are for 1971 (Jehl 1977). 



OSPREY POPULATION BAJA CALIFORNIA AND GULF 



93 




Mr 



&^*a>**»<*s&u&w*w'^v*jgr=*~' 







Photograph 1. Occupied ground nest typical of those found in the Scammon's Lagoon and Santa 
Ynez areas. This nest was found on Stony Island in Scammon's Lagoon. 

Note: All photographs by D. W. Anderson. 



the occupied nests on cliffs and in cacti were missed in areas with both air and 
ground counts. Nests occupied at the time of the survey, but believed to have 
been missed by both surveys, are accounted for in estimates presented here. 
However, occupied nests abandoned before the survey, or initiated after the 
survey, are not included in the population estimates. No visibility rates were 
available for nests in mangroves; therefore, the pooled value for cacti and cliff 
nests was used. We believe the nests in mangroves were more difficult to locate 
from the air. Thus, the nesting pairs in mangroves in the Magdalena Bay area 
and coastal Sinaloa were probably underestimated. 

Generally, in more northern latitudes where the survey has been conducted in 
the past (Henny et al., 1974; Henny and Noltemeier, 1975), the nesting cycle was 
synchronized; however, this synchrony does not occur in Mexico. Jehl's 
(1977:243) statement regarding ospreys in Mexico is typical, "nests there con- 
tained all stages from fresh eggs to flying young." To provide some insight into 
the synchrony of nesting activity among locations at the time of the survey, we 
used as an index the ratio of occupied to unoccupied nests recorded during the 
aerial counts (Table 2). Remnants of obviously very old nests were not counted. 
The percentage of occupied nests was lower in Baja California below Lat. 26°N, 
although the difference was not significant (.20 < P > .10). The population es- 
timate presented in this paper for Baja California below Lat. 26°N may, in par- 
ticular, be underestimated if breeding birds (possibly unsuccessful nesters) aban- 
doned some nest sites before the survey was conducted. 



94 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Photograph 2. Nest on a beached trawler as seen from the air north of Puertocitos. It is not 
difficult to note the tracks around the nest made by recreationists in all-terrain vehicles. 



A further complication concerns the presence of nonbreeding birds. One-year 
olds and many of the 2-yr olds are generally not present on the breeding grounds 
of the more northern migratory population in the United States (Henny and Van 
Velzen, 1972). An attempt was made to record these birds as singles during the 
study; however, this was difficult where nesting was concentrated. Locations 
where apparent nonbreeding birds were concentrated will be discussed. 

Results 
Present Distribution and Abundance 

To facilitate presentation of the findings, we have divided the survey into seven 
regions: Northwest Baja California, Southwest Baja California, Northeast Baja 
California, Southeast Baja California, Midriff Islands, Coastal Sonora, and Coast- 
al Sinaloa. A minimum estimate of the total nesting population within the survey 
area was 810 ± 55 pairs (95% C.I.). 

Northwest Baja California. — The area extends from the U.S. -Mexico border 
south to and including Scammon's Lagoon, and west to Punta Eugenia, including 
Natividad and Cedros islands. The San Benitos Islands were not surveyed in 
1977, although recent population data are provided. The total nesting population 
in the region is estimated at 138 pairs (Table 3). No nesting pairs were observed 
between the border and Desembarcadero de Santa Catarina. However, from this 
point south to Morro Santo Domingo, an estimated 20 pairs were nesting, pri- 
marily on cliffs, although one pair nested in a yucca {Yucca spp.). The nesting 
pairs associated with Scammon's Lagoon were concentrated on three small is- 
lands (27 pairs), plus towers, platforms, pilings, channel markers, and debris that 
drifted ashore near the mouth of the lagoon and at Estero de San Jose (23 pairs). 



OSPREY POPULATION BAJA CALIFORNIA AND GULF 



95 




Photograph 3. A typical cardon nest near Kino Bay. Note the large pile of old nesting debris 
below the nest and on the ground, indicating the tendency of ospreys to reuse nest sites from year 
to year. 



Nesting pairs were not observed along Scavenger's Beach between Scammon's 
Lagoon and Punta Eugenia. 

The rocky cliffs on Natividad Island contained an estimated 23 pairs (concen- 
trated on the west side); the west side of Cedros Island contained an estimated 
20 pairs. The east side of Cedros was not surveyed due to high wind conditions, 
but probably contained additional pairs. Jehl (1977) reported 23 to 29 pairs nesting 
on the three Benitos islands in 1971. 

Southwest Baja California. — An estimated 35 pairs nested in this region that 
includes the coastal area between Punta Eugenia and Cabo San Lucas (Table 3). 
Only one pair was observed nesting between Punta Eugenia and Punta Abreojos; 
the nest was on a tower at Tortugas Bay. In San Ignacio Lagoon, an estimated 
27 pairs were nesting on the ground on two small islands (Ballenas). South of San 
Ignacio Lagoon, we estimated that only seven pairs were nesting and they were 
concentrated in the Magdalena Bay region near Santa Margarita Island and on 
the mainland near San Carlos. One pair was of particular interest because it 
nested on the mast of a boat sunk in about 5 m of water near Santa Margarita 
Island. 

Anderson (unpublished data) noted five or six pairs of ospreys nesting on power 
poles and towers in the town of San Carlos in previous years. Apparently, 20-25 
birds in the town have since been shot (J. E. Mendoza, pers. comm.); and no 
nesting pairs were seen in San Carlos in 1977. 

Northeast Baja California. — An estimated 117 pairs of ospreys were nesting 
along the gulf from the mouth of the Colorado River south to Santa Rosalia (Table 



96 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




f 

? 
f 



. 



Photograph 4. Occupied osprey nest on a power pole near Topolobampo. Where power poles and 
other towers have been constructed, osprey are quick to utilize these structures as nesting platforms. 



4). The terrain from the Colorado River south to Puertecitos was very flat and 
contained few cardon cacti (Pachycereus spp.) suitable for nesting sites. Only 
two nesting pairs were observed from the air in the extreme northern portion of 
the region: one nesting on a boat that was aground, and one on a cardon cactus. 
As mentioned earlier, we possibly missed about a dozen nests which were a 



OSPREY POPULATION BAJA CALIFORNIA AND GULF 



97 



Table 4. Distribution and abundance of nesting ospreys on the Gulf side of Baja California and 
the Midriff Islands in 1977. 



Location 



Type of nest substrate 



Max. 



Total 



Cliff 



Cactus Ground Other Observed Estimate 



Northeast Baja, L.C. 
Colorado River to 



Punta Remedios 


34.2 


1.6 


Los Angeles Bay 






and Las Animas Bay 


57.1 


— 


Punta de las Animas 






to Santa Rosalia 


16.3 


6.5 


Subtotal 


107.6 


8.1 


Southeast Baja, L.C. 






Santa Rosalia to Loreto 


58.7 


— 


Loreto to Tambibiche 


42.4 


— 


Tambibiche to 






Cabo San Lucas 


27.7 


1.6 


Subtotal 


128.8 


1.6 


Midriff Islands 






Guardian Angel 


40.8 


— 


Tiburon 


19.6 


52.2 


San Lorenzo, San Lorenzo 






Norte, Salsipuedes, 






Raza, and Partida 


52.2 


— 


San Estaban, Turner, 






and Cholla 


22.8 


— 


Subtotal 


135.4 


52.2 


Grand Total 


371.8 


61.9 



5.3 



5.3 



1.6 



1.6 



1.6 



1.6 



23 


37.4 


50 


57.1 


14 


22.8 


87 


117.3 


44 


65.6 


31 


42.4 


23 


29.3 


98 


137.3 


26 


40.8 


44 


71.8 



43 



52 2 



5.3 



3.2 



25 


22.8 


138 


187.6 


323 


442.2 



considerable distance inland near Puertecitos. About half of the 37 pairs estimated 
nesting between the Colorado River and Punta Remedios were concentrated on 
a group of small islands (Miramar, Lobos, Encantada, San Luis, Pumice). 

The Los Angeles and Las Animas Bay vicinity between Punta Remedios and 
Punta de las Animas contained an estimated 57 pairs. More than half of the pairs 
were nesting on small islands of Los Angles Bay (Smith, la Ventana, Cabeza de 
Caballo, Mitlan, Islas de los Gemelos). From Punta de las Animas south to Santa 
Rosalia the estimated 23 pairs were scattered along the mainland on cliffs or 
adjacent rocks; a few pairs nested in cacti. 

Southeast Baja California. — The coastal area south of Santa Rosalia contained 
an estimated 137 pairs (Table 4). An estimated 66 pairs nesting between Santa 
Rosalia and Loreto were concentrated along the coastline at Concepcion Bay and 
between Punta Pulpito and Loreto. Islands used in the same area included Co- 
ronado, Ildefonso, San Marcos, and Santa Ynez. Five occupied nests on Santa 
Ynez were on the ground, and an additional nest was on a fishing shelter. Tortuga 
Island was not surveyed. 

Between Loreto and Tambibiche an estimated 42 pairs were nesting; an esti- 



98 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Photograph 5. Typical nest on an island in the Midriff region of the Gulf of California with feeding 
habitat in the background. 

mated 16 pairs along mainland cliffs or on immediately adjacent rocks, and the | 
remainder of Santa Catalina, Monserrate, Danzante, and Carmen islands. The 
southernmost portion of the region (Tambibiche to Cabo San Lucas) contained 
an estimated 29 pairs, with about half of the nests on islands and the rest along 



Table 5. Distribution and abundance of nesting ospreys in coastal Sonora and Sinaloa in 1977. 







Type of nest substrate 




Max. 
Ob- 
served 


Total 
Estimate 


Location 


Cliff 


Cactus 


Ground 


Other 


Coastal Sonora 














Colorado River to 
Punta Sargento 
Behind Punta Sargento 3 
Punta Sargento to Guaymas 


8.2 
13.0 


34.2 

36 

32.6 


— 


— 


26 
36 
30 


42.4 

36 

45.6 


Subtotal 


21.2 


102.8 


— 


— 


92 


124.0 


Coastal Sinaloa 














Sonora border to Topolobampo 
Topolobampo to Punta Baradito 
Punta Baradito to Mazatlan 


— 


1.6 
39.1 


— 


4.9 

22.8 

1.6 


4 
38 

1 


6.5 

61.9 

1.6 


Subtotal 


— 


40.7 


— 


29.3 


43 


70.0 


Grand Total 


21.2 


143.5 


— 


29.3 


135 


194.0 



a F. N. Hamerstrom (pers. comm.) ground count in 1977 of area not surveyed from the air. 



OSPREY POPULATION BAJA CALIFORNIA AND GULF 99 

Table 6. Locations where single ospreys were noted during in the survey in greater numbers than 
occupied nests. 

Aerial Counts 







Occupied 




Region 


Location 


Nests 


Singles 


Northwest Baja, L.C. 


Scavenger's Beach 





3 


Southwest Baja, L.C. 


Magdalena and Almejas bays 


4 


36 


Coastal Sonora 


Northern coast 


12 


20 


Coastal Sonora 


Guaymas to Sinaloa border 





6 


Coastal Sinaloa 3 


Topolobampo to Punta Baradito 


38 


14 


Coastal Sinaloa 


Punta Baradito to Mazatlan 


1 


6 



a Possibly a more typical ratio of singles to nesting pairs, although numbers could not be determined 
with accuracy in high density nesting areas. 



the shoreline. The more important nesting islands were Cerralvo and Espiritu 
Santo; a few pairs nested at San Jose and Santa Cruz. 

Midriff Islands. — An estimated 188 pairs of ospreys nested on the islands lo- 
cated at about Lat. 29°N. in the Gulf of California (Table 4). Sixty percent of the 
nesting pairs inhabit Guardian Angel and Tiburon islands. Most of the ospreys 
on the islands nest on cliffs, but a large population nests in cardon cacti on the 
east side of Tiburon. The chain of islands that include Partida, Raza, Salsipuedes, 
San Lorenzo Norte, and San Lorenzo contained an estimated 52 pairs. An esti- 
mated 23 pairs nested on San Estaban, Turner, and Cholla, smaller islands in the 
vicinity of Tiburon. (Note: The air and ground counts show 25 pairs actually 
observed. With smaller numbers involved, occasionally the estimate is slightly 
smaller than the actual count.) San Pedro Martir Island was not surveyed. 

Coastal Sonora. — An estimated 124 pairs of ospreys were nesting in coastal 
Sonora (Table 5). The extreme northern coastal area is flat with no cacti, then 
cardon began appearing sporadically, which attracted both nesting great blue 
herons (Ardea herodias) and ospreys. Moving south along the coast, there are a 
few sandy cliffs and, eventually, some rocky cliffs. An estimated 42 pairs were 
nesting between the Colorado River and Punta Sargento; the majority of the birds 
nested nearer to Punta Sargento where cardon cacti were more common. Some 
36 occupied nests behind Punta Sargento in 1977 were not surveyed from the air, 
but ground counts were made by F. N. Hamerstrom (pers. comm.). The area 
behind Punta Sargento, including also the nearby flats along the eastern shore of 
Tiburon Island, was found to contain perhaps the greatest concentrations of 
breeding ospreys in the Gulf of California region. 

An estimated 46 pairs nested between Punta Sargento and Guaymas, with pairs 
commonly found at Kino Bay, Alcatraz Island, and along the coastline from Kino 
to Guaymas. No nesting pairs were located in the generally flat region between 
Guaymas and the Sinaloa border. In the more northern portion of Sonora the 
cardon cacti provided suitable sites for ospreys to nest in flat country without 
cliffs, but the species of cactus changed below Guaymas to a type that was not 
suitable for nesting. 

Coastal Sinaloa. — An estimated 70 pairs of ospreys nested along coastal Si- 
naloa (Table 5). The few ospreys nesting between the Sonora border and Topo- 



100 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 







Photograph 6. An unoccupied nest on a pinnacle at Puerto Refugio. 



lobampo were found on the islands and peninsulas associated with San Esteban 
and San Ignacio bays, and on a power pole near the town of Topolobampo. South 
of Topolobampo to Punta Baradito, the estimated 62 pairs of ospreys were also 
nesting on the islands and peninsulas associated with the several large bays. 
Mangroves and other brushy trees were abundant and the ospreys readily nested 
on them. The species of cactus changed again in this area to a type that branches 



OSPREY POPULATION BAJA CALIFORNIA AND GULF 101 

Table 7. Types of nest sites selected by ospreys in Baja California and the Gulf of California in 
1977. 







Type of Nest Substrate 














Mangroves 




Location 


Cliffs 


Cactus 


Ground 




Trees 


Others 3 


Northwest Baja, L.C. 


86 


2 


27 




— 


23 


Southwest Baja, L.C. 


— 


5 


27 




— 


3 


Northeast Baja, L.C. 


108 


8 


— 




— 


2 


Southeast Baja, L.C. 


129 


2 


5 




— 


2 


Midriff Islands 


135 


52 


— 




— 


— 


Coastal Sonora 


21 


103 


— 




— 


— 


Coastal Sinaloa 


— 


41 


— 




24 


5 


Total No. Nests 


479 


213 


59 




24 


35 


Percent of Total 


59% 


26% 


7% 




3% 


4% 



a Towers, pilings, channel markers, debris washed ashore, boats (sunk and aground), fishing shelter, 
or power poles. 

about 1.5 m above the ground. These cacti were also used by the nesting ospreys. 
In fact, more ospreys were observed nesting in the cacti than in the mangroves 
or other trees. 

The southernmost nesting pair of ospreys observed during this survey was seen 
at Ensenada del Pabellon (Lat. 24°38'N) in a mangrove. Mazatlan was the south- 
ern terminus of the study. 

Migratory Characteristics 

Most Gulf of California ospreys are year-round residents, after adulthood is 
reached (DWA, unpublished data), a situation similar to that found in Florida at 
approximately the same latitude (Ogden, 1977). This contrasts with the highly 
migratory populations in northern latitudes of the United States, where yearlings 
do not return to the natal area (Henny and Van Velzen, 1972). 

Single birds were observed throughout the study area and in concentrations 
(presumably subadults) at Magdalena and Almejas bays (Table 6). Where nesting 
sites were limited, there was an above-average preponderance of single birds: in 
northern Sonora, in extreme southern Sonora, and in southern Sinaloa which is 
near the southern extreme for the breeding range. Generally we believe the non- 
breeding component of the population was probably scattered throughout the 
study area; however, the single birds were more apparent in areas with few nesting 
pairs. Many of these singles were apparently moulting and looked unkempt. Suit- 
able substrates on which to build nests would not, of course, limit the distribution 
of nonbreeding birds. 

Nesting Sites 

Dead trees or trees with dead tops are the traditional sites for nesting ospreys 
in the western United States (Henny et al., 1978a, b); ospreys are using more and 
more man-made structures along the mid-Atlantic coast of the eastern United 
States (Henny et al., 1974; Henny et al., 1977). Trees and man-made structures 
were rare in the present study area, consequently the ospreys found alternative 
nest sites. 



!02 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Numerically, the most important nesting substrate is large cliffs adjacent to the 
sea, both on the mainland and on the islands (Table 7). Sometimes the rocky 
cliffs consist of large pinnacles or stacks upon which the ospreys build their 
nes ts — some of which are of tremendous bulk. However, small sandy cliffs con- 
tain nesting ospreys in a few areas. An estimated 59% of the nesting pairs were 
associated with cliffs. In the flat terrain, the ospreys nest primarily in various 
species of tall cacti. 

Nesting above ground level is critical for the ospreys on the mainland 
where predator access is likely. Coyotes (Canis latrans) are found throughout 
much of the study area, and it was not uncommon to see several each day during 
the aerial survey. Both Kenyon (1947) and Jehl (1977) mentioned coyote depre- 
dation of ground nests, even when nests were located on small islands. Coyote 
depredation of ground nests occurred again at Scammon's Lagoon in 1977 (San- 
dalio Reyes, pers. comm.). Ground nesting ospreys were restricted to very small 
islands at three locations: Scammon's Lagoon, San Ignacio Lagoon, and Santa 
Ynez Island. 

Miscellaneous man-made structures were used throughout much of the study 
area, but the overall importance of these structures was very small (4%). Most 
of the pairs nesting on these structures were found at the mouth of Scammon's 
Lagoon. Mangroves and other trees were generally available only in the southern 
portion of the study area. 

Status of Population 

Few osprey populations in Mexico have been studied in the past; thus, it is 
difficult to evaluate their current status. A notable exception is northwest Baja 
California. 

Northwest Baja California. — Jehl (1977) recently reviewed the work of Kenyon 
(1947) and earlier workers in northwestern Baja and a few summary statements 
are warranted. The findings for the islands or groups of islands from north to 
south may be summarized as follows: 

(1) Los Coronados — no records of ospreys. 

(2) Todos Santos — once common: greatly reduced by 1910 (1 pair); may have 
been entirely gone by 1923. 

(3) San Martin — about 30 pairs ground nesting in 1913; only 3 pairs in 1946; 1 
pair in 1969-71. 

(4) San Geronimo — common in 1897; reduced to 1 pair by 1912; entirely gone 
by 1926. 

We saw no ospreys nesting on any of these four northernmost islands, or groups 
of islands during this study. 

(5) Natividad — common in 1927 (8 nests within half mile walk); some decline 
suggested by 1946; a minimum of 4 pairs and probably several more present 
in 1969-71. We saw 14 occupied nests from the air in 1977; an adjustment 
for aerial visibility provides an estimate in excess of 20 pairs. 

(6) Cedros — very common in 1882; abundant (13 nests in short distance) 1897; 
abundant in 1912; the west side was not observed in recent years, although 
several nests were located on the east side in 1946 and 1969-71. We saw 
12 occupied nests from the air in 1977 along the west side; an adjustment 



OSPREY POPULATION BAJA CALIFORNIA AND GULF 103 

for aerial visibility provides an estimate of about 20 pairs. The east side 
was not surveyed due to high winds. 
(7) San Benitos — nested in abundance in 1896; an estimated 23-29 pairs in 
1969-71. Not surveyed during this study. 

In summarizing the history of the island osprey populations in northwest Baja 
California, we must agree with the general conclusions of both Kenyon (1947) 
and Jehl (1977) that the populations on the northernmost islands were decimated 
early in the century and have not returned. The decline probably resulted from 
human disturbance, including shooting of adults, and fisherman eating the eggs 
and "squabs" as suggested by both Kenyon and Jehl. The populations on several 
of the more southern islands in the region (Natividad, Cedros, and San Benitos), 
which are generally more distant from human populations, have not become 
extirpated, and appear to be maintaining themselves in good numbers. 

The mainland population in northwest Baja California has been studied very 
little except for the small islands in Scammon's Lagoon. Kenyon (1947) reported 
seeing his first mainland pair of ospreys at Santa Rosalia Bay in 1946, although 
he could not find the nest. He stated (p. 154) "These were the only ospreys up 
to this point observed on the mainland, even though we had run close to shore 
most of the way down the coast." Rowley (1935) also recorded a recently oc- 
cupied nest in a cardon cactus near San Xavier on Santa Rosalia Bay. Jehl (1977) 
reported observations of birds during the nesting season at several locations in- 
cluding a nest at Arroyo San Jose, but indicated that many of the more southern 
locations were inaccessible. 

The first osprey that we saw along the mainland was a single bird at San Carlos 
Bay. We saw 12 nesting pairs from the air (total estimate 20 nesting pairs) between 
Desembarcadero de Santa Catarina and Morro Santo Domingo. Most of the birds 
were on rock, gravel, or sandy cliffs; a few pairs were nesting in cacti and one 
pair nested in a yucca at Santa Rosalia Bay. It has not escaped our attention that 
the main road in this region is away from the coastline, in contrast to the coastal 
road system to the north where ospreys are no longer present. 

The size of Scammon's Lagoon population has been documented for a number 
of years. Kenyon (1947) found 27 pairs (16 at Shell Island) in Scammon's Lagoon 
in 1946. Jehl (1977) reported 25 nests (apparently not all occupied?) on Shell 
Island in 1957, 22 active nests on Shell Island in 1970; and 20 nests (17 with eggs) 
on Shell Island with a total population for the lagoon estimated at about 30 pairs 
in 1971. We located 27 pairs of ospreys nesting on three islands in the lagoon in 
1977. However, 14 were observed from the air (for a population estimate of 23 
pairs) nesting on man-made structures near the mouth of the lagoon and at Estero 
de San Jose. This portion of the lagoon has probably not been censused in the 
past because it can be searched efficiently only from an aircraft. Therefore, the 
27 pairs on the small islands in the lagoon in 1977 compare favorably with the 27 
pairs observed in 1946. Jehl (1977) also concluded that the population seemed 
stationary. 

Other locations. — Long series of population estimates overtime, from the other 
regions, are nonexistent to our knowledge. Most of the early workers only noted 
the presence or absence of the species in general terms. However, some local 
populations are in jeopardy. For instance, the population that nested in the town 
of San Carlos in Magdalena Bay a few years ago was not present in 1977. Shooting 



104 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 8. Eggshell thickness and pollutant residues of Gulf of California ospreys. 





Thickness 
(mm) b 


% Fat 




Pollutants (ppm) 






Wet-wt. 
Mercury 




Lipid-wt. 




Year (n) 


DDE 


TDE 


PCB 


1971 (8) 
Mean ± S.D. 
Range 

1972 (7) 
Mean ± S.D. 
Range 


0.49 ± 0.05 
0.43-0.59 

0.48 ± 0.03 
0.42-0.52 


3.27 ± 0.74 
2.3^.8 

4.94 ± 0.50 
4.4-5.8 


ND C 
ND 

0.2 ± 0.2 
0.1-0.9 


71.2 ± 47.3 
29.0-116.9 

20.0 ± 7.4 
6.1-27 6 


ND 
ND 

<2.5 


0-7.4 

20.2 ± 11.2 
3.1^0.8 



a Analyses conducted by Denver Wildlife Research Center according to methods described by 
Haegele et al. (1974); no adjustment was made for moisture or lipid loss. 
b Normal eggshell thickness is about 0.50 mm (Anderson and Hickey, 1972). 
c ND = no data. 



probably occurs locally throughout the study area wherever the birds closely 
associate with man. 

Pesticides. — It is now generally accepted that persistent pesticides — particu- 
larly DDE, the major metabolite of DDT — have contributed to eggshell thinning, 
poor productivity, and population declines in some birds of prey (Lincer, 1975). 
And, the osprey was among the first species of fish-eating and raptorial birds in 
North America discovered to show indications of a regional pattern of eggshell 
thinning in local populations (Hickey and Anderson, 1968; Anderson and Hickey, 
1972). Local osprey population declines have been documented in many areas 
where production has been extremely poor (see review, Henny, 1977). Spitzer et 
al. (1977) presented some residue data from osprey eggs collected in three general 
areas of Mexico: (1) Scammon's Lagoon, (2) San Benitos Islands, and (3) Gulf 
of California. Additional data from the Gulf of California are presented in Table 
8. One clutch of eggs from the San Benitos Islands showed moderately high DDE 
and PCB residues; however, generally the residues were among the lowest re- 
corded for ospreys in North America. Eggshell thickness appeared to be about 
normal (Spitzer et al., 1977; Table 8). 

Discussion and Recommendations 

This survey yielded the first estimate of the size of the nesting osprey popu- 
lation in the Baja and Gulf of California area, but the survey had some limitations. 
Although this survey was timed to coincide with the peak of nesting activity, the 
lack of a highly synchronized nesting season confounds our approach. Some of 
the pairs may not have been associated with nests at the time the survey was 
conducted; therefore, our estimate of the nesting population's size is small. More 
research on the chronology of the nesting season is needed throughout the study 
area to evaluate the percent of the total breeding population that is nesting at a 
certain time. Additional data may allow a further refinement of the population 
estimate obtained from this study. Nesting chronology studies can also be com- 
bined with productivity studies which can be used to indirectly evaluate the status 
of the population (see Henny, 1977). Osprey eggs from only a few locations have 



OSPREY POPULATION BAJA CALIFORNIA AND GULF 105 

been collected and analyzed for pesticides to date, but the residues appear to be 
low. In view of the general stationary populations over the last three decades 
(modern pesticide era) in the few areas where historical data are available, it 
appears likely that pesticides have not posed a threat to these segments of the 
osprey population. The population declines noted in extreme northwestern Baja 
California occurred before modern pesticides were used. 

Although substantial numbers of ospreys remain today in Baja California and 
the Gulf of California, the extirpation of the population in the extreme northwest 
portion of Baja California should serve as a warning that ospreys require some 
protection if they are to be maintained as a part of the fauna. We know from 
experience in other parts of the species' range that man and ospreys can survive 
together, e.g., in Chesapeake Bay (the largest nesting osprey population in North 
America) where approximately two-thirds of the population is nesting on man- 
made structures and some are very close to human dwellings (Henny et al., 1974). 
In fact, some residents build nesting platforms for ospreys in Chesapeake Bay 
and consider nesting birds near their homes a status symbol. 

In regard to specific management practices, we believe the potentially serious 
problem of coyotes depredating ground nests on the islands in Scammon's Lagoon 
can be solved. Apparently, strong tidal currents constantly change the configu- 
ration of the sandy islands allowing access by coyotes. Nesting platforms about 
3 m above the ground could easily be built on the islands to eliminate this potential 
problem. We recommend initially placing platforms on only one of the islands as 
a test. Ospreys now nest on towers, channel markers, and pilings at the mouth 
of Scammon's Lagoon. Thus, they would probably accept nesting platforms on 
the small islands in the lagoon. 

We believe the ground nesting populations in Scammon's Lagoon and San 
Ignacio Lagoon are the most vulnerable to human disturbance. Protective mea- 
sures should be taken to limit human access to the small islands in these lagoons 
during the nesting season. Cliff nests, especially those on all gulf islands, are also 
vulnerable to the increasing human disturbance (Anderson et al., 1976). 

Acknowledgments 

The study was conducted under the auspices of the Migratory Bird Treaty 
between Mexico and the United States, and approved by Antonio Landazuri, 
Director General of the Fauna Silvestre. Ray Glahn, U.S. Fish and Wildlife Ser- 
vice pilot, guided us safely through areas where landing strips were rare, airports 
with gas were rarer, and radio communication was nonexistent. Ground counts 
were made by James Keith, Chris Mitchell, Alan Jenkins, Richard Porter, and 
Charles Stone of the U.S. Fish and Wildlife Service, who were in Mexico working 
on studies of their own; Debra Judge, graduate student at the University of Cal- 
ifornia, Davis; and Sandalio Reyes, Departmento de al Fauna Silvestre, Guerrero 
Negro, Lower California. F. N. Hamerstrom kindly provided ground counts from 
the Punta Sargento region. The manuscript was reviewed by Eugene Dustman, 
Stana Federighi, and Lawrence Blus. 

Literature Cited 

Ames, P. L. 1966. DDT residues in the eggs of the osprey in the northeastern United States and 
their relation to nesting success. J. Appl. Ecol., 3(suppl.):87-97. 



10 6 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

; and G. S. Mersereau. 1964. Some factors in the decline of the osprey in Connecticut. Auk, 

81:173-185. 
Anderson, D. W., and J. J. Hickey. 1972. Eggshell changes in certain North American birds. Int. 

Ornithol. Cong., 15:514-540. 
, j. e. Mendoza, and J. O. Keith. 1976. Seabirds in the Gulf of California: A vulnerable 

international resource. Nat. Resour. J., 16:483-505. 
Friedmann, H., L. Griscom, and R. T. Moore. 1950. Distributional check-list of the birds of Mexico. 

Pacific Coast Avifauna 29. 
Grinnell, J. 1928. A distributional summation of the ornithology of Lower California. Univ. Calif. 

Publ. Zool. 32. 
Haegele, M. A., R. K. Tucker, and R. H. Hudson. 1974. Effects of dietary mercury and lead on 

eggshell thickness in mallards. Bull. Environ. Contam. Toxicol., 11:5-11. 
Henny, C. J. 1977. Research, management, and status of the osprey in North America. Pp. 199-222 

in Proc. World Conf. of Birds of Prey (R. D. Cancellor, ed.), Int. Counc. Bird Preservation, 

Vienna, xii + 442 pp. 
, M. A. Byrd, J. A. Jacobs, P. D. McLain M. R. Todd, and B. F. Halla. 1977. Mid-Atlantic 

coast osprey population: present numbers, productivity, pollutant contamination, and status. 

J. Wildl. Manage., 41:254-265. 
, J. A. Collins, and W. J. Deibert. 1978b. Osprey distribution, abundance, and status in 

western North America: II. The Oregon population. Murrelet, 59:14-25. 
, D. J. Dunaway, R. D. Mallette, and J. R. Koplin. 1978a. Osprey distribution, abundance, 

and status in western North America: I. The northern California population. Northwest Sci., 

52:261-272. 
, and A. P. Noltemeier. 1975. Osprey nesting populations in the coastal Carolinas. Am. Birds, 

29:1073-1079. 
, M. M. Smith, and V. D. Stotts. 1974. The 1973 distribution and abundance of breeding 

ospreys in the Chesapeake Bay. Chesapeake Sci. 15:125-133. 
, and W. T. Van Velzen. 1972. Migration patterns and wintering localities of American os- 
preys. J. Wildl. Manage., 36:1133-1141. 
Hickey J. J., and D. W. Anderson. 1968. Chlorinated hydrocarbons and eggshell changes in raptorial 

and fish-eating birds. Science, 162:271-273. 
Jehl, J. R., Jr. 1977. History and president (sic) status of ospreys in northwestern Baja California. 

Pp. 241-245 in Trans. N. Am. Osprey Res. Conf. (J. C. Ogden, ed.), U.S. Natl. Park Serv., 

xiii + 258 pp. 
Kenyon, K. W. 1947. Breeding populations of the osprey in Lower California. Condor, 49:152-158. 
Lincer, J. L. 1975. DDE-induced eggshell-thinning in the American kestrel: a comparison of the field 

situation and laboratory results. J. Appl. Ecol., 12:781-793. 
Ogden, J. C. 1977. Preliminary report on a study of Florida Bay ospreys. Pp. 143-151 in Trans. N. 

Am. Osprey Res. Conf. (J. C. Ogden, ed.), U.S. Natl. Park Serv., xiii + 258 pp. 
Rowley, J. S. 1935. Notes on some birds of Lower California, Mexico. Condor, 37:163-168. 
Spitzer, P. R., R. W. Risebrough, J. W. Grier, and C. R. Sindelar. 1977. Eggshell thickness- 
pollutant relationships among North American ospreys. Pp. 13-19 in Trans. N. Am. Osprey 

Res. Conf. (J. C. Ogden, ed.), U.S. Natl. Park Serv., xiii + 258 pp. 

Accepted for publication January 4, 1979. 

Addendum: Mazatlan Harbor was the terminus of the 1977 study; however, Morlan Nelson 
(pers. comm.) reported an occupied osprey nest on an island there one year later (March, 1978). 



Bull. Southern California Acad. Sci. 
78(2), 1979, pp. 107-115 

A New Species of Leptodactylid Frog, Genus Eleutherodactylus, 
from the Cordillera de Talamanca, Costa Rica 

Jay M. Savage and James E. DeWeese 

Abstract. — Eleutherodactylus rayo a distinctive new species sharing features 
of the fitzingeri and rugulosus groups occurs along the Pacific slopes of the 
Cordillera de Talamanca of Costa Rica. Karyologically the new form most closely 
resembles El. vocalis of northwestern Mexico and certain populations of frogs 
from eastern Mexico, formerly referred to El. rugulosus but for which the name 
El. berkenbuschii W. Peters, 1870, is revived. Unlike other members of the ru- 
gulosus group which have 2N = 20, berkenbuschii, rayo and vocalis have 2N = 
22 and approach El. talamancae of Coast Rica and Panama, a member of the 
fitzingeri group, in karyotypes. 

Allan Hancock Foundation and Department of Biological Sciences, University 
of Southern California, Los Angeles, California 90007 and Department of Biology, 
California State University, Long Beach, California 90840. 



During the past 20 yr of work in Costa Rica, the University of Southern Cali- 
fornia field teams have accumulated a number of distinctive or unique frogs that 
appeared to be representative of species not previously known to science. In 
most cases formal description has been delayed pending collection of additional 
material. A single example of the genus Eleutherodactylus falling into this cate- 
gory was originally collected by Roy W. McDiarmid in 1964 from a remote area 
on the Pacific slope of the Cordillera de Talamanca. Subsequently other speci- 
mens of this form were taken from 1972-1976 in the same mountain range along 
the Carreterra Interamericana. 

The new species belongs to the fitzingeri group (sensu Lynch, 1976) and is 
somewhat intermediate in characteristics between the fitzingeri and rugulosus 
groups as used by Savage (1975, 1976). It is called: 

Eleutherodactylus rayo, new species 
' Fig. 1 

Holotype. — LA 127669, an adult male from the second sabana on the trail from 
Finca El Helechales to Sabanas Esperanza, 5 km, airline, east of Finca El He- 
lechales, Canton de Buenos Aires, Provincia Puntarenas, Costa Rica, 1640 m; 
collected by Roy W. McDiarmid, October 6, 1964. 

Diagnosis. — The new form superficially resembles several lower Central Amer- 
ican species of the fitzingeri and rugulosus groups. It differs from members (audi, 
fitzingeri and talamancae) of the former group, that share with it the feature of 
having the webs between toes III— IV extending at most only slightly distal to the 
proximal subarticular tubercle in having a well-developed calcar. All other mem- 
bers of the group have substantially more toe webbing, which extends nearly to 
distal subarticular tubercle (III) between toes III— IV and to halfway between 
proximal and penultimate subarticular tubercles (IV). 



108 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 1. Eleutherodactylus rayo, male paratype (CRE 3980) from Quebrada Fortuna at Carreterra 
Interamericana, Provincia de San Jose, Costa Rica, 1750 m. DeWeese photograph. 



Within this cluster of slightly webbed forms it may be distinguished further by 
differences in posterior thigh coloration which is uniform reddish brown (tala- 
mancae), brown with discrete small light spots (fitzingeri) or dark chocolate with 
very large light spots {audi). The posterior thigh surfaces of rayo are essentially 
a uniform dark purple in life and preservative. 

El. rayo is distinguished from members of the rugulosus group in having a well- 
developed calcar and in having greatly enlarged emarginate disks on fingers III— 
IV, while the disks are rounded, not greatly enlarged and never emarginate in the 
latter group. The dark purplish coloration of the hindlimb surfaces uniquely dis- 
tinguishes El. rayo from all other known members of the fitzingeri and rugulosus 
groups in Central America. 

Summary of characteristics. — General: Head about as wide as long. Nostril 
closer to tip of snout than to eye. Loreal outline slightly sloping. Snout profile, 
and canthus rostralis round. Dorsal outline of snout subelliptical. Choanae ovoid; 
vomerine teeth located between but behind level of choanae in two transverse 
series separated at the midline. Vocal slits and internal vocal sacs in males. 
Tympanum approximately Vi height of orbit; internal, indistinct; round in males 
oval in females. Skin of head and dorsum smooth; upper eyelid with a single 
tubercle. Finger II longer than I. Finger disks on I— II rounded and expanded, on 
III— IV emarginate; about 2 times as wide as finger on III-IV. All fingers have 
well developed lateral fringes. Whitish nuptial pads on thumbs of male. Subar- 
ticular tubercles flattened, round to ovoid in outline, globular; no supernumerary 
tubercles; thenar tubercle large, elongate; palmar tubercle large ovate; no acces- 



NEW SPECIES OF FROG FROM COSTA RICA 



109 



PERCENT GENOME 

* » 


















1 2 3 4 5 6 7 8 9 10 11 
CHROMOSOME NUMBER 



Fig. 2. An idiogram of the karyotype of Eleutherodactylus rayo. 



sory palmar tubercles. Calcar present. Toe disks palmate on I, emarginate on II- 
IV and rounded and expanded on V; about 1.5 times as wide as toe on IV. Toe 
fringes well developed. Toes webbed only basally. Modal toe webbing formula: 
I 2"-2 + II 2-3 III 3-4 + IV 4 + -3~ V. Subarticular tubercles projecting, ovoid in 
outline, obtuse to conical in profile; supernumerary tubercles lacking; outer plan- 
tar tubercle round, !4 size of oval inner tubercle. Inner tarsal fold well developed. 
Venter smooth. 

Coloration. — Base color of top of head, body and limbs a deep dark brown 
with a definite bluish purple cast; in preservative; in life deep bluish purple, adult 
females slightly lighter and tending toward tan as compared to the deep purple 
males; upper surface of snout light gray in some examples; often an interocular 
dark bar bordered anteriorly by a light area; upper lips with three dark bars in 
males; bars suggested by dark areas along mouth margin in females; a definite 
narrow supratympanic dark mark runs from middle of eye backward above tym- 
panum and curves downward to shoulder. Dorsum uniform or with dark supra- 
scapular spots or blotches in some examples; sometimes with a narrow white 
light line along median raphe; one adult female with a broad mid-dorsal light 
stripe bordered on either side by a broad dark area. Males often with some 
blotches of olive green in life. Dorsal and anterior surfaces of limbs uniform or 
with broad crossbars of dark pigment. Posterior surface of thigh dark purplish 
with obscure small light punctations. Throat heavily marked with dark pigment; 
with a narrow median light line. Venter light with a heavy mottling of dark pig- 
ment. Ventral surface of hind limbs marked like posterior thigh surface: plantar 
surface uniform purple. Groin and flanks similar in color to dorsum. 

Measurements. — In this section the notation gives the mean followed by the 
range in parentheses. Standard lengths (distance from snout to vent) are given in 
millimeters; other measurements as percentages of standard length. 

Standard length, adult males (N = 10) 40.0 (37.4-45), adult females (N = 10) 
53.6 (38.2-70.9); head length, males 38.0 (34.9-41.0), females 37.3 (35.1-38.6): 



110 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




B 



## 



o*.v>X*%*** 



c% $> ***i «**•»*• *» 



Fig. 3. A representative metaphase spread (A) and karyotype (B) for Eleutherodactylus rayo. 



head width, males 38.2 (35.8-40.5), females 41.0 (39.0^12.7); snout length, males 
18.4 (17.6-19.3), females 18.6 (17.5-19.1); loreal length, males 10.6 (9.1-11.9) 
females 8.1 (7.4-8.9); length of orbit, males 14.7 (13.3-16.4), females 12.9 (11.7- 
15.2); height of tympanum, males 8.6 (7.3-9.8), females 5.9 (5.1-6.5); hindlimb 
length, males 210.7 (203.1-217.5), females 212.5 (209.6-215.6); tibia length, males 
65.3 (62.7-66.4), females 57.4 (55.8-58.9). Note non-overlapping in tympanum 
height and tibia length to produce strong sexual dimorphism. 

Karyotype. — Seven paratypes (CRE 3184[2], 3236-39, 9770) were examined 
karyologically using the technique of Patton (1967) as modified by Lowe et al. 
(1966). 122 spreads were counted, three were photographed and the individual 
chromosomes measured (Table 1), and an idiogram constructed (Fig. 2). 

The diploid number (2N) is 22 and the nombre fundamental (N.F.) is 36. The 
chromosomes form a gradually descending series with no obvious size groupings. 
Chromosome pairs (centromere placement according to the system of Levan et 
al. 1964), 1,3,5,6 and 7 are metacentric; 4 is submetacentric; 2 is subacrocentric 
and 8-11 are acrocentric (Fig. 3). 



NEW SPECIES OF FROG FROM COSTA RICA 1 1 1 

Table 1. The centromeric indices (C.I.) and percent genome values (G) for the 11 pairs of chro- 
mosomes in the karyotype of Eleutherodactylus rayo. 

Chromosome 

Number C.I. G. 

1 1.64 16.2 

2 3.92 12.2 

3 1.44 12.0 

4 1.72 11.8 

5 1.27 10.5 

6 1.34 9.3 

7 1.12 7.6 

8 7.00 6.8 

9 7.00 5.2 

10 7.00 4.5 

11 7.00 3.7 



Habitat. — The male holotype was captured during the day while calling from 
a bromeliad during a heavy rain. Most of the paratypes were collected along the 
margins of a small stream during the daytime. They were found under debris or 
were apparently frightened out of their hiding places by the collector's activity 
and were hopping along the stream banks or across the shallows. 

Distribution. — Rainforests of the upper portion of the premontane and lower 
portion of the lower montane slope of the Pacific face of the Cordillera de Tala- 
manca of southwestern Costa Rica, between 1600-1850 m (Fig. 4). 

Localities.— COSTA RICA: PUNTARENAS: 5 km E Finca El Helechales, 
1640 m (LA 127669); SAN JOSE: Quebrada Fortuna at Carreterra Interameri- 
cana, 1750 m-1840 m (CRE 3184, 3232-33 3236-39, 3979-80, 6599, 8710-12, 9769; 
KU 65986); Rio Payner at Carreterra Interamericana, 1480 m (CRE 9770). All 
examples from San Jose Province are paratypes. 

The name rayo is an arbitrary combination of letters that happens to mean 
lightning in Spanish. The name is an allusion to Roy W. McDiarmid, who first 
collected the species, as recognition for his work on the Costa Rican herpeto- 
fauna. Those who know Roy well will discern other reasons why this name is 
appropriate. 

Relationships 

The speciose genus Eleutherodactylus has always presented a serious problem 
to the systematic herpetologist interested in establishing relationships among 
morphologically similar forms. Currently about 350-400 species are recognized 
in the genus which ranges throughout the Neotropical region. The morphology 
of the group ranges from huge, stream-adapted toad-like species through mod- 
erate-sized frog-like terrestrial forms, arboreal treefrog-like species with huge 
finger and toe disks to large toad-like forest floor burrowers. While distantly 
related forms seem very distinct, the features of external morphology within the 
genus are recombined over and over again in a mosaic pattern that tends to make 
recognition of natural subdivisions difficult, while closely related forms often 
differ only slightly in these same features. One recourse in this situation has been 
to establish a series of species groups within the genus by clustering species that 



112 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 4. Distribution of Eleutherodactylus rayo. The dotted line indicates the 1500 m contour. 



are extremely similar morphologically and are probably phylogenetically from the 
same evolutionary lineage (e.g., Savage, 1976). Another alternative (Lynch, 1976) 
has been to group the species at a higher level (subgeneric) without giving the 
groups formal taxonomic status. Unfortunately the first method does not provide 
a sound framework for establishing the evolutionary history of the genus, while 
the second creates infrageneric divisions based on "key" characteristics of ex- 
ternal and skeletal morphology that are of dubious evolutionary cogency. 

It is clear from the study of jaw musculature (Starrett, 1968), serum proteins 
(Harris, 1973) and karyology (DeWeese, 1976) that a series of at least six major 
evolutionary lineages are subsumed under the genus Eleutherodactylus. In most 
cases these lineages do not correspond to the groupings established on external 
and skeletal morphologic grounds (Lynch, 1976) and suggest that the time is 
premature for an attempt to establish higher-level divisions based solely on tra- 
ditional taxonomic (key) characters. 

El. rayo morphologically is allied to the fit zing eri and rugulosus groups (Sav- 



NEW SPECIES OF FROG FROM COSTA RICA 113 

age, 1975, 1976). Lynch (1976) proposed to place thase groups together as the 
fitzingeri group. While this is not the place to consider fully the weaknesses of 
Lynch's (1976) system, it must be noted that both El. agidi of Costa Rica and El. 
rayo within this group have emarginate finger disks to negate one feature (ungual 
flap not indented) used to characterize this division. In addition one population 
referred to El. rugulosus (Savage, 1975) has the first finger shorter than the sec- 
ond, while Lynch (1976) characterizes the fitzingeri as having the converse sit- 
uation. 

Nevertheless there can be little doubt that the two groups recognized by Savage 
(1976) are closely allied. This relationship is further supported by the condition 
of the jaw muscles: depressor mandibulae with a single slip originating on the 
dorsal fascia but a few fibers coming from the squamosal (dfsq) or with a single 
slip originating on the dorsal fascia but a few fibers coming from the squamosal 
and annulus tympanicus (dgsqat) and adductor mandibulae externus superficialis 
present (e). Formulae for jaw muscles after Starrett (1968), are inadvertantly 
misstated for the rugulosus group in Savage (1975). Karyologically the two groups 
are also similar with the fitzingeri group having 2N = 22 and and N.F. = 36^40 
and the rugulosus group with 2N = 20-22, and a N.F. = 36. 

In terms of morphology El. rayo is distinct from all other members of the 
fitzingeri and rugulosus groups in having large emarginate finger and toe disks 
and a well-developed calcar. The only other species in these groups with similar 
disks, El. andi of Costa Rica, lacks the calcar and has a bold contrasting thigh 
pattern of large light spots on a dark brown background. 

The jaw muscle formula for the new form is dfsqat + e. It seems likely that 
the differences between dfsq and dfsqat are trivial, since only a few muscle fibers 
are involved. The latter condition is not currently known to occur in the fitzingeri 
group while it does appear in some rugulosus group members (e.g., El. brocchi 
of Guatemala). 

Karyologically El. rayo agrees in diploid number (2N = 22) and nombre fun- 
damental (36) with El. talamancae of Costa Rica and Panama (in the fitzingeri 
group) and within the rugulosus group with El. vocalis of northwestern Mexico 
and the rugulosus-like populations of eastern Mexico (populations 1-2 of Savage, 
1975). Other leptodactylids with the same numbers are: El. decoratus of Mexico 
{alfredi group, Lynch, 1976); and Hylactophryne augusti, a close Eleutherodac- 
tylus relative; Leptodactylus podicipinus and L. wagneri of South America. Of 
these forms the karyotype of El. rayo most closely resembles that of the rugu- 
losus-\ike populations of eastern Mexico. 

These data confirm the uniqueness of the new species, which does not appear 
to be closely allied to any known member of either the fitzingeri or rugulosus 
groups. 

The Status of Hylodes berkenbuschii 

Savage (1975:271) regarded the eastern Mexico populations (1-2) of the ru- 
gulosus population system as morphologically distinct from other members of the 
rugulosus stock. Nevertheless he adopted a conservative position, since the dis- 
tinctive populations were allopatric to typical El. rugulosus (Cope, 1870) and 
retained them within the latter species. 

Subsequently, karyologic analysis of the rugulosus population system (De- 



114 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Weese, 1976) discerned that the eastern Mexico populations differed significantly 
from all other units in the rugulosus population system. The eastern Mexico 
populations have a diploid number (2N) of 22 and a nombre fundamental of 36. 
All other known members in the system have 2N = 20. In this regard the eastern 
Mexico populations resemble El. vocalis of northwestern Mexico but differ from 
the latter form in chromosomal definition. These two forms are amply distinct 
morphologically (Savage, 1975). A more detailed discussion of karyologic features 
and evolution in the rugulosus group is in preparation by DeWeese. 

The combination of morphologic differences supported by the unique karyology 
of the eastern Mexico populations force us to conclude that they represent a 
species distinct from El. rugulosus. The first available name for this form is 
Hylodes berkenbuschii W. Peters, 1870 (holotype: Berlin 6666; Mexico: Puebla: 
nr. Izucar de Matamoras). The species now to be called Eleutherodactylus ber- 
kenbuschii (W. Peters, 1870) is characterized by Savage (1975:270) and the dis- 
tribution reviewed (pp. 271, 291-292). El. berkenbuschii may be most easily dis- 
tinguished from El. rugulosus morphologically as follows (characteristics of the 
latter form in parentheses): canthus rostralis sharp (rounded) and first finger usu- 
ally shorter than second finger (first finger usually longer than second). 

Synonyms of El. berkenbuschii include Eleutherodactylus natator Taylor, 1939 
(holotype: CM 1000014; Mexico: Veracruz: Tlilapam) and Eleutherodactylus vul- 
cani Shannon and Werler, 1955 (holotype: Mexico: Veracruz: Volcan San Mar- 
tin.) 

Conventions and Acknowledgments 

In order to reduce the number of times the long name Eleutherodactylus is 
spelled out in this and subsequent papers and to clearly distinguish it from other 
anuran groups when the generic name is abbreviated, we have adopted the form 
El. to represent the generic name. Those who know Spanish will note that this 
also provides an euphonious neatness to discussions of species in the genus since 
el is the Spanish definite article meaning "the/' We have eschewed the use of 
diacritical marks on Spanish localities for the reasons given by Stuart (1963). 

Material used in this study has been made available through the kindness of 
the authorities at the Museum of Natural History, University of Kansas (KU) 
and the Los Angeles County Museum of Natural History (LA). Field as- 
sistance was provided by several workers from the University of Southern 
California: Rosemarie DeWeese, Ronald T. Harris, Carl S. Lieb, Sandra Lim- 
erick, James J. Talbot and Nancy D. Savage. Dr. P. H. Starrett examined the 
jaw area of the new form to establish its jaw muscle formula. Our work in Costa 
Rica was expedited by aid from the Organization for Tropical Studies and the 
Facultad de Biologia, Universidad de Costa Rica. To all of the above we express 
our deep appreciation. 

Literature Cited 

Cope, E. D. 1870. Seventh contribution to the herpetofauna of tropical America. Proc. Amer. Philos. 

Soc, 11:147-169. 
DeWeese, J. E. 1976. The karyotypes of Middle American frogs of the genus Eleutherodactylus 

(Anura: Leptodactylidae): A case study of the significance of the karyologic method. Ph.D. 

diss., Univ. Southern California: 1-210. 
Harris, R. T. 1973. Comparative serum-protein electrophoresis and protein characterization in the 



NEW SPECIES OF FROG FROM COSTA RICA 115 

systematics of Costa Rican frogs genus Eleutherodactylus (Anura: Leptodactylidae). Ph.D. 

diss., Univ. Southern California: 1-247. 
Levan, A., K. Fredga, and A. A. Sandberg. 1964. Nomenclature for centromere positioning on 

chromosomes. Hereditas, 52:201-220. 
Lowe, C. H., J. W. Wright, and C. J. Cole. 1966. Chromosomes and karyotypes of sceloporine 

iguanid lizards in the North American Southwest. Mamm. Chroms. Newslt., Houston, 

2(2): 20 1-205. 
Lynch, J. D. 1976. The species groups of the South American frogs of the genus Eleutherodactylus 

(Leptodactylidae). Occ. Pap. Mus. Nat. Hist., Univ. Kansas, 61:1-24. 
Patton, J. L. 1967. Chromosome studies of certain pocket mice, genus Perognathus (Rodentia: 

Heteromyidae). J. Mamm., 48(l):27-37. 
Savage, J. M. 1975. Systematics and distribution of the Mexican and Central American stream frogs 

related to Eleutherodactylus rugulosus. Copeia, 1975(2):254-306. 
. 1976. A preliminary handlist of the herpetofauna of Costa Rica. 2nd ed. Editorial Univ. de 

Costa Rica: 1-19. 
Shannon, F. A., and J. E. Werler. 1955. Notes on amphibians of the Los Tuxtlas range of Veracruz, 

Mexico. Trans. Kansas Acad. Sci., 58:360-386. 
Starrett, P. H. 1968. The phylogenetic significance of the jaw musculature in anuran amphibians. 

Ph.D. diss., Univ. Michigan: 1-179. 
Stuart, L. C. 1963. A checklist of the herpetofauna of Guatemala. Misc. Publ. Mus. Zool., Michigan, 

122:1-150. 
Taylor, E. H. 1939. New species of Mexican tailess Amphibia. Univ. Kansas Sci. Bull., 25:385-405. 

Accepted for publication February 1, 1979. 



Bull. Southern California Acad. Sci. 
78(2), 1979, pp. 116-121 



New Species and Records of Polychaetous Annelids from the 

Tetraclita (Cirripedia: Crustacea) Zone of the Northern 

Gulf of California, Mexico 

Jerry D. Kudenov 

Abstract. — Two new species of polychaetes belonging to the families Phyllo- 
docidae and Nereidae are described from the Tetraclita zone of Bahia Cholla, 
Puerto Penasco, Sonora, Mexico. The ranges of Syllis elongata (Johnson, 1901), 
Typosyllisfasciata (Malmgren, 1867) and Perinereis monterea (Chamberlin, 1918) 
are extended into the Gulf of California. 

Marine Studies Group, Ministry for Conservation, 605 Flinders St. Ext., Mel- 
bourne, Victoria 3000, Australia. Present Address: Allan Hancock Foundation, 
University of Southern California, Los Angeles, California 90007. 



Over 500 species of polychaetes are currently known from the Gulf of California 
(Reish, 1968; Fauchald, 1972; Kudenov, 1975a, b, c and in press). A small col- 
lection of polychaetes was kindly made available by Prof. J. R. Hendrickson, 
University of Arizona. These were collected during 10-24 October 1976 by his 
student, Mr. R. Dougherty, from under the tests of Tetraclita squamosa (Bru- 
guiere) at Bahia Cholla, Puerto Penasco, Sonora, Mexico. This collection site is 
hereafter referred to as the "study area." T. squamosa occurs in the high inter- 
tidal zone and is rather common in the Gulf of California (Brusca, 1973). The 
ecological succession and distribution of invertebrate species in vacant tests of 
T. staliactifera panamensis was investigated in Panama by Reimer (1976a, b). 
The entire collection and types are lodged at the Allan Hancock Foundation, 
University of Southern California. 

Family Phyllodocidae 

Eumida uschakovi, n. sp. 

Fig. 1 

Material examined. — Study area, Holotype, AHF POLY 1217. 

Description. — Holotype complete, with 256 setigers, measuring 25 mm long, 
1 mm wide with parapodia, 0.6 mm wide without. Body elliptical in cross section, 
lacking midventral groove; uniformly pale rose in color in alcohol with traces of 
dark dorsolateral pigmentation. 

Prostomium elliptical, longer than wide, with 5 conspicuous antennae (Fig. la). 
Anterior dorsal pair of antennae conical, longer than wide; ventral pair cirriform, 
about 2 times longer than dorsal pair; median antenna inserted on anterior pro- 
stomium, well ahead of eyes. One pair of dark, circular, lenticulate eyes present 
middorsally; nuchal papilla absent. Proboscis not everted, not examined. 

First segment reduced, not visible dorsally. Tentacular formula of 1 + Sf + S& 
with tentacular cirri cirriform; with dorsal cirri of segments 2 and 3 longest, and 
ventral cirrus of segment 2 shortest; ceratophores of each tentacular cirrus with 
3-4 annuli. 



NEW SPECIES AND RECORDS OF POLYCHAETOUS ANNELIDS 



117 




Fig. 1. a-e. Holotype, AHF POLY 1217, Euinida uschakovi. a. anterior setigers, dorsal view, 
x27; b. right parapodium, setiger 10, anterior view, x70; c. same, setiger 75, anterior view, x70; d. 
left parapodium, setiger 125, x70; e. composite spiniger, x340. 



Dorsal cirri 2-3 times longer than wide; broadly lance-shaped, distally blunt in 
anterior, medial setigers; becoming narrow, distally pointed in posterior setigers 
(Fig. lb-d). Dorsal cirri of anterior, posterior setigers oval in cross section; those 
of medial setigers flattened. Ventral cirri distally rounded, projecting beyond 
parapodial lobes only of anterior, posterior setigers. Parapodial lobes distally 
notched, with aciculum penetrating notch in anterior setigers (Fig. lb-d). 

All setae as composite spinigers with spinous distal shafts and short appendages 
(Fig. le). Appendages obliquely sculptured, with a row of denticles on cutting 
edge becoming indistinct distally; number of setae gradually decreasing from 16 
in setiger 10; 13 in setiger 75; 10-11 in setiger 125. 

Pygidium with 1 pair of stout, distally blunt anal cirri equalling length of last 
5-6 prepygidial setigers. 

Discussion. — Eumida uschakovi is similar to E. fusigera (Malmgren) sensu 
Uschakov (1972), E. parva St. Joseph and E. granulosa (Verrill) in having dorsal 
cirri at least 2 times longer than wide. E. uschakovi differs from these species in 
having inflated, instead of flattened dorsal cirri; in having the median antenna 
inserted on the anterior prostomium, instead of between the eyes; and in lacking 
both prolonged superior parapodial processes and pointed ventral cirri. This 
species is named in honor of Prof. P. V. Uschakov in recognition of his mono- 
graphic study of phyllodociform polychaetes. 



1 18 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Family Syllidae 
Syllis elongata (Johnson, 1901) 

Pionosyllis elongata Johnson, 1901:403^04, pi. 6, figs. 67-70. 

Syllis elongata. Hartman, 1968:461^62, figs. 1-3; Banse and Hobson, 1974:61. 

Material examined. — Study area, 2 specimens, AHF 000366-0. 

Description. — These specimens generally agree with the original description, 
but differ in the segmental distribution of the pharnyx and proventriculus, and in 
the number of articles per dorsal cirrus. The pharynx extends through setigers 
13—14, while the proventriculus is present in setigers 14/15-22. The long and short 
dorsal cirri of setigers 10 and 11 have 45 and 26 articles, respectively; those of 
setigers 190 and 191 with 15 and 12. 

Discussion. — S. elongata normally has the pharynx and proventriculus through 
setigers 11, and setigers 11-19/20, and dorsal cirri with 16-20 articles anteriorly, 
and 14-15 posteriorly. The mexican specimens differ, and may represent a new 
subspecies, which I hesitate doing until additional materials are available. 

Distribution. — Western Canada to southern California; newly reported from 
the Gulf of California, Mexico. 

Typosyllis fasciata (Malmgren, 1867) 

Typosyllis fasciata. Berkeley and Berkeley, 1948:74-75, fig. 109; Hartman, 
* 1968:485-486, figs. 1-3; Banse and Hobson, 1968:64. 

Material examined. — Study area, 2 specimens, AHF 000366-02. 
Discussion. — These specimens agree well with previous descriptions. 
Distribution. — Western Europe; western Canada to southern California; newly 
reported from the Gulf of California, Mexico. 

Family Nereidae 
Perinereis monterea (Chamberlin, 1918) 

Perinereis monterea. Hartman, 1968:557-558; Banse and Hobson, 1974:71. 

Material examined. — Study area, 2 specimens, AHF 000366-03. 
Discussion. — These specimens agree well with previous descriptions. 
Distribution. — British Columbia to western Mexico; newly reported from the 
Gulf of California, Mexico. 

Neanthes cortezi, n. sp. 
Fig. 2 

Material examined. — Study area, Holotype, AHF POLY 1218; 20+ Paratypes, 
AHF POLY 1219. 

Description. — A small species up to 25 mm long, 1.5 mm wide with parapodia, 
1 mm wide without, for 80 setigers. Body elliptical in cross section, midventrally 
grooved; prostomium, palps, palpistyles, antennae with brown pigment. 

Prostomium longer than wide, distally rounded, with conical frontal antennae 
(Fig. 2a). Palpi very large, about as long as prostomium; palpistyles nearly spher- 
ical. Two pairs of eyes present with anterior pair farthest apart, crescent-shaped; 
posterior pair elliptical. 

Proboscis with dark brown, conical paragnaths on both rings as follows: I, 



NEW SPECIES AND RECORDS OF POLYCHAETOUS ANNELIDS 



119 




Fig. 2. a-h. Holotype, AHF POLY 1218, Neanthes cortezi- a. anterior segments, dorsal view, 
x27; b. maxillary ring, pharynx, frontal view, x27; c. right parapodium, setiger 10, anterior view, 
x70; d. left parapodium, setiger 40, anterior view, x70; e. right parapodium, setiger 70, anterior view, 
x70; f. homogomph spiniger, x250; g-h. superior and inferior heterogomph falcigers, x340. 



diamond patch of 4 cones; II triangular patch of 25 cones in 5 rows; III, oval 
patch of 45 cones in 5 rows; IV, trapezoidal patch of 56 cones in 8 rows; V, 
diamond patch of 14 cones; VI, single high conical paranath; VII-VIII, 91 large, 
small cones in 3 continuous rows (Fig. 2a, b). Jaws dark brown in color each with 
11 triangular teeth. 

Peristomium Vt, as long as prostomium, with 4 pairs of cirriform tentacular cirri; 
anterior and posterior ventral pairs extending to setiger 1; anterior dorsal pair 
extending to setiger 3; posterior dorsal pair extending to setigers 7-8. 

Anterior biramous parapodia with trilobed noto- and neuropodia (Fig. 2c); 
rounded pre- and postsetal lobes, a small round intermediate lobe plus a rect- 
angular inferior lobe. Ventral cirri clavate in all setigers, decreasing in length 
posteriorly (Fig. 2c-e). Dorsal cirri finger-like, inserted on superior notopodial 
lobe. 

Posterior parapodia with bilobed noto- and neuropodia. Dorsal cirri becoming 
reduced; situated distally on elongated superior notopodial lobes (Fig. 2e). 

Notosetae all composite homogomph spinigers with denticulate appendages 



120 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

(Fig. 2f). Neuropodia include composite homogomph spinigers in superior posi- 
tions; composite heterogomph spinigers in intermediate positions; and thick and 
thin shafted heterogomph falcigers in intermediate and inferior positions. Superior 
heterogomph falcigers with shafts 2 times wider than inferior ones (Fig. 2g, h), 
each with 5-6 hairs on cutting margin of a short falcate appendage. 

Discussion.— Neanthes cortezi belongs to group IIB2c (Fauchald, 1972) to 
which the following species are assigned: N. noodti Hartmann-Schroder, N. 
seridentata Hartmann-Schroder, N. ruficeps (Ehlers) and N. pseudonoodti Fau- 
chald. Fauchald (1977) compared the latter 4 species. N. cortezi is most closely 
related to N. noodti and N. pseudonoodti in having paragnaths present in all 
pharyngeal regions. N. cortezi differs in having a patch of 14 cones in area V 
while the other 2 species each have a single cone. N. cortezi and N. noodti each 
have a single cone on area VI, but the paragnaths differ greatly in size; A/. 
pseudonoodti has 2 cones on area VI. 

A/, cortezi, A/, noodti and A/, pseudonoodti are also related ecologically and 
zoogeographically in that all are associated with barnacles in rocky intertidal 
habitats in the eastern Pacific. It is probable that A/, cortezi evolved from a 
widespread ancestor of both A/, noodti and N. pseudonoodti through isolation in 
the Gulf of California. 

Distribution. — Bahia Cholla, Puerto Pehasco, northern Gulf of California. 

Acknowledgments 
I am indebted to J. R. Hendrickson, University of Arizona, for allowing me to 
examine this collection, and to James A. Blake, Pacific Marine Station, for read- 
ing and commenting on this paper. This is An Allan Hancock Foundation Con- 
tribution No. 370 of Environmental Studies, Ministry for Conservation, Victoria. 

Literature Cited 

Banse, K., and K. Hobson. 1974. Benthic errantiate polychaetes of British Columbia and Washing- 
ton. Bull. Fish. Res. Bd. Canada, 185:1-111. 
Brusca, R. C. (ed.). 1973. A Handbook to the Common Intertidal Invertebrates of the Gulf of 

California. University of Arizona Press, Tucson, 427 pp. 
Berkeley, E., and C. Berkeley. 1948. Annelida. Polychaeta Errantia. Can. Pac. Fauna, 9b(l): 1-100. 
Chamberlin, R. V. 1918. Polychaetes from Montery Bay. Proc. Biol. Soc. Wash., 31:173-180. 
Fauchald, K. 1972. Benthic polychaetous annelids from deep water off western Mexico and adjacent 

areas in the eastern Pacific Ocean. Allan Hancock Monogr. Mar. Biol., 7:1-575. 
. 1977. Polychaetes from intertidal areas in Panama, with a review of previous shallow-water 

records. Smith. Contrib. Zool., 221:1-81. 
Hartman, O. 1968. Atlas of Errantiate Polychaetous Annelids from California. Allan Hancock 

Found., Univ. of So. Calif., Los Angeles, 828 pp. 
Johnson, H. P. 1901. The Polychaeta of the Puget Sound region. Proc. Boston Soc. Nat. Hist., 

29:381-437. 
Kudenov, J. D. 1975a. Errant polychaetes from the Gulf of California, Mexico. J. Nat. Hist., 9:65- 

91. 

. 1975b. Sedentary polychaetes from the Gulf of California, Mexico. J. Nat. Hist., 9:205-231. 

. 1975c. Two new species of errant polychaetes from the Gulf of California, Mexico. Bull. S. 

Calif. Acad. Sci., 74(2):75-80. 
. Annelida Polychaeta (Bristle worms). In A Handbook to the Common Intertidal Invertebrates 

of the Gulf of California. (R. C. Brusca, ed., 2nd edition), Univ. Arizona Press. In press. 
Malmgren, A. J. 1867. Annulata Polychaeta Spetsbergiae, Gronlandiae, Islandiae et Scandinaviae 

hactanus cognita. Ofvers. K. Vetensk. Akad. Forh., 24:127-235. 



NEW SPECIES AND RECORDS OF POLYCHAETOUS ANNELIDS 121 

Reimer, A. A. 1976a. Description of the Tetraclita stalactifera panamensis community on a rocky 
intertidal Pacific shore in Panama. Mar. Biol., 35:225-238. 

. 1976b. Succession of invertebrates in vacant tests of Tetraclita stalactifera panamensis. 

Mar. Biol., 35:239-251. 

Reish, D. J. 1968. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. II. 
Benthic polychaetous annelids. Trans. San Diego Soc. Nat. Hist., 15:67-106. 

Uschakov, P. V. 1972. (Polychaetes. Vol. I. Polychaetes of the suborder Phyllodociformia of the 
Polar Basin and the northwestern Part of the Pacific. Families Phyllodocidae, Alciopidae, 
Tomopteridae, Typhloscolecidae, and Lacydoniidae.) (Translated from the Russian by the 
Israel Program for Scientific Translations, 1974). Fauna SSSR, 102:1-272. 

Accepted for publication February 8, 1979. 



Bull. Southern California Acad. Sci. 
78(2), 1979, pp. 122-135 

A Survey of the Littoral and Sublittoral Ascidians of 
Southern California, Including the Channel Islands 

Rimmon C. Fay and James A. Vallee 1 

Abstract. — The ascidian populations of the eight Channel Islands, as well as 
three islets, the Cortez Bank and the Coronado Islands were surveyed. Thirty- 
eight species were found offshore compared to the 48 identified species which 
have been found above a depth of 60 m in southern California. It was observed 
that viviparous ascidians are specifically more abundant than oviparous ascidians 
on the Channel Islands and the possible significance of this mode of reproduction 
is discussed as an explanation for their local distribution. The habitat preference 
of southern California ascidians is presented. Nine species of ascidians of unre- 
solved identification are compared to the most closely related, described North 
American species. 

1 Pacific Bio-Marine Laboratories Inc., P.O. Box 536, Venice, California 90291. 



The ascidians of southern California have been surveyed by Ritter and Forsyth, 
1917; Van Name, 1945; and Fay and Johnson, 1971. Those reported by Ritter and 
Forsyth were collected intertidally, while those listed by Van Name were col- 
lected by dredge as well as intertidally. Those reported by Fay and Johnson, 
1971, were collected primarily with the aid of SCUBA, although some intertidal 
and trawl sampling was conducted. 

The present study is a continuation of Fay and Johnson's 1971 report on the 
ascidians of the mainland coast of southern California and extends the study area 
to the Channel and Coronado Islands. 

Little has been reported on the ascidians of the Channel Islands. Van Name, 
1945, summarizing information to that date, noted the occurrence of Trididemnum 
opacum, Pyura mirabilis, and Molgida pugetiensis at or near San Nicholas Island, 
and the occurrence of Clavelina hunts mani, Archidistoma psammion, Ritterella 
aequalisiphonis, Styela truncata, Molgida regularis, and Eugyra arenosa around 
Santa Cruz Island. He also reported that Halocynthia hdgendorfi igaboja was 
found around Santa Catalina Island. Given and Lees, 1967, reported that Eu- 
herdmania claviformis, Diplosoma macdonaldi, Metandrocarpa taylori, and 
Styela plicata also occur around Santa Catalina Island. With these few exceptions 
the distribution of ascidians of the Channel Islands is unreported in the literature 
and the present study is the first attempt to provide specific information about 
the ascidians of all the Channel Islands. To the best of the authors' knowledge 
this is the first comprehensive survey of any hard bottom invertebrate group in 
southern California including collecting sites on all of the Channel Islands. 

Methods 

Collection sites were usually visited using 18' and 24' dive boats, however, 
occasionally commercial dive boats were used. Trawling was conducted from the 
26' F/V PISCES using a 2 m beam trawl. Collection sites included the north, 



SURVEY OF SOUTHERN CALIFORNIA ASCIDIANS 123 




Fig. 1. Southern California Channel Island Collection Sites. Arrows around islands and rocks 
indicate collecting sites. 



east, south and west sides of each of the islands. In addition to the Channel 
Islands, sites on Richardson Rock, Wilson Rock, Begg Rock, the Coronado Is- 
lands and Cortez Bank were surveyed to a depth of about 30 m with the aid of 
SCUBA (Fig. 1). 

The period of study extended from 17 August 1972 to 11 February 1979. De- 
tailed records of the dates and locations of the collection sites are maintained at 
Pacific Bio-Marine Labs, Inc. 

Specimens collected were placed in 15% magnesium sulfate sea water along 
with a few crystals of menthol for 12 to 16 h to narcotize them. They were then 
transferred to 5% sea water formalin. Zooids were examined under a dissecting 
microscope and identifications were made using Van Name, 1945; Berrill and 
Abbott, 1949; and Abbott, 1975. 

Results and Discussion 

The mainland, island, and bank distribution of each species of ascidian col- 
lected is listed in Table 1. This table also indicates those species which brood 
their tadpoles (viviparous species) and those which release their gametes into the 
water (oviparous species). This distinction in mode of reproduction is noted be- 
cause the Channel Islands appear to have a higher percentage of viviparous 
species when compared with the ascidian biota of the mainland (Table 2). No 
oviparous species were found on the three rocks visited. This suggests that the 
short larval period of the viviparous ascidians is essential to the maintenance of 
these species at the isolated island collection sites where active current movement 
is observed. Viviparous ascidians usually produce tadpoles which are able to 
settle and metamorphose within minutes to a few hours after being released from 



124 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



pUBJUIEJM 

spuBjsi opeucuo3 

pU^lSJ 3JU3UI30 U^S 
pUBjSI BUIJBJB3 B1UBS 

pirejsi SB|oqoi{v[ ires 
>poy 8899 

pirejsi BcreoEuy 
pirejsi zriJ3 Bjires 

pUBJSI BSOy BlU^S 

pirejsi pnSijM ires 



iS O [J 

« a; pq 



Q Q .QM 

C/3 CO Q CO o 

00 to!' 

*j *-> „ *- On 

PQ PQ Q PQ PQ O 

C/3 CO Ph CO CO U 



X XX 

X 

X XXX 

X XX 



PL, (J 

J z 



tin 



Q Q Ph Q Q 



J Q 

Ph Q Q Q Q Q Q Q Q CO Q _ _ . _ 

i/)(n[/)WM«(/itrt qW ?JQ" ! *- )c/5 m -coco 

^oooooooo^o SchZo 0^00 

p LI< _.«_.. kJ *-. < -.. tJ *H._PL l< _. __ _ __ ._ ^ ._; .^ *_, 

OpQpQPQQpapQpQpQOpQ QfflfflCQQCQ CfflCQ 

UCOCOCOPhCOCOCOCOOcO PmCOCOCOPhCOciCOCO 



X X X X X 
X X X X X 

XXXXXXXX X XX 



XXX X X X X 



XX XXXX XX XXX X X XX X 



XX XXXX 
X 

X XXX 

X X X X X 

X X X X X X 

* 
* 

I^N ^i ^S N^s l"^>< 

XXX X 

X X 



XXX X XXXXX XX X 

XX X 

XX XXX X X XX X 

XXXXXXXXXXX X X 

XXXXX xxxxxxx X 

xxxxxxxxxxxxxx X 

XXXXXXXX X X X 
XXXX 



~ g ^J ^ * 



s g ^S 



§ » § £ c % e e ■« 'C ^ -a •& 






41 



■=> .2 a, k 



5 a t> 



-a o. 






~ 3 ^ 



?! 5 



5* %> .2 o, -2 a 



u a a a a -s 



""?: ^3 -3 ^3 S CJ t_i "-?* ^:"^3 rs ^ ^j 



^2 ^ rg 
^ ^ ^ 



~G Sj 



© o ?*, -~ 



t*J Oh «h ^ 



<3 ^ 



S O O ^ 2 <3 

w ""r* >> r* 71 *- 



a « g s 

<4j s a 
a ©, 



■2 <j 



a -5 



%j ^ s: ":? *) 5 



*-t3<3<30>5£ 



<*i <n 05 "5 --2 -— ~~ 
.*- ir- 1— 1— cj ~r- ^r - 



©, ?2 ?2 ?2 Is S 



.0, 

5 3 



-c -s -c > 



0000 



<o <J Q ^ ^ ^ ^ 



PS .3 -~ «5 
ft. Oh Oh ^ 



o !u ^ 5 c 



SURVEY OF SOUTHERN CALIFORNIA ASCIDIANS 



125 



pUB|UIBI\ 
SpUEJSI OpBUOJ03 

>juBg Z3JJ03 

pUBJSI 3JU3LU3IJ UB§ 

pUB|SJ BUJIBJB3 BJUBS 

PUB|SJ BJBqjBg biubs 

puBjsi SBjoqaiN ub§ 
>po^i SSag 

pUBJSI BdBDBUy 

puBjsi znj^ EJUB§ 

pUB|SJ BSOy B}UE<$ 

puB|S] pnSipv ubs 
Von uosjiav 

>poy uosp-imprg 



12 ^ ^ 

5 ° <= 

J5 02 QQ 







U 






,0 JQQQQQ 

PQ 0T) OO Oh OO OO C/3 C/3 00 




pq" 

00 CL, 




Q 

OO Q 
o 00 


^ o o 222222 




PQ 2 J 




cl, m 2 cq cq 


qQcqcqqQpqcqoqcqcq 

>(/)Cfl&<OH[/](/5[/5(/]ZI 


Q 
00 


S CQ J 
OO 00 CU 


< 


U 00 OO OO OO 



X 



X X 

X 

X 

X 



X X 



X X 



25 s 



3 -2 

-2 3' 



X 



X 



X X 

X XX 



X XX 



X 



X X 



X X 



X 



X X 

XX XX XX 

X X X X X X 



X X XXX 

X 



<£ 



■S _^ W O Q 



-2 -2 25 2i 



£ :3 -2 ~ * 







# 




* 


-2 


* 


* 


# 

2 


a 


* 


-2 


•2 


_3 


* 


t; 


C 




V 


-2 




3 


2 


2 




~ 




Sc 


O, 


ij 




5 




a 



-3 -2 c Z 

2= h > « 



5e 5? c 



2 c -e 5 a 






-2-2-2-2-2-2 a 3 



2 ^2? -S 2 



Qj ^ ^ ^ 



su 5" S cs ~^ -^ ^ ^ ^ ^ 



^CQCQ^^^^^^^^^^' ^'^CQQ3 





7i 




3- 


it 


E 


.-3 
— 


o 


'S 

c 


<u 


G 


a 


oc 


CQ 


J 


CQ 


<u 


2 


U 


,,_, 


a 


b 


a 


u 


c 


c 


C 


c 


'S 


'3 


si 


cd 


s) 


qj 


CL, 00 OO OO > 








CQ 




- CQ 


QS2 


Oh OO OO OO > 



c 
o 

SK-a 5 > 

z? c ™ o U 

a- « u a 

s s z c2 £ 






E 


Bay 
Cove 
Point 

Point 


E 


o 


c 
o 

CO 


lamitos 
balone 
oal Oil 
a Jolla 
echuza 


u 

Q. 


< < U -J -J 


>,~j 


Mill 


Q. U 




ctf <" 






CL 




DQUO-,o. 


u o 


<<UJJ 



■- > 3 « 

> o as -c 



12 6 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 2. 


The number and percentage of oviparous and viviparous species. 




Viviparous Oviparous 




# % # % 



6 


100 


16 


100 


22 


85 


20 


80 


20 


83 


15 


94 


17 


85 


4 


100 


21 


81 


15 


75 


18 


86 


9 


82 


11 


92 















4 


15 


5 


20 


4 


17 


1 


6 


3 


15 








5 


19 


5 


25 


3 


14 


2 


18 


1 


8 



All of Southern California 32 67 16 33 

Island, Islet or Bank 
Richardson Rock 
Wilson Rock 
San Miguel 
Santa Rosa 
Santa Cruz 
Anacapa 
Santa Barbara 
Begg Rock 
San Nicholas 
Santa Catalina 
San Clemente 
Cortez Bank 
Coronado 



the parent. This greatly reduces the chances of their being swept away from 
suitable habitats by currents. The eggs and larvae of oviparous ascidians on the 
other hand, must develop in the plankton for a day or more before they are able 
to settle and metamorphose; during this time they would be subject to transport 
away from isolated habitats thereby making it difficult to maintain stable popu- 
lations of oviparous species at small isolated sites. The channel rocks are ex- 
amples of this circumstance. 

The very short larval period that characterizes most of the ascidian populations 
of the Channel Islands and rocks, as well as their isolation, suggest that these 
populations are self dependent for their larval recruitment and therefore are en- 
demic. 

Nine species of ascidians known to occur along the mainland of southern Cal- 
ifornia were not found on the Channel Islands by the authors, five of these are 
oviparous. Seven species, all viviparous, were found on all of the Channel Islands 
(Table 1). These observations again suggest that viviparity is of advantage at the 
island habitats. 

Some Channel Island ascidians seem to be quite restricted in their distribution. 
Ten species have been collected on only one or two of the islands by the present 
authors (Table 1). It should be pointed out that Styela coriacea, Halocynthia 
hilgendorfi igaboja, and Bathypera ovoida from Santa Catalina Island were col- 
lected with a beam trawl which was not used around the other islands. 

Since the publication of Fay and Johnson in 1971, the present authors have 
gathered additional information on the distribution of ascidians on the mainland 
coast of southern California. This is summarized in Table 1. Noteworthy additions 
include the occurrence of Pyura mirabilis at Point Loma, Lechuza Point, Mission 
Bay and Paradise Cove; Molgula regularis (=M. verrucifera, in Fay and Johnson, 
1971) from Santa Barbara to San Diego; Molgula pugetiensis and Bathypera 
ovoida in Santa Monica Bay; Polyzoa translucida (= unidentified Styelid in Fay 



RVEY OF SOUTHERN CALIFORNIA ASCIDIANS 127 

i Johnson, 1971) on the Venice breakwater, off San Pedro, and in Alamitos 
y; Corella willmeriana, three miles south of Point Fermin; and Ascidia ver- 
"ormis at Point Dume. Recently (February, 1977) Styela partita has been col- 
ted in San Diego Bay (C. Lambert, pers. comm.). 

t should be noted that several specimens of Mogula regularis were observed 
have tadpoles in their atrial chambers. This is of importance because Van 
me (1945) indicated that a closely related ascidian, Mogula cooperi (Hunts- 
n, 1912) from off British Columbia is distinguished from M. regularis (Ritter, 
.7) only because M. cooperi broods its tadpoles while M. regularis does not. 
e brooding of tadpoles by M. regularis reported here suggests that M. cooperi 
>uld be synonymized with M. regularis. 

During this and earlier work, it was subjectively observed that many of the 
idians were associated with protected waters or vigorous wave and current 
ion. For example, Styela plicata, S. clava, and Ciona intestinalis are found 
he calm waters of protected bays while Clavelina huntasmani is found at sites 
)osed to vigorous currents or wave action. Table 3 lists the ascidians of south- 
California subjectively according to the relative vigor of water movement in 
habitats where they are found. Some of them, such as Distaplia occidentalis 
1 Pyura haustor were found in all three types of circulation: calm water, mod- 
te exposure to currents and wave action, and exposure to vigorous currents 
1/or wave action. 

t may be noted, except for the ascidians discussed in the introduction, the 
tribution of Channel Island ascidians reported in the present study all consti- 
z range extensions. The following are of particular interest. Polyclinum laxum, 
ich was previously reported only from the Gulf of California by Van Name in 
•5, is here reported from Santa Barbara and San Nicholas Islands. The previous 
ithernmost occurrence of Archidistoma /nolle was Pacific Grove (Van Name, 
■5). In the present study it was found at Santa Rosa and San Miguel Islands, 
n Name also noted that only one specimen of Pyura mirabilis had been col- 
ted off southern California. It is here reported from Santa Rosa Island, Santa 
iz Island and Cortez Bank, as well as on the mainland coast. Van Name also 
'e only one collection site for Bathypera ovoida (1000 fathoms, 22 miles south 
San Nicholas Island). It is here reported to be rather common at Santa Catalina 
md. Corella willmeriana was reported by Van Name to occur as far south as 
day Harbor, Washington. It is apparently also found off central California 
ce it is listed in Abbott's key to the ascidians of central California in Light's 
nual (Abbott, 1975); this report extends its range to 3 mi south of Point Fermin, 
s Angeles County. 

n addition to the 48 species of southern California ascidians listed above, we 
it collected nine species which cannot be identified as species of ascidians 
)wn from southern California. 

iO describe and name these species is a task beyond the scope of the present 
dy, however, they may be characterized as follows: 

Aplidium sp. A 

rhe colony consists of sand-covered, club-shaped heads similar to Aplidium 
<pinquum and A. arenatum. The branchial sac has 8 to 13 rows of stigmata 
npared to 5 rows in A. arenatum and up to 21 in A. propinquum. The stomach 



128 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 3. Water surge association of southern California ascidians. 



Species 



Calm Water 



Moderate Water Vigorous Water 
Movement Movement 



Aplidium californicum 
Aplidium solidum 
Aplidium propinquum 
Aplidium arenatum 
Euherdmania claviformis 
Polyclinum planum 
Polyclinum laxum 
Synoicum parfustis 
Didemnum carnulentum 
Trididemnum opacum 
Diplosoma macdonaldi 
Lissoclinum caulleryi 
Clavelina huntsmani 
Cystodytes lobatus 
Archidistoma diaphanes 
Archidistoma psammion 
Archidistoma ritteri 
Archidistoma molle 
Distaplia occidentalis 
Pycnoclavella stanleyi 
Ritterella aequalisiphonis 
Ritterella pulchra 
Ascidia ceratodes 
Ascidia vermiformis 
Chelyosoma productum 
Corella willmeriana 
Ciona intestinalis 
Perophora annectens 
Polyzoa translucida 
Botrylloides diegense 
Metandrocarpa dura 
Metandrocarpa taylori 
Styela plicata 
Styela clava 
Styela truncata 
Styela gibbsii 
Styela coriacea 
Styela partita 
Styela montereyensis 
Botryllus tuberatus 
Pyura haustor 
Pyura mirabilis 

Halocynthia hilgendorfi igaboja 
Boltenia villosa 
Bathypera ovoida 
Molgula regularis 
Molgula pugetiensis 
Eugyra arenosa 



X 



X 



X 



SURVEY OF SOUTHERN CALIFORNIA ASCIDIANS 129 

usually has five to eight folds compared to five folds for A. arenatum and 18 to 
21 for A. propinquum. Apildium sp. has been found at San Nicholas, San Cle- 
mente, Santa Catalina, San Miguel, and Anacapa Islands, at Wilson Rock, and 
off Nicholas Canyon, Los Angeles County. 

Rhopalaea sp. A 

This species attains a height of 25 mm and has about 80 rows of stigmata. The 
arrangement of the muscle bands in the mantle differs from that reported for R. 
abdominalis by Van Name (1945) and for R. birkelandi by Tokioda (1971), both 
from American waters. The species has been taken in Santa Monica Bay by trawl 
and by SCUBA at Point Dume. 

Trididemnum sp. A 

This species is very similar to T. opacum. It differs in that there are so few 
spicules that the test is very soft and translucent as compared to the tough, 
opaque test of T. opacum. Trididemnum sp. has been found at Santa Rosa and 
San Miguel Islands, Begg Rock, and off Nicholas Canyon, Los Angeles County. 

Botryllus sp. A 

This species is similar to B. tuberatus but the branchial sac has eight rows of 
stigmata instead of four. The systems are round or oval and usually consist of 6 
to 14 zooids. Botryllus sp. occurs in Mission Bay on floats and on Styela clava. 

Botrylloides sp. A 

The zooids of this species appear to be very similar to those of Botrylloides 
diegense, however, the tadpoles are distinctively different. The body of B. die- 
gense tadpoles is about 1.2 mm long and has about 32 ampullae, while the body 
of Botrylloides sp. tadpoles is about 0.6 mm long and has eight ampullae. The 
colony tends to mass and fold in a golden yellow complex larger than B. diegense. 
Botrylloides sp. has been found at San Nicholas, Santa Cruz, Santa Rosa, and 
San Miguel Islands. 

Ascidia sp. A 

This species is similar to A. ceratodes. However, unlike A. ceratodes, the 
present species has a thick, opaque, cartilaginous test, and is not laterally flat- 
tened. Five specimens 4.0 to 4.5 cm tall were collected in San Diego Bay. 

Styela sp. A (probably partita) 

One individual has been collected from Los Angeles Harbor, and a few from 
San Diego Bay. 

Styela sp. B 

This is a very contractile form. When fully extended the internal body struc- 
tures, especially the two pairs of gonads, are similar to S. gibbsii. However, 
externally the posterior portion of the test is covered with a dense mat of branch- 
ing papillae, similar to Styela hemicaespitosa described by Ritter in 1913, and 
synonymized with Styela coriacea by Hartmeyer in 1923. However, the present 
species is distinct from S. coriacea, which has only one pair of gonads. Styela 



130 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

hemicaespitosa should probably be re-established for it. This species has been 
trawled from Santa Monica Bay in 24-35 fathoms. 

Polyclinum sp. A 

This is an encrusting species of Polyclinum. The colony examined was about 
45 cm long, 30 cm wide and 2-3 cm thick, with a smooth upper surface lightly 
coated with sand. It was collected from San Nicholas Island, where it was found 
living under a rocky ledge at a depth of 35 ft. 

In addition to the above species, there is also a form of Polyclinum that is egg- 
shaped and has a slender stalk. This form of Polyclinum has been taken several 
times in the Point Dume area. It is here included as a form of Polyclinum planum, 
which is usually laterally flattened, and has a thick peduncle. 

During the course of this and previous studies it was recognized that there is 
a need for a key to the Southern California ascidians. Therefore the following 
artificial key was prepared. It is based upon preserved material and zooid char- 
acteristics are used extensively, thus a dissecting microscope will be required. 
Zooid size, gonad description and the characteristic number of rows of stigmata 
described in the following key refer to adult zooids only. 

A Preliminary Artificial Key to the Ascidians 
Found in Southern California 

1 . Simple ascidians 33 

- Colonial ascidians 2 

2. Body entire, not divided into 2 or 3 regions 26 

- Body divided into 2 regions 14 

-. Body divided into 3 regions 3 

3. Atrial languet present 7 

- Atrial languet absent 4 

4. Each zooid with its own test (although sharing a common base) 5 

- Zooids embedded in a common test, colony composed of lobes or club 
shaped heads 6 

5. Twelve or thirteen rows of stigmata, zooids up to 35 mm tall, color in 
life, translucent grey to slightly greenish Either dmania claviformis 

- Seven rows of stigmata, zooids up to 30 mm tall, color in life, golden 
yellow Pycnoclavella stanleyi 

6. Lobes incrusted with sand, and up to 20 mm tall, color in life, transparent 
encrusted with sand Ritterella aequalisiphonis 

- Lobes incrusted with sand, up to 25 mm in height, color in life, trans- 
parent test with red zooids Ritterella pulchra 

7. Colony composed of distinct lobes or club shaped heads 8 

- Colony surface rather even, not composed of distinct lobes or club 
shaped heads 11 

8. Branchial sac with 5 rows of stigmata, lobes of the colony up to 25 mm 
tall, sand encrusted Aplidium arenatum 

- Branchial sac with 8 or more rows of stigmata 9 

9. Stomach smooth, without distinct longitudinal folds (except for typhlo- 
sole), lobes of the colony up to 11 cm tall, color in life, salmon 

Synoicum parfustis 



SURVEY OF SOUTHERN CALIFORNIA ASCIDIANS 131 

- Stomach with 5 to about 20 distinct longitudinal folds 10 

10. Branchial sac with 8 to 13 rows of stigmata, lobes of the colony up to 35 
mm tall, sand encrusted Aplidium sp. 

- Branchial sac with 16 to 21 rows of stigmata, lobes of the colony up to 

5 cm in height, sand encrusted Aplidium propinqaum. 

11. Entire colony supported by a distinct peduncle, colony may be up to 20 
cm tall, color in life, red wine to yellow Polyclinum planum 

- Entire colony attached by a broad base, not supported by a distinct 
peduncle 12 

12. Stomach wall with many (8-23) longitudinal folds 13 

- Stomach wall smooth, without many longitudinal folds, colony up to 8 

cm across, mahogany colored in life Polyclinum laxum 

13. Eight to twelve rows of stigmata, colony up to 20 cm across, Vi to 3 cm 
thick, color in life, white to salmon pink Aplidium californicum 

- Thirteen to fifteen rows of stigmata, colony up to 16 cm across and IVi 
cm thick, color in life, white Aplidium solid urn 

14. Each zooid with its own test (although sharing a common base) 15 

- Zooids embedded in a common test 16 

15. Zooids with about 80 rows of stigmata, zooids (including test) up to 35 
mm tall, color in life, jade green Rhopalaea sp. 

- Zooids with 16 to 20 rows of stigmata, zooids (including test) up to 40 

mm tall, color in life, white or vivid pink Clavelina huntsmani 

Atrial siphon present and tube-like 17 

Atrial siphon absent, atrial aperture an opening on the dorsal surface . . 23 

Spicules present in the test 18 

No spicules present in the test 20 

Spicules disk-shaped, or occasionally in the form of amorphous calcar- 
eous deposits, colony may be 25 cm or more across and up to 2 cm 

thick, color in life, orange-tan, whitish or grayish Cystodytes lobatus 

Spicules stellate 19 

Surface of test completely opaque due to the abundance of spicules, 
zooids not visible through the test, colony up to 15 cm across and 4 mm 

thick, flesh color in life Trididemnum opacum 

Surface of test transluscent due to the scarcity of the spicules, zooids 
clearly visible through the test, colony up to 8 cm across and 3 mm thick, 

color in life, brown Trididemnum sp. 

Test tough and hard, upper surface even, without lobes, colony up to 15 

mm thick, color in life, burgundy Archidistoma psammion 

Test soft with even surface, or with projecting lobes 21 

Colony soft, with an even surface, without projecting lobes 22 

Colony with projecting lobes or elongate heads which may be up to 40 

mm tall, color in life, turquoise Archidistoma ritteri 

Zooids average about 3 mm long or less, colony about 1 cm thick color 

in life, white to pale vermillion Archidistoma diaphanes 

Zooids 5 to 8 mm long, colony about 2 cm thich, color in life, light 

grey Archidistoma molle 

Minute spicules present in the test 24 

Minute spicules absent 25 



13 2 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

24. Atrial aperture a plain round opening restricted to the dorsal surface, 
with no languet, colony up to 4 mm thick, color in life, white to orange 

Didemnum carnulentum 

- Atrial aperture very large, extending around to the sides of the branchial 
sac, atrial languet present, colony about 3 mm thick, color in life, grey 

Lissoclinum caulleryi 

25. Atrial aperture with a languet, each row of stigmata crossed by a trans- 
verse vessel, colony flat and about 1 cm thick or club shaped and up to 
3 1 /2 cm tall, color in life, variable: white, grey, green, brown, and 
mixed Distaplia occidentalis 

- Atrial aperture without a languet, the rows of stigmata not crossed by 
a transverse vessel, colony about 2 mm thick, gelatinous, color in life, 

tan Diplosoma macdonaldi 

26. Four to 8 rows of stigmata 27 

- Nine to 13 rows of stigmata 29 

27. Each zooid with its own test or zooids fused, but not arranged in systems, 
atrial siphon tube-like, and opening directly to the outside, maximum 
height (including test) about 3 mm, color in life, emerald green 

Perophora annectens 

- Zooids arranged in systems, atrial aperture opening into a common cloa- 
ca 28 

28. Four rows of stigmata, colony up to 2 mm thick, color in life, dark grey 

Botryllus tuberatus 

- Eight rows of stigmata, colony up to about 2 mm thick, colorless, has 
the appearance of fish eggs Botryllus sp. 

29. Zooids in systems, atrial aperture provided with a languet and opening 
into a common cloaca, colony up to about 5 mm thick 30 

- Zooids not in systems, atrial aperture without a languet, and opening 
directly to the outside 31 

30. Tadpoles with about 32 small papillae surrounding the anterior end of 
the body, color in life, orange, purplish Botrylloides diegense 

- Tadpoles with 8 large papillae surrounding the anterior end of the body, 
color in life, golden yellow Botrylloides sp. 

31. Branchial sac with 3 longitudinal vessels, zooids up to 6 mm in height, 
color in life, grey Polyzoa translucida 

- Branchial sac with 5 longitudinal vessels 32 

32. Zooids close together in a common test, colony up to about 4 mm thick, 
color in life, crimson Metandrocarpa dura 

- Zooids separated, each with its own test, but interconnected with at least 
a film of test, zooids up to 5 mm tall, color in life, crimson 

Metandrocarpa taylori 

33. Branchial sac with internal longitudinal folds 39 

- Branchial sac flat, without internal longitudinal folds 34 

34. Anterior end of the test a flattened disk provided with thin horny plates, 
reaches a maximum height of 50 mm, color in life, jade green, yellowish 

or brownish Chelyosoma productum 

- Anterior end not flattened nor provided with thin horny plates 35 

35. Ascidian resembling a ball of mud, stigmata arranged in perfect double 



SURVEY OF SOUTHERN CALIFORNIA ASCIDIANS 133 

spirals, maximum test diameter about 15 mm color in life, muddy grey 
Eugyra arenosa 

- Ascidian not resembling a ball of mud, stigmata not arranged in perfect 
double spirals 36 

36. Branchial sac extends posteriorly beyond the stomach in a long narrow 
pouch which is as long or longer than the distance from the stomach to 
the branchial siphon, maximum test length 14 cm, appearance in life, 
transluscent, tinged with green Ascidia vermiformis 

- Branchial sac does not extend posteriorly in a long narrow pouch 37 

37. Stigmata spiral, stomach on the right side of the body, intestine curves 
ventrally under the stomach, maximum length of the test 45 mm, color 

in life, jade green to yellowish Corella willmeriana 

- Stigmata straight, stomach on the left side of the body, intestine curves 
dorsally over the stomach 38 

38. Atrial siphon located on the dorsal side near, or posterior to the middle 

of the body 39 

- Atrial siphon located near the anterior end of the body, maximum test 
length 25 cm usually 10 cm or less color in life, yellowish green 

Ciona intestinalis 

39. Test transparent or transluscent, flexible, laterally flattened, test up to 

50 mm long, color in life, jade green Ascidia ceratodes 

- Test opaque, rigid, cartilaginous, not laterally flattened, up to 45 mm 
long, color in life, jade green Ascidia sp. 

40. Large tentacles branched 41 

- Tentacles always simple 47 

41. A closed renal sac on the right side of the body 42 

- No renal sac present on the right side of the body 43 

42. An "S" shaped ovary on each side of the body, maximum test diameter 

4 cm, usually 15 mm or less sand encrusted Molgula regularis 

- An oblong ovary on each side of the body, maximum test length 15 mm, 
sand encrusted Molgula pugetiensis 

43. Test provided with precisely intersecting rows of small papillae, maxi- 
mum diameter 4 cm, usually 2 cm or less, color in life, pearly white . . . 

Bathypera ovoida 

- Test not provided with precisely intersecting rows of small papillae (but 
may be provided with spines exhibiting no precise pattern) 44 

44. Body of test provided with spines, and usually relatively free of encrust- 
ing debris or organisms 45 

- Body of test lacking spines (although minute spines may be present on 
the siphons), test may or may not be covered with debris or encrusting 
organisms 46 

45. Body supported by a stalk, total length of the test up to 9 cm, color in 
life, dull rose red Boltenia villosa 

- Body attached directly by the posterior end, no stalk present, up to 10 
cm in diameter, but usually 3 cm or less, color in life, reddish about the 
apertures, otherwise brown Halocynthia hilgendorfi igaboja 

46. Siphons located at opposite ends of an elongate body and directed in 
nearly opposite directions, test free of encrusting debris or organisms. 



134 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

up to 65 mm long, color in life, dull white Pyura mirabilis 

- Both siphons directed upwards, test usually encrusted with debris or 
organisms, up to 120 mm long, color in life, rose red at ends of siphons 

Pyura haustor 

47. Body supported by a narrow stalk, the upper part of which is hollow and 
contains a tubular prolongation of the mantle 48 

- Body attached directly by the posterior end, no hollow narrow stalk 
present 49 

48. Conspicuous tubercles anteriorly, longitudinal folds of the test restricted 
to the posterior body and stalk, 4 or more ovaries on the right side of 
the body, test up to 20 cm tall, color in life, rose brown to brown 

Styela clava 

- Tubercles few and inconspicuous, longitudinal folds extend anteriorly 
nearly to the siphons, 3 or fewer ovaries on the right side of the body, 
test up to 30 cm tall, color in life, tan to reddish brown 

Styela montereyensis 

49. Branchial sac with only one internal longitudinal vessel between the 
branchial folds, test up to 30 mm tall, color in life, reddish brown 

Styela truncata 

- Branchial sac with 2 or more internal longitudinal vessels between the 
branchial folds 50 

50. Two ovaries on each side of the body 51 

- Other than 2 ovaries on each side of the body 53 

51. Posterior region of the test provided with slender branching papillae 
which are usually obscured by silt, test up to 25 mm in height, color in 
life, greyish yellow Styela sp. 

- Test not provided with such papillae 52 

52. Ovaries clearly sinuously curved, test up to 30 mm tall, color in life, 
yellow posteriorly, purplish anteriorly Styela partita 

- Ovaries only slightly sinuously curved, nearly straight, test up to 40 mm 
tall, color in life, reddish brown Styela gibbsii 

53. One ovary on the right side of the body, test up to 20 mm long, color in 
life, reddish brown Styela coriacea 

- At least three ovaries on the right side of the body, test up to 10 cm tall, 
color in life, white and brown Styela plicata 

Acknowledgment 

The authors would like to thank Dr. Donald Abbott for his interest and en- 
couragement during the course of this study and for his preliminary review of this 
report. Thanks are also due to Richard C. Rhode for his aid in collecting 
specimens of ascidians. 

Literature Cited 

Abbott, D. P. 1975. Phylum Chordata: Introduction and Urochordata. Pp. 638-655. In Lights Man- 
ual, Intertidal Invertebrates of the Central California Coast. University of California Press, 
Los Angeles, iii + 685 pp. 

Berrill, N. J., and D. P. Abbott. 1949. The morphology and larvae of the ascidian Pycnoclavella 
stanleyi, n. sp. Can. Jour. Res. d, 27:43-49. 

Fay, R. C, and J. V. Johnson. 1971. Observation on the distribution and ecology of the littoral 



SURVEY OF SOUTHERN CALIFORNIA ASCIDIANS 135 

ascidians of the mainland coast of southern California. Bull. So. Cal. Acad. Sci., 70(3): 114- 

124. 
Given, R. R., and D. C. Lees. 1967. Santa Catalina Island biological survey. Survey report no. 1. 

Allan Hancock Foundation, University of Southern California, 1-126. 
Hartmeyer, R. 1923. Ascidiacea, Part I. In Danish Ingolf Expedition. Copenhagen, 2(6): 1-368. 
Ritter, W. E. 1913. The simple ascidians from the northeastern Pacific in the collection of the United 

States National Museum. Proc. U.S. Nat. Mus., 45:439-512, pis. 38^6. 
and R. A. Forsyth. 1917. Ascidians of the littoral zone of southern California. Univ. Calif. 

Publ. Zool. 16:439-512. 
Tokioka, T. 1971. A new species of Rhopalaea from the Pacific coast of Costa Rica (Tunicata, 

Ascidiacea). Publ. Seto Mar. Biol. Lab. 19:119-122. 
Van Name, W. G. 1945. The North and South American Ascidians. Bull. Amer. Mus. Nat. Hist., 

84:1^176, pis. 1-31. 

Accepted for publication January 31, 1979. 



Bull. Southern California Acad. Sci. 
78(2), 1979, pp. 136-140 

A Redescription of Pettiboneia sanmatiensis Orensanz 

(Polychaeta: Dorvilleidae) and a Revised Key to the 

Genera of the Dorvilleidae 

James A. Blake 

Abstract. — Pettiboneia sanmatiensis Orensanz originally described from Ar- 
gentina, has been discovered in California and British Columbia. The new spec- 
imens are compared with the type collection and the species is redescribed. The 
jaw apparatus consists of seven pairs of denticle rows and is unique to the family 
Dorvilleidae. A new key to the genera of the Dorvilleidae is provided which 
includes Pettiboneia, but excludes Apophryotrocha Jumars which has been found 
to be a juvenile onuphid and not a dorvilleid. 

Tomales Bay Marine Laboratory, Marshall, California 94940. 



During the course of intertidal surveys in the White Gulch area of Tomales 
Bay, California, a small meiofaunal dorvilleid was discovered which appeared to 
be identical to Pettiboneia sanmatiensis Orensanz, 1973, from Argentina. The 
species had not been reported since its original description. The identification 
was confirmed following a comparison of the Tomales Bay specimens with the 
type collection from Argentina. Two additional specimens from British Columbia 
also proved to be the same species. Study of these materials has revealed that P. 
sanmatiensis possesses a unique jaw apparatus, unlike any other known for the 
Dorvilleidae. A complete redescription of the species is provided herein along 
with a revised key to the genera of the Dorvilleidae. 

I am grateful to Prof. Elena Martinez Fontes of the Museo Argentino de Cien- 
cias Naturales (MACN), Buenos Aires for the loan of the type collection and to 
the late Katharine D. Hobson for the specimens from British Columbia. The 
Tomales Bay specimens were collected by Debrah L. Armitage and are deposited 
in the National Museum of Natural History, Smithsonian Institution (USNM). 

Pettiboneia Orensanz, 1973 

Type-species. — Pettiboneia sanmatiensis Orensanz, 1973, by original designa- 
tion. 

Diagnosis. — Palps well developed, biarticulate; antennae simple, clavate, dor- 
sal cirri small, elongated, bearing notoaciculum; ventral cirrus present; setae 
including serrated capillaries, furcate and composite setae; jaws including a pair 
of mandibles and seven pairs of denticle rows, carriers lacking. 

Remarks. — Pettiboneia is monotypic and appears to be most closely related to 
Schistomeringos Jumars (=Stauronereis Verrill) in having an elongated dorsal 
cirrus, a notoaciculum and furcate setae. Pettiboneia, however, has small, simple 
antennae and 14 denticle rows while Schistomeringos has long, multiarticulate 
antennae and the typical dorvilleid arrangement of four denticle rows. Jumars 
(1974) and Oug (1978) have reported the presence of extra denticle rows in some 



REDESCRIPTION OF PETTIBONEIA SANMATIENSIS 



137 




DEF G 



Fig. 1. Pettiboneia sanmatiensis: A. Anterior end in dorsal view; B, Middle parapodium in anterior 
view, dorsal cirrus to the right, ventral cirrus to the left; C. Furcate seta; D-F. Composite setae; E. 
Serrated capillary seta. 



of their species of Schistomeringos. These authors consider them to be replace- 
ment maxillae since they are located directly under the ordinary set. The extra 
10 denticle rows found in Pettiboneia, on the other hand, are located well forward 
of the first four rows and show no evidence of being superimposed by them. 

Pettiboneia sanmatiensis Orensanz, 1973 
Figs. 1-2 

Pettiboneia sanmatiensis Orensanz, 1973: 338-339, pi. 5, figs. 1-8. 

Material examined.— ARGENTINA: N.W. Golfo San Matias, 15-16 m, sand 
and gravel, Campana exploratoria, SAO- 1/71 Sta. 22 and 24, SAO-V/73. Sta. 228. 
five paratypes and holotype (Type series number, MACN 27818), one paratype 
on two slides (USNM 56509). CALIFORNIA: Tomales Bay, White Gulch, in- 
tertidal, sand-mud, 18 March 1974, coll. D. L. Armitage, five specimens (USNM 
56498), three specimens on five slides (USNM 56510-1). BRITISH COLUMBIA: 
Chemainus Bay, subtidal, 11-12 m, sand-gravel, 30 September 1974, coll. K. D. 
Hobson, two specimens (USNM 56498). 



138 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. Pettiboneia sanmatiensis: A. Mandibles; B. Maxillary apparatus consisting of seven pairs 
of denticle rows. Left side mostly omitted. Each pair denoted by numerals I-VII. 



Description. — A small, slender, thread-like species, measuring up to 7.0 mm 
long and 0.4 mm wide for 55 setigers. Color in life: tan with brown granular spots 
scattered over cuticle; color in alcohol: opaque white. 

Prostomium pear-shaped, broadly rounded on anterior margin, bluntly trun- | 
cated on posterior margin; one pair of black granular eyes located at bases of 
antennae (Fig. 1A); antennae small, simple, clavate; palps well developed, biar- 
ticulate. Segments 1-2 achaetous; setiger 1 lacking dorsal cirrus, this beginning 
on setiger 2, small at first, increasing to full size by setiger 4 (Fig. 1A-B); dorsal 
cirri elongated, each bearing a thin notoaciculum; ventral cirri short, stubby; 
neurosetae including a thick neuroaciculum, sometimes protruding through cu- 
ticle, a single furcate seta (Fig. 1C), a single long serrated capillary seta (Fig. 
1G), and 3-4 heterogomph composite setae grading in a series from short falcigers 
to long spinigers (Fig. 1D-F); each composite seta with finely serrated cutting 
edge on blade, small spinelets on end of shaft and a thin membrane connecting 
shaft and blade. Body terminating in four short anal cirri, each weakly clavate. 

Jaw apparatus consisting of a pair of mandibles (Fig. 2A) and seven pairs of S 
denticle rows (Fig. 2B); carriers lacking. Basal denticles of pair I not fused, 



REDESCRIPTION OFPETTIBONEIA SANMATIENSIS 139 

narrow pointed with a single tooth gradually developing over subsequent denti- 
cles, distal denticles with smaller spines and spinelets; pair II with 6-1 1 denticles, 
each covered with numerous spinelets; pairs III— VII with numerous spinelets on 
denticles; pair I: dark brown; pairs II— VII: light tan. 

Ecology. — Pettiboneia sanmatiensis occurs in coarse intertidal sands in To- 
males Bay and is associated with dense populations of Pygospio elegans Cla- 
parede. In life, the specimens coil in a corkscrew fashion when removed from 
the sand. This behavior is similar to that of some archiannelids and demonstrates 
their dependence upon the sand grains for adhesion. In British Columbia this 
species was taken on a sand-gravel bottom in 1 1-12 m and in Argentina was taken 
on a sand-gravel bottom in 15-16 m. 

Distribution. — Argentina, California, British Columbia. 

Remarks. — The systematics of the genera of the Dorvilleidae have been con- 
fused in the past owing to several problems of nomenclature and varying inter- 
pretation of the weight of taxonomic characters used by different workers. As a 
result of revisions by Pettibone (1961), Fauchald (1970) and Jumars (1974) most 
of these problems would appear to be resolved. Oug (1978), however, has de- 
scribed new species of Schistomeringos and Ophryotrocha which differ from 
Jumars' (1974) definitions, and may eventually require further revisionary work 
on the genera. Until such time, the generic arrangement proposed by Jumars is, 
with one exception, accepted. An examintion of the holotype (AHF Poly 1081) 
and paratype (AHF Poly 1082) of Ap ophryotrocha mutabiliseta Jumars, 1974, 
has revealed that the species is actually a post-larval onuphid and not a dorvilleid. 
These juveniles possess five occipital tentacles or their scars and are similar in 
many respects to the larvae of Nothria elegans Johnson as described by Blake 
(1975). Pettiboneia was not included in the generic keys provided by Jumars 
(1974) and Fauchald (1977). A revised key to the genera of the Dorvilleidae is 
presented (see below). 

No differences have been detected between the specimens of Pettiboneia san- 
matiensis from Argentina, California and British Columbia. The first two pairs of 
denticle rows (I — II) approximate those observed in most other dorvilleid genera, 
except that maxillary carriers are lacking. In each of the slide preparations, the 
basal denticles of the first two pairs have separated easily, suggesting that there 
is no fusion of these parts. Denticle rows III— VII are more distally positioned and 
widely separated basally. The possible function of such a large number of denticle 
rows in P. sanmatiensis is probably related to a meiofaunal mode of life. When 
protruded, these numerous denticles could provide a broad rasping surface to 
work on the coarse sand grains and gravel found in its habitat. Diatoms and other 
small microflora are probably removed by their action. 

The lack of meiofaunal collecting along the Pacific Coast of North and South 
America may account for the apparent disjunct range of P. sanmatiensis. Very 
fine mesh screens are necessary to retain this species, and despite a thorough 
survey of the White Gulch area of Tomales Bay by Dr. Ralph Johnson in the 
1960's (Pacific Marine Station, unpublished data), this species was not found. 
Very small cores, individually sorted alive were required before the species was 
discovered. It is probable that P. sanmatiensis will be found elsewhere when 
such methods are employed. 



140 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Key to the Genera of the Dorvilleidae 

1. Parapodia uniramous, without elongate dorsal cirri and notoacicula 4 

- Parapodia sub-biramous, with elongate dorsal cirri and enclosed notoac- 
iculum 2 

2. Furcate setae present 3 

- Furcate setae absent Dorvillea 

3. Antennae small, simple; jaw pieces with 14 denticle rows Pettiboneia 

- Antennae long, multiarticulate; jaw pieces with 4 denticle rows 

Schistomeringos 

4. Furcate or geniculate setae present 5 

- Furcate and geniculate setae absent 6 

5. Palps well developed, with at least one article; jaws with four denticle 
rows Protodorvillea 

- Palps very small, simple or biarticulate; jaws with two denticle rows . . . 

Meiodorvillea 

6. Setae all simple, acicular Parophyrotrocha 

- Setae include both simple capillaries and composite setae 7 

7. Setiger 1 modified, with two types of large specialized setae: (1) a simple 
recurved spine, (2) a large, thick, sharpely recurved composite falciger; 
antennae and palps biarticulate Exallopus 

- Setiger 1 not modified; antennae and palps reduced, papilliform 

Ophryotrocha 

Literature Cited 

Blake, J. A. 1975. The larval development of Polychaeta from the northern California coast. 

II. Nothria elegans (Family Onuphidae). Ophelia, 13:43-61. 
Fauchald, K. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitim- 

idae, Arabellidae, Lysaretidae and Dorvilleidae from western Mexico. Allan Hancock 

Monogr. Mar. Biol., 5:1-335. 
. 1977. The polychaete worms. Definitions and keys to the orders, families and genera. 

Natural History Museum, Los Angeles County, Sci. Ser., 28:1-190. 
Jumars, P. A. 1974. A generic revision of the Dorvilleidae (Polychaeta), with six new species 

from the deep North Pacific. Zool. Jour. Linn. Soc, 54:101-135. 
Orensanz, J. M. 1973. Los anelidos poliquetos de la provincia biogeographica Argentina. III. 

Dorvilleidae. Physis, Sec. A, 32:325-342. 
Oug, E. 1978. New and lesser known Dorvilleidae (Annelida, Polychaeta) from Scandinavian 

and northeast American waters. Sarsia, 63:285-303. 
Pettibone, M. L. 1961. New species of polychaete worms from the Atlantic Ocean, with a 

revision of the Dorvilleidae. Proc. Biol. Soc. Wash., 74:167-186. 

Accepted for publication March 22, 1979. 



RESEARCH NOTES 141 

Research Notes 

Bull. Southern California Acad. Sci. 
78(2), 1979, pp. 141-143 

New Mammalian Records from the Late Pleistocene of 
Rancho La Brea 



Procyonids, talpids, or chiropterans have not previously been recorded from 
the late Pleistocene Rancho La Brea asphalt deposits in Hancock Park, Los 
Angeles, California (Stock, 1956). Re-examination of the existing collection and 
continuing excavation have produced specimens referrable to Bassariscus astu- 
tus, Procyon lot or, Scapanus latimanus, and Lasiurus cinereus', all of which have 
modern distributions including the Rancho La Brea area. 

The authors recently located an isolated upper left first molar referrable to 
Bassariscus astutus (Fig. IB) stored with the Ranch La Brea collections of the 
Los Angeles County Museum of Natural History (LACM). The specimen was in 
a homemade Riker mount type of box together with isolated teeth of other small 
mammalian taxa common in the La Brea fauna such as Microtus californicus, 
Neotoma sp., Spermophilus beecheyi, Sylvilagus auduboni, ILepus californicus, 
and Mephitis mephitis. The label on the outside of the box was destroyed by 
silverfish but the contained specimens are undoubtedly from Rancho La Brea. 
The preservation of all specimens in the box is typical of La Brea materials, and 
we know of no Riker boxes in the LACM collections which contain fossil ver- 
tebrate materials from other sites. Most of these homemade Riker type boxes in 
the La Brea collection contain specimens from pit A (the various excavations at 
La Brea are commonly called "pits")- The remnants of the original label and its 
position on the box are the same as was used for labelling other Riker boxes from 
pit A. The history and nature of pits A, B, C, and D remain somewhat enigmatic. 
These excavations were begun in the spring of 1929 by field parties from the 
LACM and continued perhaps to 1931, but only fragmentary records can be 
located. The excavations were disrupted in the early summer of 1929 when the 
field parties were sent to the newly discovered late Pleistocene Conkling Cavern 
site in eastern New Mexico. The age and stratigraphic relationships of pit A with 
other La Brea pits are uncertain but the avian assemblage appears to indicate a 
later period of accumulation than the typical late Pleistocene pits (Howard, 1962). 

The tooth, LACM V-5205, is that of a young adult animal. The root canals are 
hollow and the cusps are sharp and little worn. V-5205 agrees in every detail with 
the left M/l of living Bassariscus astutus. Measurements: greatest anterior-pos- 
terior diameter measured parallel to labial margin of tooth is 5.6 millimeters (mm); 
transverse diameter measured at right angles to above is 6.5 mm. It appears to 
be closest in both morphology and measurements to B. astutus willetti but ade- 
quate series of modern subspecies were not examined. 

In June of 1969, the LACM, supported by grants and monies from the LACM 
Foundation, National Science Foundation (GB 24819), Los Angeles County, and 
numerous private corporations together with volunteer assistance, embarked on 
a long-range excavation program at pit 91, Rancho La Brea. The remaining new 
mammalian records from Rancho La Brea come from pit 91. On the basis of the 



142 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




I 





i** : -m 




1 v 




Fig. 1. A, LACM R-36926, Procyon lotor left ulna xl. B, LACM V-5205, Bassariscus astutus 
left M/l x5. C, LACM R-21363, Scapanus latimanus right humerus x2.5. D, LACM R-18437, Las- 
iurus cinereus distal left humerus x9. All stereographic pairs. A and' D coated with ammonium 
chloride and photographed with a Baird Stereo-Bar. B and D photographed with a Nikon SMZ-10 
trinocular microscope. 



few preliminary radiocarbon dates, pit 91 seems to be well stratified and has a 
late Pleistocene history from about 25,100 years before present (B.P.) to greater 
than 33,000 years B.P. Detailed stratigraphic data for the following specimens are 
on file at the George C. Page Museum. 

LACM R-36926 is the left ulna of a large juvenile raccoon lacking proximal and 
distal epiphyses (Fig. 1A). Although immature, the diaphysis and semi-lunar ar- 
ticulation are fully formed and comparable with adult specimens of Procyon lotor. 
Measurements: midshaft antero-posterior diameter 6.7 mm; midshaft transverse 
diameter 4.4 mm; length of semi-lunar notch 12.0 mm; width of upper portion of 
semi-lunar notch 8.9 mm. 

A complete right humerus, LACM R-21363, of an adult mole (Fig. 1C), agrees 



RESEARCH NOTES 143 

in details and measurements with modern specimens of Scapanus latimanus. The 
measurements for R-21363 are: Total length (distance from most proximal point 
on the humerus to the center of a line connecting the two most distal points) — 
13.3 mm; proximal width (greatest distance between the teres tubercle and the 
greater tuberosity, measured at right angles to long axis) — 9.7 mm; minimum 
shaft width (narrowest mediolateral distance across the shaft of humerus, mea- 
sured at right angles to the long axis) — 3.9 mm; shaft thickness (antero-posterior 
distance from surface to surface, measured at same level as, and at right angles 
to, the shaft width) — 3.7 mm. 

LACM R-18437 is an incomplete left distal chiropteran humerus (Fig. ID). Part 
of the spinous process and much of the posterior surface are missing. The fossil 
was compared with a number of modern specimens and the illustrations of J. D. 
Smith (1972). The tubercle on the proximal anterior margin of the lateral epicon- 
dyle is well developed and sharp; the distal spinous process is well developed 
and closely appressed to the medial epiconcdyle. These features appear to be 
characteristic only of Lasiurus. The relatively large size (2.6 mm across the most 
distal portion of the articulation) is comparable only to L. cinereus of the genus. 
The preserved portions of the fossil are identical to a modern specimen of L. 
cinereus, 46141 ', in the mammalogy collections of LACM. 

Acknowledgments 

The authors wish to thank George T. Jefferson of the George G. Page Museum, 
Theodore Downs of the Los Angeles County Museum of Natural History, and 
David E. Fortsch of the Idaho State University Museum of Natural History for 
critically reading this manuscript. 

Literature Cited 

Howard, H. 1962. A comparison of avian assemblages from individual pits at Rancho La Brea, 

California. Los Angeles County Mus. Contrib. Sci., No. 58. 24 pp. 
Smith, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Univ. Kans. Mus. Nat. Hist. 

Misc. Publ., 56:132 pp. 
Stock, C. 1956. Rancho La Brea. A record of Pleistocene life in California (sixth ed.) Los Angeles 

County Mus. Sci. Series, No. 20. 81 pp. 

Accepted for publication May 7, 1979 

William A. Akersten, Richard L. Reynolds, and Antonia E. Tejada-Flores, 
George C. Page Museum, Los Angeles County Museum of Natural History, 5801 
Wilshire Boulevard, Los Angeles, California 90036. 



144 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Bull. Southern California Acad. Sci. 
78(2), 1979, pp. 144-147 



Post-larval Polychaetes in Sandy Beaches of 
Tomales Bay, California 



Although many macrobenthic organisms spend part of their lives as transient 
members of the meiofauna (Thorson, 1966), relatively little quantitative data are 
available on their interactions in the meiobenthos. For example, Muus (1973) 
presents data on macrobenthic larval settling and mortality rates, and the duration 
of their stay in this habitat. 

This paper presents preliminary quantitative data on the distribution and set- 
tling rates of post-larval polychaetes from sandy beaches of Tomales Bay, Cali- 
fornia. Research was conducted at the Pacific Marine Station, Dillon Beach, 
California, from 18 February to 18 May 1970. Sediment samples were collected 
along transects in Tomales Bay at Lawsons Flat, White Gulch and Walker Creek 
delta, and were taken either with a core (5.4 cm dia) or a petri dish (10 cm dia). 
Core samples were taken to a depth of 4 cm; dishes to 1.5 cm. In the laboratory, 
samples were divided into 4 subsamples which were not preserved or anaesthe- 
tized. A modified Boisseau apparatus was used in which a standard distillation 
flask was substituted for the cylinder (see Hulings and Gray, 1971). Filtered sea 
water was used throughout this experiment. Organisms were retained on a 70 fxm 
mesh which was examined for entangled animals after each elutriation. Eight 
control samples were examined for post-larval polychaetes in which the number 
of larvae was known prior to elutriation. All control polychaetes were removed 
from the sediment after 9 min. 

A total of 1 1 species of polychaetes belonging to 7 families was found: Anaitides 
williamsi Hartman; Platynereis bicanaliculata (Baird); Nephtys caecoides Hart- 
man; Hemipodus borealis Johnson; Glycinde armigera Moore; Lumbrineris 
Izonata Johnson; Polydora socialis (Schmarda); P. caulleryi Mesnil; Pseudo- 
polydora kempi (Southern); P. paucibranchiata (Okuda) and Pygospio califor- 
nica Hartman. The overall mean larval density for the entire sampling period was 
468.7 ± 377.8 worms -m -2 , and ranged from 56.8 to 1055.8 worms -m -2 (N = 36 
samples). Periods of heaviest settling occurred during late March and late April 
(Fig. 1). Species settling heavily during the first period are H. borealis, P. kempi 
and N. caecoides; those of the second period are G. armigera, A. williamsi and 
N. caecoides. Settling data for all species are presented in Table 1. Pseudopo- 
lydora kempi had the highest settling rate (103.67 ± 15.45 individuals -m -2 - day), 
followed by G. armigera, H. borealis, N. caecoides, A. williamsi and P. bican- 
aliculata. Habitat partitioning is shown by H. borealis and G. armigera where 
dense settling periods were temporally separated. 

All of these polychaetes were taken within the upper 2 cm of the sediment 
column. Two transects parallel to the shoreline at Lawsons Flat revealed a density 
of 488.3 worms -m" 2 above and 233.5 worms -m -2 below the waterline. A. wil- 
liamsi, N. caecoides and G. armigera were generally taken on the exposed beach, 
whileP. bicanaliculata was collected below the waterline. 



RESEARCH NOTES 



145 



1000 



CM 

I 



LU 

< 

o 



^500" 



o 



< 
> 

< 



CO 

o 

Q_ 




27 3 6 



Sampling dates (1970) 

Fig. 1. Settling trends for all species of post-larval polychaetes from Tomales Bay, California. 



The temporally most widespread species was Neplitys caecoides, which had 
a mean density of 97.1 individuals m 2 . I calculated that this species had a com- 
bined summer density of 21.2 adult worms- m 2 at White Gulch and Lawsons Flat 
(see Johnson, 1967, data for 1963). Although these data are restricted in time, the 
mortality rate of N. caecoides must be substantial. 

The presence of certain post-larval polychaetes in beach sediments corresponds 
well to their periods of reproduction (see Blake, 1975; Blake and Woodwick, 
1975). All sampling for this experiment was performed in regions where adult 
populations of these species were low (see Johnson, 1970). Muus (1966) states 
that, "vast numbers of larvae settle where they do not belong."' This is apparently 
reflected in the patchy settling pattern of Pseudopolydora kempi. It is well known 
that many larvae can postpone settling and metamorphosis. However, when such 
delays are coupled with adverse climatic conditions, for example, massive mor- 



146 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Table 1. Densities (no-m 2 ) and mean settling rates (no- m -2 - day -1 ) for post-larval polychaetes 
from Tomales Bay, California. 



tu ~ S ^ fe - 

r-> -~ .y a o v. 

o ^ ^ ^ a: d 



28.4* — ___ ____ 

28.4 28.4 — — — — — — — 

38 56.6 — — — — 38 19 — 

227 397.5 — — — — — — — 

142 28.4 — — 742 — — — 28.4 

_ 63.6 — — 573 — — — — 

_ — 128 — — ____ 

172 — 513 — — — — — — 

125 — 250 — — — — — — 

16 — 95.3 80 — 16 — — — 



27 Feb 


— 


28 


3 Mar 


— 


— 


6 Mar 


— 


— 


18 Mar 


— 


171 


26 Mar 


86 


29 


31 Mar 


— 


— 


13 Apr 


86 


43 


29 Apr 


171 


114 


4 May 


125 


— 


18 May 


— 


— 



±s 


93.6 
62.9 


48.2 
62.3 


97.1 
79.9 


114.9 

158.8 


246.6 
189.7 


657.5 — 
119.5 — 








— 


noirr 2 day~ 

X 

±s 


i 

12.8 
8.6 


8.7 
8.5 


14.7 
9.7 


15.1 
11.6 


24.5 
20.4 


103.7 — 
15.5 — 


— 


— 


— 



Data not included in computation of mean. 



talities may be inevitable. The increased density of worms on an exposed beach 
is probably a behavioral adaptation by raptorial feeders, rather than being the 
result of prevailing climatic conditions. The species typically encountered in this 
habitat may gain a feeding advantage through decreased motility of prey species. 
The nereid species typically collected below the waterline is basically omnivo- 
rous, and may be disadvantaged in exposed habitats. 

Literature Cited 

Blake, J. A. 1975. The larval development of Polychaeta from the northern California coast. III. 

Eighteen species of Errantia. Ophelia 14:23-84. 
, and K. H. Woodwick. 1975. Reproduction and larval development of Pseudopolydora pau- 

cibranchiata (Okuda) and Pseudopolydora kempi (Southern). (Polychaeta: Spionidae). Biol. 

Bull. 148:109-127. 

Hulings N. CandJ. S. Gray (eds.). 1971. A manual for the study of meiofauna. Smithsonian Contr. 

Zool. 78:1-83. 
Johnson, R. G. 1967. The vertical distribution of the infauna of a sandflat. Ecology 48:571-578. 

. 1970. Variations in diversity within benthic marine communities. Amer. Nat. 104:285-300. 

Muus, K. 1966. A quantitative 3-year survey of the meiofauna of known macrobenthic communities 
in the 0redsund. Veroff. Inst. Meeresforsch. Bremerh. (Sonderbd.) 2:289-292. 

. 1973. Settling, growth and mortality of young bivalves in the 0resund. Ophelia 12:79-116. 

Thorson, G. 1966. Some factors influencing the recruitment and establishment of marine benthic 
communities. Neth. Jnl. Sea Res. 3:267-293. 



RESEARCH NOTES 147 

Accepted for publication May 7, 1979. 

Jerry D. Kudenov, Allan Hancock Foundation, University of Southern California, 
Los Angeles, Calif. 90007. Allan Hancock Foundation Contribution No. 371 . 



Bull. Southern California Acad. Sci. 
78(2), 1979, pp. 147-150 



The Calcanea of Members of the Hyaenidae 



The calcanea of all currently recognized species of living members of the 
Hyaenidae were examined. These are the calcanea of Crocuta crocuta (15 spec- 
imens), Hyaena brunnea (2 specimens, 1 malformed), H. hyaena (5 specimens), 
and Proteles cristatus (4 specimens). All calcanea of hyaenas have the same 
articular surfaces: posterior (PAS), medial (MAS), anterior (AAS), and cuboid 
(CAS) (Fig. 1); the proximal end of the medial articular surface folds over the 
proximal end of the sustentaculum astragali (SA); and the outline of the outer 
surface lacks a distinct trochlear process (TP) and tends to dip inward near the 
cuboid facet (Fig. 1). See Stains (1959, 1962, 1973) or Robinette and Stains (1970) 
for terminology. 

Genera and Species of the Family Hyaenidae 
Crocuta crocuta — Spotted Hyaena 

Specimens examined. — American Museum of Natural History (AMNH) 21542, 
27767, 35358, 36390, 163101, 163105; U.S. National Museum of Natural History 
(USNM) 161909, 162920, 163100, 163102-4, 163299, 163344, 164506, 164834, 
172685 (USNM 163344 illustrated, Fig. la). 

Crocuta crocuta has the largest calcaneum of modern hyaenas; more than 55 
mm long and 25 mm wide (Table 1). The body of the calcaneum is robust and the 
greater tuberosity (GT) is twisted toward the inner side (Fig. la). The anterior 
articular surface is large and distinct, and is separated from the medial articular 
surface. A ridge of unarticulated bone sometimes connects the two surfaces (in 
2 of 15 specimens examined). 

Hyaena spp. 

Calcanea next in size to those of Crocuta, but slightly smaller, are those of the 
genus Hyaena which range between 45 to 54 mm long and 22 to 25 mm wide 
(Table 1). The anterior articular surface is more elongate in the proximo-distal 
direction than in Crocuta and, in most cases, is joined with the medial articular 
surface. 

Hyaena brunnea — Brown Hyaena 

Specimens examined.— AM 83690; USNM 221088 (deformed) (AM 83690 il- 
lustrated, Fig. lc). 



148 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 1. Calcanea of the Hyaenidae. a. Crocuta crocuta, b. Hyaena hyaena, c. Hyaena brunnea, 
d. Proteles cristatus. GT — greater tuberosity, S — sustentaculum, TP — trochlear process, CS — cuboid 
surface, A AS — anterior articular surface, MAS — medial articular surface, PAS — posterior articular 
surface. 



The shape of the greater tuberosity of H. brunnea is closer to that of C. crocuta 
than to other hyaenas. The junction of the anterior and medial articular surfaces 
is more obvious in H. brunnea than in H. hyaena (Fig. 1) although the young of 
H. hyaena also show a distinct connection. 

One specimen of H. brunnea is malformed (USNM 221088). The malformation 
appears to be a deletion of part of the main distal dorsal surface of the one 
opposite the anterior articular surface and just above the cuboid facet. Exami- 
nation of the calcaneum from the opposite foot in the same specimen, reveals an 
exact copy of the malformation indicating that the condition is genetic and not 
due to some previous injury. 

Hyaena hyaena — Striped Hyaena 

Specimens examined.— AMNH 54512; USNM 14403, 99626, 155455, 163111 
(USNM 14403 illustrated, Fig. lb). 

Calcanea of adults of H. hyaena show a slight separation of the anterior and 
medial articular surfaces. H. hyaena calcanea are most easily distinguished from 
H. brunnea by the greater width of the greater tuberosity of H. hyaena when 
viewed from the dorsal aspect. The calcanea of H. hyaena are larger (52 mm 



RESEARCH NOTES 149 

Table 1. Measurements and Indices for Calcanea of Species of the Family Hyaenidae. 



Hyaenid Species 


No. 


Length (mm) 


Width (mm) 


W/L Index 


Range 


Avg 


Range 


Avg 


Range Avg 


Crocuta crocuta 
Hyaena hyaena 
Hyaena brunnea 
Proteles cristatus 


15 
3 
1 
3 


56.5-62.2 
50.2-53.9 

29.0-31.9 


58.9 
51.8 
44.6 
30.2 


26.9-33.9 
23.1-25.0 

14.1-16.3 


29.5 
23.9 
22.1 
15.5 


.46-.56 .50 

.45-.52 .48 

.50 

.49-.54 .51 



length, 23.9 mm width than those of H. brunnea (44.5 mm length, 22.1 width). 
Table 1 gives additional measurements. 

Proteles cristatus — Aardwolf 

Specimens examined.— AMNH 27768, 70261; USNM 164503, 181523 (USNM 
181523 illustrated, Fig. Id). 

Proteles cristatus possesses the smallest calcaneum (total length 30.2 mm, 
width 15.5 mm), and resembles H. brunnea (Fig. lc) most closely in shape than 
other members of the family. In most respects, the calcaneum of P. cristatus 
(Fig. Id) is a miniature of H. brunnea. The cuboid facet is nearly oval in shape 
medio-laterally in P. cristatus; more circular in shape in other hyaenas. In ad- 
dition, the more proximal half of the posterior articular surface is sigmoid in P. 
:ristatus\ in other hyaenas this surface is slightly sigmoid to rounded. In this 
feature, H. brunnea again approaches P. cristatus more closely than do the other 
hyaenas. 

Conclusions 

H. brunnea and P. cristatus have calcanea with joined anterior and medial 
irticular surfaces and similar shapes. Calcanea of C. crocuta are the most distinct 
►vith an inward flaring of the greater tuberosity. 

The hyaenas are regarded as being offshoots of viverrid ancestors according 
o the fossil evidence (Beddard, 1902; Romer, 1966; Simpson, 1945). Of all cal- 
;anea examined (Stains, 1959) in other familes of the Carnivora, some members 
)f the Viverridae (Stains, J. Morph.) show the closest resemblance to the hyae- 
las. Within the viverrids, calcanea of the Herpestinae resemble the hyaenas in 
ihape of the greater tuberosity in particular. In viverrids, the trochlear process 
s usually distinct and often well above the distal end of the calcaneum. However, 
n the Herpestinae this process is slight and at the distal end of the calcaneum. 

Most dogs (Stains, 1975) and cats also lack a distinct trochlear process although 
here is usually an enlargement near the distal end of the bone. In general, a 
eature of the calcaneum such as the trochlear process remains rather uniform 
hroughout members of a family such as the bears and procyonids (Stains, 1973), 
nustelids (Stains, 1976a, 1976b), sciurids (Stains, 1962) and viverrids (Stains, J. 
vlorph.). Robinette and Stains (1970) discuss terminology in seals. Of all the 
iving genera of carnivores I have examined, the cheetah of the Felidae has a 
:alcaneum with an outline closest to the hyaenas. The cheetah has a small trochl- 
ear process near the distal end of the bone, an elongation of the calcaneum, a 
mall swelling between the minute trochlear process and distal to the posterior 



150 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

articular surface, a smooth curved posterior articular surface, and a grooved 
greater tuberosity in which it differs from the hyaenas. 

Literature Cited 

Beddard, F. E. 1902. Mammalia. Macmillan and Co., Ltd., N.Y. 605 pp. 

Robinette, H. R. and H. J. Stains. 1970. Comparative study of the calcanea at the pinnipedia. J. 

Mamm., 51:527-541. 
Romer, A. S. 1966. Vertebrate paleontology. Univ. Chicago Press. 468 pp. 

. 1968. Notes and comments on Vertebrate paleontology. Univ. Chicago Press. 304 pp. 

Simpson, G. G. 1945. The principles of classification and a classification of mammals. Bull. American 

Mus. Nat. Hist., 85:1-350. 
Stains, H. J. 1959. Use of the calcaneum in studies of taxonomy and food habits. J. Mamm., 40:392- 

401. 

-. 1962. Osteological data used in mammal classification. Syst. Zool., 11:127-130. 

1973. Comparative study of the calcanea of members of the Ursidae and Procyonidae. Bull. 

So. California Acad. Sci., 72:137-148. 

-. 1975. Calcanea of members of the Canidae. Bull. So. California Acad. Sci., 74:143-155. 
1976a. Calcanea of members of the Mustelidae. Part I, Mustelinae. Bull. So. California Acad. 

Sci., 75:237-248. 
1976b. Calcanea of members of the Mustelidae. Part II, Melivorinae, Melinae, Mephitinae, 

and Lutrinae. Bull. So. California Acad. Sci., 75:249-257. 
. Calcanea of the Viverridae. Submitted to Journal of Morph. 

Accepted for publication February 13, 1979. 

Howard J. Stains, Department of Zoology, Southern Illinois University, Carbon- 
dale, Illinois 62901 . 



INSTRUCTIONS FOR AUTHORS 

s BULLETIN is published three times each year (April, August, and November) and includes articles in English 
any field of science with an emphasis on the southern California area. Manuscripts submitted for publication 
>uld contain results of original research, embrace sound principles of scientific investigation, and present data 
i clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide 
contributors. Consult also recent issues of the BULLETIN. Authors should strive for directness and lucidity, 
lieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous 
jrence of pronouns, and logically placed modifiers. 

MANUSCRIPT PREPARATION 

t is strongly recommended that, before submitting a paper, the author ask qualified persons to review it. The 
hor is requested to submit at least two additional copies with the original, on 8V2 x 11 opaque, nonerasable 
>er, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. 
atnotes should be avoided. Manuscripts which do not conform to the style of the BULLETIN will be returned to 
author. 

Iin abstract summarizing in concise terms the methods, findings, and implications discussed in the paper must 
ompany a feature article. Abstract should not exceed 100 words. 

\ feature article comprises approximately five to thirty typewritten pages. Papers should usually be divided into 
following sections: abstract, introduction, methods, results, discussion and conclusions, acknowledgments, and 
rature cited. Avoid using more than two levels of subheadings. 

1 research note is usually one to six typewritten pages and rarely utilizes subheadings. Consult a recent issue 
he BULLETIN for the format of notes. Abstracts are not used for notes. 

abbreviations: Use of abbreviations and symbols can be determined by inspection of a recent issue of the 
LLETIN. Omit periods after standard abbreviations: 1.2 mm, 2 km, 30 cm, but Figs. 1-2. Use numerals before 
ts of measurements: 5 ml, but nine spines (10 or numbers above, such as 13 spines). The metric system of 
ghts and measurements should be used wherever possible. 

"axonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International 
ie of Botanical Nomenclature (Lawjouw et al.. 1956), the International Code of Nomenclature of Bacteria and 
uses (Buchanan et al.. 1958), and the International Code of Zoological Nomenclature (Stoll et al., 1961). 
rcial attention should be given to the description of new taxa, designation of holotype, etc. Reference to new 
a in titles and abstract should be avoided. 

he literature cited: Entries for books and articles should take these forms. 
lcWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. 

lolmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm.. 54: 452-458. 
Irattstrom, B. H. 1969. The Condor in California. Pp. 369-382 ;'/; Vertebrates of California. (S. E. Payne, ed.), 
Jniv. California Press, xii + 635 pp. 

ables and figures (line drawings, graphs, or black and white photographs) should not repeat data contained in 
text. The author must provide numbers and short legends for tables and figures and place reference to each 
hem in the text. Legends should be typed on a separate sheet of paper and placed at the end of the manuscript. 
strations and lettering thereon should be of sufficient size and clarity to permit reduction to standard page size; 
inarily they should be no more than twice the size of intended reduction and should not exceed 8' 2 by 1 1 inches 
ize. Photographs must be printed on glossy paper. Submit one photoduplicated copy of each illustration. All 
Orations accompanying Research Notes will be reduced to one column width. All half-tone illustrations will have 
|t screen (grey) backgrounds. Special handling such as dropout half-tones, special screens, etc., must be requested 
»nd will be charged to authors. 

cover illustration pertaining to an article in the issue or one of general scientific interest will be printed on the 
?er of each issue. Such illustrations along with a brief caption should be sent to the Editor for review. 

PROCEDURE 

II manuscripts should be submitted to the Editor, Jeffrey I. Chapman. School of Public Administration, Uni- 
fcity of Southern California, Los Angeles, California 90007. Evaluation of a paper submitted to the BULLETIN 

tins with a critical reading by the Editor; several referees also check the paper for scientific content, originality, 
1 clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent 
)he author at which time he or she may be requested to rework portions of the paper considering these rec- 
mendations. The paper then is resubmitted and may be re-evaluated before final acceptance. 

roof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He or she 
".ild promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations, legends. 
r bibliographical references. He or she marks corrections on the galley (copy editing and proof procedures in 
ie Manual) and promptly returns both galley and manuscript to the Editor. Manuscripts and original illustrations 
' not be returned unless requested al this time. All changes in galley proof attributable to the author (misspellings, 
insistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with 
• printer, not the Editor. 



CONTENTS 

Osprey Distribution, Abundance, and Status in Western North America: III. 

The Baja California and Gulf of California Population. By Charles J. 

Henny and Daniel W. Anderson 89 

A New Species of Leptodactylid Frog, Genus Eleutherodactylus , from the 

Cordillera de Talamanca, Costa Rica. By Jay M. Savage and James 

E. De Weese 107 

New Species and Records of Polychaetous Annelids from the Tetraclita 

(Cirripedia: Crustacea) Zone of the Northern Gulf of California, 

Mexico. By Jerry D. Kudenov 116 

A Survey of the Littoral and Sublittoral Ascidians of Southern California, 

Including the Channel Islands. By Rimmon C. Fay and James A. 

Vallee 122 

A Redescription of Pettiboneia sanmatiensis Orensanz (Polychaeta: Dor- 

villeidae) and a Revised Key to the Genera of the Dorvilleidae. By 

James A. Blake 136 

Research Notes 

New Mammalian Records from the Late Pleistocene of Rancho La Brea. By William A. Aker- 

sten, Richard L. Reynolds, and Antonia E. Tejada-Flores 141 

Post-larval Polychaetes in Sandy Beaches of Tomales Bay, California. By Jerry D. Kudenov 144 

The Calcanea of Members of the Hyaenidae. By Howard J. Stains 147 



COVER: Eleutherodactylus rayo, a new species of frog from Costa Rica. 



UTHERN CALIFORNIA ACADEMY OF SCIENCES 




ULLET1N 



rAN^CAL GARDEN' 



me 78 



Number 3 



' 1 '*'^ B -"• ^K^FaF^^» • fir HK* ^» . . . » . 


^2£^ftr^Csl'' £lf i Jr/« 


■--aSLaH <»SSil^ ^^^r^ ^"^^^ 

Wfmm^^M "JETS fSSStSVi f ^^BPLjMl 



S-A78(3) 151-206 (1979) 



DECEMBER 1979 



Southern California Academy of Sciences 

Founded 6 November 1891, incorporated 17 May 1907 



OFFICERS 

Takashi Hoshizaki, President 
Fred G. Hochberg, Vice President 
Richard E. Pieper, Secretary 
Joseph E. Haring, Treasurer 
Robert J. Lavenberg, Editor 
Gretchen Sibley, Assistant Editor 



1978-1980 

Kristin H. Berry 

Robert A. Georges 

Joseph E. Haring 

Jerry D. Kudenov 

Donald J. Reish 



BOARD OF DIRECTORS 

1979-1981 

John Baird 

Jules Crane 

Fred G. Hochberg 

Richard E. Pieper 

Gloria Takahashi 



1980-1982 

Takashi Hoshizaki 
Alan J. Mearns 

Steven N. Murray 
Camm C. Swift 

Robert G. Zahary 



Membership is open to scholars in the fields of natural and social sciences, and to any pers< 
interested in the advancement of science. Dues for membership, changes of address, and requests f 
missing numbers lost in shipment should be addressed to: Southern California Academy of Science 1 
the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 900C 

Annual Members $ 12.; 

Life Members 150.j J 

Fellows: Elected by the Board of Directors for meritorious services. 



The Bulletin is published three times each year by the Academy. Manuscripts for publication shou 
be sent to the appropriate editor as explained in "Instructions for Authors' ' on the inside back cov, 
of each number. All other communications should be addressed to the Southern California Acader 
of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, L 
Angeles, California 90007. 



Date of this issue 7 March 1980 



Bull. Southern California Acad. Sci. 
78(3), 1979, pp. 151-162 



Seventieth Anniversary of Academy Excavations at 
Rancho La Brea 

Gretchen Sibley 

Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los 
Angeles, California 90007. 



The Southern California Academy of Sciences was one of the early organiza- 
tions permitted to collect fossil material from the Pleistocene beds at Rancho La 
Brea in Los Angeles, California. The Academy work began in June, 1909, though 
the first recognition of the importance of the fossils had come 34 years before. 
In 1875 William Denton visited Major Henry Hancock at the rancho, collected 
a few fossil bones and, two years later, published a description of them in the 
Proceedings of the Boston Society of Natural History. The article apparently 
created no appreciable notice and was soon forgotten (Stock, 1956). 

It was not until 1901 that a sustained interest brought effectual activity. Mr. 
W. W. Orcutt, while studying the geology of the area for the Union Oil Company, 
found a number of bones which he transferred to his office. In 1905 Mr. F. M. 
Anderson, a geologist with the Southern Pacific Company, noticed the material 
and realized its importance. He contacted his friend, Prof. John C. Merriam of 
the University of California, who immediately obtained permission from Mrs. 
Henry Hancock to excavate at the ranch during the years 1906 to 1913. Merriam 
made a fine collection for the university and published the first public announce- 
ment in the Sunset Magazine, October, 1908. He carried on the work with some 
of his colleagues and students but was never able to finance an extensive project. 

Shortly after Merriam began his work, both Occidental College and the Los 
Angeles High School were granted permission to collect the fossil material. James 
Zacchaeus Gilbert, professor of biology at Los Angeles High School, became 
intensely interested, headed up the dig and made a small collection for the high 
school. The bones remained at the school until the late sixties when they were 
given to the Los Angeles County Museum of Natural History. 

Gilbert used interested students to help excavate, but he realized the impor- 
tance of the fossils and the necessity for securing money to carry on the work on 
a larger scale. He turned to the Southern California Academy of Sciences of 
which he was a long-standing member and fellow. The progress can be followed 
from the excellent minutes and editorials provided by Holdridge Ozro Collins, 
secretary, and editor of the Bulletin. The first mention of the work at Rancho La 
Brea appears in the minutes of a meeting of the directors held on Wednesday, 
March 17, 1909. 

"A communication was received from Mr. Samuel Fox, relating to a proposed 
excavation in the Rancho La Brea, west of the City of Los Angeles, which is 
owned by Mrs. Ida Hancock, and the Secretary was instructed to inform Mr. 
Fox that while the directors are in full sympathy with this project, there are no 
funds subject to their control which are available for this purpose." 



152 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

April and May passed with no further information, but Gilbert's enthusiasm 
began to kindle interest. At a section meeting held at the residence of Mr. S. J. 
Keese on Monday, June 12, 1909, "Prof. Gilbert gave a statement of the progress 
in the excavations at the Rancho La Brea, made immediately possible by means 
of the generous donation to the Section by Mr. John D. Hooker. He placed upon 
exhibition several fossils, in excellent condition of preservation, of long-extinct 
animals, which he had excavated at this place, and he outlined the plan for future 
work under the auspices of the Academy." 

On June 28 the project was continued by the directors. "Professor J. Z. Gilbert 
was authorized to organize a Zoological Section, and the Secretary was instructed 
to communicate with Mrs. Erskine M. Ross [formerly Mrs. Henry Hancock], and 
request from her an allotment in her Rancho La Brea for excavation under the 
auspices of this Academy, in a search for fossil and geological specimens." 

It is interesting to note that the work was under the new zoological section 
though there was a geological section in the Academy. 

Mrs. Ross allowed the Academy to begin excavation near the old Hancock 
home. Following John D. Hooker's initial donation, Los Angeles City also con- 
tributed to the fund, and the Academy canvassed members to take care of other 
expenses. 

Holdridge Collins published the details in the following editorial in the Bulletin 
of July, 1909. 

The organization of a Section of Zoology marks a new era in the life of the 
Academy. Professor James L. [sic] Gilbert, Chairman of the Section, well 
known in this community for his scientific attainments, is at the head of the 
Biological Department of the Los Angeles High School, and during the short 
period of the life of this Section he has accomplished wonders. It has been 
known for some time that in a depression on the Rancho La Brea, are buried 
the remains of early fauna of California. A communication from the Academy 
to Mrs. Erskine M. Ross, owner of the rancho, requesting permission to 
excavate at this place, brought a most gracious consent, and, owing to the 
liberality of Mr. John D. Hooker, Prof. Gilbert was able to proceed at once 
with the work. An opportunity so rare comes but once, and we are fortunate 
that we have been able to delve in so rich a bed of extinct fauna at the very 
door of the Academy. 

On Monday, July 26, 1909, Prof. Gilbert placed upon exhibition in the City 
Hall the results of his work since the commencement of the public school 
summer vacation. A Buffalo skull, with a sweep of horns of 31 inches from 
tip to tip, the largest ever found; the bones of the Camel, Horse, Antelope, 
Sloth and Elephant in their most gigantic forms; a skull of the sabre-toothed 
tiger, complete in all its parts; the head and part of the bones of a huge lion — 
not the mountain lion of California, but of the African type; smaller cats, 
rodents and a large variety of birds, great and small, were shown in an 
unusually excellent state of perfection. None of these are petrified, and their 
preservation from a time before man made his appearance is explained by 
their burial in a grave of brea. 

This exhibit was inspected by the Mayor and the gentlemen of the City 
Council, who were so impressed with the value of these discoveries and fear 



SEVENTIETH ANNIVERSARY 



153 



->' r- 




James Z. Gilbert (black hat and beard) explains fossils to visitors and student workers in the 
Academy dig at Rancho La Brea, 1909. 



that the numerous foreign Scientific Bodies, which have applied for permis- 
sion to excavate in this wonderful deposit, will carry away fossils which 
should forever remain in a home Museum, that, on the same day an appro- 
priation of five hundred dollars was made to the Academy for the continuance 
of this work. 

Academy members received their first detailed account of the work in October. 
A summary is given in the minutes. 

The first lecture of the season of 1909-1910 was given by Professor James 
Z. Gilbert on Monday evening, October 4, 1909, in Symphony Hall. 

His subject was "The extinct fauna of the Southwest, as represented by 
numerous fossils of extinct animals discovered at La Brea Rancho. 

A large attendance was in presence and a very pleasant episode of the 
evening was the music of the young ladies' Glee Club of the Los Angeles 
High School. 

In introducing his subject. Prof. Gilbert paid a tribute of thanks to Mrs. 
Erskine M. Ross for her gracious allotment of a tract on her ranch, and the 



154 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

liberality of Mr. John D. Hooker and the Council of Los Angeles, which had 
enabled him to carry on the excavations without cessation throughout the 
entire summer. 

Upon the screen he presented views of the small lake in the eucalyptus 
grove and explained the geological and paleontological conditions of the dis- 
trict surrounding this unique deposit of fossils. 

The ancient spring of water, gushing through a bed of asphaltum, was a 
fatal trap to hold to their death the numerous animals of earth and air, which 
resorted to this place to quench their thirst. 

A most interesting collection, in an excellent condition of preservation, of 
the fossils of the lion, the sabre-toothed tiger from infancy to old age, the 
wolf, mastodon, giant sloth, camel, horse, antelope, small rodents of nu- 
merous species, the peacock, the great condor, and the ancestors of the 
eagle, and many other animals whose descendants are now found only in 
tropical lands, was placed upon exhibition, and their characteristics ex- 
plained by the Professor, and at the close of the address these remains were 
examined by the intensely interested audience. 

Later in the month, 'The Secretary was instructed to extend to Professor W. 
H. Housh, Principal of the High School, an expression of the appreciation of this 
Board for his kindness in furnishing room in the High School building for the 
work of cleaning the fossils lately excavated and giving them a secure place for 
storage/' 

At the director's meeting on October 15, the following was recorded. "In view 
of the valuable acquisition of fossils from Rancho La Brea; the tender of geolog- 
ical and zoological collection; the necessity for the proper housing and care of 
these valuable contributions to science, and the increasing needs of the Academy, 
it was resolved to place before the members and the public generally, a formal 
statement of the financial requirements of the Academy, and solicit subscriptions, 
to enable us to continue a work which has been so fruitful of results. 

"A note of thanks was unanimously given to Mr. John D. Hooker for an 
additional gift of $500.00 for the prosecution of the excavations in Rancho La 
Brea." 

Collins reported later in the year. "The work of cleaning and mounting the 
fossils from Rancho La Brea is proceeding with gratifying success, under the 
direction of Professor Gilbert. At this writing, a sabre-tooth tiger and a giant wolf 
stand in all the perfection of their skeleton ferocity, and bones of a giant ground 
sloth have been assembled, and work has been commenced on their mounting. 
A finely mounted and perfect skull of a sabre-tooth tiger, with sabres, eight inches ; 
in length, has been placed in the office of the Secretary where it can be seen at 
any time by members of the Academy and their friends." 

At this time the Academy had been in existence for eighteen years. On No-; 
vember 6, 1891 it was established as the Southern California Science Association 
by an enthusiastic group of men and women interested in science. Five years 
later the rapidly growing organization adopted its present name. By 1909, when 
excavations at La Brea were undertaken, the membership had grown to around 
one hundred and eight (Howard, 1957). And the decision to aid Gilbert at Rancho 
La Brea led the Academy into a period of its greatest activity. The new Hall of 



SEVENTIETH ANNIVERSARY 155 

Records had just been completed and it was hoped that an area might be secured 
in the new building for a display of La Brea fossils. Hector Alliot, Academy 
member, met with the County Board of Supervisors and reported to William 
Spalding, president of the Academy. 

Following our recent conversation I met the county Supervisors, by ap- 
pointment, this morning and (unofficially) [sic] explained to the board the 
civic importance of the work of the Academy of Science [sic] of Southern 
California in various fields, particularly that of paleontology. 

The necessity of their recognizing the efforts of the Society in preserving 
the valuable specimens from the La Brea ranch deposits, by providing a 
suitable space for exhibition purposes in the new Hall of Records building 
was urged upon them. 

While unwilling to bind themselves by a resolution, the Supervisors unan- 
imously agreed that they would see that an exhibition room, or rooms, for 
the display of zoological and other collections owned by the Academy of 
Science [sic] would be provided in the projected Museum Building at Agri- 
cultural Park. The first installment of county funds will be forthcoming in 
September, and the architects will be able to commence work on this Mu- 
seum in a short time. 

Pending the completion of the Museum, Supervisor Eldredge has kindly 
offered the Academy of Science space for the public exhibition of their col- 
lections in the County Court House, when certain portions of that building 
shall have been vacated by departments which are to move to the new Hall 
of Records. The entire space in the latter building has already been appor- 
tioned. 

The Academy of Science can therefore feel certain that in the near future 
temporary rooms will be at their disposal for public exhibition purposes in 
the County Court House; and that they are assured permanent quarters in 
the Museum, when it shall have been erected in Agricultural Park. 

The "projected Museum Building at Agriculture Park" would become the Los 
Angeles Museum of History, Science and Art (now the Los Angeles County 
Museum of Natural History), and the park would soon be named Exposition 
Park. This cultural area was the dream of Judge William M. Bowen. In 1898 he 
had become interested in the park then given over to rabbit coursing, gambling, 
drinking and related activities. When he discovered the land belonged to the State 
of California, he spent years in the courts clearing title to the property which had 
fallen into private hands. He was convinced that the park was to become a cultural 
center for the City and County of Los Angeles. The new museum, state exposition 
building (now the State Museum of Science and Industry), the armory (now the 
Space Museum), the new race track (later replaced by the Memorial Coliseum) 
and the gardens and lawns, were a part of his general plan for the park (Sibley, 
1978). His friend, Howard Robertson (1953) shared his dream and recalled Bow- 
en's first plan made in the fall of 1908: "We spread out a piece of wrapping paper 
an the desk, sketched in the park boundaries, and began laying plans for putting 
the land to good use as a cultural center. At last, here was the place for a mu- 
seum!" 



156 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Several organizations were interested in a building in which they might exhibit 
their collections. On January 16, 1910, the board of directors of the Historical 
Society of Southern California met with Judge Bowen who showed them prelim- 
inary plans for a museum. They decided to invite several groups to meet with the 
Board of Supervisors to request room to house their collections (Guinn, 1909- 
10). 

On February 7, 1910, representatives from the Historical Society, the Southern 
California Academy of Sciences, the Fine Arts League and the Cooper Ornitho- 
logical Club, Southern Division, met with the County Board of Supervisors to 
discuss the proposed museum. Judge Bowen outlined an agreement that was 
drawn up and signed by the supervisors and representatives of each of the or- 
ganizations. 

According to the contract — from the Academy files — the County agreed "to 
erect at Agricultural Park a Historical Museum and Art Gallery for the purpose 
of collecting and exhibiting therein a collection of fine arts, specimens, and data 
of biology and zoology, and historical matter relative to the Pacific Coast, more 
particularly Southern California, with a view of promoting and encouraging sci- 
entific art and historical education and investigation;" It would also maintain and 
operate the building, handle the exhibits and pay salaries of employees. The four 
organizations "shall have the care, supervision, control and management of said 
building and the collection, installation and supervision of said exhibits therein, 
for a period of fifty years, from and after the 7th day of February, 1910, through 
a Board of Governors." 

The Board was set up as follows on the next day: 

Chairman of the Board of Supervisors: 

Mr. C. J. Nellis 
Two representatives from the Fine Arts League: 

Mrs. Henrietta House 

Mr. T. E. Gibbon 
Two representatives from the Historical Society: 

Dr. George Bovard 

Mr. J. M. Guinn 
Two representatives from the Southern California Academy of Sciences: 

Mr. William A. Spalding 

Dr. Anstruther Davidson 
One representative from the Cooper Club: 

Mr. Howard Robertson 
One representative selected at large: 

Judge William M. Bowen 

To the members on December 5, 1910, Collins reported in the Bulletin, "The 
President, Mr. Spalding, reported progress the erection of the new County Mu- 
seum Building in Agricultural Park, and, in a view thrown upon the screen, he 
pointed out the wing which will be the home of this Academy, and explained the 
conveniences which will be placed at our disposal. He invited all present to attend 
the ceremonies of the laying of the Corner Stone on the afternoon of December 
17, 1910." 

At the Academy Director's meeting five days later, 






SEVENTIETH ANNIVERSARY 



157 



I 




The cornerstone of the Museum was laid on November 17, 1910. 



The Secretary reported that he had been requested to take charge of the 
selection of records to be sealed in the Copper Box which will be deposited 
in a cavity of the Corner Stone of the new County Museum Building, to be 
laid by the Most Worshipful Grand Master of Ancient, Free and Accepted 
Masons of California, on Saturday, December 17, 1910, and he asked for 
instructions from the Directors as to what documents of this Academy should 
be selected for that collection. 

He was directed to have the Articles of Incorporation, Certificate of the 
Secretary of State and List of Members of the Academy, printed in a suitable 
form, and with Volumes VIII and IX of the Bulletin and a short typewritten 
History of the Academy, to be prepared by Mr. Knight, placed in the Box. 
The Secretary was also instructed to send to all members, a circular, inform- 
ing them of the time and place of the ceremonies connected with the laying 
of the Corner Stone and to include with said Circular a silk badge upon which 
shall be printed the words "Academy of Sciences. " 

The secretary must have had a busy six days accomplishing so much. Today 
there would not be a printer available to set all the material in type on such short 
notice. 

As scheduled, the cornerstone was laid on December 17, 1910 and the park 

was christened Exposition Park. After the ceremony Collins wrote at the bottom 

of the list of contents placed in the cornerstone: "The box containing the above 

temized documents was retained in my possession and by me placed in the 



158 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Cavity of the Corner Stone as it was sealed and laid by the Grand Lodge F. and 
A.M. of California." It would have been interesting to have the full account of 
happenings at the ceremony because on December 22 Collins wrote to Spalding: 
"I hereby tender my resignation as Secretary of the Southern California Academy 
of Sciences.' 1 The letter was filed with a notation by Collins at the bottom: "The 
resignation was tendered by reason of certain transactions connected with the 
laying of the Corner Stone of the County Museum Building in Agricultural Park 
on Saturday, December 17, 1910. The Board of Directors refused to consider it 
and it was returned to me. December 24, 1910, Holdridge O. Collins." 

Apparently members of the Academy were working diligently on their collec- 
tions for the following editorial appeared in the Bulletin in January, 1911: 

The Directors are particularly gratified in being able, at this early 
date, to inform our members that immediately upon the completion of that 
Building, most valuable, interesting, beautiful and, in some respects, unique 
collections in Geology, Ichtheology [sic], Conchology, Ethnology, Ornithol- 
ogy, Botany, Entomology and Zoology will be speedily placed in position in 
the large exhibition room, for inspection by the curious and examination by 
the student. Some of these collections have been purchased, some have been 
gathered at the expense of the Academy, some have been presented and a 
few have been loaned. Professor Gilbert has been most assiduous in his labor 
of cleaning and mounting the fossils from Rancho La Brea, and he reports 
that he now has enough skeletons to occupy the space of one side of the hall. 
Among them are two complete giant ground-sloths, several sabre-tooth ti- 
gers, a lion, giant wolf, foxes, coyotes, probably a complete mastodon and 
camel, a turtle, deer, the giant ox; and, of the creatures of the air, numerous 
skeletons of the giant condor, vultures, hawks, eagles, owls, herons, geese, 
peacocks and an innumerable quantity of creeping, crawling, walking and 
flying smaller fry, which possessed this region as least two hundred thou- 
sands of years ago, and which nature has embalmed and preserved in their 
air tight graves. 

The Museum building was completed early in 1912 and exhibits were quickly 
moved in. Art occupied the west wing and rotunda, history was assigned the 
north wing and science was installed in the south wing. By July the following 
description of science exhibits appeared in the Bulletin: 

The progress made in cleaning, assembling and mounting the fossils from 
La Brea Rancho is most gratifying, and by September of this year we expect 
to present for public inspection an exhibit, which will surprise even those 
who have followed our work and which will convince the public of the great 
treasures we have exhumed from the zoological graves of the antiduluvium 
[sic] past. 

Mr. Raymond D. Jewett and Mr. Eugene J. Fischer have devoted most 
faithful, conscientious and skilled labor to the mounting of these fossils and 
the excellent taste and supervision of Mrs. Daggett have furnished most 
appropriate and beautiful desks, standards and cases for the mounts. 

At the present time, there stands in all its skeleton ferocity a sabre-tooth 
tiger six feet two inches in length by three feet in height; the gigantic ground 



SEVENTIETH ANNIVERSARY 



159 




The Museum was finished and opened in November, 1913. 



sloth measuring fourteen feet from its snout to the end of its tail standing 
over nine feet high; a mastodon, thirteen and one-half feet long and eight feet 
high with the cores of its tusks complete, its skull being seven and one-half 
feet in circumference; a female bison eleven feet two inches in length and 
five and one-half feet high; a giant wolf, about the size of the present Alaska 
timber wolf and resembling in its general build the German and Russian wolf; 
the skeleton of an African lion of the present day standing close to the sabre- 
tooth and showing by comparison the great size and strength of the latter. 
Work upon the male bison is progressing rapidly and, at this time, the spinal 
column is mounted thirteen feet in length, and with ribs eighteen inches 
longer than those of the female. Its skull measures forty-two inches between 
the cores of the horns, from which an estimate may be made of the enormous 
spread of these weapons of offense and defense as they once were in all their 
glory. The elephantus imperialis or imperiator, which exceeded in size the 
Mastodon and Mammoth, is represented by an enormous humerus and fe- 
mur, and hopes are entertained that among the tons of unassorted fossils 
remaining to be examined, enough of the other parts will be found to furnish 
an entire skeleton. 

In the cases are displayed the complete skeletons of a sabre-tooth tiger 
and giant wolf articulated, with the bones spaced and labelled for examina- 
tion, and a series of wolf bones of adults and young for study of comparative 
anatomy. 

There are also to be seen the humerus and pelvis of an extinct lion of ihe 



160 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




La Brea fossils dominated exhibits in the science wing of the Museum, 1913 



African type and we did have the skull, excavated from our concession in 
the Brea beds, but it has mysteriously disappeared. 

It would be tedious to enumerate everything now shown in our Exhibition 
room, but the cases contain several skulls of the sabre-tooth, the giant wolf, 
four distinct species of the extinct coyote, skulls and bones of birds and 
fowls of the air and water and parts of the camel and giant horse. 

Although this hall has not been thrown open to the public, Mr. Daggett, 
the general superintendent, is always glad to welcome any member of our 
Academy for an inspection of these treasures. 

Science and History wings of the Museum were opened in March, 1913, but 
the gala opening was on November 6. A great parade of city officials, marching 
bands and industrial floats slowly passed through the flower strewn streets of 
central Los Angeles and traveled to Exposition Park. Both the Museum and State 
Exposition Building were officially opened, the cornerstone for the Armory was 
laid and water from the new Owens River Aqueduct rose thirty feet in the air 
from the pipe that would soon supply the fountain in the sunken garden. That 
evening the Museum was filled with important citizens invited to a private opening 
(Los Angeles Times and Los Angeles Tribune, 1913; Sibley, 1978). This was the 
climax of many months of hard work by the Academy, begun in a tar pit at 
Rancho La Brea. 

On October 17, 1914 a gavel made from La Brea fossils was presented to 
President Arthur B. Benton, Dr. Astruther Davinson, second president of the 
Academy, designed the gavel using a ground sloth vertebra for the head, and 



SEVENTIETH ANNIVERSARY 



161 




Since 1976, re-mounted fossils, such as the saber-tooth cat, have been housed in Hancock Park in 
the new George C. Page satellite of the Natural History Museum. 



wood from the fossil McNab Cypress for the handle. And because the Academy 
had been so closely associated with Rancho La Brea, in 1915 the skull of the 
saber tooth cat was included in the seal of the organization. 

As the years passed the Academy continued its work at the Museum through 
representatives on the Board of Governors. However, curators and preparators 
took over the work of developing collections and exhibits. 

Two additions were added to the Museum in the twenties and then The Depres- 
sion arrived. The Academy suffered but survived with a membership reduced to 
less than 50 (Howard, 1957). 

Beginning in the early forties, meetings were held at the Museum on the ground 
floor beneath the old science wing. Here a potluck dinner preceded each monthly 
gathering. In the early sixties the growing Academy moved into the new Audi- 
torium where it met for a number of years before adopting its present pattern of 
meetings. 



162 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

In all those years few changes were made in the old Rancho La Brea exhibit 
in the south wing. Numerous plans were made for a museum to house the animals 
in Hancock Park. Allan Hancock had deeded the fossil bearing area to the County 
to preserve the animals and he hoped for a small museum in which to exhibit the 
fossils. Bond issues and budget requests failed so many times that the project 
seemed hopeless. Finally in the late sixties the fossil exhibit at the Museum was 
modernized with what little money was available. 

In 1973 Mr. George Page, a wealthy philanthropist, offered to build a museum 
at Hancock Park. The County accepted his gift and today an unique building 
stands in the midst of the area where struggling animals once died and early 
Academy members excavated for their remains. The Museum presents the story 
with film, paintings and reconstructions (Sibley, 1978). But the important exhibits 
are the skeletons — many are the same collected and mounted by the Academy 
for the first Museum exhibits. Now the animals stand in more natural positions 
and the remounting has removed most of the old iron bars and pipes. Thus the 
work of the Academy remains, evidence of the foresight of the early scientists, 
men and women who realized the significance of the fossils and secured public 
support for their preservation. 

Acknowledgments 

The author would like to express her appreciation to Dr. Hildegarde Howard, 
Chief Curator Emeritus, Los Angeles County Museum of Natural History, for 
her suggestions in the critique of this paper. Thanks also are extended to Mr. 
Henry Wylde and Dr. Harry Kelsey of the Museum for reading the material. 

Literature Cited 

Collins, Holdridge Ozro. 1909-1912. Minutes, correspondence, documents and early Bulletins of the 

Southern Academy of Sciences. On file in the Academy office, Los Angeles County Museum 

of Natural History, Los Angeles. 
Denton, William. 1877. On the Asphalt Beds Near Los Angeles, California. Proc. Boston Soc. Nat. 

Hist., 18:185-186. 
Guinn, J. M. 1909-1910. The Museum of Science and Art. Hist. Soc. of S. Calif., 8: Parts 1 and 2: 

5-8. 
Howard, Hildegarde. 1957. A History Commemorating the Fiftieth Anniversary of Incorporation, 

1907-1957: 20 pp. Booklet in Academy Files, Los Angeles County Museum of Natural History, 

Los Angeles. 
Los Angeles Tribune. 1913. March 17. 
Los Angeles Times. 1913. Part II, November 7. 

Merriam, John C. 1908. Death Trap of the Ages. Sunset Magazine, 21, 6:465-475. 
Robinson, Howard. 1953. The Beginning of the Museum. Los Angeles County Museum of History, 

Science and Art Alliance Quarterly, 10, 3:2-6. 
Sibley, Gretchen. 1978. 65 and Still Growing, Anniversary Annals of the Museum, from the First 

Bird's Nest to the New North Wing. The Natural History Museum Alliance, Terra, 16, 3:3- 

30. 
Stock, Chester. 1956 Revision. Rancho La Brea, a Record of Pleistocene Life in California. Los 

Angeles County Museum of Natural History, Sixth Edition, Science Series No. 20, Paleon. 

No. 11, 81 pp. 

Accepted for publication August 15, 1979. 



Bull. Southern California Acad. Sci. 
78(3), 1979, pp. 163-182 

The Salt Marsh and Transitional Vegetation of 
Bahia de San Quintin 

Leon F. Neuenschwander, Ted H. Thorsted, Jr., and Richard J. Vogl 

Abstract. — This study describes a relatively pristine salt marsh and ecotones 
that could serve as a comparison for understanding disturbed West Coast marsh- 
es. The salt marsh and terrestrial transitional vegetation of Bahia de San Quintin 
is composed of a low number of species with only a few dominants, and it is 
similar to southern California marshes. Species react to minor elevational differ- 
ences to create vegetation patterns or gradients, with each species reaching its 
highest frequency in the higher drier portions in a particular part of the marsh. 
Salicornia virginica has the broadest amplitude of occurrence. Plant cover in- 
creases with elevation which corresponds to decreases in tidal inundation, satu- 
rated soils, and salinity. Frequencies of species were low and variable in the 
marsh-upland transitions, but were high and consistent in the marsh proper. The 
presence of capillary extensions of tidal waters appears to be a critical factor in 
controlling the distribution of species in the ecotone areas. The establishment of 
submerged Zostera marina beds blocks the open body of water and initiates 
successive development that may culminate in terrestrial vegetation by contrib- 
uting to its stabilization and sedimentation. Piles of detached Zostera mat down 
the emergent vegetation which results in open pockets that are eventually reveg- 
etated. 

Department of Forestry Resources, University of Idaho, Moscow, Idaho 83843. 



In general, enclosed bays and lagoons of semiarid eastern Pacific regions be- 
tween Pt. Conception, California and Acapulco, Mexico have not been exten- 
sively studied (Johnson, 1958; Barnard, 1961; Mudie, 1970). 

California salt marshes are concentrated around San Francisco, Los Angeles, 
and San Diego where human population expansion has caused an alteration, rapid 
destruction (Darby, 1964; Anonymous, 1971), or pollution (Barnard, 1961; Vogl, 
1966). Around 1900 there were 28 sizeable estuaries between Morro Bay, Cali- 
fornia, and Ensenada, Mexico (Orme, 1973). Since then, 15 have been modified 
either slightly or moderately, 10 have been altered drastically and 3 have been 
destroyed (Speth, 1969; Macdonald, 1977a). Remaining marshes are threatened 
by highways or development in the U.S., and farming or settlement in Mexico, 
and the alterations of natural drainages and tidal prisms. Intact undisturbed salt 
marshes complete with their transition zones do not exist in southern California. 

The closest undisturbed salt marsh with an intact transitional vegetation to the 
southern California marshlands is Bahia de San Quintin. This extensive bay (Fig. 
1) is located at 116°00' longitude and 30°31' latitude which is approximately 273 
km (170 miles) south of the U.S. border on the Pacific side of Baja California. 
Mexico (Fig. 2). 



164 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 1. Location map showing Bahia de San Quintin, Baja, California. 



VEGETATION OF BAHIA DE SAN QUINTIN 



165 




Fig. 2. Map of Bahia de San Quintin. Local landmarks are given their Spanish names. Dotted 
areas represent the marsh and/or the transition areas studied. 



The primary objective of the study of Bahia de San Quintin was to provide the 
baseline information to make possible the comparisons with disturbed salt marsh 
ecosystems. Hence, this study could be of use in understanding and countering 
problems that arise from disturbance and pollution. 

The invertebrates of Bahia de San Quintin have received considerable attention 
(Barnard, 1962, 1964; Keen, 1962; King, 1962; Menzies, 1962; Reish, 1963; Mac- 
donald, 1967), but the emergent vegetation has been only briefly described (Daw- 
son, 1962). Vonder Haar (1972) studied the evaporite environment at Laguna 
Mormona adjacent to Bahia de San Quintin. Rundel et al. (1972) reported on the 



166 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

upland lichen community surrounding the Bay. Barnard (1962) and Dawson (1962) 
described the benthic flora of Bahia de San Quintin. 

Salt marshes located south of Bahia de San Quintin remain largely unstudied. 
The salt marsh vegetation of Laguna Guerro Negro and Ojo de Liebo has been 
briefly described by Phleger and Ewing (1962). An overview of Pacific Coast salt 
marshes (Macdonald and Barbour, 1974; Macdonald, 1977b; Breckon and Bar- 
bour, 1974) includes some general information regarding these Baja California 
salt marshes. 

Of all the studies mentioned, only those done in Newport Bay (Vogl, 1966) and 
Tijuana Marsh (Zedler, 1977) utilized quantitative methods (Thorsted, 1972). The 
remaining studies are based largely on partial floral lists or general observations 
of the vegetation. Except for Newport Bay (Vogl, 1966), the peripheral or tran- 
sitional vegetation bordering West Coast salt marshes has not been published 
(Neuenschwander, 1972). 

Description of Area 

Two types of salt marshes occur in Bahia de San Quintin: 1) extensive littoral 
marshes occurring in bay cul-de-sacs, large coves, and silted shallows, and 2) 
limited shoreline marshes occupying eroded shoreline edges, minor bay inden- 
tations, and the wet fringes of small valleys (Fig. 1). 

Littoral marsh physiography. — The large marshes are fairly level and charac- I 
terized by extensive drainage channels that dissect the marshes in a snake like ; 
fashion with numerous sharp bends and oxbows. The meandering main channels I 
or tidal cuts are usually 2 m or more deep and range from 3-10 m wide. The j 
vertical mud banks are honeycombed with crab (Hemigrapsus sp.) burrows and I 
the lower portions support small yellow sponges. The larger channel bottoms are 
flat and covered with firm muds that support vigorous stands of Zostera marina. I 
Water remains in these channels during normal low tides, and crabs, small fishes, I 
and bubble snails are abundant. The outside banks of channel curves are often I 
undercut by tidal flow, and short sections of bank are frequently slumped with I 
the marsh vegetation intact. Just offshore the emergent vegetation forms large 
areas. Extensive mud flats occur in a few of the marshes. 

The smaller, higher elevation channels are up to 3 m wide and 2 m deep and : i 
also meander dramatically. Bank sides are similar to those of main channels, but 
the bottoms tend to be U-shaped and consist of soft gray-brown muds and black 
ooze with a strong hydrogen sulfide odor. These channel bottoms are exposed 
during low tides and are generally devoid of vegetation but support numerous 
California horn snails (Cerithidea calif ornica). Some secondary channels in the 
shallow marshes at the base of Punta Azufre have completely slumped banks and 
support stands of Spartina foliosa, a condition not found elsewhere in the study 
area. 

Primary and secondary channel banks have shoulders that are usually elevated 
a few cm above the adjacent marsh surface. These mud levees are usually 0.25- 
1 m in width and are apparently produced as waterborne silts and clays precipitate 
in the dense Spartina foliosa stands that typically line the raised banks. 

In some marshes, channel curves have occasionally been cut off, forming short- 
er channels and leaving dead slough curves and oxbow bends, which appear to 



VEGETATION OF BAHIA DE SAN QUINTIN 167 

remain devoid of vegetation for long periods before being invaded by Batis ma- 
ritima, Spartina, and Salic ornia virginica. 

The uppermost drainage channels are shallow, short, and fairly straight. These 
vary from just a few cm to about 30 cm in depth and are completely vegetated. 
These tertiary drains usually enter the secondary channels at right angles and at 
some height above the secondary channel bottoms. 

The lower and middle elevations of the marshes contain occasional "rotten" 
spots, small open pockets in the otherwise dense vegetation usually created by 
suffocating mats of detached Zostera. These "rotten" spots are most abundant 
in the Punta Azufre marshes which are downwind from the prevailing northwest 
winds that push huge drifting quantities of Zostera into these marshes. Small 
vegetation free depressions or salt pans with impaired drainage are less common, 
being widely scattered in the middle zone of the most extensive marshes. 

Exposed mudflats (Doty, 1946) amidst Zostera beds and the emergent Spartina 
are usually covered with dense green algal mats composed of species of Cladoph- 
ora, Enteromorpha, Vaucheria, and patches of Ulva. Large numbers of California 
horn shells were observed on and beneath the extensive algal mats. These algae 
extend throughout the lower littoral and into the middle littoral zone wherever 
the emergent vegetation is not dense. 

The large littoral marshes have eroded mud banks along their open- water edges. 
In places these mud banks rise vertically 1.5-2 m above the open bay mudflats. 
They contain numerous invertebrate burrows, often worn smooth by tides and 
waves, and occasional slumped bank sections. In other places extensive bank 
erosion has taken place, resulting in Spartina covered banks that slope sharply 
or more gently toward open water. In the shallow marshes at the base of Punta 
Azufre, these open-water ledges are the lowest, consisting of a hard clay curb 
averaging about 30 cm in height. These clay banks are devoid of invertebrate 
burrows and are often free of plant life for 3 m or more inland, whereas other 
marshes support dense Spartina or, less commonly, Salicornia to the immediate 
edges of the eroded banks. 

The large marshes are underlain by muds that contain varied mixtures of clay, 
silt, volcanic ash, sand, and organic matter that are at least 2 m or more deep. 
The marsh surfaces are firm, except for "rotten spots," salt pans, and channel 
bottoms, and can be traversed on foot with ease during low tides. The marshes 
backed by playas and the shoreward portions of marshes contain particularly 
stable substrates. 

An exception to the firm substrates was found in the lava cove marsh in the 
north end of Bahia Falsa, the west bay arm. Portions of the middle zone of this 
marsh support sparse and chlorotic stands of annual Salicornia bigelovii plants 
and abundant "rotten spots" and shallow salt pans covered by dark platelets of 
salt and algae underlain by a spongy substrate. These peat like muds remain 
saturated or charged with water even when the tides have dropped the surround- 
ing water levels. 

The ever wet soil surface is firm enough to support the weight of a man. but 
quakes with each step. A stake can be plunged into the substrate 1-2 m and 
removed with ease, a feat that cannot be accomplished elsewhere. These boggy 
sites might relate to subterranean lava dikes that impair drainage, water blockage 



168 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 3. Bahia de San Quintin showing the typical littoral zones of the salt marsh. 

created by offshore mud flats, a particularly peaty substrate that has a high water 
retaining capacity, or to the subsurfacing of fresh waters draining the adjacent 
volcanic cone and lava beds. 

Shoreline marsh physiography. — The fringe marshes are discontinuous and lim- 
ited to shoreline edges. They tend to be less than 25 m wide and are generally 
devoid of tidal channels, although they are readily inundated during high tides. 
Some of the marshes are level, others grade gently toward the Bay, and still 
others are somewhat terraced. The mud and silt substrates are shallow and firm, 
usually less than 25 cm thick, and are often overridden by erosional deposits of 
sand, loam, volcanic ash, and lava from adjacent uplands. In many places these 
marshes have become established on inundated upland soils. 

The bayward edges of these narrow marshy strips are abrupt and consist of 
water worn clay ledges, exposed sand bars or lava rock rubble, stratified beach 
rock created by the freshwater deposition and lamination of calcium carbonate 
in sand and loam, or in a few places, beaches stabilized by accumulations of 
mollusk shells. Shoreline marshes which are exposed to constant wave action or 
strong tidal currents along the open water edges and generally lack Spartina. 

Transition zone physiography. — The most common ecotones between marsh- 
lands and upland vegetations occur on variable slopes in stabilized sandy loam 
hills. A few marshes are backed by abrupt aa' lava flows or sand dunes. Recently 
disturbed dunes have spread directly into the marshlands as a result of strong 
northwest winds. Most of the transition zones occur on relatively steep slopes, 
and vegetation changes are abrupt. 

The transition in Punta Azufre marsh extends onto a salt playa. The slope is 
nearly level and the transition zone is about 50 m wide. This transition is under- 
lain, at about 21 cm depth, with a salt layer or pan. 



VEGETATION OF BAHIA DE SAN QUINTIN 



169 




Fig. 4. Bahia de San Quintin showing the typical transitional vegetation with the capillary fringe 
and upland influence. 

Geology and climate. — The marine and quaternary geology of Bahia de San 
Quintin has been reported by Gorsline and Stewart (1962). 

A dry Mediterranean climate characterizes most of Baja California, but the 
presence of cold upwelling water along the coast (Dawson, 1951) results in a high 
frequency of fog (Meigs, 1966). Few climatic records are available, but temper- 
atures at Santa Maria del Mar, just inland from the Bay, range from 13°-23°C 
annually (Hastings, 1964, 1965; Hastings and Humphrey, 1969). The annual rain- 
fall is reported by Gorsline and Stewart (1962) to be between 5-10 cm per year, 
usually occurring in the winter. 

The wind is generally present in the mornings and early evenings, and is pre- 
dominantly from the northwest. These winds often move offshore fog clouds 
rapidly over the Bay. Winds peak from March through June with 20-30 knot 
velocities (Barnard, 1961). Tropical hurricanes occasionally make their way up 
the coast from the southwest during late summer and produce unusual amounts 
of rainfall. 



Methods 

The vascular plants present in the salt marshes of Bahia de San Quintin were 
identified according to Munz (1959) and Shreve and Wiggins (1964). Voucher 
specimens are on file in the California State University, Los Angeles (CSULA) 
herbarium. 

Seven littoral salt marshes and 14 upland transition zones were selected for 
quantitative study. Each of the selected marshes was divided into three littoral 
(Fig. 3) or marsh zones and one transitional zone (Fig. 4). Seven additional tran- 
sition zones were sampled to include the variability of the upland vegetation. The 



170 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

lower littoral zone is characterized by alternating periods of daily inundation and 
exposure and is always bound on the bayside by open water. The middle littoral 
zone is slightly higher and drier than the lower zone, but has characteristically 
soggy or water logged soils due to capillary action from tidal waters. It is inun- 
dated regularly during neap tides, storms, and windy weather. The upper littoral 
zone borders the uplands and is flooded only by unusually high or storm tides 
and lies above mean high tide. Soils are wetted during the higher tides, and soils 
and plants are subject to salt spray during rough weather. The transition or mar- 
itime zone (Vogl, 1966) is the ecotone between the upland vegetation and the 
lowland littoral zone vegetation. This zone is not inundated by tidal or storm 
waters. The lower limits of this zone are clearly delimited by a line of debris, 
flotsam, and dead Zostera marina deposited by the highest water of the year 
(referred to as the debris or storm line). The transition zone was arbitrarily limited 
to a 30 m wide perimeter inland of the debris line or the salt marsh edge. 

Each zone in each study marsh was sampled with 160 (25 x 25 cm 2 ) quadrats. 
This quadrat size was found to be the maximum usable to obtain meaningful 
frequencies (less than 100%) for the common salt marsh species (Vogl, 1966). 
Percent plant cover was visually estimated for 16 quadrats in each zone. 

Transition zones were sampled differently to help determine the vegetation 
position and zonation within each transition zone, four representative line inter- 
cepts were taken on different slopes. The line intercept samples were obtained 
by placing a 30 m tape at right angles to the shoreline, starting at the debris line. 
The horizontal distance each species occurred above this high tide mark, the 
amount of the line covered by each species, and the slope angle was recorded. 

Depth to soil moisture was taken in each transect of the transition zone. Depth 
to the capillary fringe of the salt marsh water table was obtained by boring soil 
cones at 1 m intervals along 30 m lines placed at right angles across the transition 
zone. Each hole was dug to a maximum of 2 m. 

Water retaining capacities were obtained for the various transitional soil types 
using the method presented in Curtis and Cottam (1962). 

Results 

The phytogeographic area of Bahia de San Quintin (Wiggins, 1960) encompass- 
es coastal salt marsh, alkali flat, coastal strand, and coastal desert scrub (Shreve 
and Wiggins, 1964; Munz, 1968). The last two communities comprise the upland 
vegetation around the Bay. The transition zone between the salt marsh and the 
surrounding uplands contains floral elements of all four communities. 

The marsh flora is composed of only 17 emergent phanerogam species (Table 
1). All of the species, with the possible exception of Mesembryanthemum nodi- 
florum, are indigenous (Moran, 1950). The transitional flora is represented by 33 
species from 28 genera and 19 families. Of these species, 12 are characteristically 
found in salt marsh, and 21 are normally associated with desert and coastal up- 
lands. 

Frequency -Cover. — Average frequency and cover values (Table 1 and 2) show 
that only eight species are common in the marshes. Salicornia virginica, Spartina 
foliosa, Batis maritima, and Monanthochloe littoralis are dominants, with Fran- 
kenia grandifolia, Salicornia subterminalis , Suaeda calif ornica and Salicornia bi- 



VEGETATION OF BAHIA DE SAN QUINTIN 



171 



Table 1. Average percent frequencies for all species in each zone. Frequency range for the seven 
stands for the littoral zones and 14 stands in the transition zone are in parentheses. 1 







Littoral 






Species 


Lower 


Middle 


Upper 


Transition 


Spartina foliosa 


90.8% 
(86-97) 


6.9% 

(0-21) 


— 


— 


Salicornia bigelovii 


0.8% 
(0-6) 


21.5% 
(0-96) 


0.1% 
(0-1) 


— 


Triglochin concinna 


0.8% 
(0-6) 


9.5% 
(0-33) 


0.1% 
(0-1) 


— 


Salicornia virginica 


34.4% 
(20-65) 


86.7% 
(71-93) 


52.2% 
(20-86) 


3.7% 
(0-21) 


Batis maritima 


27.0% 
(13-36) 


55.2% 
(9-86) 


4.9% 
(0-16) 


0.0% 
(0-0) 


Frankenia grandifolia 


6.8% 
(1-16) 


29.0% 
(14-46) 


43.3% 
(24-84) 


4.6% 
(0-16) 


Suaeda californica var. californica 


4.1% 
(1-8) 


20.4% 
(5-31) 


25.8% 
(9-50) 


1.4% 
(0-8) 


Jaumea carnosa 


1.3% 
(0-5) 


3.3% 
(6-18) 


6.3% 
(0-28) 


0.2% 
(0-2) 


Salicornia subterminalis 


0.1% 
(0-1) 


0.9% 

(0-4) 


46.5% 
(5-81) 


38.7% 
(13-74) 


Monanthochloe littoralis 


— 


27.4% 
(0-12) 


75.1% 
(22-97) 


42% 
(7-49) 


Limonium californicuin 
var. mexicanum 


— 


4.0% 
(0-12) 


13.7% 
(0-34) 


7.3% 
(0-62) 


Cressa truxillensis 
var. vallicola 


— 


— 


3.5% 
(0-16) 


0.9% 

(0-7) 


Atriplex watsonii 


— 


— 


1.5% 
(0-4) 


3.9% 
(0-18) 


Cuscuta salina 


— 


0.4% 
(0-3) 


0.1 
(0-1) 


— 


Meseinbryantheinum nodiflorum 


— 


— 


0.1% 

(0-1) 


2.6% 
(0-25) 


Frankenia palmeri 


— 


— 


— 


31.7% 
(4-68) 


Lycium brevipes 


— 


— 


— 


6.2% 

(3-11) 


Atriplex julacea 


— 


— 


— 


8.8% 

(1-24) 


Haplopapptts venittits 


— 


— 


— 


0.195 

(0-1) 


Euphorbia inesera 


— 


— 


— 


0.7% 

(0-4) 


Diulleya brittonii 


— 


— 


— 


0.495 

(0-2) 



172 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Table 1. Continued. 



Littoral 
Species 



Aes cuius parry i 

Distich lis spicata 

Juncus acutus 

Allenrolfea occidentalis 

Mammillaria dioica 

Machaerocereus gummosus 

Echinocereus maritiinus 

Lichens 

Green algae mat 

Bare ground 



Lower 


Middle 


Upper 


Transition 


— 


— 


— 


0.1% 
(0-1) 


— 


— 


— 


0.4% 
(0-5) 


— 


— 


— 


0.1% 
(0-1) 


— 


— 


— 


0.3% 
(0-3.2) 


— 


— 


— 


0.1% 
(0-1) 


— 


— 


— 


0.1% 
(0-1) 


— 


— 


— 


0.1% 
(0-1) 


— 


— 


— 


6.9% 

(1-17) 


67.8% 


48.6% 


— 


— 


(6-90) 


(0-94) 






0.6% 


0.6% 


0.2% 


10.4% 


(0-3) 


(0-1) 


(0-1) 


(3-26) 



1 Each value in the littoral zones is based on 1120 quadrats and each value in the transition zone 
is based on 2240 quadrats. 



gelovii less frequent. Frequency Cover of the common species varied in the 
different zones within the marsh zonation. 

The lower littoral zone contains only 9 species, and is dominated by Spartina 
(90.8% -47.0%), Salicornia virginica (34.4% -33.1%), and Batis (27.0%-6.4%). 
Spartina characterizes the lower littoral zone. Its growth is particularly green 
and robust in comparison to stands farther north, perhaps indicating the absence 
of toxic pollutants in the Bay (Mobberley 1956). Spartina pioneers the algae 
covered mudflats and extends to the upper or landward limits of daily inundation. 
Salicornia virginica and Batis also occur throughout this zone. Sometimes they 
are thinly dispersed throughout the understory of the uniformly tall (1-1.35 m) 
Spartina grass stems, but they may occur in localized dense colonies where the 
Spartina is sparse or they may grow on the open mudflats beyond the typically 
dense band of Spartina. 

The middle littoral zone contains 12 species. Salicornia virginica (86.7% -22.7%) 
dominates, with lesser amounts of Batis (55.2%- 16.8%) (Table 1 and 2). These 
most widespread littoral species transcend the middle zone in both directions. 
Salicornia virginica dominates this zone with its robust and dense clonal growth. 
Batis typically pioneers poorly drained pans and occupies low flats, slight depres- 
sions, and old drainage channels. 



VEGETATION OF BAHIA DE SAN QUINTIN 



173 



Table 2. Average percent cover for the common species in each zone. The range for all seven 
stands is given in parentheses. 1 



Species 



Littoral 



Lower 



Middle 



Upper 



Transition 



Spartina foliosa 

Salicornia virginica 

Batis maritima 

Frankenia grandifolia 

Jauinea carnosa 

Suaeda californica 
var. californica 

Triglochin concinna 

Limonium californicum 
var. mexicanum 

Salicornia bigelovii 
Monanthochloe littoralis 
Salicornia subterminalis 
Atriplex watsonii 
Frankenia palmeri 
Lycium brevipes 
Atriplex julacea 
Lichens 

Green algae mat 
Bare ground 
Zostera litter 



47.0% 
(42-53) 

20.2% 
(6-43) 

6.4% 
(4-8) 

2.3% 
(0-5) 

1.1% 
(0-8) 

1.0% 
(0-3) 



81.8% 
(59-100) 

3.0% 
(0-11) 

3.0% 
(0-10) 



1.0% 
(0-4) 

33.1% 
(27-61) 

16.8% 

(3-35) 

9.9% 

(2-17) 

1.0% 
(0-4) 

6.9% 
(3-11) 

5.2% 
(0-25) 

2.1% 
(0-6) 

6.9% 
(0-41) 

1.0% 
(0-8) 



22.4% 
(0-56) 

14.0% 
(3-30) 



22.7% 
(5-58) 

1.2% 
(0-6) 

13.8% 
(0-35) 

4.3% 
(0-24) 

8.3% 
(0-22) 



6.0% 
(0-17) 



45.7% 
(1-81) 

22.7% 
(0-45) 

1.6% 
(0-7) 



8.7% 
(0-24) 

1.2% 
(0-9) 



2.5% 
(0-21) 



1.5% 
(0-7) 



1.6% 
(0-13) 



22.0% 
(0-55) 

20.5% 
(6-62) 

2.2% 
(0-16) 

19.0% 
(6-30) 

2.7% 
(0-6) 

2.0% 
(0-6) 

2.2% 
(0-9) 



29.2% 
(12-42) 



1 Each value is an average based on 
transition zone. 



1 120 quadrats for the littoral zones and 2240 quadrats in the 



174 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Of the less frequent species, Frankenia grandifolia occurs in almost pure stands 
in soggy areas, while Monanthochloe littoralis and Suaeda californica are most 
frequent on slightly higher and drier ground. Salicornia bigelovii is confined to 
low, wet flats and "rotten spots," sometimes occurring in pure stands or sharing 
the sites with Triglochin concinna and Batis. Salicornia virginica often occurs 
on the edges of, or occasionally within, these poorly drained flats. Cuscuta salina 
parasitizes Salicornia virginica in this zone, particularly, growing on drier sites. 
Green algae have a frequency of occurrence of nearly 50% (Table 1). 

The upper littoral zone has 14 species, with Monanthochloe (75.1% -45.7%) 
dominant (Table 1 and 2). Salicornia subterminalis (46.5% -2.7%) attains its high- 
est frequency cover in the upper and drier sections, while Salicornia virginica is 
most common in the lower and more moist regions. Frankenia frequently occurs 
in pure aggregations on slight rises, or intermingles with Monanthochloe and Sal- 
icornia virginica in wetter areas. Suaeda and Limonium calif ornicum occur as 
scattered individuals throughout this zone, particularly on drier sites. Jaumea 
carnosa occurs in wet and dry areas, spreading by vegetative runners with weak 
upright branches. Exceptionally dense and vigorous aggregations of Jaumea grow 
on rough lava sites overlain with sandy silt. Cressa truxillensis mixes with Mon- 
anthochloe on the driest places. Cuscuta parasitizes Cressa and Salicornia vir- 
ginica growing on dry sites, but was not observed on any other species. Atriplex 
watsonii and Mesembryanthemum nodiflorum were only rarely encountered 
along the extreme landward edge of this zone. The upper limits of this zone are 
marked by a band of debris and decomposing Zostera litter. 

The transition zone is dominated by Salicornia subterminalis (38.7% -20.5%) 
and Monanthochloe (42.7% -22.0%). Nine other salt marsh species have frequen- 
cies of occurrence of less than 8% (Table 1), and are concentrated in the lowest 
portions of this ecotone zone near the debris line. Salicornia subterminalis also 
occurs in alkaline substrates around the Bay and in alkali flats near the coast. 

The dominant species in the upland parts of the transition zone are Frankenia 
palmeri (31.7%- 19.0%), Atriplex julace a (8.8%-2.0%), and Lycium brevipes 
(6.2% -2.7%). Frankenia is the most widespread species, and consistently occurs 
lower in the transition zone than the other upland species. An additional 13 upland 
species have frequencies less than 2% (Table 1). Mesembryanthemum nodiflorum 
(2%) only occurs in the disturbed portions of upper sandy loam transitions. The 
coastal scrub species Euphorbia mesera, is found in ash, cinder, and sandy loam 
substrates associated with volcanic outcrops. Lichens are attached to rocks, soils, 
and some plants in the upper portions of the transition zone (Rundel et al., 1972). 

Average percent cover values generally correspond to frequency values in the 
littoral and transition zones. Total cover is 74.5% in the transition zone, 78.0% 
in the lower littoral zone, 83.9% in the middle littoral zone, and 126.3% in the 
upper littoral zone. Both cover and the number of species increased from the 
lower or wetter to the higher or drier portions of the marsh, as found in Newport 
Bay (Vogl, 1966). However, in the transition zone, cover decreased from the 
lower portion near the marsh toward higher elevations. Combined salt marsh 
species account for 48.3% of the cover in the lower part of the transition which 
is more than the cover of the combined upland species (25.8%). 

The dominant upland species are scattered throughout the upper portion of the 
transition. Frankenia palmeri extends the lowest toward the marsh (Fig. 5). Fran- 



VEGETATION OF BAHIA DE SAN QUINTIN 



175 



DEBRIS LINE 

TRANSITION 



SALT MARSH 



Species : 


Suaeda californica — 

Salicornia subterminalis — 

Monanthochloe littoralis — 

Frankenia grandifolia -I 

Atriplex watsonii -| 

Limonium californicum 

Cressa truxillensis 

Frankenia palmeri 

Lycium brevipes 

Atriplex julacea 

Abronia umbellata 




Fig. 5. Dominant position for each species within the transition zone (30 m) on the mean slope. 
Cover is expressed by standard error of the mean as a solid bar and the ranges are the dotted lines. 



kenia palmeri and Lycium have approximately the same range (10.5-18.0 m from 
debris line) and approximately the same cover (15.8%). Atriplex julacea's cover 
(7.3%) is about half that of Frankenia palmeri and Lycium. Atriplex julacea 
ranges higher in the transition zone (15.9-19.8 m) than the other upland domi- 
nants. 

The line intercept cover in the transition zone is graphically represented in 
Figure 5. Species cover, composition and vertical stratification vary with the 
slope. Slopes on sandy loam substrates range from 2-30%. 

The width of the transition is a result of slope. Steep slopes produce narrow 
transitions. Species overlap and vertical stratifcation are confined to lower por- 
tions of the transition zone. Steep slopes also support more extensive upland 
cover, whereas gentle slopes produce long transitions extending species overlap 
and vertical stratification farther inland. Gentle slopes contain more cover of salt 
marsh species. Salt marsh species are essentially confined to the lower portion 
of the transition defined by the capillary fringe of the Bay, while upland species 
occupy the upper portion. Plants in the lower portion of the transition are low in 
stature, dense in cover, and luxuriant in growth. Plants in the upper portion of 
the transition are sparse but larger in stature. 

Upland cover values taken by line intercepts (Fig. 5) do not correspond to 
those taken by quadrats (Table 2), indicating variability in the upland vegetation. 
Cover values for the dominant salt marsh species are approximately the same for 
line intercepts as for quadrats, indicating less variability. 

Three common salt marsh species are present in line intercepts, and are re- 



176 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



+ 2m- 



TRANSITION 



SALT MARSH 




SALT MARSH 



V. 



25m 



J 



CAPILLARY FRINGE 

V 



J 



MARITIME 

Fig. 6. Average depth to wet, moist and dry sandy loam soil taken by soil cores. The vegetation 
position of Fig. 5 can be overlaid on the soil moisture characteristics of Fig. 6 so that the transition 
can be subdivided into the capillary fringe and maritime zones. 



stricted to the first 12 m inland, the marsh debris line. Salt marsh species on the 
lowest sandy loam slopes extend a maximum of 30 m above the debris line. This 
maximum range is seldom attained. 

Transition soils. — Depth to wet or moist soil is proportional to the slope. Depth 
to moisture increased with the horizontal and vertical distance from the Bay (Fig. 
6). The lower portion of the transition, occupied by salt marsh type species, is 
an area in which roots of these plants receive moisture from the Bay, at least 
during high tide. 

A wetted soil surface is a visual expression of the capillary fringe zone around 
the Bay. The inland distance from the Bay for the capillary fringe is about 4.75 
m. At that distance, the soil moisture suddenly drops to a depth of .75 m where 
it then extends horizontally at that depth for 6 m. Beyond 10 m from the debris 
line, the soil moisture suddenly drops off below the rooting depth of most upland 
species. 

Exceptions to the above soil moisture regime are promoted by formations of 
salt pans. A salt pan was formed in one area on a shallow slope (3%) beginning 
about 5 m inland from the Bay and extending the entire length of the transition 
at .2 m below the soil surface. The salt pan induces the extension of salt marsh 
vegetation into the transition area normally occupied by upland species. 

The extension of salt marsh vegetation inland appears to be a result of the soil 
particle-size distribution and water holding capacity. Sandy loam soil near the 
Bay had a water holding capacity of 33.4%, but at an elevation above the Bay, 



VEGETATION OF BAHIA DE SAN QUINTIN 177 

it was higher (40.8%). Dune and volcanic substrates have lower water holding 
capacities with 32.4 and 24.1%, respectively. The higher the water holding ca- 
pacity, the more well developed and the farther inland the capillary fringe ex- 
tends. Also, the higher the water holding capacity, the richer and more abundant 
is the upland vegetation. However, with lower water holding capacity, the upland 
vegetation is more frequently mixed with salt marsh vegetation. 

Discussion 

The vegetation of Bahia de San Quintin is of interest because of its almost 
pristine state and its unique position within a fog-desert. Local upwellings of 
relatively cool waters off the Bay produce frequent fog, reduce rain along the 
immediate coastline, and produce enriched waters. Strong and persistent onshore 
winds remove water, soils, sediments, and salts (Brongerama and Sanders, 1948; 
Dawson, 1951; Barnard, 1961; Phleger and Ewing, 1962). As a result, the bay side 
vegetation is sparse and has a large number of lichens (Rundel et al., 1972). Field 
observations suggest that the prevailing rapidly moving fogs contribute moisture 
to the transitional and upland plants and soils in the form of condensation and 
fog drip, but that reproduction in these uplands is only observed after the infre- 
quent winter rains. Except for annual species, seed germination and new plant 
establishment in the littoral zone is probably confined to the unusual wet periods 
produced by tropical hurricane rains, since many salt marsh plants need abundant 
fresh water for successful sexual reproduction (Barbour, 1970; Barbour and Da- 
vis, 1970). 

The vegetation of salt marshes characteristically has few species with large 
numbers of individuals per species, illustrating the principle that extreme envi- 
ronments tend to produce simple systems. In Bahia de San Quintin, the salt 
marshes are dominated by the same few families and genera that dominate salt 
marshes worldwide. 

Of the 17 salt marsh species found in Bahia de San Quintin, five represent 
tropical floristic elements that occur as far north as Santa Barbara, California. 
Monanthochloe littoralis and Batis maritima are the most important tropical 
elements. The remaining 12 species are widespread with ranges extending to San 
Francisco Bay, and some occur as far north as Alaska. When Pacific Coast marsh- 
es are compared (Macdonald and Barbour, 1974; Barbour et al., 1975; Macdonald, 
1977), those of Bahia de San Quintin and Southern California contain more 
species than the more northern marshes. 

Marsh gradients and patterns. — The zonal patterns apparent in the littoral 
environment are closely tied to elevational gradients. Where the gradients are 
gentle, the zones are expansive and diffuse as in the large littoral marshes, and 
where the elevation changes are sharp, the zones are narrow and distinct as in 
some of the shoreline marshes. Small changes in elevation can result in large 
changes in exposure or inundation, salinity, pH, temperature, and the vegetation 
(Stevenson and Emery, 1958). Numerous studies have demonstrated that littoral 
vegetational responses are largely a result of the inundation-exposure regime 
working in concert with physical gradients (Purer, 1942; Doty, 1946; Reed, 1947; 
Bourdeau and Adams, 1956; Chapman, 1960; Adams, 1963; Vogl, 1966; Zedler, 
1977). 

These frequency distributions (Fig. 7) illustrate that most of the common 



178 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



DOMINANT SPECIES 




-£ 80 

<D 
O 

<D 

0- 60 



40 



20 



Monanthocloe 
ittoralis 




LOWER 



MIDDLE 



UPPER 



TRANSITION 



Littoral Zones 



Fig. 7. Average percent frequencies of the dominant species plotted by zones. 



species are not restricted to one zone. Salicornia virginica, has the broadest 
amplitude as in Newport Bay, California (Vogl, 1966). In some places Salicornia 
virginica even thrives on open mudflats below the Spartina belt; it also grows 
vigorously in some of the highest and driest parts of the upper littoral zone. 



VEGETATION OF BAHIA DE SAN QUINTIN 179 

Batis maritima is also a widespread species, dominating all low, wet, and poorly 
drained areas, regardless of zone. Frankenia grandifolia is present in all three 
zones but favors higher and drier sites. Spartina and Monanthochloe represent 
low and high extremes within the emergent vegetation. Frankenia palmeri is 
restricted to the transition or maritime zone. 

Apparently, the vegetational composition of the salt marsh is not a cohesive 
entity explained simply by zonation, but is a complex of individual species re- 
sponding to environmental gradients. Relatively simple elevational gradients are 
altered by rain runoff, sediment deposition, tidal action, wave action, storms, 
erosion, and changing water levels. The resulting micro-topographic differences 
create variable vegetational patterns as each species selects favorable sites. 

Vegetational patterns dictated by various environmental gradients are compli- 
cated by the vegetative reproduction common to almost all salt marsh species. 
Huge clones sometimes give the appearance of zones. Individuals often spread 
vegetatively into less favorable sites. 

Transition zone. — Dominant upland species found in the upland-marsh ecotone 
are succulent halophytes, and all species appear to be salt tolerant (Boyce, 1954). 
It appears that the salt marsh species present in the lower transition zone are 
controlled by the same factors that affect them in the salt marsh proper, namely 
salt concentrations, soil aeration, and saltwater inundation (Chapman, 1960; Vogl, 
1966). Inundation in the transition zone is, however, reduced to the capillary 
fringe activity of saltwater, and salt concentrations are largely products of evap- 
oration, capillary action, salt spray, and fog drip. Hence, distribution of species 
at Bahia de San Quintin in transitional areas is not random and species are con- 
sidered to be regulated by these environmental factors. 

The presence of capillary extensions of tidal waters appears to be a critical 
factor controlling the distribution of species in the ecotones. Near-surface cap- 
illary moisture promotes the growth of salt marsh species, particularly Salicornia 
subterminalis and Monanthochloe, or perhaps excludes the growth of upland 
species, thereby allowing the salt marsh species to extend above the debris line 
and salt marsh proper. Steep slopes reduce the sub-irrigation properties of the 
tidal waters and permit the growth of upland species which are usually widely 
spaced Frankenia palmeri and Atriplex julacea. Slightly higher transitional areas 
support Lycium brevipes with lichen covered branches. The capillary action ap- 
pears to extend slightly higher in the finer silty ash and sandy loam soils than in 
the sand and lava substrates. 

Investigation of this upland marsh ecotone indicates that the sub-irrigation and 
capillary action of tidal waters is a significant factor regulating species distribu- 
tions. Therefore, the transition should be divided into a capillary fringe zone and 
a maritime zone. 

The transition appears to be the focal point of animal activity for both the 
terrestrial and salt marsh fauna of Bahia de San Quintin. Numerous animal trails 
cross the transition. During high tides and severe storms, the capillary fringe may 
provide critical cover-habitat for salt marsh birds. Also, some upland animal 
species apparently feed in the marsh. Preliminary observation suggests that the 
transition is an important area for animal use, but this area has not been studied 
in any Pacific West Coast salt marsh. 

Role o/Zostera. — Zostera marina occurs in deeper water beyond the emergent 



180 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

species. Zoster a beds in the Bay have been described and mapped by Dawson 
(1962). Contrary to the usual way that Zostera grows on the Pacific Coast, San 
Quintin Zostera beds are exceptionally dense and grow in unusually shallow 
waters that expose plants and mud during lowest tides. 

A factor thought to be responsible for the biotic initiation of salt marshes is the 
establishment of Zostera marina beds in the open water of enclosed bays, es- 
tuaries, and lagoons. Beds of Zostera retard the erosional tidal flux and result in 
a continual and ever increasing accretion of sediments. This accumulation of 
sediments usually resists storm excavation because of the extensive perennial 
Zostera roots and initiates the successional development of an open body of 
water that may culminate in terrestrial vegetation. 

Zostera also contributes to the enrichment of coastal waters by the annual 
productivity of above ground plant parts that die back each winter and form 
detritus (Redfield, 1965). Often, during violent storms or die-back periods, huge 
windrows of dead Zostera are carried landward onto the emergent vegetation, 
particularly in the Spartina belt. The windrows mat down the vegetation and 
eventually kill the underlying plants causing bare spots in the emergent vegeta- 
tion. These openings are eventually revegetated, usually by Batis and Salicornia 
virginica. These gap phase perturbations and their resulting re vegetation are prob- 
ably responsible for the slightly larger number of species in the lower littoral zone 
of Bahia de San Quintin compared to Newport Bay where Zostera is uncommon 
(Vogl, 1966). In some locations, e.g., the downwind Punta Azufre marshes, Zos- 
tera debris piles completely kill off the bayside emergent vegetation, thus expos- 
ing extensive mudflat edges to erosional forces of tides and waves. 

Occasionally dried Zostera leaves resist decay and become incorporated into 
mud and silt deposits, enhancing their ability to withstand erosion. The annual 
increments of decomposed Zostera products must also contribute to the enrich- 
ment and siltation of the emergent salt marsh vegetation. Windrows of Zostera 
litter persist year-round at the upper debris line, and were noted to serve as 
protective cover for insects, marine invertebrates, small mammals, and birds. 
Previous salt marsh studies have been conducted in locations where Zostera beds 
were absent or severely disturbed, and therefore, have overlooked the role that 
Zostera plays in the ontogeny of Pacific Coast salt marshes. The role of Zostera 
needs further study to determine these relationships. 

Acknowledgment 

Manuscript preparation was supported by the College of Forestry, Wildlife and 
Range Sciences, University of Idaho, Forest, Wildlife and Range Experiment 
Station, Contribution No. 173. 

Literature Cited 

Adams, D. A. 1963. Factors influencing vascular plant zonation in North Carolina salt marshes. 

Ecology 44:445-456. 
Anonymous. 1971. Southern California estuaries and coastal wetlands, endangered environments. 

Bureau of Sport Fisheries and Wildlife, U.S. Dept. Int. Portland, Ore. 15 p. 
Barbour, M. G. 1970a. The flora and plant communities of Bodega Head, California. Madrono 

20:289-336. 
. 1970b. Is any angiosperm an obligate halophyte? Amer. Midland Nat. 84:105-120. 



VEGETATION OF BAHIA DE SAN QUINTIN 181 

, and C. B. Davis. 1970. Salt tolerance of five California salt marsh plants. Amer. Midland 

Nat. 84:262-265. 
, DeJong, T. M., and A. F. Johnson. 1975. Additions and corrections to a review of North 

American Pacific Coast beach vegetation. Madrono 23:130-134. 
Barnard, J. L. 1961. Benthic marine exploration of Bahia de San Quintin, Baja, California, 1960-61. 

Pacific Nat. 3:249-274. 
. 1962. Benthic marine exploration of Bahia de San Quintin, Baja, California, 1960-61. No. 

6. Pacific Nat. 3:250-274. 

. 1964. Marine amphipod(s) of Bahia de San Quintin, Baja, California. Pacific Nat. 4:55-139. 

Bourdeau, P. F., and D. A. Adams. 1956. Factors in vegetational zonation of salt marshes near 

Southport, N.C. Bull. Ecol. So. Amer. 37:68. 
Boyce, S. G. 1954. The salt spray community. Ecol. Monogr. 24:29-67. 
Bradshaw, J. S. 1968. Report on the ecological relationships in Los Penasquitos Lagoon and salt 

marsh areas of Torrey Pines Reserve. California State Division of Beaches and Parks, Contract 

No. 4-05094-033. 113 p. 
Breckon, G. J., and M. G. Barbour. 1974. Review of North American Pacific Coast beach vegetation. 

Madrono 22:333-360. 
Brongersma-Sanders, M. 1948. The importance of upwelling water to vertebrate paleontology and 

oil geology. Verh. Akad. Wet., Amst., Afd. Nat. (tweede sect.) 45:1-112. 
Chapman, V. J. 1960. Salt marshes and salt deserts of the world. The University Press, Aberdeen. 

Leonard Hill Ltd., London. 393 p. 
Clark, J. R. 1977. Coastal ecosystem management. John Wiley and Sons. 928 p. 
Curtis, J. T., and G. Cottam. 1962. Plant ecology workbook. Burgess Co. Minneapolis, Minn. 94 p. 
Darby, R. 1964. The Last Chance Lagoon. Westways 56:30-31. 
Dawson, E. Y. 1951. A further study of upwelling and associated vegetation along Pacific Baja 

California, Mexico. J. Mar. Res. 10:39-58. 
. 1962. Benthic marine exploration of Baja de San Quintin, Baja, California, 1960-61. No. 7. 

Marine and marsh vegetation. Pacific Nat. 3:275-280. 
Doty, M. S. 1946. Critical tide factors that are correlated with the vertical distribution of marine 

algae and other organisms along the Pacific Coast. Ecology 27:315-328. 
Gorsline, D. S., and R. A. Stewart. 1962. Benthic marine exploration of Bahia de San Quintin, Baja, 

California, 1960-61. No. 8. Marine and Quaternary Geology. Pacific Nat. 3:282-319. 
Hastings, J. R. 1964. Climatological data for Baja California. Technical reports on the meteorology 

and climatology of arid regions. No. 14. Tucson. Univ. of Arizona Institute of Atmospheric 

Physics, p. 204-223. 
. 1965. Seasonal precipitation regimes in Baja California, Mexico. Geografiska Annaler 

47A:205-223. 
, and R. R. Humphrey. 1969. Climatological data and statistics for Baja California. Technical 

reports on the meteorology and climatology of arid regions. No. 18. Tucson. Univ. of Arizona 

Institute of Atmospheric Physics. 196 p. 
Hinde, H. P. 1954. The vertical distribution of salt marsh phanerograms in relation to tide levels. 

Ecol. Monogr. 24:209-225. 
Johnson, B. H. 1958. The botany of the California Academy of Science expedition to Baja California 

1941. Wasmann J. Biol. 16:217-318. 
Keen, A. M. 1962. A new west Mexican subgenus and new species of Montacitidae (Mollusca: 

pelecypoda), with a list of Mollusca from Bahia de San Quintin. Pacific Nat. 3:321-328. 
King, Roy E. 1962. A new species of Parahemiorus and notes on Tobolovesicula lindbergi (Trem- 

atoda: Hemioridae) from fishes of Bahia de San Quintin. Pacific Nat. 3:329-336. 
Macdonald, K. B. 1967. Quantitative studies of salt marsh mollusc faunas from the north Pacific 

coast. Ph.D. thesis. Scripps Institution of Oceanography, Univ. California at San Diego. 
. 1977a. Coastal Salt Marsh, p. 263-293. In M. G. Barbour and J. Major (Eds.). Terrestrial 

Vegetation of California. John Wiley and Sons, Inc., N.Y. 
. 1977b. Plant and animal communities of Pacific North American salt marshes. V. J. Chapman 

(Ed.) Wet Coastal Ecosystems: Chapt. 8 p. 167-191. Elsevier Scientific Pub. Co.. Amsterdam. 
, and M. G. Barbour. 1974. Beach and salt marsh vegetation North American Pacific Coast. 

pp. 175-233. In R. J. Reinold and W. H. Queen. (Eds.) Ecology of Halophytes. Academic 

Press, N.Y. 



182 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Meigs, P. 1966. Geography of Coastal deserts. United Nations Educational, Scientific and Cultural 

Organization, Place de Fontenoy Paris. 110 p. 
Menzies, R. J. 1962. The marine isopod fauna of Bahia de San Quintin, Baja California, Mexico. 

Pacific Nat. 3:337-348. 
Mobberley, D. G. 1956. Taxonomy and distribution of the genus Spartina. Iowa State College. J. 

Sci. 30:474-574. 
Moran, R. 1950. Mesembryanthemum in California. Modrono 10:161-163. 
Mudie, P. J. 1970. A survey of the coastal wetlands vegetation of north San Diego County. Calif. 

State Res. Agency. Wildl. Manage. Admin. Rep. 70-4. 18 p. 
Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley. Calif. 1681 p. 
Neuenschwander, L. F. 1972. A phytosociological study of the transition between salt marsh and 

terrestrial vegetation of Bahia de San Quintin. M. A. thesis. California State Univ., Los An- 
geles. 70 p. 
Orme, A. R. 1973. Coastal salt marshes of northwest Baja California, Mexico. Office of Naval 

Research, Technical Report 0-73-2, Contract No. N 000-1 4-69- A-0200-5003. 
Phleger, C. F. 1971. Effect of salinity on growth of a salt marsh grass. Ecology 52:908-911. 
Phleger, I. B., and G. C. Ewing. 1962. Sedimentology and oceanography of coastal lagoons in Baja 

California, Mexico. Bull. Geol. Soc. America. 73:145-181. 
Purer, E. A. 1942. Plant ecology of the coastal salt marshlands of San Diego County, California. 

Ecol. Monogr. 12:83-111. 
Redfield, A. C. 1965. Ontogeny of a salt marsh estuary. Science. 147:50-55. 
Reish, D. J. 1963. A quantitative study of the benthic polychaetons annelids of Bahia de San Quintin, 

Baja, California. Pac. Nat. 3:397-436. 
Reed, J. F. 1947. The relation of the Spartinetum glabrae near Beaufort, North Carolina, to certain 

edaphic factors. Amer. Midland Nat. 38:605-614. 
Rundel, P.. P. Bowler, and T. W. Mulroy. 1972. A fog-induced lichen community in northwestern 

Baja California, with two new species of Pesmaziena. Bryologist 75:501-508. 
Shreve, F., and I. L. Wiggins. 1964. Vegetation and flora of the Sonoran Desert. Stanford Univ. 

Press. Stanford, Calif. 1669 p. 
Speth, J. 1969. The fuss over coastal wetlands. Outdoor Calif. 30:6-7. 
Stevenson, R. E., and K. O. Emery. 1958. Marshlands at Newport Bay, California. Allan Hancock 

Foundation Occas. Paper No. 20. 109 p. 
Thorsted, T. H., Jr. 1972. The salt marsh vegetation of Bahia de San Quintin. M. A. thesis. California 

State Univ., Los Angeles. 64 p. 
Vogl, R. J. 1966. Salt-marsh vegetation of upper Newport Bay, California. Ecology 47:80-87. 
Vonder Haar, S. P. 1972. Semi-arid Coastal Evaporite Environment at Laguna Mormona, Pacific 

Coast, Baja California, Mexico. M. S. thesis, Univ. So. Calif., Los Angeles. 72 p. 
Wiggin, I. L. 1960. The biogeography of Baja California and adjacent seas. Syst. Zool. 9:148-165. 
Zedler, J. B. 1977. Salt marsh community structure in the Tijuana Estuary, California. Estuarine 

and Coastal Marine Sci. 

Accepted for publication July 27, 1979. 



Bull. Southern California Acad. Sci. 
78(3), 1979, pp. 183-191 

Gametogenesis and Embryonic Development in the Calcareous 

Sponges Clathrina coriacea and C. blanca from 

Santa Catalina Island, California 

Marion Fischel Johnson 

Abstract. — Gametogenesis and embryonic development of Clathrina coriacea 
and C. blanca, two closely related calcareous sponges from Santa Catalina Island, 
California, are described. Oogenesis is asynchronous in both species. Spermato- 
genesis was not observed. Cleavage is total and equal, resulting in the formation 
of a blastula larva. The larva of C. coriacea contains one large posterior granular 
cell, whereas two posterior granular cells are present in the larva of C. blanca. 
Migration of the larval blastomeres into the blastocoel begins while the larva is 
in the tube of the parent sponge. 

LGL Limited, Environmental Research Associates, 103 Pleasant Street, Bryan, 
Texas 77801. 



The life histories of Clathrina coriacea (Montagu) and C. blanca (Miklucho- 
Maclay), two closely related calcareous sponges from Santa Catalina Island, Cal- 
ifornia, were studied to clarify their systematic affinity (Johnson, 1976). Minchin 
(1900), Topsent (1936) and Borojevic (1967), among others, treated C. coriacea 
and C. blanca as separate species. Burton (1963), on the other hand, believed 
that the two species names were synonymous. 

Levi (1956) utilized differences in embryological development and breeding 
period to separate two morphologically identical sponges of the genus Halisarca 
into H. dujardini and H. metschnikovi. This paper reports the findings of a com- 
parative study of gametogenesis and embryonic development in C. coriacea and 
C. blanca. 

Methods 

Clathrina coriacea and C. blanca were studied at Santa Catalina Island from 
April 1973 through October 1975. The sponges were collected at weekly intervals 
during the reproductive period in order to obtain detailed information on game- 
togenesis and embryonic development. Most of the specimens were obtained 
from a submarine grotto and the Santa Catalina Marine Biological Laboratory 
pier in Big Fishermans Cove. 

Collection of the sponges involved carefully removing them from the attach- 
ment surface with forceps and placing them in labelled plastic bags. The speci- 
mens were immediately transferred to sea water Bouin's Fixative for preservation 
of the sponge and decalcification of the spicules. The spicules were dissolved to 
facilitate observation of the reproductive structures and histological pro- 
cedures. After decalcification was complete (24 to 48 hours) the sponges were 
transferred to 70% ethanol. They were then observed under a dissecting micro- 
scope for the presence of oocytes. Observations of the location of the oocytes 
within each sponge also were recorded. 



184 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

A small piece of each sponge, or the whole sponge when it was less than 5 mm 
in size, was used for histological study. The sponges were dehydrated, stained 
temporarily with a weak solution of Fast Green in 70% ethanol, embedded in 
Paraplast (Sherwood Medical Industries) and sectioned at 8 /xm. Every fifth or 
tenth section, depending on the size of the embedded piece, was mounted on a 
glass slide with a minimum of 30 sections from each sponge. Minute specimens 
were sectioned serially. The slides were stained with Ehrlich's Hematoxylin and 
counterstained with Eosin. 

Results 

Reproductive Season. — Maturing oocytes were observed in C. blanca between 
April and June in 1973 and 1974, and between April and August in 1975. In C. 
coriacea reproductive elements were seen from July to September in 1973, from 
July to August in 1974 (specimens were not collected during September and 
October), and from July through October in 1975 (Johnson, 1978). 

Oogenesis. — The process of oocyte development is similar in C. coriacea and 
C. blanca. The young oocyte can be distinguished from other nucleolate cells by 
its strongly basophilic staining cytoplasm, large nucleus and nucleolus surrounded 
by a deeply staining nuclear membrane. As the oocyte grows it begins to push 
the surrounding layer of choanocytes into the tube of the parent sponge. Oocytes 
larger than 30 /xm apparently phagocytize the surrounding eosinophilic amebo- 
cytes (Fig. 1). These amebocytes are common in specimens with developing 
oocytes, but rare in nonreproducing sponges. The length of the eosinophilic cells 
varies from 7 to 10 ^m in C. coriacea and C. blanca. Although there is no 
distinction in the dimensions of the eosinophilic amebocytes between the two 
species, noticeable differences exist in the eosinophilic granules. In C. coriacea 
the eosinophilic amebocytes are filled with numerous highly refractile eosinophilic 
staining granules. In C. blanca, on the other hand, the granules are smaller, less 
refractile and less abundant. Continued growth of the oocyte results in a cell 
highly granular in appearance (Fig. 2). The maximum length recorded for a mature 
oocyte of C. coriacea was 90 fxm. Oocytes 100 (xm in length were seen in C. 
blanca. 

There is no apparent synchronization of oogenesis within the breeding popu- 
lation of the two species. Some of the specimens collected at the same time 
contain no oocytes, some have large oocytes and others contain embryos or 
larvae. Occasionally small and large oocytes, oocytes and embryos or larvae 
occur in the same sponge. The different stages of development are generally 
found in different regions of the sponge. 

Spermatogenesis. — All of the specimens collected during the reproductive sea- 
son were examined for stages of spermatogenesis. No sperm or spermatic cysts 
were seen in the two species. 

Fertilization. — The process of sperm transport to the oocyte was not observed 
with certainty in C. coriacea or C. blanca. Occasionally a small deeply staining 
ovoid structure enclosed within a vesicle was seen in a choanocyte near the 
oocyte. It resembled Tuzet's figure of a spermatozoan in the choanocyte 
(1947:plate 1, fig. 17). Some cells between the choanocyte layer and the oocyte 
appeared to contain two nuclei. According to Tuzet (1947:plate 1, fig. 13), one of 
the "nuclei" is actually a spermatic vesicle in the cytoplasm of a carrier cell. On 



GAMETOGENESIS AND EMBRYOLOGY IN CERTAIN CALCAREOUS SPONGES 



185 




* 



<& ■$* 



* ' 






v* «£ 





Fig. 1. Oocyte of Clathrina blanca phagocytizing eosinophilic amebocytes (arrows). Scale, 10 
/Am. 

Fig. 2. Large granular oocyte of Clathrina coriacea. Scale, 10 fim. 

rare occasions an oocyte contained two deeply staining masses within a vesicle. 
These structures resembled Tuzet's figure of the spermatozoan within its vesicle 
at the periphery of the oocyte (1947:plate 1, fig. 19). Fertilization was not seen 
in either species. 



186 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




gfvV 



T r 









VSLi^&v^i»^S9^22 






•• .* 



Fig. 3. Two cell stage of cleavage of Clathrina blanca; note the nucleus and large nucleolus. 
Scale, 10 jxm. 

Fig. 4. Young embryos of Clathrina coriacea; note the flattened, elongated cells surrounding each 
embryo and the unorganized appearance of the parent tissue. Scale, 50 /xm. 

Embryonic Development. — Cleavage in C. coriacea and C. blanca is total and 
equal. The two cell stage, illustrated in Fig. 3, shows the granular cytoplasm, and 
the nucleus and nucleolus. In the four cell and eight cell stages further equal 
divisions occur. After the eight cell stage a blastula is formed. The young blastula 



GAMETOGENESIS AND EMBRYOLOGY IN CERTAIN CALCAREOUS SPONGES 187 

embryo is composed of cells, approximately 12 /jltu in dimension, surrounding a 
large blastocoel (Fig. 4). Each embryo is enclosed within the parent tissue by 
flattened elongate cells. The sponge tissue in which the embryos develop loses 
its normal appearance and bears little resemblance to that found in specimens 
without reproductive elements (Fig. 4). The later stages of cleavage take place 
in the tubes of the parent sponge and result in the formation of a blastula larva. 
The blastula larva contains two types of cells, the abundant blastomeres and the 
rare posterior granular cells. The blastomeres are narrow columnar flagellated 
cells containing deeply staining elongate nuclei located at the periphery of the 
larva. The posterior granular cell is a large nucleolated cell. The larva of C. 
coriacea contains one large posterior granular cell (Fig. 5), whereas two posterior 
granular cells are present in the larva of C. blanca (Fig. 6). These cells are not 
seen in all the larvae in a sponge section as it depends on how the larva is 
sectioned. 

While the larvae are still in the tube of the parent sponge some of the blasto- 
meres begin to migrate into the large blastocoel, losing their flagella and columnar 
shape and becoming spherical (Fig. 6). The obliteration of the blastocoel pro- 
gresses at different rates in individual larvae. The blastocoel of some larvae re- 
mains relatively free of cells, whereas others are partially or completely filled 
with cells (Fig. 7). At the completion of larval development the tubes of the parent 
sponge are solidly packed with larvae moving towards the oscula to be expelled. 
Measurements of the larvae in the two species reveal much variation in their 
dimensions. Larvae range from 60 to 150 /xm in length in C. coriacea, and from 
70 to 160 jbtm in C. blanca. 

Discussion 

Oocytes of C. coriacea and C. blanca from Santa Catalina Island larger than 
30 /Am in length appear to phagocytize the eosinophilic amebocytes that aggregate 
around them. Similar observations were made by Tuzet (1947) in her study of 
Leucosolenia coriacea (=C. coriacea). Sara (1955ac) reported that the oocyte of 
C. coriacea forma blanca (=C. blanca) first phagocytized choanocytes and then 
later eosinophilic cells, whereas the oocyte of C. coriacea forma coriacea (=C. 
coriacea) engulfed only choanocytes. These differences between the two species 
were not observed in the California specimens. There appears to be no difference 
in the intensity of phagocytosis between C. coriacea and C. blanca from Santa 
Catalina Island. Sara (1955a), on the other hand, observed that phagocytosis was 
greater in C. blanca than in C. coriacea. The eosinophilic amebocytes are rare 
in nonreproducing individuals from Santa Catalina Island and common in speci- 
mens with developing oocytes. Sara (1955b), however, reported that the eosin- 
ophilic amebocytes decreased in specimens with developing oocytes. 

The eosinophilic amebocytes differ between the two sponges from Santa Cat- 
alina Island. In C. coriacea the granules within these amebocytes are larger, 
more numerous and more refractile than in C. blanca. Variations in the eosino- 
philic amebocytes among species of calcareous sponges also were reported by 
Minchin (1898), Duboscq and Tuzet (1936) and Borojevic ( 1969). Minchin (1898), 
in fact, believed that he could distinguish between species of Clathrina solely on 
the basis of the granular (eosinophilic) amebocytes. 



188 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 





PC 




Fig. 5. Single large posterior granular cell (PC) in larva of Clathrina coriacea. Scale, 10 /xm. 

Fig. 6. Two large posterior granular cells (PC) in larva of Clathrina blanca. Scale, 10 /j.m. 
Fig. 7. Larvae of Clathrina blanca; note the varying stages of migration of the blastomeres into 
the blastocoel (B). Scale, 50 jitm. 



The oocyte in C. coriacea and C. blanca becomes very granular as it reaches 
sexual maturity. Similar observations were made by Tuzet (1947) for C. coriacea. 

Oogenesis is asynchronous in the breeding population of the two California 
sponges. Some specimens contain developing oocytes, embryos or larvae, where- 



GAMETOGENESIS AND EMBRYOLOGY IN CERTAIN CALCAREOUS SPONGES 189 

as other individuals have no reproductive elements. Duboscq and Tuzet (1937) 
reported that oogenesis also was asynchronous in the calcareous sponge Grantia 
compressa. 

Spermatogenesis was not observed in C. coriacea or C. blanca from Santa 
Catalina Island. The lack of evidence of stages of spermatogenesis in calcareous 
sponges has puzzled sponge specialists since the latter part of the nineteenth 
century. Although Haeckel (1871) described spermatozoa in calcareous sponges, 
Dendy (1914) wrote that no one else had been able to repeat Haekel's obser- 
vations. Polejaeff (1882) collected rare male specimens of Sycon raphanus which 
he said were so completely filled with spermatic cysts that their whole develop- 
ment could be traced in a single section. Gorich (1903ab, 1904) reported the 
presence of spermatic cysts in the upper third of Sycandra raphanus (=Sycon 
raphanus) and described the early stages of development. Dendy (1914) described 
what he thought were spermatic cysts and some stages of spermatogenesis in 
Grantia compressa. Gatenby (1920, 1927) looked at many breeding specimens of 
Grantia compressa and saw only a single stage of spermatogenesis. He concluded 
that spermatogenesis must take place sporadically and very rapidly. Vacelet 
(1964) was unable to find stages of spermatogenesis in Petrobiona massiliana. 
Tuzet (1973) reported that spermatogenesis was not known in calcareous sponges. 
In the demosponges, on the other hand, spermatogenesis has been seen in many 
species (Tuzet, 1930; Levi, 1956; Tuzet and Pavans de Ceccatty, 1958; Tuzet and 
Paris, 1964; Tuzet et al., 1970, among others). 

Fertilization was not observed with certainty in C. coriacea or C. blanca 
from Santa Catalina Island. Sara (1955a) and Borojevic (1969) also had little 
success in following the process of fertilization. Borojevic (1969) reported that 
he never saw any figures of fertilization in the numerous specimens of the Cal- 
carea Calcinea he studied, whereas they were easily discernible in the Calcarea 
Calcaronea. Tuzet (1947), however, described the process of fertilization of C. 
coriacea in great detail, but she did not observe the actual fusion of the male and 
female pronuclei. 

Cleavage in C. coriacea and C. blanca is total and equal, and a blastocoel 
forms after the eight cell stage. Similar observations were made by Tuzet (1948) 
for C. coriacea and by Borojevic (1969) for other calcinean Calcarea. In the 
developing blastula larva of C. coriacea and C. blanca two cell types become 
apparent, the narrow columnar flagellated cell and the large granular cell. Tuzet 
(1948) reported that the granular cell determined the posterior region of the larva. 
Borojevic (1969) considered the posterior granular cell to be a blastomere whose 
division had been retarded. The blastula larva of C. coriacea from Santa Catalina 
Island contains one posterior granular cell, whereas C. blanca has two posterior 
granular cells. These observations agree with Minchin (1900) who reported that 
C. coriacea contains one, C. blanca has two, and C. contort a and Ascandra 
falcata have four posterior granular cells, whereas C. cerebrum and C. reticulum 
do not contain posterior granular cells. In the two California sponges some of the 
larval blastomeres migrate into the large blastocoel while still within the tubes of 
the parent sponge. This process also was observed in Leucosolenia coriacea 
(=C. coriacea) by Minchin (1896) and Tuzet (1948). 

The relationship between the blastula larva of calcareous sponges and other 
larval types of sponges was discussed by Tuzet (1948, 1973) and Borojevic (1969). 



190 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Unlike the amphiblastula larva, the blastula larva is formed directly by multipli- 
cation of the blastomeres. There is no stage of blastomeres with internal flagella, 
no inversion of the surfaces, and no formation of the typical "cellules en croix." 
Borojevic (1969, 1970) also found no similarity between the blastula larva and the 
parenchymella larva of many demosponges. In the parenchymella larva no blas- 
tocoel is formed and cellular differentiation occurs very early in the development. 
The parenchymella larva possesses at its liberation from the parent sponge the 
principal cell types of the adult sponge, whereas in the blastula larva the larval 
cells remain totipotent (Tuzet, 1948; Borojevic, 1969, 1970). 

Conclusion 

The differences in the eosinophilic amebocytes, the number of posterior gran- 
ular cells in the larvae, and the dimensions of the oocytes and larvae, in addition 
to the differences in the reproductive period between C. coriacea and C. blanca, 
reaffirm that the two sponges are separate species. 

Acknowledgments 

I wish to thank Dr. G. J. Bakus for guidance and encouragement, Dr. R. Bils 
for the use of the histology laboratory and equipment, and Dr. W. O'Day for 
photographic assistance. I also thank the many other individuals who assisted me 
during this study. 

Literature Cited 

Borojevic, R. 1967. Spongiaires d'Afrique du Sud. II. Calcarea. Trans. R. Soc. S. Afr., 37:183-226. 
. 1969. Etude du developpement et la differenciation cellulaire d'eponges calcaires calcinees 

(genres Clathrina et Ascandra). Ann. Embryol. Morphog., 2:15-36. 
. 1970. Differenciation cellulaire dans l'embryogenese et la morphogenese chez les spongiaires. 

Pp. 467-490 in The biology of the Porifera. (W. G. Fry, ed.), Symp. Zool. Soc. Lond., No. 

25. Academic Press, 512 pp. 
Burton, M. 1963. A revision of the classification of the calcareous sponges. British Museum of 

Natural History, 693 pp. 
Dendy, A. 1914. Observations on the gametogenesis of Grantia compressa. Quart. J. Microsc. Sci. 

(Ser. 2), 60:313-376. 
Duboscq, O., and O. Tuzet. 1936. Les amebocytes et les cellules germinales des eponges calcaires. 

Mem. Mus. R. Hist. Nat. Belg. (Ser. 2), 3:209-226. 
. 1937. L'ovogenese, la fecondation et les premiers stades du developpement des eponges 

calcaires. Arch. Zool. Exp. Gen., 79:157-316. 
Gatenby, J. B. 1920. Germ cells, fertilization and early development in Grantia. J. Linn. Soc. Lond. 

Zool., 34:261-297. 
. 1927. Further notes on oogenesis and fertilization of Grantia. Quart. J. Microsc. Sci., 71: 173- 

188. 
Gorich, W. 1903a. Zur Kenntnis der Spermatogenese bei den Poriferen und Colenteraten. Zool. 

Anz., 27:64-70. 
. 1903b. Weiteres uber die Spermatogenese bei den Poriferen und Colenteraten. Zool. Anz., 

27:172-174. 
. 1904. Zur Kenntnis der Spermatogenese bei den Poriferen und Colenteraten nebst Bemer- 

kungen iiber die Oogenese der ersten. Z. Wiss. Zool., 76:522-543. 
Haeckel, E. 1871. Uber die sexuelle Fortpflanzung und das natiirliche System der Schwamme. Jena. 

Z. Naturw., 6:641-651. 
Johnson, M. F. 1976. Conspecificity in calcareous sponges: Clathrina coriacea (Montagu, 1818) and 

Clathrina blanca (Miklucho-Maclay, 1868). Ph.D. Dissertation, University of Southern Cali- 
fornia, xvii + 442 pp. 



GAMETOGENESIS AND EMBRYOLOGY IN CERTAIN CALCAREOUS SPONGES 191 

. 1978. Studies on the reproductive cycles of the calcareous sponges Clathrina coriacea and 

C. blanca. Mar. Biol. 50:73-79. 
Levi, C. 1956. Etude des Halisarca de Roscoff; embryologie et systematique des demosponges. 

Arch. Zool. Exp. Gen., 93:1-184. 
Minchin, E. A. 1896. Note on the larva and post-larval development of Leucosolenia variabilis n. 

sp., with remarks on the development of other Asconidae. Proc. R. Soc. Lond., 60:42-52. 
. 1898. Materials for a monograph of the Ascons. I. On the origin and growth of the triradiate 

and quadriradiate spicules in the family Clathrinidae. Quart. J. Microsc. Sci., 40:469-587. 
. 1900. Eponges calcaires. La Clathrine coriace. Pp. 107-147 in Zoologie descriptive des 

invertebres. Vol. I. (L. M. Boutan, ed.), O. Doin, 2 vol. 
Polejaeff, N. 1882. Uber das Sperma und die Spermatogenese bei Sycandra raphanus Haeckel. 

Sber. K. Akad. Wiss. Wien (Abt. I), 86:276-298. 
Sara, M. 1955a. La nutrizione deirovocita in Calcispongie Omoceli. Annuar. 1st. Mus. Zool. Univ. 

Napoli, 7:1-30. 
. 1955b. Osservazioni sugli amebociti eosinofili di Clathrina e Leucosolenia (Calcispongiae) 

e sulla loro colorizione vitale con blu trypan. Boll. Zool., 22:43-50. 
. 1955c. Sulle cellule nutrici nell'ovogenesi delle Calcispongie Omoceli. Boll. Zool., 22:323- 

327. 
Topsent, E. 1936. Etude sur des Leucosolenia. Bull. Inst. Oceanogr. Monaco, No. 711:1-47. 
Tuzet, O. 1930. Spermatogenese de Reniera. C. R. Seances Soc. Biol., 103:970-973. 
. 1947. L'ovogenese et la fecondation de l'eponge calcaire Leucosolenia (Clathrina) coriacea 

Mont, et de l'eponge siliceuse Reniera elegans Bow. Arch. Zool. Exp. Gen., 85:127-148. 
. 1948. Les premiers stades du developpement de Leucosolenia botryoides Ellis et Solander 

et de Clathrina {Leucosolenia) coriacea Mont. Ann. Sci. Nat. (Ser. 11), 10:103-114. 
. 1973. Eponges calcaires. Pp. 27-132 in Traite de zoologie. Vol. Ill (1). Spongiaires. (P-P. 

Grasse, ed.), Masson et Cie., 690 pp. 
Tuzet, O., R. Garrone, and M. Pavans de Ceccatty. 1970. Origine choanocytaire de la lignee ger- 

minale male chez la demosponge Aplysilla rosea Schulze. C. R. Hebd. Seances Acad. Sci. 

(Paris) (Ser. D), 270:955-957. 
Tuzet, O., and J. Paris. 1964. La spermatogenese, l'ovogenese, la fecondation et les premiers stades 

du developpement chez Octavella galangani. Vie Milieu, 15:309-327. 
Tuzet, O., and M. Pavans de Ceccatty. 1958. La spermatogenese, l'ovogenese, la fecondation et les 

premiers stades du developpement d'Hippospongia communis LMK (= H. equina O. S.). 

Bull. Biol. Fr. Belg., 92:331-348. 
Vacelet, J. 1964. Etude monographique de l'eponge calcaire pharetronide de Mediterranee Petro- 

biona massiliana Vacelet et Levi. Les pharetronides actuelles et fossiles. Rec. Trav. Stn. Mar. 

Endoume, 50:1-125. 

Accepted for publication July 27, 1979. 



Bull. Southern California Acad. Sci. 
78(3), 1979, pp. 192-196 



Research Notes 
Observations of a Gray Whale Birth 



Observations of births of gray whales (Eschrichtius robustus) have been made 
by Balcomb (1974), Leatherwood and Beach (1975) and Storro-Patterson and 
Kipping (1977). We report a fourth observation of a gray whale birth; this from 
Estero de la Soledad in the Bahia Magdalena complex, Baja California Sur, Mex- 
ico (Fig. 1). We learned from Storro-Patterson that both his (1977) and Balcomb's 
(1974) observations were also made in this area; Storro-Patterson' s at a location 
near the cannery at Puerto Adolfo Lopez Mateos, and Balcomb's nearby to the 
north (per. coram.). Leatherwood 's observations were made in the open sea off 
Mission Bay, San Diego. 

On Sunday 19 February 1978 at approximately 1755 hours we approached the 
arm of the lagoon leading to the fishing village of Matancitas when the senior 
author noticed a solitary gray whale maintaining a stationary position with its tail 
flukes held vertically and motionless above the surface. Approximately 2 m of 
caudal peduncle and the flukes were exposed, holding for one to two minutes 
(Fig. 2a). The flukes then were lowered closer to the surface, held a few moments, 
and descended until the posterior margins were awash (Fig. 2b). After another 
minute the flukes were again raised vertically exposing a meter of caudal peduncle 
and flukes. The flukes then rotated slowly as the whale moved a few degrees on 
the vertical. 

The flukes were again lowered, this time horizontal to the surface, and we 
caught our first view of the calf s emerging snout. From a distance of 50 to 100 
m the only visible portions of the whales were the mother's flukes barely splashing 
at the water's surface and the calf s snout. At this point the calf s snout sank 
momentarily from view, but reappeared almost immediately. 

The female's flukes next were positioned nearly vertical just above the surface 
as the calf was again submerged. This fluke movement was slow, unlike either 
the thrashing described by Balcomb (and shown in his photographs) or the lurch- 
ing and violent tail spasms described by Leatherwood and Beach. Within a minute 
the flukes were again lowered horizontal to the surface as the calf reappeared. At 
this time we discerned its wrinkled nostril area. As the extrusion of the calf 
reached this point which, with the shoulders, forms the greatest girth, the calf 
appeared to be positioned with its left latero-dorsal aspect facing posteriorly. 
Balcom's photographs (1974) show the calf being presented with its dorsal aspect 
facing anteriorly. At this point in our observation the birth seemed essentially 
completed. The calf appeared to wobble in its near vertical position as though 
adding its own movements. The pectoral fins, or flippers, had not yet cleared the 
genital orifice as the couple again sank beneath the surface with the mother's 
flukes only slightly visible. 

Within 30 seconds the calf surfaced independent of its mother. Due to poor 
water clarity and failing light we were unable to determine what, if any, assistance 



RESEARCH NOTES 



193 



LOCATIO 
OF BIRTH 



J.G.M. 
1979 




Fig. 1. Locale of birth observations. Configurations of channels determined from Earth Resources 
Technology Satellite photography. 



194 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. a. Elevated position of mother's tail flukes at early stage of parturition, 
positioning before first appearance of emerging calf. 



b. Subsequent 



the female might have rendered, e.g., by way of nudging the newborn to the 
surface to breathe. The calf did not give any appearance of being held above the 
water's surface as shown clearly in a Storro-Patterson photograph (1977). It was 
not for another 30 seconds that the female appeared at the surface and blew. 

The whole episode, from our first sighting of the elevated tail flukes until the 
female's first blow, lasted less than 10 minutes (1755 to 1804). We did not see the 
female blow until after the birth was completed. 

At no time during the birth did we see any portion of the female's body anterior 
to the genital aperture. This suggests that she kept her head positioned below the 
surface with her back arched in order to keep the head of the emerging calf above 
the water (Fig. 3). 

The whale reported by Storro-Patterson and Kipping (1977) executed a slow 
360° roll at the surface with the calf s rostrum protruding from the vagina as the 
female's ventral surface became visible. They noted that the newborn calf dis- 
played a lack of coordination in both its early swimming efforts and in initial 
lunging behavior associated with breathing. We also observed lunging behavior 
but paid less attention to it since we had observed it before in older calves usually 
during windy conditions as they encountered some difficulty in clearing the sur- 
face of the water with their nostrils. 

Poor viewing conditions precluded our observing the calf s use of its tail flukes. 
As we moved within 25 m the calf appeared a light grayish-white in contrast to 



RESEARCH NOTES 



195 







'&? 



r^' 



Fig. 3. Position during final stages of birth with calf maintained above surface of water. 



the nearly black female. The calf swam quite well as the pair moved purposefully 
away from us. An unsuccessful search was made for any placental material. No 
attempt was made to follow the pair which had slowed and was milling around. 

The female exhibited a high tolerance to our continued noisy presence. In what 
must have been trying circumstances for her, not once did she display any hint 
of aggressive recognition of our presence. Two other females with calves passed 
within 50 m during this female's parturition with no indication or inclination to 
"assist" with the birth. 

Gilmore (1961) noted the likelihood of the tail-first underwater delivery but told 
us of a verbal report of a cephalic birth at Laguna San Ignacio in very shallow 
water during the week preceding Easter, 1976 (per. comm.) 

While the birth of the whale reported by Leatherwood apparently took place 
below the surface and out of sight, the remaining three reports in the literature 
of the gray whale are of cephalic births at the surface. 

The possibility remains that all reported cephalic births including Lindsay 
(1978) are abnormal births; that gray whales, and perhaps by extension, baleen 
whales, normally deliver their young by breech, or tail-first presentation, just 
below the surface. This is the case with Tursiops and other genera of smaller 
odontocetes whose births have been observed in various marine aquaria, a ma- 
jority of which has been of tail-first births. 

Additional support for tail-first births has been obtained from the whaling in- 
dustry. 

Slijper (1962) cites records of female baleen whales of several species taken 
during pregnancy, particularly advanced pregnancy, in which the great majority 
of fetuses was found with the caudal section oriented toward the cervix. 

Additionally, Rice and Wolman (1971) report that all of 55 near-term fetuses 
of gray whales taken during the course of their study were found to be oriented 
for caudal presentation, tending to confirm the caudal birth as the normal delivery 
method. 

For the present, it can be stated that births of gray whales, in spite of their 



196 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

annual abundance in relatively restricted locations and time periods, are rarely 
observed. 

An additional point, possibly relating to the hazardous conditions of the birth 
process of a mammal in an entirely marine environment, came to our attention. 
A Sr. Modesto, a fisherman resident at Matancitas, reported to us that three dead 
calves had been encountered in Estero de la Soledad during the season of 1977 
and none up to the date of our observations in 1978. 

Acknowledgment 

We wish to acknowledge with gratitude the generous assistance and comments 
of Dr. Raymond M. Gilmore during the preparation of this manuscript. 

Literature Cited 

Balcomb, K. C, III. 1974. The birth of a gray whale. Pac. Disc. 27:28-31. 

Gilmore, R. M. 1961. The story of the gray whale. Privately published. San Diego. 17 pp. 

Leatherwood, S. L., and D. W. Beach. 1975. A California gray whale calf (Eschrichtius robustus) 

born outside the calving lagoons. Bull. So. California Acad. Sci., 74:45-46. 
Lindsay, G. 1978. The friendly whale. Pac. Disc. 31:1-9. 
Rice, D. W., and A. A. Wolman. 1971. The life history and ecology of the gray whale (Eschrichtius 

robustus). Amer. Soc. Mamm. Spec. Publ. No. 3, viii + 142 pp. 
Slijper. E. J. 1962. Whales. Basic Books, New York, 475 pp. 
Storro-Patterson, R. 1977. Gray whale protection. Oceans. 4:45-49. 
, and Kipping. 1977. Unpublished abstract, Proc. Second Conf. Biol. Marine Mammals, San 

Diego. 12-15 Dec. 1977, p. 84.) 

Accepted for publication August 1, 1979. 

James G. Mills and James E. Mills, 5326 Berkeley Rd., Santa Barbara, California 
93111. 



Bull. Southern California Acad. Sci. 
78(3), 1979, pp. 196-199 



Range Extension and Notes on the Habitat of the Isopod 
Munna halei Menzies 



The small asellote isopod Munna halei Menzies, 1952, was described from 
Tomales Point, Marin County, California, from under stones and in kelp (Mac- 
rocystis) holdfasts in the lowest intertidal zone (Menzies, 1952). It remained un- 
known elsewhere until Iverson (1974) reported it at El Capitan Beach, San Luis 
Obispo County, California, from among the spines of the purple sea urchin Stron- 
gylocentrotus purpuratus (Stimpson) from mid-intertidal rocks. 

Munna halei has now been discovered at Cape Arago, near Coos Bay, Oregon. 
The isopods were living among the spines of the sea urchin S. purpuratus which 
inhabited a relatively open, wave swept, rocky section of the coast. The isopods 



RESEARCH NOTES 197 

were collected from four of five study areas around Cape Arago: the South Cove 
of Cape Arago, the Middle Cove of Cape Arago, Squaw Island, and the rocky 
intertidal zone to the west of Squaw Island. They were not found at Sunset Bay 
near Cape Arago, but this may be due to the small sample size of urchins taken 
from that area. 

Sea urchins both in urchin holes and "free-living" were collected from all five 
areas, placed in individual plastic bags without water and immediately transported 
to the laboratory where the plastic bags were placed in a seawater drip table in 
order to maintain them at the proper temperature. Within a twelve hour period 
the urchins were removed from their bags and individually submerged in a bowl 
of seawater where they were examined under a dissecting microscope for isopods. 
The number of isopods that had been dislodged from the urchin and had subse- 
quently fallen into the bag was recorded. The number of isopods that remained 
on the urchin and their positions were also recorded. 

A total of 50 sea urchins was collected and 147 isopods were recovered from 
25 of the urchins. The number of M. halei per urchin ranged from zero to 18 with 
a mean of 2.94 per urchin (standard deviation 4.70). At the Middle Cove of Cape 
Arago, where the majority of samples were collected (27 urchins), the mean 
number of M. halei per urchin was 3.77 (standard deviation 5.51). M. halei 
showed no preference for sea urchins occurring in holes; 74 were found on urchins 
in holes and 72 were found on urchins not in holes. There was no correlation 
between the size of the urchin and the number of isopods present on it. 

The isopods were observed usually clinging to the spines of the sea urchins 
rather than crawling upon the test of the urchin (96% on spines, 4% on the test). 
There appeared to be no oral-aboral distinction in their position on the urchins, 
with 51% occurring on the oral side of the urchins and 49% on the aboral side. 

M. halei exhibited several interesting behavioral traits with regard to its mi- 
crohabitat. Typically a M. halei clung to an urchin spine by wrapping its per- 
eopods around the spine, with no region of the spine appearing to be frequented 
more than any other. In moving, an isopod, while holding on with its posterior 
pairs of pereopods, would lean off the spine it was on and grab another spine 
with its anterior pereopods, pulling itself over and onto the second spine. Oc- 
casionally M. halei was observed crawling off a spine and along the test surface 
a short distance before climbing another spine. 

The isopods were usually able to move without appearing to disturb the urchin. 
On occasion, however, the isopods set off a "spine reaction," indicating a local 
disturbance to the sea urchin. The sea urchin spines would fold down at the point 
of the disturbance in an apparent attempt to protect the surface. While the spines 
were folded down the M. halei normally was trapped between spines or under 
a "pile" of spines. If the isopod was unable to squeeze through the moving spines 
it remained still until the spines became erect and the isopod would crawl away. 

On several occasions the sea urchin's pedicellaria were actually observed 
clutching an isopod's legs. The pedicellaria would hold tightly to the leg and 
occasionally tug on it as the isopod clung precariously to a nearby spine. The 
pedicellaria would eventually release its hold and the isopod would crawl away 
without any apparent harm to the appendage. Usually, however, the pedicellaria 
in the vicinity of an isopod would not react to the isopod's presence. 

On one occasion a single M. halei was observed perched approximately half- 



198 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

way up a spine, waving its antennae in the water, and then wiping the antennae 
across the mouth parts in a possible feeding motion. 

In this study two other invertebrates were found to occur regularly in associ- 
ation with S. purpuratus. 97 specimens of an unidentified flatworm (Platyhel- 
minthes, Turbellaria) were observed, as were 107 specimens of an undescribed 
species of the purple amphipod, Pontogeneia sp. This same amphipod had pre- 
viously been collected from S. purpuratus at Cape Blanco, Oregon (Barnard, 
1954), as Pontogeneia inermis. Other invertebrate epizoics of purple sea urchins 
have occasionally been noted in the literature. The shrimp Betaeus macginitieae 
Hart, 1964, occurs in pairs underneath both S. purpuratus and S. franciscanus 
from Monterey Bay, California, south to Santa Catalina Island (Hart, 1964). Fla- 
belligera commensalis Moore, 1909, a polychaete worm, occurs among the spines 
of purple sea urchins (Moore, 1909; Light, 1978). Johnson and Snook (1927: 291) 
reported that the isopod Colidotea rostrata (Benedict, 1898), "lives among the 
spines of sea urchins, and its coloring resembles that of the sea urchin." 
MacGinitie and MacGinitie (1949:265) specifically indicate that Colidotea occurs 
with S. purpuratus. 

It is of interest to speculate that there may be a protective advantage for M. 
halei to live in association with S. purpuratus. If a potential isopod predator, 
such as a small fish, were to attempt to feed on a M. halei clinging to an urchin 
spine, almost invariably a "spine reaction" would be set off, and the isopod 
would immediately be covered under moving spines out of the predator's reach. 
Some indirect evidence for this was provided by laboratory attempts to remove 
M. halei from the urchin spines by using forceps, a procedure that repeatedly set 
off a spine reaction making the isopods inaccessible. The protective array of erect 
spines on the sea urchin may be a further deterrent to the predators of M. halei. 
The urchins may also provide a low energy microhabitat for the isopods, pro- 
tecting them from the strong wave action of their environment. 

Acknowledgments 

This work was performed as part of a summer project at the University of 
Oregon's Institute of Marine Biology (OIMB) at Charleston, in Dr. James T. 
Carlton's invertebrate zoology class. I thank the director, Dr. Paul Rudy, and 
the OIMB staff for providing facilities, laboratory space, and transportation. Er- 
nest Iverson of the University of Southern California, Los Angeles, and John 
Chapman, of the University of California, Santa Barbara, graciously identified 
the isopods and amphipods, respectively. Dr. Robert Jonas of Washington State 
University kindly read and commented on the manuscript. Special thanks go to 
James Carlton for his assistance and advice on the project and for critically 
reading and commenting upon the manuscript. 

Literature Cited 

Barnard, J. L. 1954. Marine Amphipoda of Oregon. Oregon State Monog. Studies in Zoology 8:1- 

103. 
Benedict, J. E. 1898. Two new isopods of the genus Idotea from the coast of California. Proc. Biol. 

Soc. Wash. 12:53-55. 
Hart, J. F. L. 1964. Shrimps of the genus Betaeus on the Pacific coast of North America with 

descriptions of three new species. Proc. U.S. Nat. Mus. 115:431-466. 



RESEARCH NOTES 199 

Iverson, E. W. 1974. Range extensions for some California marine isopod crustaceans. Bull. So. 

Calif. Acad. Sci. 73(3): 164-169. 
Johnson, M. E., and H. J. Snook. 1927. Seashore Animals of the Pacific Coast. New York: Macmillan 

Co. 659 pp. 
Light, W. J. 1978. Reexamination of the species referred to the genus Flabelliderma Hartman (Po- 

Iychaeta: Flabelligeridae and Acrocirridae) Proc. Biol. Soc. Wash. 91(3):681— 690. 
MacGinitie, G. E., and N. MacGinitie. 1949. Natural History of Marine Animals. New York: 

McGraw-Hill Book Co. Inc. 473 pp. 
Menzies, R. J. 1952. Some marine asellote isopods from northern California with descriptions of 

nine new species. Proc. U.S. Nat. Mus. 102:117-159. 
Moore, J. P. 1909. Polychaetous annelids from Monterey Bay and San Diego, California. Proc. Acad. 

Nat. Sci. Phil. 61:235-295. 

Accepted for publication May 7, 1979. 

Rebekah Harty, Oregon Institute of Marine Biology, Charleston, Oregon 97420. 
Present address: 811 Birch, Richland, Washington 99352. 



Bull. Southern California Acad. Sci. 
78(3). 1979, pp. 199-206 



Species of Demersal Zooplankton Inhabiting a Kelp Forest Eco- 
system off Santa Catalina Island, California 



Demersal zooplankton are those animals that migrate at various times between 
the benthic and pelagic environments. Previous studies have concentrated on 
coral reef habitats (e.g. Emery, 1968; Porter, 1974; Alldredge and King, 1977; 
Porter and Porter, 1977; Porter et al., 1977; Hobson and Chess, 1979; Ohlhorst 
and Hutchinson, 1979). Inverted funnel traps were used by Thomas and Jelley 
(1972) to study demersal zooplankton in an estuarine ecosystem. King and All- 
dredge (1978) reported on the emergence patterns of demersal zooplankton from 
a subtidal sand-flat. Diel changes in zooplankton composition over an intertidal 
eelgrass flat were monitored by Robertson and Howard (1978). 

Demersal zooplankton studies are of considerable interest to kelp forest ecol- 
ogy because these organisms may have substantial impact on food availability, 
feeding strategies, and behavior patterns of other community members, especially 
the ecologically and economically important kelp forest fishes. The only published 
study of kelp forest zooplankton is that of Hobson and Chess (1976), based on 
integrated plankton net collections. In contrast, emergence traps provide discrete 
samples of demersal zooplankton populations inhabiting specific substrates. The 
following study provides data from emergence traps on the species composition 
of demersal zooplankton inhabiting six substrates in a kelp forest ecosystem off 
Santa Catalina Island, California. 

The study site was located on Harbor Reefs, approximately 1 km northwest of 
the Catalina Marine Science Center, Santa Catalina Island, California (Fig. 1). 



200 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



1 






ENLARGED AREA 


-N- 




[ BIRD ROCK\J) 




~^— X ("—"J 


SANTA CATALINA ISLAND ^- 


f 




J 
x-^ HARBOR REEFS V * • 




( ISTHMUS COVE ( 

i 


/CATALINA L 
y # MARINE ^\ 
•^ SCIENCE \ 
CENTER \ 



118°30'W 118° 29' W 

Fig. 1. Map of the study site at Harbor Reefs, Santa Catalina Island, California. 



Each station was selected as representative of the various substrates within the 
kelp forest. Station 1 was composed of algal turf and Cystoseira sp. at a depth 
of 3 m. For the purposes of this study, algal turf is defined as those species of 
algae, primarily members of the Rhodophycophyta, that do not form a canopy 
but instead produce a thick mat that covers the bottom to a height of 10 cm or 
less. The habitat at Station 2 consisted of sand and pebbles in water of 3 m depth. 
Station 3 had similar sand and pebble substrate but was in 5 m of water and the 
southern sea palm, Eisenia arborea, was present. The depth at Station 4 was 6 
m and the substrate was composed of algal turf located under a canopy of the 
giant kelp, Macrocystis pyrifera. Station 5 consisted of M. pyrifera holdfasts at 
a depth of 9 m. The primary stipe was cut and removed from the holdfast just 
before the traps were set in position. The region around the holdfasts was com- 
posed of large rocks, sand, and pebbles. Station 6 was located outside the kelp 
canopy at a depth of 11 m in a bed of Dictyopteris undulata and Sargassum 
muticum on a sand and pebble substrate. 

Cone-shaped emergence traps designed by King and Alldredge (1978) were 
used to capture demersal zooplankton emerging from each substrate (Fig. 2). 
Samples of demersal zooplankton were collected with 2 emergence traps per 
substrate type. Emergence traps were set by SCUBA divers at 1930 h on 5 July 
1978 and samples were collected after a 24-h new-moon period. Samples were 
preserved in a solution of 5% formaldehyde in sea water and returned to the 



RESEARCH NOTES 



201 



COLLECTING JAR 



INVERTED FUNNEL 



RUBBER STOPPER — 




RETAINING LINE 

HOSE CLAMP 
SNAP SHACKLE 

POLYETHYLENE SKIRT 



FRAME 

HOSE CLAMP 



-SUBSTRATE 



Fig. 2. 0.25 m 2 emergence trap for collecting demersal zooplankton (modified from the basic design 
provided by Dr. Alice L. Alldredge). 



laboratory for analyses. All specimens within each sample were sorted, identified, 
and counted using dissecting microscopes. 

This study provides the first demonstration of the existence of demersal zoo- 
plankton in a kelp forest ecosystem and the first list of species caught in emer- 
gence traps (Table 1). The number of species per substrate is as follows: 59 — 
algal turf and Cystoseira; 79 — sand and pebbles; 79 — Eisenia on sand and peb- 
bles; 61 — algal turf under Macrocystis canopy; 78 — Macrocystis holdfasts; and 
58 — Dictyopteris and Sargassum on sand. A total of 134 species were collected 
from the six substrates, 114 of which were crustaceans. Thirty six species of 
gammaridean amphipods were captured, many more than in any other taxon. The 
gammaridean and caprellid amphipods, most of the harpacticoid copepods, the 
ostracods, isopods, caridean shrimp, mysids, cumaceans, tanaids, nebaliaceans, 
polychaetes, cephalopods, and fish are all generally considered to be true de- 
mersal zooplankton (see references cited above). The gastropod mollusc Tricolia 
pulloides probably entered the traps by crawling rather than by swimming. The 
remaining groups of organisms collected in the traps are presently considered as 
possible contaminants because they are usually regarded as being planktonic all 
day and night (Hobson and Chess, 1979). The emergence traps are being modified 
to determine whether these organisms are actually contaminants or true demersal 
forms. 

The quantitative data generated from this study were highly variable. A more 
comprehensive quantitative sampling program with additional replicate samples 
collected over short time intervals is in progress to determine the abundance, 
biomass, and diurnal migratory patterns of kelp forest demersal zooplankton. 



202 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Table 1. Species collected from the six kelp forest substrates. + = presence. Station 1 = algal 
turf and Cystoseira substrate, 2 = sand and pebbles, 3 = Eisenia on sand and pebbles, 4 = algal turf 
under Macrocystis canopy, 5 = Macrocystis holdfasts, 6 = Dictyopteris and Sargassum on sand 
and pebbles. 



Station Number 



Taxon 



POLYCHAETA 

Alciopidae, species A + 

Autolytus sp. + + 

Diopatra ornata + 

Eunoe sp. + 

Exogone sp. + 

Lumbrineris sp. + + 

Odontosyllis sp. + + + + + 

Platynereis bicanaliculata + + + + + 

Schistomeringos longicornus + 

Syllidae, species A + + + 

MOLLUSCA 
Gastropoda 

Gastropoda, Larva + + + + + + 

Tricolia pulloides + + + + 

Cephalopoda + + 

CRUSTACEA 

Cladocera 

Penilia avirostris + 

Ostracoda 

Cycloleberis sp. + 

Cxlindroleberididae, species A + + + + + + 

Cylindroleberididae, species B + + + 

Cylindroleberididae, species C + + + 

Cypridae, species A + + 

Neonesidea phlegeri + 

Rutiderma, species A + + + + 

Rutiderma, species B + 

Rutiderma, species C + 

Sarsiella sp. + 

Vargula tsujii + + + + + + 

Copepoda 
Calanoida 

Acartia tonsa + + + + 

Candacia bipinnata + 

Candacia sp., juvenile + 

Clausocalanus furcatus + + + + + + 

Clausocalamts sp. + + + + + + 

Copepodite + + + + 

Paracalanus parvus + + + + + 

Cyclopoida 

Corycaeus amazonicus + + + + + 

Corycaeus anglicus + 

Farranula curta + 

+ 
+ + + + + + 



Oithona similis 
Oncaea sp. 



RESEARCH NOTES 



203 



Table 1. Continued. 









Station 


Number 






Taxon 


1 


2 


3 


4 


5 


6 


Harpacticoida 














Altheutha langi 


+ 


+ 


+ 


+ 


+ 


+ 


Diosaccus spinatus 


+ 


+ 


+ 


+ 


+ 


+ 


Ectinosomatidae, species A 


+ 


+ 


+ 


+ 


+ 


+ 


Euterpina acutifrons 










+ 


+ 


Laophonte sp. 




+ 








+ 


Microsetella rosea 












+ 


Paraltheutha sp. 




+ 






+ 




Porcellidium spp. 


+ 


+ 


+ 


+ 


+ 


+ 


Tisbe spp. 


+ 


+ 


+ 


+ 


+ 


+ 


Monstrilloida 




+ 


+ 


+ 


+ 


+ 


Cirripedia 














Cypris Larva 


+ 


+ 


+ 


+ 


+ 


+ 


Nebaliacea 














Nebalia sp. 






+ 




+ 




Mysidacea 














Acanthomysis sculpta 




+ 


+ 


+ 


+ 




Erythroponii, species A 


+ 


+ 


+ 


+ 


+ 




Siriello pacifica 




+ 


+ 


+ 




+ 


Cumacea 














Cumella sp. 


+ 


+ 


+ 


+ 


+ 


+ 


Tanaidacea 














Leptochelia sp. 




+ 


+ 


+ 


+ 


+ 


Pancolus californiensis 


+ 






+ 






Isopoda 














Antias sp. 


+ 


+ 










Bathura luna 




+ 










Cirolana harfordi 




+ 










Cirolana parva 


+ 


+ 


+ 


+ 


+ 


+ 


Cryptoniscus Larva 




+ 




+ 






Eurydice can data 




+ 


+ 


+ 


+ 




Gnathia sp. 


+ 


+ 


+ 


+ 


+ 


+ 


Ida tea resecata 






+ 








Idotea sp., juvenile 


+ 




+ 








Paracerceis cordata 


+ 




+ 




+ 


+ 


Paranthura elegans 








+ 






Sphaeromatidae, species A 


+ 


+ 


+ 


+ 


+ 




Sphaeromatidae, species B 




+ 


+ 




+ 




Amphipoda 














Gammaridea 














Ampelisca lobata 


+ 


+ 


+ 


+ 


+ 


+ 


Ampithoe plea 


+ 


+ 


+ 






+ 


Ampitlioe polle.x 




+ 










Ampithoe simulans 


+ 


+ 


+ 




+ 


+ 


Amphithoe sp., juvenile 




+ 


+ 




+ 


+ 


Aoroides columbiae 




+ 


+ 




+ 




Batea transversa 


+ 


+ 


+ 


+ 


+ 


+ 


Ceradocus sp. 










+ 





204 

Table 1. Continued. 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Station Number 



Taxon 



Chevalia aviculae 
Erichthonius brasiliensis 
Gammaridae, species A 
Gammaridae, species B 
Gammaridae, juvenile 
Gammaropsis thompsoni 
Gitanopsis vilordes 
Heterophlias seclusus 
Hyale frequens 
Lysianassa dissimilis 
Lysianassa macromerus 
Lysianassidae, juvenile 
Maera similie 
Microjassa litotes 
Paraphoxus obtusidens 
Paraphoxus oculatus 
Paraphoxus spinosus 
Parapleustes oadatus 
Parapleustes pugettensis 
Photis californica 
Photis sp. 
Pleustes platypa 
Podocerus cristatus 
Polycheria osborni 
Pontogeneia intermedia 
Pontogeneia quinsana 
Pontogeneia rostrata 
Stenothoe sp. 

Hyperiidea 
Hyperella sp. 
Parathemisto sp. 
Unidentified 

Caprellidea 

Caprella angusta 
Caprella californica 
Caprella mendax 
Caprella verrucosa 
Caprella sp. 
Mayerella banksia 
Tritella laevis 
Unidentified 

Euphausiacea 

Euphausia recurva 
Calyptopis Larva 

Natantia 

Alpheus bellimanus 
Alpheus clamator 
Alpheus sp. 
Betaeus sp. 
Caridea Zoea Larva 





+ 


+ 


+ 


+ 




+ 


+ 


+ 


+ 


+ 


+ 


+ 




+ 


+ 



+ 






+ 








+ 


+ 




+ 


+ 
+ 




+ 




+ 


+ 


+ 



+ 


+ 
+ 
+ 


+ 
+ 
+ 








+ 


+ 


+ 


+ 


+ 
+ 


+ 


+ 


+ 


+ 


+ 


+ 
+ 


+ 


+ 


+ 


+ 






+ 


+ 


+ 


+ 


+ 


+ 


+ 




+ 


+ 


+ 


+ 




+ 


+ 


+ 


+ 


+ 


+ 


+ 




+ 









RESEARCH NOTES 












205 


Table 1. Continued. 




















Station Number 






Taxon 


1 


2 


3 


4 


5 


6 


Heptacarpus stimpsoni 


+ 




+ 


+ 


+ 


+ 


Hippolyte clorki 


+ 


+ 


+ 


+ 


+ 


+ 


Lysmata californica 


+ 


+ 


+ 


+ 


+ 


+ 


Brachyura 














Megalopa Larva 


+ 


+ 




+ 


+ 


+ 


Nauplii Larva 




+ 










BRYOZOA 














Cyphonautes Larva 


+ 




+ 




+ 




LARVACEA 














Oikopleura sp. 






+ 




+ 





PISCES 

Chromis punctipinnis 
Gobies ox rhessodon 
Lythrypnus dalli 
Oxyjulus californica 
Paraclinus integripinnis 



Acknowledgments 

The authors would like to thank Alice L. Alldredge for providing the basic 
emergence trap design and Alice L. Alldredge, Richard C. Brusca, John P. 
Chapman, James A. Coyer, Kristian Fauchald, Jon Kastendiek, Geraldine 
Knatz, Eugene M. Kramer, Robert J. Lavenberg, James H. McLean, Barry R. 
Wallerstein, Richard N. Winn, Mary K. Wicksten, and Russel L. Zimmer for 
taxonomic identifications, readings of the manuscript, and stimulating conversa- 
tions. We thank Robert R. Given and his staff at the Catalina Marine Science 
Center for making facilities available. This research was supported in part by 
Richard C. Brusca, Bernard C. Abbott, and a Grant-in-Aid of Research from 
Sigma Xi, The Scientific Research Society. Allan Hancock Foundation Contri- 
bution No. 375. Catalina Marine Science Center Contribution No. 41. 



Literature Cited 

Alldredge, A. L.. and J. M. King. 1977. Distribution, abundance, and substrate preferences of 

demersal reef zooplankton at Lizard Island Lagoon, Great Barrier Reef. Mar. Biol.. 41:317- 

333. 
Emery, A. R. 1968. Preliminary observations on coral reef plankton. Limnol. Oceanogr., 13:293- 

303. 
Hobson, E. S., and J. R. Chess. 1976. Trophic interactions among fishes and zooplankters near 

shore at Santa Catalina Island, California. Fish. Bull.. 74:567-598. 
. 1979. Zooplankters that emerge from the lagoon floor at night at Kure and Midway Atolls. 

Hawaii. Fish. Bull., 77:275-280. 
King, J. M., and A. L. Alldredge. 1978. Effects of moonlight on the emergence patterns of demersal 

zooplankton. Am. Soc. Limnol. Oceanogr. Abstracts, 41st Ann. Meeting. 



206 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Ohlhurst, S. L., and G. E. Hutchinson. 1979. Abundance and distribution of demersal plankton on 

coral reefs at Jamaica, W.I. Am. Soc. Limnol. Oceanogr. Abstracts, 1st Winter Meetings. 
Porter, J. W. 1974. Zooplankton feeding by the Caribbean reef-building coral Monastrea cavernosa. 

Proc. Intern. Symp. Coral Reefs, 1:111-125. 
, and K. G. Porter. 1977. Quantitative sampling of demersal zooplankton migrating from 

different coral reef substrates. Limnol. Oceanogr., 22:553-555. 
, , and Z. Batac-Catalan. 1977. Quantitative sampling of Indo-Pacific demersal reef 

zooplankton. Proc. 3rd Intern. Coral Reef Symp., 1:105-112. 
Robertson, A. I., and R. K. Howard. 1978. Diel trophic interactions between vertical migrating 

zooplankton and their fish predators in an eelgrass community. Mar. Biol., 48:207-214. 
Thomas, M. L. H., and E. Jelley. 1972. Benthos trapped leaving the bottom in Bideford River, 

Prince Edward Island. J. Fish. Res. Bd. Canada. 29:1234-1237. 

Accepted for publication August 7, 1979. 

Richard M. Hammer and Richard C. Zimmerman, Allan Hancock Foundation, 
University of Southern California, Los Angeles, California 90007. 



INSTRUCTIONS FOR AUTHORS 

'e BULLETIN is published three times each year (April, August, and November) and includes articles in English 
i any field of science with an emphasis on the southern California area. Manuscripts submitted for publication 
ould contain results of original research, embrace sound principles of scientific investigation, and present data 
ia clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide 
J- contributors. Consult also recent issues of the BULLETIN. Authors should strive for directness and lucidity, 
iihieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous 
rerence of pronouns, and logically placed modifiers. 

MANUSCRIPT PREPARATION 

Jilt is strongly recommended that, before submitting a paper, the author ask qualified persons to review it. The 
;thor is requested to submit at least two additional copies with the original, on SVi x 11 opaque, nonerasable 
]per, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. 
jotnotes should be avoided. Manuscripts which do not conform to the style of the BULLETIN will be returned to 
h author. 

An abstract summarizing in concise terms the methods, findings, and implications discussed in the paper must 
.company a feature article. Abstract should not exceed 100 words. 

A feature article comprises approximately five to thirty typewritten pages. Papers should usually be divided into 
e following sections: abstract, introduction, methods, results, discussion and conclusions, acknowledgments, and 
erature cited. Avoid using more than two levels of subheadings. 

A research note is usually one to six typewritten pages and rarely utilizes subheadings. Consult a recent issue 
the BULLETIN for the format of notes. Abstracts are not used for notes. 

Abbreviations: Use of abbreviations and symbols can be determined by inspection of a recent issue of the 
(JLLETIN. Omit periods after standard abbreviations: 1.2 mm, 2 km, 30 cm, but Figs. 1-2. Use numerals before 
lits of measurements: 5 ml, but nine spines (10 or numbers above, such as 13 spines). The metric system of 
sights and measurements should be used wherever possible. 

Taxonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International 
ade of Botanical Nomenclature (Lawjouw et al., 1956), the International Code of Nomenclature of Bacteria and 
iruses (Buchanan et al., 1958), and the International Code of Zoological Nomenclature (Stoll et al., 1961). 
fecial attention should be given to the description of new taxa, designation of holotype, etc. Reference to new 
xa in titles and abstract should be avoided. 

The literature cited: Entries for books and articles should take these forms. 
McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. 

Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54: 452-458. 
Brattstrom, B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), 
'Univ. California Press, xii + 635 pp. 

Tables and figures (line drawings, graphs, or black and white photographs) should not repeat data contained in 
ie text. The author must provide numbers and short legends for tables and figures and place reference to each 

them in the text. Legends should be typed on a separate sheet of paper and placed at the end of the manuscript. 
lustrations and lettering thereon should be of sufficient size and clarity to permit reduction to standard page size; 

dinarily they should be no more than twice the size of intended reduction and should not exceed SV2 by 1 1 inches 
I size. Photographs must be printed on glossy paper. Submit one photoduplicated copy of each illustration. All 
ustrations accompanying Research Notes will be reduced to one column width. AH half-tone illustrations will have 
!»ht screen (grey) backgrounds. Special handling such as dropout half-tones, special screens, etc., must be requested 
\ and will be charged to authors. 

A cover illustration pertaining to an article in the issue or one of general scientific interest will be printed on the 
Dver of each issue. Such illustrations along with a brief caption should be sent to the Editor for review. 

PROCEDURE 

All manuscripts should be submitted to the Editor, Robert J. Lavenberg, Los Angeles County Natural History 
luseum, 900 Exposition Blvd., Los Angeles, California 90007. Evaluation of a paper submitted to the BULLETIN 
egins with a critical reading by the Editor; several referees also check the paper for scientific content, originality, 
nd clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent 
) the author at which time he or she may be requested to rework portions of the paper considering these rec- 
mmendations. The paper then is resubmitted and may be re-evaluated before final acceptance. 

Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He or she 
hould promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations, legends, 
nd bibliographical references. He or she marks corrections on the galley (copy editing and proof procedures in 
tyle Manual) and promptly returns both galley and manuscript to the Editor. Manuscripts and original illustrations 
/ill not be returned unless requested at this time. All changes in galley proof attributable to the author (misspellings, 
iconsistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with 
ie printer, not the Editor. 



CONTENTS 

Seventieth Anniversary of Academy Excavations at Rancho La Brea By 

Gretchen Sibley 151 

The Salt Marsh and Transitional Vegetation of Bahia de San Quintin 
By Leon F. Neuenschwander , Ted H. Thorsted, Jr., and Richard J. 
Vogl 163 

Gametogenesis and Embryonic Development in the Calcareous Sponges 
Clathrina coriacea and C. blanca from Santa Catalina Island, Cali- 
fornia By Marion Fischel Johnson 183 



Research Notes 

Observations of a Gray Whale Birth. By James G. Mills and James E. Mills 192 

Range Extension and Notes on the Habitat of the Isopod Munna halei Menzies. By 

Rebekah Harty 196 

Species of Demersal Zooplankton Inhabiting a Kelp Forest Ecosystem off Santa Cata- 
lina Island, California. By Richard M. Hammer and Richard C. Zimmerman 199 



COVER: Southern California Academy of Sciences dig at Rancho La Brea, 1909. 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




ULLETIN 




R/V P V 

NOV 1 2 ]%0 Number 1 

BOTANICAL GARDEN 





*CAS-A79(1) 1-^44(1980) 



APRIL 1980 



Southern California Academy of Sciences 

Founded 6 November 1891, incorporated 17 May 1907 



OFFICERS 

Takashi Hoshizaki, President 
Fred G. Hochberg, Vice President 
Richard E. Pieper, Secretary 
Joseph E. Haring, Treasurer 
Robert J. Lavenberg, Editor 
Gretchen Sibley, Assistant Editor 



1978-1980 

Kristin H. Berry 

Robert A. Georges 

Joseph E. Haring 

Jerry D. Kudenov 

Donald J. Reish 



BOARD OF DIRECTORS 

1979-1981 

John Baird 

Jules Crane 

Fred G. Hochberg 

Richard E. Pieper 

Gloria Takahashi 



1980-1982 

Takashi Hoshizaki 
Alan J. Mearns 

Steven N. Murray 
Camm C. Swift 

Robert G. Zahary 



Membership is open to scholars in the fields of natural and social sciences, and to any persoi 
interested in the advancement of science. Dues for membership, changes of address, and requests fo 
missing numbers lost in shipment should be addressed to: Southern California Academy of Sciences 
the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 90007 



Annual Members 
Life Members . . . 



12.01 
150.01 



Fellows: Elected by the Board of Directors for meritorious services. 



The Bulletin is published three times each year by the Academy. Manuscripts for publication shouk 
be sent to the appropriate editor as explained in "Instructions for Authors" on the inside back covei 
of each number. All other communications should be addressed to the Southern California Academy 
of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, Lo; 
Angeles, California 90007. 



Date of this issue 31 October 1980 



Bull. Southern California Acad. Sci. 
79(1), 1980, pp. 1-5 

Discovery of the Male of the Katydid Idiostatus viridis 

Rentz, with Descriptions and Biological Notes 

(Orthoptera: Tettigoniidae: Decticinae) 

Charles L. Hogue 

Abstract. — Discovery of the male of the katydid Idiostatus viridis Rentz, with 
descriptions and biological notes (Orthoptera: Tettigoniidae: Decticinae) by 
Charles L. Hogue, Bull. Southern California Acad. Sci., 79(1): 1-5, 1980. The 
male of Idiostatus viridis, hitherto unknown only for this species in the genus, 
has been discovered. Two specimens from the type locality are now available 
and serve as the basis for description. Color photographs of the live female also 
permit redescription of this sex. Some biological notes are made on the basis of 
the author's collecting experiences. 

Curator of Entomology, Natural History Museum of Los Angeles County, 900 
Exposition Boulevard, Los Angeles, California 90007. 



At the time of Rentz' (1973:134) original description of Idiostatus viridis only 
females were available, the one species in the genus for which this deficiency 
persisted. Thanks to the perseverance of my collecting companions David Turner 
and James Hogue, two males were discovered at the type locality in September 
1979. Also a few additional female specimens and color photographs of living 
material are now available that permit complete descriptions of both sexes. Some 
fragmentary notes can also be made on the species' biology, although host plants, 
microhabitat and other details still elude us. 

Idiostatus viridis Rentz 
Figures 1-2 

Idiostatus viridis Rentz, 1973:130-4. Holotype female, California, San Bernardino 
County, San Bernardino Mountains, Camp O'Ongo, near Running Springs, 
Elev. 6200 feet, Natural History Museum of Los Angeles County. 

Description of the male. — The following is based upon the plesioallotype and 
color photographs of same; unfortunately, the second individual was badly dam- 
aged by yellow jackets (Vespula pensylvanica) before it was found, including 
complete removal of the genitalia, and is not adequate for descriptive purposes. 
The format and terminology follow that of Rentz' (1973) generic revision. 

Size medium for genus, form slender. HEAD with fastigium of vertex low, 
distinct, broad, sides of latter slightly concave, margin broadly rounded, without 
distinct knob. Third segment of antennal flagellum distinctly longer than second. 
Eye situated high, distinctly dorso-ventrally elongate. PRONOTUM smooth, de- 
planate, no structural evidence of median carina. Anterior margins slightly pro- 
duced laterad, concave in dorsomedial area; posterior margin truncate, barely 
concave medially. Posterior and ventral margins of lateral lobe straight; submar- 
ginal portion of posterior sector with conspicuous oval swelling defined dorsally 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 





Figs. 1, 2. Idiostatus viridis, male. 1. Terminalia, dorsal aspect. 2. Right titillator. 



by transverse sulcus. A shallow, U-shaped median sulcus on disc. TEGMEN 
broad, protruding beyond apex of pronotum for a distance of about one-half the 
mid-dorsal pronotal length. APPENDAGES. Hind femur projecting beyond apex 
of abdomen for a distance of about one-half the femur length. Fore tibia unarmed 
on anterodorsal margin, posterodorsal margin with four widely-spaced spines. 
Dorsal margins of hind tibia with two subapical spurs, distinctly more erect and 
longer than adjoining spines. Ventral margins with two apical uncinate spurs, 
slightly larger than dorsal pair opposite. Femora without teeth. Plantula of hind 
tarsus distinct, as in female. All legs hispid. Mid-dorsal carina of abdomen eva- 
nescent. TERMINALIA (Figs. 1-2). Tergite ten modified, U-shaped medially 
with distinct lateral projections without soft mesal area. Cercus extending for 
one-quarter or less their length beyond apex of tergal projection. Cercus uncinate, 
single internal hook apical. Titillator (Fig. 2) with well-developed triangular base; 
arm elongate, longer than base, without teeth, apex of latter only slightly broader 
than shaft. Subgenital plate with distinct V-shaped, moderately deep median in- 
cision; styles elongate, length about equal to side of median incision. COLOR- 
ATION (photograph of living specimen). General color and pattern similar to that 
of female, basic green somewhat lighter. Vertex of head pale brown, remaining 
portions of head capsule pale green with violaceous intrusions. Antennal scape 
purplish, flagellum medium-brown. Eye deep reddish brown, eye ring black. Disc 
of pronotum light reddish brown, yellowish marginally and dark along median 
carinal line. Ventral margin of lateral lobe creamish yellow ventral to black sub- 
marginal sulcus. Remainder of pronotum green, no stripes on humeral angle. 
Labium, maxillae (including palps), ventral portions of pleural sclerites, coxae, 
trochanters and ventrolateral portions of abdomen all purplish-light brown. A 



IDIOSTATUS VIRIDIS 3 

broad, white spiracular band separates this color from green dorsopleural area of 
abdomen. Dorsum of abdomen with two distinct, parallel creamish-yellow lines 
continuing from beneath tegmina to abdominal apex, the submedian dorsal areas 
between these lines pale reddish brown adjoining a broad median dorsal line. Leg 
segments all green, except for yellowish articular areas; all tarsi medium brown, 
segment three on all tarsi of legs black basally. Tegmen translucent yellow an- 
teriorly, the rest brown except for distinctly dark-brown median vein; no dark 
subapical spot. Apices of all spines and inner tooth of cercus dark. MEASURE- 
MENTS (in mm). Length: Body, 21.0; pronotum, 5.6 x 4.5; left tegmen, 4.0; 
fore femur, 5.7; fore tibia, 7.0; middle femur, 6.4; middle tibia, 7.6; hind femur, 
17.5; hind tibia, 17.5; titillator (apex of arm to base extreme), 1.60. 

Partial re description of female. — Rentz' description of the female coloration 
was based on dry specimens and admittedly incomplete. The following utilizes 
a series of color transparencies that I made of specimens from the type series 
while in a live state and which require modification as reflected by live material 
as follows: COLORATION. Head dorsally same green as rest of body (not 
brownish). Hind femora entirely green, except for yellow at extreme apex (no 
yellowish cast). Longitudinal yellow stripes dorsally are intermittently reddish 
brown on the anterior third of each tergite, the yellow following posteriorly pale. 
Ovipositor green basad (not totally reddish brown). 

An error is corrected in describing one appendage structure (Rentz 1973:131): 
the clause reading, "... apex of dorsal margin of hind femur with two slender 
apical spurs . . ." , (p. 131, lines 33-4), should read, "apex of dorsal margin of 
hind tibia with two slender apical spurs . . . ." 

Material. — To the two females known to Rentz from the type locality, I can 
add the following four specimens: 

CALIFORNIA: San Bernardino County, San Bernardino Mountains, Camp 
O'Ongo, near Running Springs, 20-25 August 1978, C. & J. Hogue No. 254 (2 
females:LACM); 23 August 1979, C. Hogue et al. (designated here PLESIOAL- 
LOTYPE male and one additional fragmentary male in alcohol, LACM). 

Identification. — The male of viridis will run to aberrans in Rentz' key, page 
36, couplet 22. The following modifications, if inserted into the key at that point, 
will allow correct identification of the species: 

22. Dorsal projections of abdominal tergite ten acute. Cercus attenuate. 
Tegmen light straw brown; overall coloration greenish with two yel- 
low, longitudinal stripes dorsally. Mountains of southern, central and 

northern California 22a 

Dorsal projections of abdominal tergite ten obtuse, rounded. Cercus 
with apex rounded, varying geographically. Tegmen reddish brown; 
overall coloration greyish or brownish. Owens Valley and adjacent 

areas of eastern California inyo Rehn & Hebard 

22a. Apex of cercus bifurcate, outer tooth present. Styles of subgenital plate 
short, knob-like, hardly distinguishable from plate itself; indistinct 

median incision. Mountains of central and northern California 

aberrans Rentz 

Apex of cercus simple, outer tooth absent. Styles of subgenital plate 
elongate, length about equal to side of median incision: median in- 



4 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

cision moderately deep. San Bernardino and San Jacinto Mountains 
viridis Rentz 

No changes in the key to females are necessary. 

Biological notes. — I have spent a week during the late summer (20 August to 
as late as 12 September) at the type locality every year since 1966. During this 
period I have been able to find only six specimens of viridis, four females and 
two males, and this in spite of intensive searching by myself and my companions 
on vegetation and around the general area. Such a score in collecting success 
indicates that the species is indeed rare or occupying a niche largely out of reach 
to the ordinary searcher. 

All of the specimens have been found on the ground directly beside or on the 
concrete and wood porches of mountain cabins. All but the one battered male 
last taken were discovered during the night, between approximately 2000 and 
2400 hours, PDST, when they were presumably normally active and attracted to 
lights which are customarily kept burning inside and outside the structures. The 
second male was found after sunrise in the morning after it had suffered severe 
attacks from yellow jackets. 

I and my collecting associates named above have made intensive efforts to find 
specimens on vegetation near the cabins but so far have been completely unsuc- 
cessful. I have hypothesized that the green ground color pattern, broken only by 
thin longitudinal yellow stripes, is cryptic to match the general appearance of a 
bundle of pine needles and therefore have searched most ardently among the 
branches of Pinus jeffreyi and coulteri at the locality, but to no avail. At least 
some trophic compatibility with Pinus is indicated by the complete acceptance 
of the plant as food in captivity. The male plesiotype was kept alive for six days 
and fed unhesitatingly on the needles and needle scales of Pinus jeffreyi. The 
specimen produced normal-appearing fecal pellets, which further implies that the 
plant was suitable food. Idiostatus aberrans, the species most closely related to 
viridis, has been found on fir {Abies) in nature (Rentz 1973:128). 

The apparent rarity of viridis may otherwise possibly be due to an activity 
season that peaks earlier or later than the time of year when our collecting was 
done. A similar situation has been suggested by Rentz with the closely similar /. 
aberrans. Therefore, collectors are urged to watch for this poorly known species 
in the Yellow Pine Forest belt (5000-8000 feet elevation) of southern California's 
Transverse Range mountain system from early August to late September. 

We are familiar with the stridulatory sounds of the katydids of the type locality, 
which include those of Neduba gurneyi Rentz and Birchim and Platylyra cali- 
fornica Scudder, but have not been able to identify any sonification associated 
with Idiostatus viridis, nor for /. aequalis Scudder and Neduba morsel Caudell 
which also occur in the area. 

Taxonomy. — The characteristics of the male that are now available show clear- 
ly that Rentz' original placement of viridis next to aberrans was correct. The 
genitalia of the two species are very similar and only show the specific differences 
cited in the key supplement given above. The distribution pattern of the members 
of the Aberrans Group suggests that they are geographically disjunct sister species 
whose origins are probably associated with orogenic events in California's mid- 
Cenozoic history. 



I DIO STATUS VIRIDIS 5 

Acknowledgments 

The help of David Turner and James Hogue in finding viridis deserves my 
special thanks. I also appreciate the contributions of both Dr. David Rentz who 
kindly reviewed this paper and offered comments and Eugenia Paul who rendered 
the figure of the male terminalia. 

Literature Cited 

Rentz, D. C. 1973. The shield-backed katydids of the genus Idiostatus. Amer. Entomol. Soc., Mem. 
29:1-211. 

Accepted for publication March 30, 1980. 



Bull. Southern California Acad. Sci. 
79(1). 1980. pp. 5-13 

Postfire Seedling Reproduction of Adenostoma fasciculatum 

H. and A. 

George F. Howe and Linn E. Carothers 

Abstract. — Postfire seedling reproduction of Adenostoma fasciculatum H. and 
A. by George F. Howe and Linn E. Carothers, Bull. Southern California Acad. 
Sci., 79(1):5— 13, 1980. Seedlings and crown sprouted plants of Adenostoma 
fasciculatum H. and A. were studied at various times after fires in the chaparral 
near the Newhall-Castaic area in California. In contrast to certain reports in 
which it was stated that chamise seedlings seldom contribute to mature chaparral 
cover, eighty-six percent of all chamise plants were seedlings, fourteen percent 
crown sprouts. Mean seedling height was significantly shorter than the crown 
sprout height six years after fire but not at eight or nine years postfire. 

Another area which burned twice in four years had two types of crown sprouted 
plants but no seedlings following the second fire. 

Differences in numbers of taxa in certain genera are discussed in relation to 
reproductive strategies following fire. 

Division of Natural Sciences and Mathematics, Los Angeles Baptist College, 
Newhall, California 91322. 



Introduction 



Certain species of chaparral shrubs such as Arctostaphylos glauca and Cea- 
nothus greggii may be classified as "obligate seeders" because they are non- 
sprouting and must reproduce after fire by seedlings alone — Keeley and Zedler 
(1978). Arctostaphylos glandulosa and Ceanothus leucodermis may be called 
"sprouters' ' as they are able to regenerate after fire by sprouts from the burl of 



6 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

old plants, as well as by seedlings. Adenostoma fasciculatum H. and A. (hereafter 
referred to as "chamise" or simply Adenostoma) is a sprouter also. 

Wells (1969) noted that chapparral genera which have the ability to regenerate 
by both crown sprouting and seedlings, such as Adenostoma, contain relatively 
few taxa. Conversely, he reported that genera in which certain species are unable 
to reproduce by crown sprouting (such as Ceanothus and Arctostaphylos) pos- 
sess many taxa. He concluded that the tempo of evolution in Arctostaphylos and 
Ceanothus was quickened by ". . . abandonment of the conservative, crown- 
sprouting mode of reproduction in favor of a non-sprouting, obligately-seeding 
response to recurrent fire that results in a greater frequency and intensity of 
selection." 

Keeley (1977) and Keeley and Zedler (1978) have developed a different model 
in which they ascribe adaptive significance to the obligate seeding mode as well 
as the sprouting habit among chaparral species — suggesting that under certain 
circumstances, one mode of reproduction or another is favored. 

In their studies of regrowth after fire in the chaparral of the San Jacinto Moun- 
tain region, Vogl and Schorr (1972) reported that seedling mortality in chamise 
was high and they concluded that chamise seedlings seldom contribute to the 
mature chaparral cover. Wells (1969) has asserted that "When maximum popu- 
lation density is attained, the crown-sprouters tend to regenerate only vegeta- 
tively." But in other studies by Hanes and Jones (1967), Hanes (1971) and Chris- 
tensen and Muller (1975) generous numbers of chamise seedlings were found after 
chaparral fires in various regions. 

It was our purpose to locate and study areas in which seedling survival of 
Adenostoma after fire was high. As the project developed, we also attempted to 
assess the measure of fit our data might have with Wells' model on the one hand 
or that of Keeley and Zedler on the other. 

Methods 

Regrowing populations of chaparral shrubs were observed at three different 
fire sites over a period of several years following each burn. These fires occurred 
on north or south facing slopes of ridges that run east and west. 

Dates of various fires are either from personal observation or the records of 
the Newhall Station, Los Angeles County Fire Department. The growing season 
corresponds to the rainy season which extends from approximately November 
to the following May. Plants observed in Spring 1978, for example, would have 
experienced nine growing seasons following a fire in Autumn 1969. 

One 10 x 10 m quadrat was established in the regrowing chaparral at each of 
the three postfire localities. Since it had been asserted in some of the literature 
that seedlings seldom if ever contribute to regrowth, quadrats were located in 
places where seedling survival was obvious. While such a choice of sites for 
quadrats produces an internal bias towards seedling reproduction, it was our 
intent to study reproduction under just such conditions and in just such localities. 
In actuality, the bias toward seedling reproduction was minimal in two of the 
three localities because seedlings were prevalent throughout those two entire 
slopes. 

It was possible to distinguish seedlings of chamise from old sprouted specimens 
because the seedlings are usually narrower and have no charred stem or burl at 



ADENOSTOMA FASCICULATUM 7 

ground level. A crown sprouted plant, however, has at least one burned stump 
visible amid the new, leaf-covered branches. 

All living chamise plants in each quadrat were scored as seedlings or crown 
sprouts and a count was made of dead chamise stumps that failed to sprout after 
fire. The height of each plant from the top of the tallest branch to ground level 
was recorded for each living individual. 

Site descriptions. — On 28 August 1973, a fire occurred east of Wild wood Can- 
yon Road, Newhall, CA. This area has a north-south running ridge with several 
east-west running branch ridges. Approximately six growing seasons after the 
fire, 15 February 1979, a quadrat was established on the south facing slope near 
the crest of an east-west running lateral ridge at an elevation of 440 m. 

On 2 June 1969, a fire burned a mixed stand of vegetation composed largely of 
chamise and Ceanothus crassifolius Torr. (Hoary-leaved Ceanothus) covering 
several east-west running ridges north of Oak Orchard Lane, Newhall, CA. The 
study quadrat was established on the north facing slope of one ridge at an altitude 
of 410 m. 

A large area of chaparral burned in Autumn of 1970 on a gentle south facing 
slope in the vicinity of Lake Hughes Road, Castaic, CA. at an elevation of 595 
m. Most of this same Castaic region was burned once again in Autumn 1974. But 
one large south facing slope area did not burn a second time. On 26 April 1978 
(8 growing seasons after fire) a 10 x 10 m quadrat was established in the midst 
of this section that burned in Autumn 1970 but not in 1974. 

At the Castaic site, a second quadrat was established on a steep east facing 
slope (about 400 m south and down the ridge from the previously mentioned 
quadrat) at an elevation of 549 m. This east facing slope was burned during both 
the Autumn 1970 and Autumn 1974 fires and will be called the "Castaic double- 
6^6" quadrat. 

Statistical procedures. — Means and standard deviations of height data from 
each quadrat were calculated. The Mann-Whitney U-test was used to evaluate 
the null hypothesis that any two samples came from the same population. The 
Mann- Whitney sample statistic was tested against two-tailed critical values of 
Student's T for a 5% level of significance at infinite degrees of freedom. The 
nonparametric Mann- Whitney test was used instead of a Student's T-test because 
of failure to meet assumptions of normality — see Sokal and Rohlf 1969. 

Results 

An analysis of seedling survival and regrowth on five circular quadrats within 
the Wildwood Canyon Road fire zone has been reported elsewhere (Howe 1976). 
Data for chamise regrowth on the Wildwood Canyon Road 10 x 10 m quadrat 
and data for subsequent quadrats of the present study are reported in Table 1. It 
will be noted that reproduction was largely by seedlings on the south facing slopes 
of the Wildwood Canyon Road fire. However, on nearby north facing slopes, 
chamise reproduction was by means of crown sprouts, no seedlings were evident 
six growing seasons after the fire. 

On the Oak-Orchard Lane quadrat, chamise reproduction by seedlings and 
sprouts was vigorous nine years after fire with seedlings far outnumbering sprout- 
ed plants. In the same stand, Ceanothus had been largely destroyed by the fire — 
Ceanothus seedlings being few in number. At this Oak-Orchard Lane fire area 



8 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Table 1. Chamise seedling and crown sprout reproduction after 6, 8, or 9 years since the last fire 
on quadrats from three areas near Newhall, CA. 















Number 












Total 


Number 




chamise 


Mean 










cha- 


chamise 


Mean 


crown 


height 










mise 


seedlings 


height 


sprouts 


crown 


Number 








on 10 


per 


seedlings 


per 


sprouts 


dead 


Location 


Eleva- 




x 10 m 


quadrat 


and 


quadrat 


cm and 


chamise 


and time 


tion 


Slope 


quad- 


and % of 


standard 


and % of 


standard 


per 


after fire 


meters 


aspect 


rat 


total 


deviation 


total 


deviation 


quadrat 


Wildwood 


440 


south 


115 


74, 64% 


33 ± 19 


41, 36% 


82 ± 22 


34 


Canyon 




facing 














Road 


















6 years 


















Castaic single 


595 


south 


131 


123, 94% 


91 ± 27 


8, 6% 


108 ± 17 


10 


fire 




facing 














8 years 


















Oak Orchard 


410 


north 


127 


123, 97% 


93 ± 30 


4, 3% 


111 ±46 


5 


Lane 




facing 














9 years 


















Total of all 


— 


— 


373 


320, 86% 


— 


53, 14% 


— 


49 


3 quadrats 



















the quadrat chosen was representative of the whole burn since crown sprouted 
plants were in low percentages over this entire north facing slope. They were 
present in slightly greater numbers in the more mesic environment at the bottom 
of the ravine, however. 

Four years after the fire at the Castaic site, seedlings outnumbered crown 
sprouts approximately 3 to 1 as reported by Howe (1976). Data for 26 April 1978, 
eight growing seasons after the fire at Castaic, are reported in Table 1. Here, as 
at the other two localities, seedlings far outnumbered the resprouted plants. 

Observations of the Castaic double-fire quadrat revealed that two distinct types 
of chamise plants existed four years after the second fire: (1) type-1 crown sprout- 
ed plants that were relatively small, having grown as postfire seedlings after the 
1970 burn and then crown sprouting after the 1974 fire and (2) type-2 crown 
sprouted plants which gave evidence of having sprouted twice — once after the 
1970 fire and again after the 1974 burn. This was apparent from the fact that the 
type-2 plants had both large and small burned stumps present. There were no 
postfire seedlings of the 1974 burn. On the double-fire quadrat 37 chamise plants 
(82%) were type-1 while 8 (18%) were type-2. There were 7 dead crowns on the 
quadrat on 26 April 1978. Statistical comparisons involving type-1 and type-2 
crown sprouts are presented in Table 3. 

Height comparisons. — After six growing seasons following fire, the mean 
height of chamise seedlings at the Wildwood Canyon Road quadrat (Table 2) was 
significantly shorter than the mean height of chamise crown sprouts on that same 
quadrat (P < 0.001). These same six-year chamise seedlings of Wildwood Can- 
yon Road were significantly shorter in mean height than the eight-year seedlings 
of Castaic (P < 0.001) and the nine-year seedlings of Oak Orchard Lane {P < 



ADENOSTOMA FASCICULATUM 9 

Table 2. Mann-Whitney U-test values for various chamise quadrat comparisons with the level of 
significance and sample size in parenthesis. N.S. indicates not significant at the 5% level. 





Type of 
plants 


Wildwood Canyon Road 
6 years 




Castaic 
8 years 

Crown 
(n = 8) 


Oak Orchard 

Lane 

9 years 


Location and 


Seed- 


time after 
last fire 


Seedlings 
(n = 72) 


Crown 
(n = 41) 


lings Crown 
(n = 123)(n = 4) 


Wildwood Canyon 
Road 
6 years 


Seedlings 
(n = 72) 


— 


2817 
(P < 0.001) 




— 


— — 


Wildwood Canyon 
Road 
6 years 


Crown sprouts 
(n = 41) 






(0.01 


259 
< P < 0.02) 


— 114 

(N.S.) 


Castaic 
8 years 


Seedlings 
(n = 123) 


8410 
(P < 0.001) 


— 




686 

(N.S.) 


7570 — 
(N.S.) 


Oak Orchard 
Lane 
9 years 


Seedlings 
(n = 123) 


8457 
(P < 0.001) 








— 310 

(N.S.) 



0.001). The mean height of chamise crown sprouts at Oak Orchard Lane (9 years), 
on the other hand, was not significantly different than the mean height of crown 
sprouts at Wildwood Canyon Road (6 years). 

Comparing the mean height of seedlings with that of crown sprouts from the 
same quadrat eight or nine years after five (Castaic and Oak Orchard Lane re- 
spectively) there were no significant differences — see Table 2. 

Where two fires had occurred within four years of each other (Castaic double- 
fire quadrat) no significant difference existed between the mean height of the 
type-1 and type-2 sprouters (Table 3). These type-1 and type-2 Castaic double- 
fire plants likewise showed no significant mean height differences when compared 
to the six-year crown sprouted plants of the Wildwood Canyon Road quadrat. 

Discussion 

Seedling survival. — Since the data of this study arise from selected quadrats 
where seedling survival was obvious, sweeping conclusions may not be legiti- 
mately drawn regarding the overall reproductive tendencies of Adenostoma 
throughout its entire range. However, the results are characteristic of postfire 
regrowth in this particular region of Southern California and they demonstrate 
that seedling reproduction of chamise is not unknown and certainly not unim- 
portant after fires in the chaparral. 

Seedlings were of greater importance than crown sprouts in the postfire repro- 
duction of chamise on all three quadrats selected for study in the Newhall-Castaic 
CA. area (see Table 1). Six years after the Wildwood Canyon Road fire, seedling 
reproduction was more vigorous than crown sprouting on the south facing slope 
but not on the north facing slope of that same ridge. 

This north facing slope where sprouters predominated doubtlessly differs in 
irradiation, available moisture, and certain other factors from the south facing 



10 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 3. Mann- Whitney U-test values for various chamise quadrat comparisons with the level of 
significance and sample size in parenthesis. N.S. indicates not significant at the 5% level. 









Castaic 


Castaic 








double fire, 


double fire, 








8 years, 


8 years, 








4 years 


4 years 


Location and 






Type-1 


Type-2 


time after 






crown sprout 


crown sprout 


last fire 


Type of plants 




(n = 37) 


(n = 8) 


Castaic double fire: 


Type- 1 




— 


199 


8 years, 4 years 


Crown sprouts (n = 


37) 




(N.S.) 


Wildwood Canyon Road 


Crown sprouts (n = 


41) 


779 


200 


6 years 






(N.S.) 


(N.S.) 



ridge where seedlings were most abundant. Our data here conform to a suggestion 
by Hanes (1971) that sprouters may be at a disadvantage on xeric sites when 
compared to seedlings. In other areas sprouters may have the advantage, de- 
pending on available moisture and drought stress after fire. Certain other factors 
such as intensity of the fire also have a bearing on which types of reproduction 
will be most advantageous. 

Vogl and Shorr (1972) concluded "... that Arctostaphylos and Adenostoma 
seedlings seldom contribute to mature chaparral cover." They indicated that 
perhaps selective herbivore activity explained the demise of seedlings. Likewise, 
Wells (1969) asserted that when a taxon can reproduce by both crown sprouting 
and seedlings after fire, crown sprouted plants will consistently be more important 
than seedlings in the establishment of the new population. 

Horton and Kraebel, however (1955) reported that chamise seedlings were still 
surviving and had reached an average height of 31.9 inches 25 years after fire in 
chaparral near San Bernardino. In citing an earlier 1944 paper by Sampson, Wells 
indicated that "... chamise possesses an equally superb capacity for reproduc- 
tion by crown-sprouting or by seed . . . ." 

Hanes and Jones (1967) found prolific germination of Adenostoma seeds pro- 
ducing seedlings after fire in the San Gabriel Mountains. In a comprehensive 
study, Hanes (1971) reported that at altitudes between 1000 and 2000 feet, 27% 
of the chamise plants regrowing after fire were known to be seedlings. This figure 
might have actually been higher as some of the 22% which he reported as "un- 
determined" in origin (seedling or sprouter) may also have been seedlings. Con- 
cerning Adenostoma he wrote that "It sprouts vigorously and also germinates 
from dormant seeds, and composes about one-third of the plant cover during the 
first decade after fire . . . ." He found that altitude has a bearing on which type 
of reproduction of chamise predominates — "Chamise, for instance, has a higher 
proportion of seedlings at 1,000-2,000 ft (ca. 300-600 m) than at 2,000-4,000 ft 
(ca. 600-1,200 m)." 

In undertaking a major study of factors which affect germination of seedlings 
in Adenostoma chaparral, Christensen and Miller (1975) found that seeds of cha- 
mise germinated at least to some extent in all four conditions they studied — 
undisturbed shrub cover, artificial clearings, the first year following fire, and in 
the second year following fire. They found that the foliar leachate from chamise 



ADENOSTOMA FASCICULATUM 1 l 

was inhibitory to the germination of certain herbs and not others. They reported 
no data on the effect such foliar leachate might have on the germination of Ad- 
enostoma seeds themselves. Yet they indicated that seedling growth was wide- 
spread after fire. 

While seedling survival may be low under certain circumstances (Vogl and 
Shorr 1972; Wells 1969) the results of the present study are in clear agreement 
with Horton and Kraebel (1955), Hanes and Jones (1967), Hanes (1971) and Chris- 
tensen and Muller (1975) in that chamise seedlings survived vigorously and con- 
tributed in large measure to the postfire stands of Adenostoma on the quadrats 
studied. 

Differences in height between seedlings and crown sprouted plants existed at 
the six-year stage but had vanished after eight or nine years of growth after fire. 
Such data fit with the observation that crown sprouted plants are larger than 
corresponding seedlings during the first few years after fire. This difference may 
be attributable to several factors. Crown sprouts form on burned plants in just 
a few weeks and shoots over 12 inches tall may exist before the first rainy season 
begins. Likewise, crown sprouted plants may have a fully developed root system 
immediately after the fire whereas seedlings generally do not begin growing until 
the rains arrive which may be a period of up to six months, depending on the 
date of a particular burn. 

Seedlings on the Castaic and Oak Orchard quadrats had grown eight or nine 
years respectively to the point where they were just as high as the corresponding 
crown sprouted individuals. It may be reasoned from such data that although 
seedlings grow more slowly during the first few years after fire, they ultimately 
achieve a height which is equal to that of the sprouters. 

In the Castaic double-fire, seedlings that had grown for four years after a fire 
were able to survive a second fire in good numbers, although there was an un- 
usually large number of dead chamise crowns on this quadrat as compared to the 
number of dead crowns on the single-fire quadrats. Some of the original plants 
survived the second fire, demonstrating the ability of chamise to sprout twice in 
four years time. 

The fact that there were no seedlings evident four years after the second fire 
may have been attributable to one or more of the following factors: 

(1) low seed production during the first four years of growth after the first fire 
(1970-1974). 

(2) different moisture regime after the second fire (1974) than after the first ( 1970). 

(3) different herbivore densities after the second than after the first fire. 

Crown sprouting, speciation, and fire adaptation. — Wells evaluated twenty 
chaparral genera regarding the number of taxa they contain and the modes of 
reproduction they manifest after fire. The two genera having the largest number 
of taxa {Arctostaphylos with 75 and Ceanothus with 58) each contain whole 
sections which do not reproduce by crown sprouting. Among the other 18 genera, 
all of which regenerate after fire by sprouting as well as by seedlings, the number 
of taxa per genus is much lower, ranging downward from Quercus with twelve, 
to genera like Pickeringia (chaparral pea) and four others having only one taxon 
per genus. Wells attributed the large number of taxa in a genus to the loss of the 
ability to crown sprout which in turn forced that particular group (Arctostaphylos 



12 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

or Ceanothus) to reproduce sexually after each fire, thus causing the rate of 
natural selection and speciation to exceed that found in sprouting genera. 

Our data have little fit with Wells' genetic hypothesis since it was quite easy 
to find whole areas in the Newhall, CA. region in which seedling germination and 
survival after various fires was vigorous. If Adenostoma produces large numbers 
of seedlings that survive (as in our data and in the results of others) then speciation 
would be expected to have gone on just as rapidly as among the obligate seeders. 
Thus Wells' theory does not explain why Adenostoma has fewer taxa than either 
Arctostaphylos or Ceanothus. 

Keeley (1977) evaluated the hypothesis of Wells as ". . . not overly compel- 
ling" because the "... nonsprouters do not appear to have any such obvious 
advantage; sprouting species are very successful." Furthermore, Keeley wrote 
that "... the sprouting species of Arctostaphylos and Ceanothus also produce 
many seedlings," as we have also shown in this one locality for Adenostoma. 

Keeley has proposed instead what he called a "Stochastic Fire Hypothesis" 
(1977) to explain how both obligate seeders and sprouters may each manifest an 
adaptive advantage under different circumstances. Since the crown sprouting 
adaptation is apparently beneficial in chaparral regions, Keeley and Zedler (1978) 
addressed the question of why certain species of Ceanothus (such as C. greggii) 
and Arctostaphylos (such as A. glauca) have no capacity to sprout — reproducing 
only by seedlings after fire. 

Based on an extensive series of their own experiments and reports of others, 
they concluded that sprouting reproduction is advantageous where there has been 
an abnormally short length of time between fires. They reasoned that such a 
situation ". . . would be only slightly damaging to a sprouting species, and re- 
production could be safely deferred or reduced for the first years after fire and 
all energy dedicated to growth." The data from our double-fire plot support this 
contention in that no seedlings of Adenostoma survived when a second fire fol- 
lowed four years after the first burn. But under such circumstances sprouters of 
two classes were present — type-1 plants which grew as seedlings after the first 
fire and sprouted after the second fire and type-2 plants which sprouted after both 
fires. 

In the stochastic fire hypothesis of Keeley and Zedler, on the other hand, the 
obligate seeders such as C. greggii and Arctostaphylos glauca have a distinct 
advantage wherever there has been an especially long fire-free period before the 
burn. After a long period, they reasoned, the sprouting individuals such as Ad- 
enostoma will be few in number because of previous thinning in the old stand 
prior to the fire and death as a result of the intensity of such a fire. Keeley and 
Zedler (1978) proposed that under such circumstances "... the longer the fire- 
free period, the larger the opening after fire." Since seedlings are well equipped 
to survive in chaparral openings, the obligate seeder adaptation is of advantage 
where fires are infrequent. 

By the same token, the sprouter species are adapted for survival and spread 
where fires come in close succession. By means of the stochastic fire hypothesis 
it is thus possible to envision how both obligate seeders and sprouters can be 
successful in the same chaparral area. 



ADENOSTOMA FASCICULATUM 13 

Acknowledgments 

We thank Rex Lohoff and Tim Troyer for assistance in field studies and Joyce 
Zaiger in statistical computations. 

Literature Cited 

Christensen, N. L., and C. H. Muller. 1975. Effects of fire on factors controlling plant growth in 
Adenostoma chaparral. Ecological Monographs, 45:29-55. 

Hanes, T. L. 1971. Succession after fire in the chaparral of Southern California. Ecological Mono- 
graphs, 41(l):27-52. January. 

, and H. W. Jones. 1967. Postfire chaparral succession in Southern California. Ecology, 

48(2):259-264. Early Spring. 

Horton, J. S., and C. J. Kraebel. 1955. Development of vegetation after fire in the chamise chaparral 
of Southern California. Ecology, 36:244-262. 

Howe, G. F. 1976. Post-fire regrowth of Adenostoma fasciculatum H. & A. and Ceanothus cras- 
sifolius Torr. in relation to ecology and origins. Creation Research Society Quarterly, 12:184- 
190. 

Keeley, J. E. 1977. Fire-dependent reproductive strategies in Arctostaphylos and Ceanothus. In 
Proceedings of The Symposium on The Environmental Consequences of Fire and Fuel Man- 
agement in Mediterranean Ecosystems. U.S. Department of Agricultural Forest Service Gen- 
eral Technical Report, WO-3:39 1-396. November. 

, and P. H. Zedler. 1978. Reproduction of chaparral shrubs afterfire: a comparison of sprouting 

and seedling strategies. The American Midland Naturalist, 99(0:142-161. January. 

Sokal, R. R., and F. J. Rohlf. 1969. Biometry. W. H. Freeman Co., San Francisco. 

Vogl, R. J., and P. K. Schorr. 1972. Fire and manzanita chaparral in the San Jacinto Mountains, 
California. Ecology, 53: 1 179-1 188. 

Wells, P. V. 1969. The relation between mode of reproduction and extent of speciation in woody 
genera of the California chaparral. Evolution, 23:264-267. 

Accepted for publication August 7, 1979. 



Bull. Southern California Acad. Sci. 
79(1), 1980, pp. 13-19 

A New Frog of the Genus Eleutherodactylus 

(Leptodactylidae) from the Monteverde 

Forest Preserve, Costa Rica 

Jay M. Savage 

Abstract. — A new frog of the genus Eleutherodactylus (Leptodactylidae) from 
the Monteverde Forest Preserve, Costa Rica by Jay M. Savage, Bull. Southern 
California Acad. Sci., 79(1): 13-19, 1980. Eleutherodactylus cuaquero from the 
Monteverde Forest Reserve in the Cordillera de Tilaran, Costa Rica, is described 
as new. The species appears to be closely allied to El. andi of central montane 
slopes of Costa Rica. Both forms appear to be members of the fitzingeri species 
group. El. cuaquero is unique in the family Leptodactylidae in the condition of 
the jaw muscles with three major slips to the depressor and only an externus 
adductor (formula: DFSQAT + e). 



14 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Department of Biological Sciences and Allan Hancock Foundation, University 
of Southern California, Los Angeles, California 90007. 



In 1964 as part of the fieldwork associated with the discovery of the unusual 
toad, Bufo periglenes (Savage 1967), collections were made of other amphibians 
in the area around the famous Quaker colony at Monteverde, Provincia de Pun- 
tarenas, Costa Rica. Included in these samples were two female Eleutherodac- 
tylus resembling El. audi of the fitzingeri group (Savage 1974, 1976) that seemed 
to represent an undescribed species. Description of this form has been delayed 
in the hope that a male example might be obtained. After 15 years the hope has 
not materialized. The forest area around Monteverde has now been established 
as a preserve to protect B. periglenes, where all collecting is strictly prohibited 
and there appears to be little likelihood that another 15 year wait will produce a 
male example. For this reason it seems wise to describe the new form at this time 
as: 

Eleutherodactylus cuaquero, new species 
Figure 1 

Holotype. — LACM 128460, an adult female from 1.75 km east southeast of 
Monteverde; Provincia Puntarenas, Canton de Puntarenas, Costa Rica; 1520 m; 
collected by Jay M. Savage and Fred S. Truxal, May 17, 1964. Paratype: CRE 
7213B, an adult female, with same data as holotype. 

Diagnosis. — El. cuaquero is a member of the fitzingeri group (Savage, 1976) 
and allied to Central American members (audi, fitzingeri, rayo, and talamancae) 
of this stock having a minimal amount of toe webbing. The other Central Amer- 
ican members of the group (crassidigitus and longirostris) have the web between 
toes III— IV extending nearly to the distal subarticular tubercle on finger III and 
to halfway between proximal and penultimate subarticular tubercles on finger IV. 
From its congeners of this group with minimal webbing (features for cuaquero 
in parentheses) it is immediately distinguished from El. rayo of Costa Rica and 
El. talamancae of Atlantic lowland Nicaragua, Costa Rica and Panama which 
have the posterior thigh surface uniform (posterior thigh surface spotted). In 
addition rayo has a heel calcar (absent) and talamancae has a distinct enamel 
white stripe or series of spots along the upper lip (absent). 

The two remaining allied forms with spotted thighs {El. fitzingeri and El. audi), 
that might be confused with the new form have toe webbing that usually extends 
to the proximal subarticular tubercles between toes I— II— III and slightly beyond 
the proximal subarticular tubercles between toes III— IV (webbing barely extend- 
ing to level of proximal margin of proximal subarticular tubercle on any toe). The 
three forms may be further separated by the following features: 

audi — finger disks greatly enlarged, III— IV emarginate, equal to or broader than 
length of inner metatarsal tubercle; posterior thigh surface dark chocolate brown 
with discrete large bright-yellow (in life) stripes or spots; groin with bright-yellow 
(in life) large spots or stripes; throat almost solid black to dark brown, under- 
surfaces of body and limbs heavily marked with dark brown spots or mottling; 
light areas on posterior undersurfaces bright yellow, almost always suffused with 



ELEUTHERODACTYLUS 



15 




' - 2 3 4 



. 



5 6 

SORYAI I CENTRIFUG 



JJo o» co r~ 

U. O <» OB — 3 






liililii 



Fig. 1. Dorsal view of adult female holotype, LACM 128460 of Eleutherodactylus cuaquero. 



bright salmon red in life; adult males 40-55 mm in standard length, adult females 
65-80 mm. 

cuaquero — finger disks greatly enlarged, 1 1 1— IV emarginate, broader than 
length of inner metatarsal tubercle; posterior thigh surface dark brown, with small 
bright yellow spots or vertical stripes; groin mottled; throat and hindlimbs heavily 
marked with dark brown pigment; posterior undersuifaces bright yellow, suffused 
with bright pink in life; adult females 33-47.5 mm in standard length. 



16 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. Distribution of Eleutherodactylus cuaquero and allied species of the fitzingeri group in 
Costa Rica. The dotted line indicates the 1500 m contour. 



fitzingeri — finger disks moderately enlarged, III— IV barely indented, narrower 
than length of inner metatarsal tubercle; posterior thigh surface dark black to 
brown, with numerous small pale yellow spots in life; groin mottled or uniform; 
throat almost immaculate to heavily mottled with dark brown; undersurfaces 
white with a yellow cast posteriorly and under thighs, usually immaculate but 
sometimes weakly mottled with dull gray and rarely strongly mottled with dark 
brown; adult males to 35 mm in standard length, adult females to 52 mm. 

Summary of characteristics. — GENERAL: Head about as wide as long. Nostril 
closer to tip of snout than to eye. Dorsal outline of snout subovoid to subelliptical; 
rounded in profile. Canthus sharp; loreal outline obtuse. Choanae ovoid, vomer- 
ine teeth located between and behind choanae in two transverse series separated 
at mid-line. Tympanum internal, indistinct, oval, a little less than Vi height of 
orbit. Skin of head and upper surfaces essentially smooth with a few scattered 
pustules. Each upper eyelid with a distinct tubercle. Finger II longer than I. 



ELEUTHERODACTYLUS 17 

Finger disks about 2 times as wide as digits on fingers III— IV: rounded and 
expanded on finger I, palmate on finger II, emarginate on fingers III— IV. No 
distinct fringes on fingers. Subarticular tubercles flattened, rounded in outline, 
globular; no supernumerary tubercles; thenar tubercle elongate, palmar large cor- 
date; no accessory palmar tubercles. No calcar, although heel with several tu- 
bercles. Toe disks emarginate or notched, about 1.5 times width of digit on toes 
I-IV. Toe fringes weak. Modal toe webbing formula: 12 + - 1 /iII2 + -3 1 /iIII3 + - 
4+IV4+_3"V. Subarticular tubercles projecting, ovoid, in outline, obtuse in pro- 
file; no supernumerary tubercles; outer plantar tubercle round, l A size of inner 
oval plantar tubercle. An inner tarsal fold. Venter smooth. 

COLORATION: Upper surfaces of head and body dark brown, limbs lighter 
brown. An obscure interocular dark bar bordered anteriorly by an obscure light 
area; upper lips with three dark bars; a definite supratympanic dark mark running 
from middle of eye above tympanum and downward toward shoulder. Iris of eye 
dark brown below, upper half silver, in life. Dorsum nearly uniform with weak 
dark spots associated with dorsal tubercles. Dorsal surfaces of limbs with dark 
crossbars. Posterior surface of thigh dark brown with distinct light spots or spots 
lined up to form vertical stripes. Throat and underside of hindlimbs heavily 
marked with dark brown pigment; distinct light longitudinal stripe running length 
of throat. Plantar surface uniform. Groin and flanks without distinctive markings. 
In life, hindlimbs and groin suffused with pale rose. 

MEASUREMENTS: In the following section the measurement for the holotype 
is given first followed by that for the paratype (CRE 7213B) in parentheses. 
Standard lengths are in millimeters; other measurements are given as percentages 
of standard lengths. 

Standard length 47.5 (33.0); head length 39.6 (40.9); head width 39.4 (38.2); 
length of orbit 15.2 (16.7); snout length 18.7 (19.7); loreal length 12.0(13.0); height 
of tympanum 6.9 (7.0); hindlimb length 230.7 (227.9); tibia length 75.4 (71.8). 

Jaw musculature. — The types have the depressor mandibulae originating as 
three distinct slips, one each from the dorsal fascia, squamosal and annulus tym- 
panicus. This is the DFSQAT condition according to the system of Starrett (1968). 
An adductor mandibulae externus superficialis is present (e). 

Distribution. — Known only from the lowermost portion of the lower montane 
rainforest 1.75 km ESE Monteverde in the Cordillera de Tilaran, Costa Rica, 
1520 m (Figure 2). 

Ecological associates. — Both known examples of El. cuaquero were taken 
early in the evening (7-9 p.m.) on herbaceous vegetation 1-1.5 m high. Ecological 
associates included the frogs: Eleutherodactylus angelicas, El. cruentus. El. di- 
astema, El. melanostictus. El. ridens, Hyla rivularis, H. pseudopuma, H. uran- 
ochroa, Centrolenella colymbiphyllum and C. prosoblepon and the lizard Norops 
tropidolepis. 

Remarks. — The name cuaquero is an arbitrary combination of letters that hap- 
pens to mean Quaker in Spanish. The name is used in allusion to the habitat of 
this frog, whose type locality is located on part of the property that formed the 
original land holdings of the Quaker colony at Monteverde. The settlement was 
established in 1951 on the Pacific slope by 15 families of North American Quakers 
whose moral and economic beliefs led them to emigrate so that their taxes would 



18 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

not be used for the construction of nuclear armaments. By 1954 the Quakers had 
established a cooperative cheese factory where the famous Monteverde brand 
cheeses continue to be made. 

The Quakers from the beginning at Monteverde strongly believed in preser- 
vation of forest tracts and a watershed preserve. The flora and fauna of the area 
is especially rich and the discovery of the unique toad, Bufo periglenes Savage, 
1967, on the continental divide above Monteverde, heightened the Quakers' in- 
terest in conservation of the local biota. George Powell, originally an American 
Peace Corps worker, conceived the idea of establishing a permanent nature pre- 
serve in the area in 1971 and led a campaign to raise funds from a variety of 
sources to make the Monteverde Forest Preserve a reality. The Quaker colony 
donated 900 hectares of the some 2800 hectares that now comprise a preserve 
that involves both Atlantic and Pacific slope areas to the north and east of Mon- 
teverde proper. The preserve includes 7 major tropical plant formations and 
ranges in elevation from 1500-1842 m. The preserve is now owned and operated 
by the nonprofit Tropical Science Center of San Jose and is enjoyed by many 
visiting nature lovers each year. A visitor's center and a recently completed field 
station supplement the accommodations at Monteverde, for those interested in 
observing and/or photographing the rich biota of upland Costa Rica in this es- 
sentially undisturbed preserve. 

Relationships. — Eleutherodactyhts cuaquero clearly belongs, on the basis of 
external morphology, to the fitzingeri group (Savage 1976). Members of this stock 
have smooth venters, an inner tarsal fold, at least basal toe webbing and enlarged 
disks on all fingers and toes. Lynch (1976) has advocated inclusion of a number 
of species usually associated with the rugulosus (Savage 1975) and gollmeri (Sav- 
age 1976) groups into a larger unit including the fit zing eri group (sensu Savage), 
on the basis of trivial external features. Savage and DeWeese (1979) pointed out 
similarities between the fitzingeri and rugulosus groups in karyology and jaw 
musculature, but refrained from attempting to undertake a full critique and eval- 
uation of Lynch' s schemata. 

Up until the present time all members of the fitzingeri group whose jaw mus- 
cles had been examined had the depressor mandibulae forming a single slip with 
an origin primarily from the dorsal fascia and a few fibers coming from the squa- 
mosal and the adductor mandibulae externus superficialis present (dfsq + e, using 
the formula of Starrett 1968). El. cuaquero, which on the basis of all other features 
seems very close to El. andi, differs not only from other members of the fitzingeri 
group in which the jaw musculature is known, but is unique in the family Lep- 
todactylidae in having a formula of DFSQAT + e. The interpretation of the sig- 
nificance of this feature must await detailed studies of interspecific and intergroup 
variation in jaw muscle features. Suffice it to say that the muscle character strong- 
ly confirms the validity of El. cuaquero as a distinctive species. 

Acknowledgments 

The curators at the Museum of Natural History of Los Angeles County 
(LACM) and the Museo Zoologia, Universidad de Costa Rica, aided the research 
in several ways. James E. DeWeese and James M. Polisini of the University of 
Southern California (CRE) prepared the photograph and map, respectively. Aid 
in the fieldwork associated with rny John Simon Guggenheim Foundation Fellow- 



ELEUTHERODACTYLUS 19 

ship (1963-1964) in Costa Rica was provided by Norman J. Scott and the Orga- 
nization for Tropical Studies. It is a pleasure to express my thanks for the support 
of these institutions and individuals. 

Literature Cited 

Lynch, J. D. 1976. The species group of the South American frogs of the genus Eleutherodactylus 

(Leptodactylidae). Occ. Pap. Mus. Nat. Hist., Univ. Kansas, 61:1-24. 
Savage, J. M. 1967. An extraordinary new toad (Bufo) from Costa Rica. Rev. Biol. Trop., 14(2): 153— 

167. 
. 1974. On the leptodactylid frog called Eleutherodactylus palmatus (Boulenger) and the status 

of Hylodes fitzingeri O. Schmidt. Herp., 30(3): 289-299. 
. 1975. Systematics and distribution of the Mexican and Central American stream frogs related 

to Eleutherodactylus rugulosus. Copeia, 1975(2):254-306. 
. 1976. A preliminary handlist of the herpetofauna of Costa Rica. II edition. Editorial Univ. 

de Costa Rica: 1-19. 
, and J. E. DeWeese. 1979. A new species of leptodactylid frog, genus Eleutherodactylus, 

from the Cordillera de Talamanca, Costa Rica. Bull. South. Calif. Acad. Sci., 78(2): 107-1 15. 
Starrett, P. H. 1968. The phylogenetic significance of the jaw musculature in anuran amphibians. 

Ph.D. Dissertaiton, Univ. Michigan: 1-179. 

Accepted for publication April 10, 1980. 



Bull. Southern California Acad. Sci. 
79(1), 1980, pp. 19-37 

Navajo Social Interactions in an Urban Environment: 
An Investigation of Cognition and Behavior 

Shirley J. Fiske 1 and J. C. Weibel 2 

Abstract. — Navajo social interactions in an urban environment: an investiga- 
tion of cognition and behavior by Shirley J. Fiske and J. C. Weibel, Bull. Southern 
California Acad. Sci., 79(1): 19-37, 1980. This paper explores the relationship 
between the way people think and what they do, among a recently migrated urban 
ethnic group. The hypothesis is that people will seek assistance (housing, em- 
ployment, friends) from other people whom they perceive as most similar to 
themselves. The paper compares two sets of data in an attempt to examine this 
relationship. Both cognitive and behavioral data on social interactions were col- 
lected from Navajo women in Los Angeles, California. The data indicate that 
urban Navajo women tend to establish their social assistance networks based on 
some of the underlying dimensions of their cognitive classifications of the urban 
social world, but that in other respects their behavior diverges from the expected 

1 School of Public Administration, University of Southern California, Los An- 
geles, California 90007. - Alcohol Research Center, Neuropsychiatric Institute, 
University of California, Los Angeles, California 90024. 



20 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

pattern of social interaction. The dimension of traditional vs. acculturated, used 
as a tribal marker in choosing assistance and friends, appears to be the strongest 
predictor of behavior. Regionalism and anti-white sentiment are not as strongly 
associated as expected with choice of assistors. The anomalies in these expec- 
tations and their implications for understanding the relationship between cogni- 
tion and behavior are discussed. 



Introduction 



In the last 30 years anthropology has demonstrated much theoretical and meth- 
odological interest in determining the "native's" point of view. This trend has 
borne many labels, namely, formal analysis, componential analysis, ethnoscience, 
the "new ethnography," and cognitive anthropology. One common assumption 
is that the goal of ethnography is to understand what the native will do in any 
given situation. Thus, as phrased by Goodenough (1964:36), a goal of ethnosci- 
ence is a description "of whatever it is one has to know or believe in order to 
operate in a manner acceptable to its [a society's] members." A more basic 
premise, then, is that culture is an ideational phenomenon; it is a system of 
categories and rules for their use. 

With the aid of 25 years of hindsight it appears that cognitive anthropology has 
focused almost entirely on discovering and organizing the ideational aspects of 
culture. The enduring criticisms of ethnoscience are that it is trivial, too devoted 
to obscure methodologies, too time consuming, and above all too isolated — it has 
ignored the relevance of cognitive structure to people's everyday behavior (see 
Berreman 1966; Burling 1964; Harris 1968). 

Recognizing both sides of the issue, Keesing has proposed that the nature of 
culture is similar to the nature of language: just as linguistic behavior includes 
both competence and performance, so too culture includes an ideational aspect 
and observable behavior (Keesing 1971). The challenge inherent in conceptual- 
ization of culture lies in refining the relationship between the two aspects. 

In recent years there have been several notable attempts to link cognitive 
analyses with measures of observable behavior. Johnson (1974) shows that the 
manner in which people categorize land types and appropriate crops predicts the 
pattern of planting (i.e., the direction of deviation from a random planting pat- 
tern). He concludes that the "cognitive model does appear to summarize or 
account for the main trends in observed planting behavior" (1974:96). Addition- 
ally, Howe and Sherzer have analyzed what they call a "practical" classification 
of crops — one based on rules of access to the crops. They discuss the relevance 
of the classification to understanding behaviors of theft, generosity, and economic 
behavior (Howe and Sherzer 1975). Sanjek (1977) analyzes urban ethnic terms in 
Accra, Ghana, and compares the classification with actual interactions; Young 
analyzes Tarascan illness categories and relates the distinctions among them to 
strategies for prevention (1978). 

This paper is a continuance of this hopeful trend. We will outline the cognitive 
configuration of social groups and the actual interactional patterns which Navajo 
demonstrate. We will examine the degree of fit between the two data sets. Two 
distinctive data types are used: the first set are cognitive data, gathered using 



NAVAJO SOCIAL INTERACTIONS 21 

ethnoscientific techniques and analyzed using nonmetric scaling. The second set 
are behavioral data, gathered from intensive interviews and informant recall of 
social and assistance resources over a 10 year period in Los Angeles. 3 

Navajo in Los Angeles 

Navajo are the most heavily represented tribal group in Los Angeles (Price 
1968; U.S. Dept. of Commerce 1970). They are a recently arrived population 
from a culturally traditional rural area where the characteristic settlement pattern 
is sparse and widely dispersed. Except for the occasional visit to a bordertown, 
trading post, or the boarding school experience, the reservation social world is 
made up almost exclusively of other Navajo. 

The incentive for migration to Los Angeles is primarily economic — to find jobs, 
higher wages, and improved living conditions. Upon arrival in Los Angeles Na- 
vajo tend to find residences (or are placed by the Bureau of Indian Affairs) either 
in the the Central City or Southeast sections of Los Angeles. Both are areas of 
high Indian concentration in which friends and relatives are well established. 
Navajo are a highly mobile and geographically dispersed population, exhibiting 
a general movement away from the Central City area into the Southeast and other 
more outlying suburban areas of the Los Angeles basin. The neighborhoods in 
which Indians choose to live were originally Anglo neighborhoods, now becoming 
increasingly Latino in influence and population. 

While Navajo women generally enter the urban area as single individuals, most 
are now married. The vast majority of Navajo women have married within the 
tribe (Weibel 1977; Fiske 1975). The majority of Navajo women do not work 
outside the home but remain at home raising their young families. Those that are 
wage-earners are employed at unskilled factory, assembly, semi-skilled, and 
piece-work garment positions with commensurate incomes. Most of the women 
who do not work have husbands in blue collar positions. 

Navajo consistently exhibit strong ties to their cultural tradition. The need to 
perpetuate certain Navajo crafts and skills, the use of traditional medicine and 
curing ceremonies and tribal language in the urban environment is still strongly 
felt by most people. Annual and seasonal trips back to the Navajo reservation 
are a way of life for a major segment of this population even after spending most 
of their adult lives in the urban environment. As the ethnographic data indicate, 
when Navajo migrate to urban areas the strong interactional ties within the tribe 
tend to be perpetuated (Ablon 1964; Fiske 1977; Hirabayashi 1972; Snyder 1971). 
Not only do they limit their interactions with Anglos, other ethnic groups, and 
Indians other than Navajos, but also their use of institutionalized public services 
is minimal. This social phenomena evokes several separate but related ques- 
tions — how do Navajo organize and classify their social world? Given a social 
environment in which alternative assistance sources and friendship choices are 



:i The collection of the cognitive data was supported by a National Institute of General Medical 
Sciences fieldwork grant (GM 01485-04), and by a Ford Foundation Dissertation Fellowship in Ethnic 
Studies. Funds for the behavioral research were provided by a Ford Foundation grant (710-0370) 
administered through the American Indian Studies Center at University of California, Los Angeles. 
We wish to express our gratitude to Professors Allen Johnson, Jim Lincoln, Tom Weisner, Walter 
Goldschmidt, and Roy D'Andrade for their useful comments on earlier drafts of this paper. 



22 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

multiplied, how do the Navajo solve the problems of living in a large cosmopolitan 
area? Is there any relationship between the social conceptualization and the peo- 
ple Navajo choose to assist them? 

Cognitive Structure 

Previous research with Navajo indicates that there is a domain of "living 
things," including "those with five fingers" or "people," and that the classifi- 
cation is used in traditional reservation areas (Perchonock and Werner 1969); but 
the classification does not include the most recent additions to the domain of 
people; namely, the tremendous variety of people experienced by Navajo when 
they reach a large metropolitan area such as Los Angeles. What happens to this 
new information? How is it integrated into peoples' cognitive organization? 

Ethnoscientific techniques of elicitation frames (see Black 1969; Metzger and 
Williams 1966) and sorting tasks were used to discover the categories and inclu- 
sion/exclusion relations among all the terms within the domain. The analysis was 
carried one step further; lexical items were tested using both paired comparisons 
and triads tests. The triads and paired comparison tests were administered to the 
set of terms in Figures 1 and 2 (see Romney and D'Andrade 1964; Berlin, Breed- 
love, and Raven 1968; Nerlove and Burton 1972). Triads testing requires a re- 
spondent to choose the most similar (or dissimilar) terms in a set of three; for 
example, in the triad of intertribal terms: Navajo, Choctaw, and Cherokee, the 
two terms Choctaw and Cherokee were judged to be most similar. Triads data 
can be arranged in similarity matrices which yield quantitative information on 
how similar terms are to each other. 

Paired comparisons require the respondent to verbalize both the similarities 
and differences between every possible pair of terms; this is a very useful (albeit 
time consuming) technique for discovering the semantic bases on which the triads 
judgments are made. Thus, for example, if the two terms Choctaw and Cherokee 
have a similarity measure of 89/100 by triad testing (that is, 89 times out of 100, 
these two tribes were judged to be most similar out of the triad of terms), the 
researcher does not necessarily have any understanding as to why they are judged 
most similar. Paired comparisons provide these data; each term is compared with 
every other term, such that the respondents generate information and distinctions 
which are important to them. For instance, Cherokee and Choctaw share the 
attributes of being 

"from the East; both show signs of a lot of education — go to college; both 
were the first to approve of their kids going to school; a lot of them are 
teachers; usually work in offices." 

The two tribes are different in "their skin shade. Cherokee are light; Choctaw 
are darker." This kind of information about the content of the terms allows the 
researcher to interpret the triads data and subsequent scaling plots more com- 
pletely. 

The results of the triads tests were arranged in similarity matrices and analyzed 
using nonmetric multidimensional scaling to assess structure in the domain. Non- 
metric multidimensional scaling indicates by arranging in multidimensional space 
the configuration which best represents the similarity data generated by the testing 
procedures (see Shepard 1962; Kruskal 1964; Burton and Romney 1975). 



NAVAJO SOCIAL INTERACTIONS 23 

The cognitive data were gathered from three female Navajo respondents in a 
series of intensive interviews, elicitation, and testing sessions from 1971-1973. 
The small sample is a limitation to generalization, but there is reason to believe 
that the respondents and data are representative of the larger Navajo population. 
The women were purposively selected because their rural-urban experiences and 
current lifestyles were similar to many of the urban migrants. This is confirmed 
by comparison with Weibers 47 respondents in her independent study (1977); the 
women in both studies are similar in length of time spent in Los Angeles (7.3 
years in 1971 vs. 9.7 years in 1975), years of education (1 1.6 years vs. 1 1.2 years), 
and age (29.3 in 1971 vs. 30.2 in 1975). The women were selected from both 
suburban and downtown population areas and they represent both traditional and 
less traditional Navajo backgrounds. In addition there was a high degree of agree- 
ment among the respondents as to the terms and their structure. The classification 
is validated by Navajo behavior in the pan-Indian arena (Fiske 1977), and by the 
nature of the inter-ethnic relations established by Navajo women (Fiske 1978). 

The multidimensional scaling results can be seen in Figures 1 and 2 in two 
separate analyses. As Burton and Nerlove point out, it is important that triads 
data be tested from the same semantic domain and be on the same level of 
contrast (1975:248). Hence, two separate testing procedures and analyses were 
carried out — one from the contrast level of ethnic groups (Fig. 2) and one for the 
intertribal contrast level under the cover term bitsi'yishttizhii ('Indians") (Fig. 
1). 

As can be seen in Figure 1, there are two main clusters of people in the domain 
"five fingered beings"'; (a) Indians and Mexicans cluster together at the extreme 
lower pole of the vertical dimension, (b) 'foreigners,' 'people from overseas,' 
'enemies,' and 'white Americans' cluster together at the extreme top of the con- 
figuration. Asians and Blacks are in intermediate positions peripheral to either 
cluster. 

In interpreting the Figure 1, the strongest dimension is the vertical one of social 
distance between Indians and Mexicans, and the constellation of concepts which 
represent Caucasians. Both Asians and Blacks are intermediate in social distance. 
Mexicans are unequivocably the most psychologically accessible group to Indians 
(from the Navajo perspective). 

Turning to the second analysis — the intertribal domain — it appears that Navajo 
women conceptualize other Indians in two major clusters which can be inter- 
preted along two dimensions. Figure 2 illustrates the configuration of the partial 
domain bitsi'yishttizhii. The cluster on the upper half of the scaling plot, which 
includes Navajo-Apache and Hopi-Zuni dyads, is considered to be a Southwestern 
group of tribes; that is, they share a regional and ecological base with Navajo. 
The clusters on the lower half of the configuration are from very different geo- 
graphical bases, and at the farthest regional extremity from Navajo are hak'az 
dine'e, 'Eskimo,' who live in a region of snow and cold — very different from the 
Navajo desert environment. Hence, the two clusters are arranged along a vertical 
dimension of regionalism, ranging from the 'Pima' (keti'ahi) in the extreme 
Southwest, through Navajo and Oklahoma Indians, and with 'Eskimo' on the 
polar end of the dimension. 

The two clusters also demonstrated a horizontal dimension of traditionalism 
and acculturation. On the extreme right-hand side of the configuration are dine 



24 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



bilagaana « 
('white Americans') 




, t<5 hinaande dine'e ("foreigner from 



anaa 1 ( enemy ) 



overseas") 



»aaj igi ("slant eye enemies") 




zhinii • 
('Black') 



•benaats' 6zl ('Asians') 



Fig. 1. 



nakaii ('Mexican') 

] 

• bitzi'yisht^izhii 
('Indian') 



Multidimensional scaling plot for triads test, ethnic categories. The clusters indicate hi- 
erarchical clustering results (see Johnson 1967; D'Andrade 1973). 



('Navajo'), who in their own opinion are the most traditional — the most "Indian" 
of all the tribes. On the extreme left-hand side of the tribal array are the Cherokee, 
a tribe who the Navajo feel have capitulated their tribal customs to become part 
of the white world. 

As mentioned previously, Navajo perceive themselves to occupy the position 
of most extreme traditionalism. Clustered closely with them are Apache, Hopi, 
and Zuni; these core traditional tribes are perceived as retaining their language, 
traditional housing, clothing, hair styles, and ceremonies. Pima and Paiute are 
seen as somewhat more acculturated than the core tribes; but they along with 
California Indians and Eskimo 4 are seen as having lost most of their traditional 
culture, retaining fewer of their religious ceremonies, and having less dependence 



4 The Eskimo are perceived as very traditional by most Navajo, but this is not reflected in their 
position on the scaling plot because of general unfamiliarity with specific traditions. 






NAVAJO SOCIAL INTERACTIONS 



25 




Fig. 2. Multidimensional scaling plot for triads test, intertribal categories. 



on traditional subsistence activities (such as growing one's own food, making 
one's own household utensils). 

Moving farther to the left-hand side along the traditional/acculturation dimen- 
sion, the Choctaw, Cherokee, and Sioux are perceived as least traditional of the 
major tribes. On the extreme pole of the acculturation dimension, Cherokee are 
characterized as "eating out of cans, like the whites," having lots of education, 
not being shy around whites, being more city-oriented and less reservation-ori- 
ented, not speaking their language, etc." 1 

To summarize, in interpreting the social-psychological classification of Navajo 
women, there is a strong affinity for Mexican-Americans, an ambivalence towards 
Blacks and Asians, and definite social distance from the constellation of concepts 
for Caucasians. In the intertribal domain the social world is clustered into two 



:> For a more detailed analysis of Navajo perceptions of the intertribal domain as it relates to 
participation in pan-Indian organizations and social interaction, see Fiske 1977, and 1975. 



26 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

main groups: those traditional tribes of the Southwest, with whom the Navajo 
feel most comfortable and share a traditional orientation; and a second cluster, 
with whom the Navajo feel they share less in common regionally and who have 
become more acculturated than the Navajo. We might expect, then, given these 
cognitive organizations of the social universe, that Navajo women would prefer 
to interact with members of those social categories with whom they share the 
most identifying traits — Mexican-Americans among major ethnic groups, and 
Navajo and Southwestern Indians among Indians. 

Research has shown that as perception of similarity between items increase 
(i.e., the more traits, dimensions, etc. which are shared in common between 
items), then those items will receive similar behavioral responses (Romney and 
D'Andrade 1964; D'Andrade 1974). In other words, if Navajo judge themselves 
to be most similar with Apache, Hopi, and Zuni, and least similar to Oklahoma 
Indians, then one would expect that they will behave toward these Southwestern 
Tribes as they would toward members of their own tribe — initiate social relations, 
friendships, seek assistance, and so on. One would not expect these kinds of 
behaviors to be directed toward people for whom there was little demonstrated 
judgment of similarity, namely the Oklahoma Indians. 

Social Interactional Patterns 

In an independent study conducted in 1975 and 1976, Weibel worked with 
Navajo, Choctaw, and Cherokee, Creek, Seminole, and Chickasaw informants' 5 
who had migrated to Los Angeles since 1950. Starting with a list of clients from 
a major Los Angeles Indian health service agency and by initiating a system of 
personal referrals, Weibel was able to interview intensively 47 Navajo and 37 
Oklahoman Indians who, at the time of the interview, were predominantly young 
mothers with pre-school and school age children still in the home. The women 
were 30.2 years old on the average, had 11.2 years of schooling, and were living 
in neighborhoods with approximately the same socioeconomic status. These areas 
are predominantly low income, older working class white or Mexican-American 
neighborhoods. For purposes of analysis here, we will utilize only the Navajo 
patterns of interaction. 

The interview consisted of both open-ended and semi-structured questions 
which elicited personal history of pre-Los Angeles life, the migratory process, 
and the problem-solving strategies they employed in the re-establishment of their 
lives in the urban environment. The respondents were asked to recall real life 
situations in which they had sought out information and assistance in the urban 
setting for such necessities as housing, employment, medical services, and gen- 
eral social services. To account for the influence of time in the city on their 
behavior, the respondents were asked to recall both their initial and their most 
recent assistance seeking situations. 

Opportunely, one of three measures of assistance patterns is ethnicity. The vari- 



6 Choctaw, Cherokee, Creek, Seminole, and Chickasaw Indians are the tribes who make up the so- 
called "Five Civilized Tribes." Historically agriculturalists indigenous to the southeastern sector of 
the United States, their ancestors were "removed" as a matter of federal policy to Oklahoma in 1830. 
In the interest of time, space, and enlightened anthropology we chose not to use the term "Five 
Civilized Tribes"; subsequently, this sub-cultural group will be referred to as the Oklahoma Indians. 



NAVAJO SOCIAL INTERACTIONS 27 

able was coded in such a way that the descriptive categories are comparable to the 
ethnic categories of Figures 1 and 2. These sets of data, then, lend themselves 
to a comparison of cognitive perceptions and actual social interaction in the urban 
milieu. We suggest that the cognitive categories and perceptions of social distance 
as identified by Fiske's work (1975, 1977) are operative and influence the selec- 
tion, out of a wide array of possible ethnic categories, of those people whom one 
cognizes as most similar. 

Given the previous discussion about the cognitive classification, we hypoth- 
esize that Navajo interact with other Navajo significantly more often than they 
do with other Indian groups; that they prefer to interact with Southwestern In- 
dians over other tribes; and that they interact least with the most acculturated 
and regionally different Indians (Oklahomans). Further, they interact more with 
other Indians and Mexican-Americans than they do with Anglos. Three behav- 
ioral areas are examined — locating housing, finding a job, and choosing friends. 
The first two are situations of involuntary action, where the ethnicity of assister 
may be a coincidental rather than purposive choice; the friendship network, how- 
ever, is a matter of voluntary choice and personal preference. These two types 
of behavior patterns are chosen in hopes of discerning a difference between them. 

Finding a Place to Live 

One of the first problem-solving situations with which a person deals in the 
migration process is the location of a permanent place of residence. Two ques- 
tions were asked to elicit this information: "How did you go about finding the 
first place you lived in Los Angeles? 1 ' and "How did you go about locating the 
house you presently live in?" 

As can be seen in Table 1, in the initial search for housing, the assistance given 
by the Bureau of Indian Affairs (BIA) field office in Los Angeles was consider- 
able; 50% of the Navajo's initial residence assisters were BIA counselors. If the 
individual had migrated to Los Angeles on her own, the most salient housing 
assister was another tribesperson or kin (44% of the assisters). Only three Navajo 
sought out housing assistance initially from anyone other than the BIA or another 
Navajo; the other assisters included a Mexican-American in-law, a Caucasian 
family for whom the participant had worked as a domestic, and the tribal news- 
paper The Navajo Times. 

Importantly, in the initial housing search no Navajo interacted purposively with 
an Oklahoma Indian, who are conceptualized as being least like Navajo, even 
though there are proportionately more Oklahoma Indians than Navajo in the Los 
Angeles area. These two tribal groups constitute the two most heavily represented 
tribal groups in Los Angeles (Price 1972). One reason for this tribal exclusivity 
is that the bulk of the "same tribe" assisters were kin. This pattern of kin as the 
initial gatekeeper is a cross-cultural phenomenon of rural-to-urban migration 
(Aldous 1962; Gulick 1965; Farsoun 1970; Gans 1962; Hauser 1965; Snyder 1971). 

Table 1 compares the initial search for housing with the strategy for finding the 
present residence. If the city exerts a sophisticating influence on interactional 
behaviors this early pattern of ethnic exclusivity should be modified in the later 
housing search. 

In fact, in their most recent move, Navajos were even more exclusive in their 
choice of residence assisters (see Table 1). Fifty-seven percent of the Navajo 



28 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



f- 



<U cS 



08 2 
o "3 



<u CO 

5 '-5 
O = 



5 I 
O 2 



£ 



$ 



~ fc\ 00 



^ 



(N w rJ 









^ 



^ C 



<n ^ 



c# IN 









^ 









8? o 

— (N 



^ 



^ 



^ 



^ Ci 



^ Ci 



^ <3 



oo m 



OO lr> 



^ <N 






^£ ^ 






^ - ^ 



o tc 



o .3 

-rr co 
Q. co 
g «3 



% 



Zro 
X) 

2 c rt " 



o 
o 
> 

co 

u -a 

uf.<5 

■ — - c co 

„i D CO 
~ C2 J5 

c5 j£ - 
o J2 & 

ra ._ co 

■is' 

0- U u 

- , c 



N 



O J2 C U ._* O 3 



c -"5 



o U 



- X 



^H 






'J 

— -1- fl) 

< U CO 



NAVAJO SOCIAL INTERACTIONS 29 

sample interacted with only other Navajos. Dependence on the BIA was greatly 
reduced (two interactions). Two women used the help of Sioux and Mexican- 
American friends or neighbors. One used the help of a Winnebago and Cherokee 
friend from an all-Indian church. Interestingly, four Navajos chose assisters who 
were Caucasian, and eight answered ads in the classified section of a newspaper 
or employed the services of a real estate agent in locating their present housing. 
This distribution suggests some sophistication in use of urban institutions on the 
part of the Navajo sample, but indicates a continuing preference on the part of 
the Navajo sample for intra-tribal assistance in problem-solving situations. 

The fact that the BIA was the most salient assister in the initial search for 
housing raises an interesting question in that Fiske's (1975) study elicited no 
equivalent cognitive social category. Where does this salient assister fit in a hi- 
erarchy of cognitive categories? If pressed, the women could remember that the 
BIA counselor was usually white. Occasionally an Indian would surface at the 
BIA field office; but that was not often the case, and we feel that the actual 
ethnicity of the BIA counselor is not an important variable since the participants 
who used the BIA relocation services exercised no personal selection in the 
ethnicity of their BIA counselor. The individual was simply assigned to the next 
available counselor when they arrived to be processed. 

We suggest that the BIA is a buffer or marginal social category (Snyder 1971). 
When an Indian decided to move to the city, the migrant had the further choice 
of either going it alone (perhaps with the assistance of friends or kin already 
settled in the target urban center) or making the transition with the sponsorship 
of a BIA relocation program. That 50% of the Navajo sample chose to avail 
themselves of BIA assistance programs in making the shift from reservation or 
rural life to an urban-industrial lifestyle is indicative of the BIA's mediating in- 
fluence and salience in the lives of the relocated Indian families. The BIA is a 
social category which serves initially as a bridge between two social interactional 
milieux (the reservation and the city); its salience decreases over time, presum- 
ably because ties with the Relocation Program are severed after housing is located 
and employment secured. 

One further characteristic of these interactional patterns should be noted. Thir- 
ty-seven percent of the Navajos 1 last house hunting strategies consisted of self 
reports something like this: "Well after a while I got to know where the cheap 
housing was and where they would take kids, so I just went to that place and 
walked up and down the streets looking for 'For Rent' signs until I found a 
place." This self-reliance or "walk and search" strategy frequently recurred as 
an effective problem-solving technique of Navajo families. It suggests that these 
families have successfully developed a cognitive map of their urban environment 
and rather than initiate extratribal interactions have elected to problem-solve 
independently. When this behavior occurred, therefore, it was coded as an ex- 
clusively Navajo interactional pattern. 

To summarize, even after an average of eight years in the city, the intra-tribal 
assistance pattern is the modal preference for Navajo women. Interaction with 
other tribes in the housing search is remarkably low. There were no cases of inter- 
action with other Southwestern tribes, and after eight years, the Navajo women 
sought help from only one Sioux and one Oklahoman Indian. Among the other 
ethnic groups, Navajo sought help from Mexicans at a low but consistent rate. 



30 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

The proportion of Navajo who contact Caucasians for help in locating housing 
increased somewhat over time, as did the percent of Navajo who used secondary 
sources to find housing. The discrepancies from expected behavior will be dis- 
cussed more fully below. 

Finding a Job 

Locating steady employment in the urban work market pulls the individual into 
more heterogeneous social situations. If the traditional interactional patterns per- 
sist over time, they would suggest a strong cultural bias in the inclination to seek 
out one specific type of assister over the vast array of assistance possibilities in 
the urban environment. 

The data indicate that essentially the same interactional patterns occur in the 
employment seeking situations as occurred in the search for housing. A third of 
the Navajo preferred the help of a Navajo friend, kinsman, or the self-reliant 
"walk and search" strategy in locating their first job in the city. Initially, 23 
percent used the BIA vocational services and another 26 percent were assisted 
in their initial job search by the employment offices of the vocational schools in 
which they had been placed by the BIA vocational training program. Indians, 
other than Navajos, constituted only 9% of the interactions. There were no in- 
teractions with Oklahoma and Sioux Indians. Only three women were assisted 
by Anglo friends and one college student found work because of a suggestion by 
a Filipino classmate. 

The general pattern of these assistance-seeking behaviors tends to persist over 
time. In the last employment search, once again, there is a very small proportion 
of interactions with Southwestern Indians; however, there are no cases of seeking 
assistance from either Sioux or Oklahoman Indians over the ten-year period. 
There is a continued reliance on a small number of Caucasians, and one quarter 
of the population use secondary sources to find employment. 

One problem in comparing initial and last job search strategies is that 63% of 
the Navajo women who ever worked in the city were no longer working at the 
time of the interview. Of those who are still working, the modal strategy is to 
handle the job hunt independently or ask for the assistance of a Navajo friend. 

One trend which appears to be borne out by both types of assistance-seeking 
behavior is that Navajo groups apparently interact in exclusion of other Indians. 
83% of all the interactions among Indians were either "self-help" or the enlist- 
ment of help from one's own tribal members. This exclusivity is underscored by 
the relative availability of contacts with other Indian tribes, and particularly with 
Oklahoman Indians, who are also present in the neighborhoods in similar pro- 
portions. We suggest that the decision to seek the assistance of members from 
one ethnic group rather than another is a selective process that may reflect the 
cognitive salience of relative ethnic affinity. 

Socio-Affective Ties 

As has been suggested, assistance patterns in the urban environment are not 
always a statement of personal preference. The circumstances around the initial 
migration process and the vicissitudes of the job market in the urban-industrial 
complex place certain constraints upon the individual's selection of an effective 
assister. Alternatively, who one chooses to associate with during one's non-work- 



NAVAJO SOCIAL INTERACTIONS 

Table 2. Tribal affiliation of five closest Indian friends by tribe of respondent. 



31 











Tribal affiliation of friends 






Tribe of 
respondent 


Navajo 


South- 
western 


Sioux 


Oklahoma 


Other* 




Navajo 



E 


115 
(65.96) 


6 
(5.95) 


5 
(4.33) 


5 
(43.79) 


-> 
(12.98) 


n = 133 


Oklahoman 



E 


7 
(56.04) 


5 
(5.05) 


3 
(3.67) 


76 

(37.21) 


22 
(11.02) 


n = 113 


Total 




122 


11 


8 


81 


24 


246 



X- = 174.53. 

df = 4. 

P = .001. 

* The category "other" includes one Chippewa and one Umatilla friend among the Navajo. The 
Oklahomans listed two friends from each of the following tribes: Arapaho. Winnebago, Athabascan, 
Mission, and one friend from each of the following tribes: Shoshone, Lummi, Menominee, Ankara, 
Cheyenne, Porno, Sac and Fox, Kiowa, Caddo, Shawnee, and Natchez. Oklahomans could not iden- 
tify the tribe of one closest friend but knew that one was from Oklahoma originally and one was from 
some place other than Oklahoma. 



ing, non-problem solving time is an exercise of relatively greater personal choice 
and control. If the social cognitive categories are translated into observable be- 
haviors, we would expect to observe more intra-tribal and intra-regional ties, and 
relatively fewer purely social interactions between the Navajo women and mem- 
bers of tribal groups outside the Southwest and ethnic groups other than Indian. 

The women were asked to identify, by tribal affiliation or ethnicity, their five 
closest friends. The distribution of closest friends across the salient cognitive 
ethnic categories is summarized in Table 2. Navajo overwhelmingly (74%) seek 
out and maintain close friendship ties with other Indians rather than other ethnic 
groups. In addition, 64% of the Navajo's Indian friends are other Navajo. Con- 
trary to our expectations, there seems to be relatively little difference in the 
frequency with which Navajo choose friends from among Southwestern, Sioux, 
and Oklahoma Indians. 

Is this pattern of Navajo friendships different from other tribal groups? How 
can we suggest that these cognitive configurations correspond with behavior 
which is specific to Navajo? Using data collected by Weibel ( 1977) on Oklahoman 
Indian women, we are able to compare Navajo and Oklahoman patterns of friend- 
ship in Table 2 and Table 3 below. 

The results show an interesting pattern. The discrepancy between the observed 
frequency and the statistically expected frequency of interactions suggests that 
Navajos do, indeed seek out other Navajos in the ethnically heterogeneous social 
milieu. They cultivate those friendships on a significantly higher level than chance 
alone would lead one to expect. In particular, there is virtual exclusion between 
Navajo-Oklahoma social networks. These data indicate that Navajo choice of 
friends is based on a quite different set of assumptions from those of the Okla- 
homans. Given the Navajo perception of Oklahoman Indians as irrevocably as- 
similated into Caucasian society, the behavior is quite understandable. 



32 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 3. Ethnicity of five closest friends by tribe of respondent. 



Tribe of 
respondent 








Ethnicity 


of friends 








Indian 


Mexican 


Asian 


Black 


Caucasian 




Navajo 





133 

(74%) 


20 
(11%) 


5 
(.03%) 


2 
(.01%) 


19 

(11%) 


n = 179 




E 


129.13 


15.22 


2.62 


2.10 


29.92 




Oklahoman 





133 

(70%) 


9 

(6%) 




(0%) 


2 
(.01%) 


38 

(23%) 


n = 162 




E 


116.87 


(13.78) 


2.38 


1.90 


(27.08) 




Total 




246 


29 


5 


4 


57 


341 



X 1 = 18.44. 
df = 4. 
P = .01. 



Table 3 reveals Navajo ethnic preferences for friends in light of comparable 
data for Oklahomans. The preference of both groups for Indian friends is clear. 
Additionally it appears that Navajos have more Mexican friends than do Okla- 
homan Indians. Friendships with Caucasians occur at higher levels among Okla- 
homan Indians, and are lower than expected for Navajo. These data again suggest 
that Navajo may be operating with a different "cognitive screen" which tends to 
favor Mexican relative to Caucasian contacts. Given the fact that Navajo and 
Oklahomans live in Los Angeles in the same areas, the fact that Navajo have twice 
as many Mexican-American friendship ties as do Oklahomans in a predominantly 
Caucasian social milieu, suggests a relative ease with which Navajo initiate and 
maintain close social ties with Mexican-Americans and their relative reluctance 
to interact socially with Caucasians. 

Discussion 

Our original hypothesis was based on a cognitive analysis of the domain of 
"five fingered beings," including the major ethnic groups and pan-tribal array. 
We hypothesized that Navajo women would seek assistance and social ties from 
Navajo, Southwestern Indian, Sioux, and Oklahoma Indians in decreasing order 
of frequency, corresponding to the social dimensions perceived among Indians 
(Fig. 1). Secondly, based on the analysis of the configuration of major ethnic 
groups (Fig. 2), we hypothesized that Navajo would interact most frequently with 
Caucasians (Asians and Blacks in intermediate positions). 

The behavioral data bear out some but not all of our expectations. Considering 
the pan-tribal data first, it appears that the polar extremes of the traditionalism 
dimension are supported: there is clear preference for Navajo assistance and 
friends, and Oklahoman Indians are excluded. An important confirmation of this 
tendency is the increase rather than decrease of the proportion of Navajo-Navajo 
contacts over time in the city and the mutual exclusion between Navajo and 
Oklahoma networks shown in Table 2. The interpretation of the interactional 
trends with other tribes is less clear. We had expected a much greater proportion 
of Southwestern Indians to be included in the Navajo assistance and friendship 



NAVAJO SOCIAL INTERACTIONS 33 

networks. In general, there is a relative flatness to the distribution of pan-Indian 
interactions that was unexpected. The finely tuned gradient of ethnic preference 
which characterizes the cognitive analysis does not appear in the behavioral data. 

Several factors must be taken into consideration in interpreting the results. 
Foremost is the fact that there are numerically fewer Southwest Indians than 
there are Oklahoma Indians in the Los Angeles area (Price 1972: 43 1). Aggregating 
Price's data on tribal representation in Los Angeles, the Indians from eastern 
Oklahoma constitute 21.1 percent, Navajo 13.7 percent, Sioux 11.7 percent, and 
Southwestern Indians other than Navajo 13.7 percent of the Los Angeles Indian 
population. Given the preponderance of Oklahoma Indians, the near abstention 
from Oklahoma contacts indicates the strength of the traditionalism dimension in 
the cognitive analysis. Navajos tend to stay away from Oklahoma Indians even 
though it would be relatively easy numerically to seek help or initiate friendship 
with them. Navajos see Oklahoma Indians as acculturated, nontraditional, and 
not like Navajo; and they infrequently seek assistance or friendship with them. 
The relatively low interactions with Southwestern Indians and other Indians in 
general may be a function of sample size, relative numbers of tribal representa- 
tives in Los Angeles, and geographical dispersion. 

Turning to the analysis of the major ethnic groups, we hypothesized that Navajo 
would interact most frequently with Mexican- Americans, and least frequently 
with Caucasians. We did find a tendency on the part of Navajo to seek out and 
maintain close ties with Mexican-Americans more often than other ethnic groups 
beside Caucasians. The proportion of interactions is low but consistently greater 
than with other ethnic groups beside Caucasian. In the case of friendship ties, 
Navajo women had more Mexican-American friends than other Indians com- 
bined. 

Given the extreme social distance between Anglos and Navajos, as expressed 
by the Navajo dimension of enemy/stranger which orders the ethnic group con- 
figuration, the frequency with which Navajos indicated close social ties with 
Caucasians was an unexpected finding. Several factors may have influenced this 
apparent anomaly. It must be remembered that Navajo live in neighborhoods 
which are, on the average, 75% Caucasian and 22% Mexican-American (U.S. 
Census 1970). 

The relatively frequent interaction with Caucasians may reflect the natural 
probability of Navajo-Anglo interactions in a predominantly urban environment. 
Additionally, over half of the Caucasian friends mentioned by Navajos are mem- 
bers of Christian churches in which Navajo women are active members. These 
fellowships provide the bulk of their social-interactional structure and mitigate or 
override the cognitive dimensions. While Navajo interactions with Anglos may 
appear high in absolute number, the Oklahoman interactional patterns shown in 
Table 3 suggest that, relative to other tribes, Navajo may initiate fewer friendships 
with Caucasians. 

In summary, the polar extremes on the traditionalism dimension of the cogni- 
tive analysis are supported — the Navajo prefer other Navajo and avoid the less 
traditional Oklahoman Indians. The regionalism dimension of Southwestern pref- 
erence is not clearly supported by behavioral data. Preference for Indians above 
all other ethnic groups is clearly supported. There appears to be a preference for 
Mexican assistance and social ties, but again it is not an overwhelming choice. 



34 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

The dimension of enemy/stranger which orders the ethnic groups appears to be 
diluted by the pragmatics of urban living, and more Navajo-Caucasian interac- 
tions are reported than expected. 

Cognition and Behavior — Situational Determinants 

The field of ethnoscience and cognitive anthropology has consistently been 
criticized as producing formally correct but inapplicable analyses of linguistic 
phenomena. A hopeful trend in recent years has been the growing interest in 
understanding the relationship between people's classificatory systems and how 
these systems influence real world behavior. Johnson (1974) compares Brazilian 
sharecroppers' cognitive categories of land types with their actual choice of land 
for planting crops. He found that beliefs about land types did predict the deviation 
of choice away from random selection; but in addition he offered the caveat that 
"no simple cognitive paradigm can predict behavioral outcomes absolutely" 
(1974b:93). Howe and Sherzer (1975) consistently elicit considerable variation 
among respondents as to their classification of rules of access to crops among the 
San Bias Cuna. 

In an article which analyzes ethnic cognitive domains in urban Ghana, Sanjek 
(1977) finds close correspondence between the "implicit underlying structure of 
the domain" and the frequency of interaction with those categories of people 
(1977:61 1). The author found that 

There is considerable correspondence between the two sets of data. While 
precise prediction of cognitive salience from behavioral data is not possible, 
a more general prediction of the first five, second five, and subsequent ten 
most frequently encountered southern Ghanaian ethnic identities can be 
made. The opposite prediction from cognitive salience to behavior is also 
possible. (Sanjek 1977:612) (authors' emphasis) 

Johnson and Bond (1974) compare the expectations for appropriate exchange 
with actual behavior using food sharing in tribal and peasant groups as examples. 
The authors conclude that behavior may contrast with cognitive expectations, 
and that the presentation of only the normative descriptions of a community may 
portray a one-sided view which "obscures certain practical aspects of social life 
in each community" (Johnson and Bond 1974:56). 

The growing number of studies in the area of cognition and behavior indicate 
that the relationship is complex, and at times only very general correlates are 
found empirically between the two. In Sanjek's investigation of ethnic terms in 
the Adabraka section of Accra, Johnson's analysis of planting rules and practices, 
and Young's (1978) study of disease categories and illness prevention strategies, 
there is a support for the argument that cognition and behavior correspond to a 
considerable degree. Johnson and Bond's work comparing ideal and real inter- 
personal exchange structures among the Boa Ventura and Muyombe, on the other 
hand, indicate very little correspondence between cognition and behavior. The 
findings reported here suggest that very general predicted correspondences be- 
tween categories and behavior are realized among Navajo women in Los Angeles, 
but there is some deviation from the expected pattern. The data suggest that to 
assume a close linear correspondence between cognitive dimensions and behavior 
may be an overstatement. 



NAVAJO SOCIAL INTERACTIONS 35 

In understanding the relationship between cognitive dimensions and behavior, 
other researchers have suggested the importance of such situational variables as 
historical and economic forces (Howe and Sherzer 1975:443; Young 1978:94). 
The data in this study suggest more specifically that the demographic profile of 
the immediate urban environment and the nature of the bureaucratic-institutional 
milieu may affect the use of cognitive dimensions. Both of these factors 
appear to influence the relationship between the cognitive dimensions of 
Navajo and the actual behavioral outcomes. These factors are considered to be 
situational variables because they are demographically specific to Los Angeles, 
to American social service delivery and public life, and to the historical relation- 
ship between the U.S. and Indians; also, they are not informant-based variability 
such as the twelve dimensions discussed by Sanjek 1977:618, or as found in Howe 
and Herzer 1975. 

Some of the demographic factors have been discussed previously. An over- 
whelming proportion of the population with whom Navajo must interact in Los 
Angeles are Caucasian, and the Indian population is geographically dispersed 
throughout the metropolitan area. In addition, Navajos realize that Anglos have 
greater access to economic rewards than other population groups. Thus, when 
Sanjek (1977:612) found that rank order salience of ethnic terms among Ada- 
brakan residents corresponded with the frequency of interaction among those 
groups, one must consider that the demographic distribution among the tribes in 
Accra was more evenly divided than is the proportion of Navajo, Caucasian, 
Mexican- American, and other ethnic groups in Los Angeles. The residents of 
Adabraka had a better opportunity based on chance alone to interact with pre- 
ferred groups than do Navajo in Los Angeles who are usually without transpor- 
tation and limited to contacts within the immediate Anglo community. Therefore, 
while the cognitive classification of Anglos is extremely distant, demographic 
factors and realities of economic stratification increase the chances that Navajo 
will interact with Caucasians — a practical choice not necessarily based on one's 
cognitive dimension. 

Finally, in order to understand the interaction between cognition and behavior, 
it is important to understand the bureaucratic-institutional milieu of the popula- 
tion. Sanjek notes the importance of educational and national cultural policy on 
the ethnic categories in Ghana ( 1977:617); Howe and Sherzer emphasize the Cuna 
classification is "neither static nor timeless ... it reflects both the situation in 
which the Cuna now find themselves and the dominant social and economic 
changes they have undergone in the last century, and we cannot fully understand 
the classification without taking these processes into account . . ." (1975:443). 
This is especially true for administered populations such as American Indians. 
The BIA is a salient category because of the historical and current relationship 
between Indians and the federal institution; and particularly because of the BIA's 
policy to encourage migration and assist the migrant in locating employment and 
housing. Cognitive categories and behavior are not in a static relationship, but 
are influenced by situational variables. It is these factors which need to be de- 
termined in future studies; what are the situational determinants in the relation- 
ship between how people think and how they behave? The findings here suggest 
that there is a complex relationship between cognition and behavior. 



36 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Literature Cited 

Ablon, J. 1964. Relocated American Indians in the San Francisco Bay Area: Social Interaction and 

Indian Identity. Human Organization, 24(4):296-304. 
Aldous, J. 1962. Morroco's Expanding Towns. Pp. 114-121 in Peasants in Cities. (W. Mangin, ed.) 

Houghton Mifflin Co., Boston vii-207. 
Berlin, B., D. Breedlove, and P. H. Raven. 1968. Covert Categories and Folk Taxonomies. American 

Anthropologist, 70:290-299. 
Berreman, G. 1966. Anemic and Emetic Analysis in Social Anthropology. American Anthropologist, 

68:346-354. 
Black, M. 1969. Eliciting Folk Taxonomy in Ojibwa. Pp. 165-189 in Cognitive Anthropology. (S. 

Tyler, ed.) Holt, Rinehart, and Winston, New York, v-521. 
Burling, R. 1964. Cognition and Componential Analysis: God's Truth or Hocus-Pocus? American 

Anthropologist, 66:20-28. 
Burton, M., and S. B. Nerlove. 1976. Balanced Designs for Triads Tests: Two Examples from 

English. Social Science Research, 5:247-267. 
, and A. K. Romney. 1975. A Multidimensional Representation of Role Terms. American 

Ethnologist, 2(3):397-407. 
D'Andrade, R. G. 1973. The All-Possible Pair Method of Cluster Analysis. Mimeo, Anthropology 

Department, University of California, San Diego. 
. 1974. Memory and the Assessment of Behavior. Measurement in the Social Sciences. (H. 

M. Blalock, ed.) Aldine, Chicago, vi-464. 
Farsoun, K. 1970. Family Structure and Society in Modern Lebanon. Pp. 257-307, in People and 

Cultures of the Middle East, Volume II. (L. Sweet, ed.) The Natural History Press, Garden 

City, New York, vii-438. 
Fiske, S. J. 1975. Navajo Cognition in the Urban Milieu: An Investigation of Social Categories and 

Use of Address Terms. Ph.D. dissertation. Stanford University, 304 pp. 

. 1977. Intertribal Perceptions: Navajo and Pan-Indianism. Ethos, 5(3):358-375. 

. 1978. Rules of Address: Navajo Women in Los Angeles. Journal of Anthropological Re- 
search, 34:72-91. 
Gans, H. 1962. Urbanism and Suburbanism as Ways of Life: A Reevaluation of Definitions. Pp. 625- 

648 in Human Behavior and Social Processes. (A. Rose, ed.) Houghton Mifflin Co., Boston, 

xv-680. 
Goodenough, W. E. 1964. Cultural Anthropology and Linguistics. Pp. 36-39 in Language in Culture 

and Society. (D. Hymes, ed.) Harper and Row, New York, v-764. 
Gulick, J. 1965. Old and New Institutions in a Lebanese Arab City. Human Organization, 1:49-52. 
Harris, M. 1968. The Rise of Anthropological Theory. New York: Crowell. 806 pp. 
Hauser, M. 1965. Application of the Ideal-type Constructs to the Metropolis in the Economically 

Less-Advanced Areas. Pp. 511-514 in The Study of Urbanization. (P. M. Hauser and L. F. 

Schnore, eds.) John Wiley and Sons, Inc., New York, viii-554. 
Hirabayashi, J., W. Willard, and L. Kemnitzer. 1972. Pan-Indianism in the Urban Setting. Pp. 77- 

87, in The Anthropology of Urban Environments. (T. Weaver and D. White, eds.) Society for 

Applied Anthropology Monograph Series No. 11, Boulder, Colorado. 134 pp. 
Howe, J., and J. Sherzer. 1975. Take and Tell: A Practical Classification from the San Bias Cuna. 

American Ethnologist, 2(3):435-460. 
Johnson, A. 1974. Ethnoecology and Planting Practices in a Swidden Agricultural System. American 

Ethnologist, 1(1):87-101. 
, and G. C. Bond. 1974. Kinship, Friendship, and Exchange in Two Communities: A Com- 
parative Analysis of Norms and Behavior. Journal of Anthropological Research, 30:55-68. 
Johnson, S. C. 1967. Hierarchical Clustering Schemes. Psychometrika, 26:173-183. 
Keesing, R., and F. Keesing. 1971. New Perspectives in Cultural Anthropology. Holt, Rinehart, and 

Winston, New York, xvi-457. 
Kruskal, J. B. 1964. Nonmetric Multidimensional Scaling: A Numerical Method. Psychometrika, 

29:115-129. 
Metzger, D., and G. Williams. 1966. Some Procedures and Results in the Study of Native Categories: 

Tzeltal "Firewood." American Anthropologist, 68:389-407. 



NAVAJO SOCIAL INTERACTIONS 37 



Nerlove, S. B., and M. Burton. 1972. A Further Examination of Cognitive Aspects of English Kin 

Terms. American Anthropologist, 74:1249-1253. 
Officer, J. E. 1971. The American Indian and Federal Policy. Pp. 8-65, in The American Indian in 

Urban Society. (J. O. Waddell and O. M. Watson, eds.) Boston: Little, Brown, vii-414. 
Perchonock, N., and O. Werner. 1969. Navajo Systems of Classification: Some Implications for 

Ethnoscience. Ethnology, 8:229-242. 
Price, J. A. 1968. The Migration and Adaptation of American Indians to Los Angeles. Human 

Organization, 27(2): 168-175. 
. 1972. Migration of Indians to Los Angeles. Pp. 428-439, in Native Americans Today: So- 
ciological Perspectives. (H. M. Bahr, A. Chadwick, and R. C. Day, eds.) Harper and Row, 

New York, v-547. 
Romney, A. K., and R. G. D'Andrade. 1964. Cognitive Aspects of English Kin Terms. American 

Anthropologist, 66(3, pt. 2): 146-170. 
Sanjek, R. 1977. Cognitive Maps of the Ethnic Domain in Urban Ghana: Reflections on Variability 

and Change. American Ethnologist, 4(4):603-622. 
Shepard, R. N. 1962. The Analysis of Proximities: Multidimensional Scaling with an Unknown 

Distance Function. I. Psychometrika, 27:125-140. 
Snyder, P. Z. 1971. The Social Environment of the Urban Indian. Pp. 206-243, in The American 

Indian in Urban Society. (J. O. Waddell and O. M. Watson, eds.) Little, Brown & Co., Boston, 

vii-414. 
U.S. Bureau of the Census. 1970. Census of Housing and Population. PHC( 1)-1 17. U.S. Government 

Printing Office, Washington, D.C. 
U.S. Department of Commerce, Bureau of the Census. 1970. Census of Population, Subject Report: 

American Indians. PC(2)-IF. U.S. Government Printing Office, Washington, D.C. 
Weibel, J. C. 1977. Native Americans in Los Angeles: A Cross-Cultural Comparison of Assistance 

Patterns in an Urban Environment. Ph.D. Dissertation, University of California, Los Angeles. 

402 pp. 
Young, J. C. 1978. Illness Categories and Action Strategies in a Tarascan Town. American Ethnol- 
ogist, 5(l):81-97. 

Accepted for publication February 5, 1980. 



Bull. Southern California Acad. Sci. 
79(1), 1980, pp. 38-41 

Range Extensions of Four Species of Crangonid Shrimps in the 
Eastern Pacific Ocean (Decapoda: Crangonidae) 

Mary K. Wicksten 

Allan Hancock Foundation, University of Southern California, Los Angeles, 
California 90007. 



Members of the family Crangonidae, often called sand shrimp or blacktailed 
shrimp, are common inhabitants of sandy, muddy, or mixed bottoms. Recent 
identification of specimens at the Allan Hancock Foundation has provided a good 
series of shrimp of this family. New records of four species have been found 
outside their recorded ranges. Sclerocrangon alata is reported for the first time 
from California. The range of Neocrangon zacae is extended south from Baja 
California to Colombia. Crangon munitellus has been found in Baja California. 
Crangon lomae is reported for the first time since the description of the species 
in 1921. 

Sclerocrangon alata Rathbun, 1902 

Sclerocrangon alata Rathbun 1902:891-892.— Rathbun 1904:134-135, fig. 72, pi. 
Ill, fig. 2. 

Previous records. — Bering Sea to Puget Sound, Washington, 11-168 m. Type 
locality: Admiralty Inlet, Puget Sound, Washington, 74 m, Albatross station 2865 
(Rathbun 1902). 

Material. — Pacific Grove, California, depth not recorded, 20 August 1937, 
Burch station 3710, 1 specimen. Friday Harbor, Washington, at surface, 27 Au- 
gust 1949, John L. Mohr, 1 specimen. 

Remarks. — There is one previous record of a specimen of the genus Sclero- 
crangon in "California." Ross and Owen (1835) recorded a specimen of Sclero- 
crangon boreas (Phipps). I have been unable to locate the original material on 
which this record was based. Holmes (1900) and Rathbun (1904) mentioned S. 
boreas from California on the basis of this one record. 

S. boreas is a circumpolar species, known from Arctic Siberia, Alaska south 
to the Strait of Georgia, the Canadian and Alaskan Arctic, eastern Canada south 
to Cape Cod, Greenland, Iceland, and northern Europe (Rathbun 1919). S. alata 
was not recognized as a species distinct from S. boreas until 1902. It seems likely 
that the shrimp taken by Ross and Owen was S. alata rather than S. boreas. 

Members of the genus Sclerocrangon are broad, heavy shrimps distinguished 
by the presence of second pereiopods, dactyls of the fourth and fifth pereiopods 
not broad and flattened, and the absence of an arthrobranch from the third max- 
illipeds (Holthuis 1955). S. alata has a carapace nearly as long as wide. The blade 
of its antennal scale does not exceed the spine. Unlike S. boreas, S. alata tends 
to be small, about 26-38 mm in total length (Rathbun 1904). 

The species of Sclerocrangon are most common in Arctic or boreal waters. S. 
alata may be a rare visitor to Monterey Bay rather than a resident species. 



CRANGONJD SHRIMPS 39 

Crangon lomae (Schmitt, 1921) 
Crago lomae Schmitt 1921:100-101, pi. 12, figs. 3 and 4. 

Previous records. — Type locality: off Point Loma, California, 929-999 m, Al- 
batross station 4334. Off Point Loma, California, 1159-1182 m. Albatross station 
4353 (Schmitt 1921). 

Material. — California: 7.3 miles, 46° true from Point Bennett, San Miguel Island 
(33°56'N, 120°33'W to 33°56'N, 120°31'W), 830-1126 m, rock dredge, 29 April 
1976, Velero IV station 24889, one male, total length 31.8 mm. 

Remarks. — C. lomae is one of the deepest species of its family in California, 
exceeded in depth range only by Pontophilus occidentalis Faxon. 

Crangon munitellus Walker, 1898 

Crangon munitellus Walker 1898:275, pi. 16, fig. 1.— Holmes 1900:176. 

C. munitella.— Rathbun 1904: 132.— Carlton and Kuris 1975:404.— Word and 

Charwat 1976:81-82. 
C. munitella.— Schmitt 1921:101-102, fig. 70. 

Previous records. — Type locality: Puget Sound, Washington (Walker 1898). — 
San Francisco Bay to Santa Catalina Island, California, 6.5-74 m (Schmitt 1921). 

Material.— BAJA CALIFORNIA, MEXICO: 3.75 miles NNW of Punta Eu- 
genia (27°54'45"N, 115°06'0"W to 27°54'20"N, 1 15°06'35"W), 37 m, 5 March 1949, 
Velero IV station 1702, 3 specimens. Between Melpomene Cove and Inner Gua- 
dalupe Island (28°52'N, 1 18°19'W), 9-28 m, 18 December 1949, Velero IV station 
1914, 4 specimens. Melpomene Cove, Guadalupe Island (28°55'23"N, 1 18°18'38"W 
to 28 o 51'0"N, 1 18°17'30"W), 92-94 m, 19 December 1949, Velero IV station 1920, 
2 specimens. 1.25 miles from Sandstone Point, Guadalupe Island (28°54'08"N, 
118°15'36"W to 28°53'57"N, 1 18°15'41"W), 46-55 m, 20 December 1949, Velero 
IV station 1924, 2 specimens. 2 miles, 142° true to Thurloe Head (27°35'45"N, 
114°49'15"W), 37 m, 7 December 1967, Velero IV station 11842, 1 ovigerous 
female. CALIFORNIA, U.S.A.: 62 specimens from 29 other stations: Santa 
Rosa, Santa Cruz, Anacapa, Santa Catalina, and San Clemente Islands; Port 
Hueneme, Redondo Beach, Point Vicente, White's Point to Portuguese Bend, 
Newport Inlet, and Corona del Mar, 0-185 m, 1926-1962. 

Remarks. — The records suggest that C. munitellus prefers shallow sublittoral 
bottoms of clean sand or sand mixed with rock and shell. Kuris and Carlton 
(1977) related the squat body forms of C. handi and Lissocrangon stylirostris to 
habitat specialization, in which their shape allows rapid escape response over 
short distances. C. munitellus, a short, broad shrimp, may have adapted in a 
similar fashion to its environment. 

Neocrangon zacae (Chace, 1937) 

Crago zacae Chace, 1937:136-138, fig. 9. 
Crangon zacae. — Word and Charwat 1976:93-94. 
Neocrangon zacae. — Kuris and Carlton 1977:554. 

Previous records. — Type locality: east of Cedros Island, Mexico (28°13'N, 
115°07'W), 81 m, mud bottom, 27 March 1936, Zaca station 125 (Chace, 1937). 
Monterey Bay to Dana Point, California (Chace 1937; Word and Charwat 1976). 



40 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Material— North of Gorgona Island, Colombia (3°01'25"N, 78°10'W), 18-37 m, 
mud and rock bottom, 24 February 1938, Velero III station 851-38, 1 damaged 
specimen. Sulphur Bay, Clarion Island, Mexico (18°20'45"N, 114°44'15"W), 9.2 
m, among coralline algae, 16 March 1939, Velero III station 915-39, 1 male and 
1 ovigerous female. 

Remarks. — N. zacae is distinguished from the closely related N. communis 
(Rathbun) by the lack of carinae on its fifth abdominal segment, the dactyl of the 
first pereiopod closing more longitudinally than horizontally, and the smaller size 
of the adults in the former species. These differences may be difficult to detect 
in damaged or poorly preserved specimens. Chace (1937) suggested that TV. zacae 
might be a southern subspecies of N. communis. 

In the collections of the Allan Hancock Foundation, there are about 700 spec- 
imens identified as N. zacae from 105 stations in southern California and Baja 
California, Mexico. These were taken in 1938-1977 at depths from the shore to 
572 m. At 75 of the 105 stations, the specimens were collected at depths of 185 
m or less. N. communis was taken at 34 stations at 21-230 m. One hundred 
eighty-eight specimens were collected in 1917-1976. At 23 stations, the species 
was taken at 92-277 m. The records suggest that N. communis prefers somewhat 
greater depths than N. zacae, although both occur over a wide range of depths 
in southern California. 

N. zacae is the only member of the family Crangonidae recorded so far from 
the continental shelf of the Panamic zoogeographic province, the region of coast 
from northern Peru to southern Baja California. Pontophilus occidentalis occurs 
in bathyal depths in the region. 

Acknowledgments 

I thank Fenner A. Chace, Jr. of the National Museum of Natural History for 
examining the specimens of Neocrangon zacae. The specimen of Crangon lomae 
was collected during the Southern California Baseline Studies and Analysis, spon- 
sored by the U.S. Bureau of Land Management contracts 08550-CT5-52 and 
AA550-CT6-40. 

Literature Cited 

Carlton, J. T., and A. M. Kuris. 1975. Keys to decapod Crustacea. In Smith, R. I. and J. T. Carlton, 
eds. Light's manual: intertidal invertebrates of the central California coast. Berkeley: Univ. 
Calif. Press, pp. 385-412. 

Chace, F. A. Jr. 1937. The Templeton Crocker Expedition. VII. Caridean decapod Crustacea from 
the Gulf of California and the west coast of Lower California. Zoologica, 22(2): 109-138. 

Holmes, S. J. 1900. Synopsis of California stalk-eyed Crustacea. Occ. Papers Calif. Acad. Sci., 7:200 
pp. 

Holthuis, L. B. 1955. The recent genera of the caridean and stenopodidean shrimps (class Crustacea, 
supersection Natantia) with keys for their determination. Zool. Verhand. Leiden No. 26, 157 
pp. 

Kuris, A. M., and J. T. Carlton. 1977. Description of a new species, Crangon handi, and new genus, 
Lissocrangon, of crangonid shrimps (Crustacea: Caridea) from the California coast, with notes 
on adaptation in body shape and coloration. Biol. Bull., 153:540-559. 

Rathbun, M. J. 1902. Descriptions of new decapod crustaceans from the west coast of North Amer- 
ica. Proc. U.S. Nat. Mus., 24:885-905. 

. 1904. Decapod crustaceans of the northwest coast of North America. Harriman Alaska 

Exped., 10:1-219. 



CRANGONID SHRIMPS 41 

. 1919. Report of the Canadian Arctic Expedition 1913-18. Vol. VII: Crustacea. Part A: 

Decapod crustaceans. Southern Party 1913-16. 14 pp. 
Ross, J. C, and R. Owen. 1835. Marine invertebrate animals. In Appendix to a narrative of a second 

voyage in search of a northwest passage, and a residence in the arctic regions during the years 

1829, 1830, 1831, 1832, 1833. London: A. W. Webster, pp. 81-100. 
Schmitt, W. L. 1921. The marine decapod Crustacea of California. Univ. Calif. Publ. Zool., 23:470 

pp. 
Walker, A. D. 1898. Crustacea collected by W. A. Herdman in Puget Sound, Pacific coast of North 

America, September 1897. Proc. and Trans. Liverpool Biol. Soc, 12:268-287. 
Word, J. Q., and D. Charwat. 1976. Invertebrates of southern California coastal waters. II. Natantia. 

El Segundo: So. Calif. Coastal Water Res. Project. 238 pp. 

Accepted for publication April 11, 1980. 



Bull. Southern California Acad. Sci. 
79(1), 1980, pp. 42-44 



INDEX TO VOLUME 78 



Acanthogobius flavimanus in Southern California, 56-61 

Akersten, William A., Richard L. Reynolds and Antonia E. Tejada-Flores: New 
mammalian records from the late pleistocene of Rancho La Brea, 141-143 
Anderson, Daniel W., see Henny, Charles J. 

Annelida: Systematics and ecology of benthic Phyllodocidae (Polychaeta), 1-19 
Arachnida: Pseudoscorpions in Florida slash pine, 32-39 
Ascidians of Southern California, 122-135 

Barbourisia rufa, otoliths and presence of whalefish in California, 61-67 

Biology of Onthophagus medorensis, 67-69 

Blake, James A.: A redescription of Pettiboneia sanmatiensis Oresanz (Poly- 
chaeta: Dorvillidae) Dorvilleidae, 136-140 

Brach, Vincent: Species diversity and distributional relationships of pseudoscor- 
pions from slash pine (Pinus elliotti Eng.) in Florida (Arachnida: Pseudo- 
scorpionida), 32-39 

: The larva of Onthophagus medorensis Brown with notes on its biology 

(Coleoptera: Scarabaeidae), 67-69 

California, Demersal zooplankton off Santa Catalina Island, 199-206; Develop- 
ment of two sponges at Santa Catalina Island, 183-191; Freshwater fishes near 
Salton Sea, 46-55; Littoral and sublittoral ascidians in Southern California, 
122-135; New mammalian records from the late pleistocene of Rancho La 
Brea, 141-143; Range extension and notes on habitat of isopod Munna halei 
Menzies, 196-199; Post-larval polychaetes in Tomales Bay, 144-147; Two 
asiatic gobiid fishes in Southern California, 56-61; Water and air tempera- 
tures around Cape Mendocino, 40-45 

Clathrina coriacea and C. blanca, gametogenesis and embryonic development, 
183-191 

Coleoptera: Larva of Onthophagus medorensis, 67-69 

Colombia River: Systematics and ecology of benthic Phyllodocidae, 1-19 

Costa Rica: Eleutherodactylus rayo, n. sp., 107-115 

Crustacea, New species and records of polychaetes from Gulf of California, 1 16- 
121; Range and notes on isopod Munna halei Menzies, 196-199 

DeWeese, James E., see Savage, Jay M. 
Dorvilleidae, Revised key to genera, 136-140 

Eleutherodactylus rayo, n. sp. from Costa Rica, 107-115 
Eumida uschakovi, n. sp. from Gulf of California, 116-121 

Fay, Rimmon C. and James A. Vallee: A survey of the littoral and sublittoral 
ascidians of Southern California, including the Channel Islands, 122-135 

Fishes, Freshwater fishes, distribution near Salton Sea, 46-55; Otoliths in whale- 
fishes, 61-67; Two asiatic gobiids in Southern California, 56-61; Velvet 
whalefish in California, 61-67 

Fitch, John E: The velvet whalefish, Barbourisia rufa, added to California's 
marine fauna, with notes on otoliths of whalefishes and possible related gen- 
era, 61-67 

Frog, Eleutherodactylus rayo, n. sp., 107-115 

Florida: Pseudoscorpions in slash pine, 32-39 



INDEX TO VOLUME 78 43 

Gellura, Joseph A., see Samaras, William F. 

Haaker, Peter L.: Two asiatic gobiid fishes, Tridentiger trigonocephalus and 
Acanthogobius flavimanus, in Southern California, 56-61 

Hammer, Richard M. and Richard C. Zimmerman: Species of demersal zooplank- 
ton inhabiting a kelp forest ecosystem off Santa Catalina Island, California, 
199-206 

Hannes, Gerald: Summer mean diurnal water and air temperature distributions 
in the Cape Mendocino area, California, 40-45 

Harty, Rebekah: Range extension and notes on the habitat of the isopod Munna 
ha lei Menzies, 196-199 

Henny, Charles J. and Daniel W. Anderson: Osprey distribution, abundance, and 
status in Western North America: III. The Baja California and Gulf of Cal- 
ifornia population, 89-106 

Hyaenidae, calcanea, 147-150 

Index: Volumes 74 through 78, 70-87 

Isopod, Habitat of Munna halei Menzies, 196-199 

Johnson, Marion Fischel: Gametogenesis and embryonic development in the cal- 
careous sponges Clathrina coriacea and C. blanca from Santa Catalina Is- 
land, California, 183-191 

Jones, Howard R., see Kravitz, Michael J. 

Kelp forest, demersal zooplankton inhabitants, 199-206 

Kravitz, Michael J. and Howard R. Jones: Systematics and ecology of benthic 

Phyllodocidae (Annelida: Polychaeta) off the Colombia River, U.S.A., 1-19 
Kudenov, Jerry D.: New species and records of polychaetous annelids from the 

Tetraclita (Cirripedia: Crustacea) zone of the Northern Gulf of California, 

Mexico, 1 16-121 
: Post-larval polychaetes in sandy beaches of Tomales Bay, California, 

144-147 

Mammals, Gray whale birth, 192-196; Hyaenidae calcanea, 147-150: New rec- 
ords of late pleistocene forms from Rancho La Brea, 141-143 

Marine terraces, Quaternary wave formation, 20-31 

Mexico, New species and records of polychaetous annelids in Gulf of California, 
116-121; Osprey in Western North America, III., 89-106; Vegetation of Ba- 
hia de San Quintin, 163-182 

Mills, James E., see Mills, James G. 

Mills, James G. and James E. Mills: Observations of a gray whale birth, 192-196 

Munna halei Menzies, Range extension and notes on habitat, 196-199 

Neanthes cortezi, n. sp. from Gulf of California, 116-121 

Neuenschwander, Leon F., Ted H. Thorsted, Jr., and Richard J. Vogl: The salt 

marsh and transitional vegetation of Bahia de San Quintin, 163-182 
New taxa: Eleutherodactylus rayo, 107-115; Eumida uschakovi, 116-121; 

Neanthes cortezi, 116-121 
Niche separation within a population of freshwater fishes in an irrigation drain 

near the Salton Sea, California, 46-55 



44 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Onthophagus medorensis Brown larva, Biology of, 67-69 

Osprey distribution, abundance, and status in Western North America: III. The 

Baja California and Gulf of California population, 89-106 
Otoliths of whalefishes, 61-67 

Pettiboneia sanmatiensis, Redescription, 136-140 

Pinus elliotti Eng., Pseudoscorpion inhabitants, 32-39 

Plants: Salt marsh and transitional vegetation of Bahia de San Quintin, 163-182 

Pleistocene, Academy excavations at Rancho La Brea, 151-162; New records of 
mammals from Rancho La Brea, 141-143 

Polychaeta, Post-larval forms in Tomales Bay, California, 144-147; Redescription 
of Pettiboneia sanmatiensis and revised key to genera of Dorvilleidae, 136— 
140; Systematics and ecology of benthic Phyllodocidae, 1-19 

Pseudoscorpions, in Florida slash pine, 32-39 

Quaternary, Wave formed marine terraces on Santa Catalina Island, 20-31 

Rancho La Brea, New mammalian records, 141-143; Seventieth anniversary of 

Academy excavation, 151-162 
Reynolds, Richard L., see Akersten, William A. 

Samaras, William F., and Joseph A. Gellura: Recognition of quaternary wave 

formed marine terraces on Santa Catalina Island, 20-31 
Savage, Jay M. and James E. DeWeese: A new species of leptodactylid frog, 

genus Eleutherodactylus , from the Cordillera de Talamanca, Costa Rica, 

107-115 
Scarabaeidae, Larva of Onthophagus medorensis, 67-69 
Schoenherr, Allan A.: Niche separation within a population of freshwater fishes 

in an irrigation drain near the Salton Sea, California, 46-55 
Sibley, Gretchen: Seventieth anniversary of Academy excavations at Rancho La 

Brea, 151-162 
Sponges, Gametogenesis and embryonic development of Clathrina coriacea and 

C. blanca, 183-191 
Stains, Howard J.: The calcanea of members of the Hyaenidae, 147-150 
Systematics and ecology of benthic Phyllodocidae, 1-19 

Tejada-Flores, Antonia E., see Akersten, William A. 
Thorsted, Ted H. Jr., see Neuenschwander, Leon F. 
Tridentiger trigonocephalus in Southern California, 56-61 

Vallee, James A., see Fay, Rimmon C. 

Vogl, Richard J., see Neuenschwander, Leon F. 

Water and air temperature around Cape Mendocino, California, 40-45 
Whalefish, Velvet in California, 61-67 
Whales, Gray whale birth, 192-196 

Zimmerman, Richard C, see Hammer, Richard M. 
Zooplankton, Demersal in kelp forest, 199-206 



INSTRUCTIONS FOR AUTHORS 

;he BULLETIN is published three times each year (April, August, and November) and includes articles in English 
I any field of science with an emphasis on the southern California area. Manuscripts submitted for publication 
:iould contain results of original research, embrace sound principles of scientific investigation, and present data 
• a clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide 
ir contributors. Consult also recent issues of the BULLETIN. Authors should strive for directness and lucidity. 
:hieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous 
ference of pronouns, and logically placed modifiers. 

MANUSCRIPT PREPARATION 

It is strongly recommended that, before submitting a paper, the author ask qualified persons to review it. The 
jthor is requested to submit at least two additional copies with the original, on SVi x 11 opaque, nonerasable 
iper, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. 
lootnotes should be avoided. Manuscripts which do not conform to the style of the BULLETIN will be returned to 
ie author. 

An abstract summarizing in concise terms the methods, findings, and implications discussed in the paper must 
.•company a feature article. Abstract should not exceed 100 words. 

A feature article comprises approximately five to thirty typewritten pages. Papers should usually be divided into 
ie following sections: abstract, introduction, methods, results, discussion and conclusions, acknowledgments, and 
erature cited. Avoid using more than two levels of subheadings. 

A research note is usually one to six typewritten pages and rarely utilizes subheadings. Consult a recent issue 
? the BULLETIN for the format of notes. Abstracts are not used for notes. 

Abbreviations: Use of abbreviations and symbols can be determined by inspection of a recent issue of the 
[ULLETIN. Omit periods after standard abbreviations: 1.2 mm, 2 km, 30 cm, but Figs. 1-2. Use numerals before 
hits of measurements: 5 ml, but nine spines (10 or numbers above, such as 13 spines). The metric system of 
;cights and measurements should be used wherever possible. 

Taxonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International 
ode of Botanical Nomenclature (Lawjouw et al.. 1956), the International Code of Nomenclature of Bacteria and 
iruses (Buchanan et al.. 1958), and the International Code of Zoological Nomenclature (Stoll et al.. 1961). 
pecial attention should be given to the description of new taxa, designation of holotype, etc. Reference to new 
xa in titles and abstract should be avoided. 

i The literature cited: Entries for books and articles should take these forms. 
; McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. 

; Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54: 452-458. 
; Brattstrom. B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), 
\ Univ. California Press, xii + 635 pp. 

| Tables and figures (line drawings, graphs, or black and white photographs) should not repeat data contained in 
ie text. The author must provide numbers and short legends for tables and figures and place reference to each 
'them in the text. Legends should be typed on a separate sheet of paper and placed at the end of the manuscript. 
lustrations and lettering thereon should be of sufficient size and clarity to permit reduction to standard page size; 
dinarily they should be no more than twice the size of intended reduction and should not exceed 8'/6 by 1 1 inches 
size. Photographs must be printed on glossy paper. Submit one photoduplicated copy of each illustration. All 
titrations accompanying Research Notes will be reduced to one column width. All half-tone illustrations will have 
^ht screen (grey) backgrounds. Special handling such as dropout half-tones, special screens, etc., must be requested 
1 and will be charged to authors. 

A cover illustration pertaining to an article in the issue or one of general scientific interest will be printed on the 
>ver of each issue. Such illustrations along with a brief caption should be sent to the Editor for review. 

PROCEDURE 

All manuscripts should be submitted to the Editor, Robert J. Lavenberg, Los Angeles County Natural History 
luseum, 900 Exposition Blvd., Los Angeles, California 90007. Evaluation of a paper submitted to the BULLETIN 
:gins with a critical reading by the Editor; several referees also check the paper for scientific content, originality, 
id clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent 
> the author at which time he or she may be requested to rework portions of the paper considering these rec- 
nmendations. The paper then is resubmitted and may be re-evaluated before final acceptance. 

Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He or she 
lould promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations, legends. 
id bibliographical references. He or she marks corrections on the galley (copy editing and proof procedures in 
tyle Manual) and promptly returns both galley and manuscript to the Editor. Manuscripts and original illustrations 
ill not be returned unless requested at this time. All changes in galley proof attributable to the author (misspellings, 
iconsistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with 
ie printer, not the Editor. 



CONTENTS 

Discovery of the Male of the Katydid Idiostatus viridis Rentz, with Descrip- 
tions and Biological Notes (Orthoptera: Tettigoniidae: Decticinae). 
By Charles L. Hogue 1 

Postfire Seedling Reproduction of Adenostoma fasciculatum H. and A. 

By George F. Howe and Linn E. Carothers 5 

A New Frog of the Genus Eleutherodactylus (Leptodactylidae) from the 

Monteverde Forest Preserve, Costa Rica. By Jay M. Savage 13 

Navajo Social Interactions in an Urban Environment: An Investigation of 

Cognition and Behavior. By Shirley J. Fiske and J. C. Weibel 19 

Range Extensions of Four Species of Crangonid Shrimps in the Eastern 

Pacific Ocean (Decapoda: Crangonidae). By Mary K. Wicksten 38 

Index to Volume 78 42 



COVER: Male of the Katydid Idiostatus viridis Rentz 



$i> 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




ULLETIN 



/olume 79 



APR 6 1981 

New YORK 
B °T ANICAL QARDm 



Number 2 




8CAS-A79(2) 45-88 (1980) 



AUGUST 1980 



Southern California Academy of Sciences 

Founded 6 November 1891, incorporated 17 May 1907 



OFFICERS 

Takashi Hoshizaki, President 
Fred G. Hochberg, Vice President 
Richard E. Pieper, Secretary 
Joseph E. Haring, Treasurer 
Robert J. Lavenberg, Editor 
Gretchen Sibley, Assistant Editor 



1978-1980 

Kristin H. Berry 

Robert A. Georges 

Joseph E. Haring 

Jerry D. Kudenov 

Donald J. Reish 



BOARD OF DIRECTORS 

1979-1981 

John Baird 

Jules Crane 

Fred G. Hochberg 

Richard E. Pieper 

Gloria Takahashi 






1980-1982 

Takashi Hoshizaki 
Alan J. Mearns 

Steven N. Murray 
Camm C. Swift 

Robert G. Zahary 



Membership is open to scholars in the fields of natural and social sciences, and to any persor 
interested in the advancement of science. Dues for membership, changes of address, and requests fo 
missing numbers lost in shipment should be addressed to: Southern California Academy of Sciences 
the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 90007 

Annual Members $ 12.0 

Life Members 150.0 



Fellows: Elected by the Board of Directors for meritorious services. 






The Bulletin is published three times each year by the Academy. Manuscripts for publication should 
be sent to the appropriate editor as explained in "Instructions for Authors" on the inside back cover 
of each number. All other communications should be addressed to the Southern California Academy 
of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, Los 
Angeles, California 90007. 



Date of this issue 20 March 



Bull. Southern California Acad. Sci. 
79(2), 1980, pp. 45-54 

A Synopsis of the Larvae of Costa 
Rican Frogs and Toads 

Jay M. Savage 

Abstract. — A synopsis of the larvae of Costa Rican frogs and toads by Jay M. 
Savage, Bull. Southern California Acad. Sci., 79(2):45-54, 1980. Of 87 species 
of Costa Rican frogs and toads with free living larvae, 61 tadpoles are known in 
sufficient detail to be distinguished. Of the remaining 16 forms, all but two will 
probably share distinctive combinations of generic features that will allow them 
to be recognized as previously undescribed tadpoles of an appropriate genus 
when discovered. A synoptic key to Costa Rican tadpoles that will distinguish 
larvae from the time the operculum closes through the time of front limb eruption 
is presented. It is supplemented by a guide to published accounts and illustrations 
of all described tadpoles of Costa Rican forms. 

Department of Biological Sciences and Allan Hancock Foundation, University 
of Southern California, Los Angeles, California 90007. 



The characteristics of the aquatic free-living larvae (tadpoles) of frogs and toads 
have long been known to be distinctive for a spectrum of taxonomic categories 
ranging from suborders to species (Starrett 1973). Generally the larvae show a 
series of special adaptations for a particular kind of ecologic role superimposed 
upon a basic morphology that is consistent for a number of related frog families. 
Because of this mixture of plastic and conservative features, respectively, in well- 
studied areas (e.g., the United States, Altig 1970; or Southern Africa, Wager 
1965) or taxonomic groups (e.g., the mecoamerican Hylidae, Duellman 1970) most 
tadpoles may be distinguished at the specific level. As pointed out by Altig and 
Brandon (1971) an essential precursor to any detailed study of the ecology and 
behavior of the immature forms in any area must be the ability to identify and 
associate them with adult species populations. In reality, attempts to fully un- 
derstand the evolution and ecology of the species require this same ability. 

As part of my long-term studies of the herpetofauna of Costa Rica (Savage 
1976), I have accumulated specimens and data for a synopsis of the larvae of the 
Anura of the republic. The present paper is a summary of the available knowledge 
presented in a simple and convenient format. 

Costa Rica has a known amphibian fauna of 150 species, 3 caecilians, 27 sal- 
amanders and 120 frogs and toads. Unlike the situation in areas east of central 
Panama (caecilians) and north of the Isthmus of Tehuantepec (for salamanders) 
no free-living larvae of caecilians or salamanders or known or are likely to be 
discovered in Costa Rica. All species of the genera of caecilians represented in 
Costa Rica retain the developing eggs in the oviduct and give birth to living young 
(M. H. Wake 1977). All species of the genera of salamanders known from Central 
and South America have land-laid encapsulated eggs that undergo direct devel- 
opment (D. B. Wake and J. F. Lynch 1976). 

Of the 120 known species of frogs and toads that occur in Costa Rica, 33 belong 



46 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

to the leptodactylid genus Eleutherodactylus . Insofar as known (J. D. Lynch 
1976) all members of this genus lay terrestrial encapsulated eggs that undergo 
direct development into small frogs. Free-living tadpoles of 87 species of anurans 
in the families Rhinophrynidae (1), Microhylidae (3), Leptodactylidae (6), Bufon- 
idae (14), Hylidae (39), Dendrobatidae (7), Centrolenidae (13) and Ranidae (4) 
occur in Costa Rica. Available data make it possible to include 62 of these species 
in the synoptic key presented below. 

The eggs of most of these species are deposited in aquatic situations. Excep- 
tions include: 1) two species of Leptodactylus, fragilis and poecilochilus, that 
lay their eggs in a foam-filled burrow (Heyer 1969); 2) the tree-frog Hyla ebrac- 
cata that lays its eggs on the leaves of broad-leafed herbaceous plants emergent 
from ponds (Duellman 1967); 3) all members of the family Centrolenidae (Starrett 
1960), the hylid genera Agalychnis and Phyllomedusa and Hyla lancasteri (Duell- 
man 1970) lay eggs on vegetation, usually leaf-surfaces, a considerable distance 
above a stream or other body of water; and 4) all members of the family Den- 
drobatidae lay their eggs in moist terrestrial sites and transport the hatched tad- 
poles on the adult's back to water (Savage 1968). 

Larval Structure and Terminology 

The characteristics of anuran larvae that may be used in taxonomic description 
have been detailed by Orton (1952), Altig (1970), Duellman (1970) and Starrett 
(1973). Distinctions among preserved tadpoles generally can only be made in 
developmental stages 25 (operculum closure) through 40 (immediately before 
front leg eruption) using the Gosner (1960) system. The synopsis that follows is 
based on tadpoles of these stages and the characterizations provided by the cited 
authors. Points of minor departure or preference include: 

1. denticle is used instead of tooth since larval Anura have no true teeth but 
only a series of non-homologous keratinized structures; 

2. the jaws, beaks, labia and denticle rows are referred to as upper and lower 
because of their association with the upper and lower jaw structures common to 
all vertebrates; Altig (1970) uses anterior and posterior, respectively, since most 
tadpoles have a ventrally located mouth; 

3. a complete oral disk refers to an unindented disk; 

4. description of mouth position and size and body form follows the terminology 
of Duellman (1970). 

Identification of Costa Rican Tadpoles 

The tadpoles of 61 Costa Rican species have been described and/or illustrated 
in considerable detail. Reasonably accurate predictions as to the basic features 
of several other species can also be made as follows: 

1. Microhylidae; the features of the only other four species of the genus Gas- 
trophryne are known; Glossostoma tadpoles will probably agree in general with 
Gastrophryne and Hypopachus when discovered; 

2. Bufonidae; tadpoles of all known members of the genus Atelopus have the 
peculiar enlarged ventral disk found in varius (Starrett 1967; Duellman and Lynch 
1969); all known Central American Bufo agree in having 2/3 denticle rows, an 
indented oral disk and oral papillae incomplete across both upper and lower labia; 
it seems likely the unknowns from Costa Rica will share these features; 



LARVAE OF COSTA RICAN FROGS AND TOADS 



47 



3. Centrolenidae; it is anticipated that other Costa Rican tadpoles of the genus 
Centrolenella will agree with known species in having 2/3 denticle rows, the oral 
disk complete, a median anus, the spiracle posterior in position and the denticle 
row just above the mouth restricted to two short segments. 

The guide to identification of the tadpoles is organized into two parts. First is 
a synoptic key based upon available specimens and published data. It is designed 
to lead to an identification when well-preserved material conspecific with de- 
scribed tadpoles is available. In most cases it is anticipated that material of as 
yet undescribed tadpoles of known species will be correctly identified at the 
generic level. Tadpoles of the genera Agalychnis, Atelopus, Bufo, Centrolenella 
and Phyllobates fall into this category. 

The second portion of the guide consists of an index to published illustrations 
of the tadpoles. In cases where species are very similar or where material may 
be of a form whose tadpole has not been described reference to the illustration 
should be the final authority. 

The following species of Costa Rican frogs do not have their tadpoles described 
or illustrated and remain cryptic. In cases where material does not conform to 
the synopsis and/or cited illustrations the possibility that one has discovered a 
previously missing immature stage must be given serious consideration. 



Agalychnis calcarifer 
Atelopus chiriquiensis 
Atelopus senex 
Bufo coccifer 
Bufo coniferus 
Bufo fastidiosus 
Bufo haematiticus 
Bufo luetkenii 
Bufo melanochloris 
Centrolenella albomaculata 
Centrolenella chirripoi 
Centrolenella colymbiphyllum 
Centrolenella euknemos 



Centrolenella ilex 
Centrolenella pulverata 
Centrolenella talamancae 
Centrolenella valerioi 
Centrolenella vireovittata 
Crepidophryne epioticus 
Dendrobates granuliferus 
Glossostoma aterrimum 
Gastrophryne pictiventris 
Hyla lythrodes 
Hyla miliaria 
Hyla xanthosticta 
Phyllobates lugubris 



A SYNOPSIS OF COSTA RICAN TADPOLES 

I. NO DENTICLES 

A. NO BEAKS 



Two Ventral Spiracles 
barbels present 
Rhinophrynus 



One Ventral Spiracle 
no barbels 

i) oral flaps without scallops or 
papillae; spiracle opens im- 
mediately ventral to anus 
Gastrophryne 
ii) oral flaps with scallops or 
papillae; spiracle opens near 
anus 

Hypopachus variolosus 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



B. BEAKS PRESENT 

and 
One Sinistral Spiracle 



Tail terminating in a thin filament 

a) one row of papillae under 
mouth; median anus 

Hyla ebraccata 

b) no papillae; dextral anus 
Hyla microcephala — no postor- 

bital dark 
stripe 
Hyla phlebodes — a distinct 
postorbital 
dark stripe 



2. Tail not forming a thin terminal 
filament 

a) no oral papillae 

Colostethus nubicola 

b) oral papillae complete around 
mouth 

Hyla picadoi 



II. DENTICLES AND BEAKS PRESENT 

A. AN ENLARGED VENTRAL DISK 

Atelopus varius 

B. NO ENLARGED VENTRAL DISK 



Denticle Rows: 0/1 1/1 1/2 



2/2 



2/4 



1. 



Anus dextral; denticle rows 
complete; labial papillae 
rows complete 

Hyla zeteki 
Anus median; denticle 
rows very short; no labial 
papillae above mouth 

Dendrobates pumilio 



1. Oral disk bordered 1. No pre orbital 



by large papillae 
Anotheca spi- 
nosa 
2. Oral disk bordered 

by minute papillae 
Hyla rufioculis 
Hyla urano- 
chroa 

2/3 
1. Oral disk indented laterally 



stripe 

Hyla rosenbergi 
A dark preorbital 
stripe 

Hyla rufitela 



a) oral papillae incomplete on upper 
and lower labia 

i) body black or brown above and 
below 
Bufo marinus — 2 rows of pa- 
pillae 

Bufo periglenes — 1 row of 
papillae for most of disk 
ii) body dark above, light below 
Bufo holdridgei — 6-7 large 
papillae 

Bufo valliceps — numerous 
small papilla 



b) oral papillae complete across lower 
labium 

i) 1 row of papillae below mouth 
uniform body and tail 

Physalaemus pustulosus 
tail spotted 
Rana "pipiens" 
ii) 2 rows of papillae below mouth 
uniform dark body and tail 

Phyllobates vittatus 
body black, tail spotted 
Colostethus talamancae 



LARVAE OF COSTA RICAN FROGS AND TOADS 



49 



2. Oral disk complete 
a) median anus 



i) spiracle about halfway between 
eye and posterior margin of 
body; denticle row just above 
mouth complete or barely in- 
terrupted medially 
x) tail with dark spots and 
blotches a distinct light 
spot posterior to oral disk; 
mouth anteroventral 
Leptodactylus fragilis — 

body uniform 
Leptodactylus poecilo- 
chilus — body with 
dark flecks 
no distinct light spot 
posterior to oral disk: 
mouth terminal 
Leptodactylus penta- 
dactylus 
xx) tail without dark spots and 
blotches, uniformly dark or 
with a few light spots 
Leptodactylus bolivi- 
anus — all rows of 
denticles complete; 
papillae extending well 
onto upper lip 
Leptodactylus melano- 
notus — denticle row 
above mouth usually 
with a median gap; 
papillae essentially 
restricted to sides of 
oral disk 
Dendrobates auratus — 
denticle row above 
mouth complete; oral 
papillae incomplete 
across upper labium 



ii) spiracle much closer to poste- 
rior margin of body than to eye; 
denticle row just above mouth 
restricted to two short segments 
near corners of mouth 
y) lowermost denticle row 
much shorter than other 
two lower rows 

Centrolenella proso- 
blepon 
yy) lowermost denticle row 
equal to or only slightly 
shorter than other two 
lower rows 

Centrolenella granu- 
losa — lower beak with 
large blunt serrations 
Centrolenella spinosa — 
lower beak with large 
sharp serrations 
Centrolenella fleisch- 
manni — lower beak 
with small fine 
serrations 
Centrolenella vale- 
rioi — lower beak with 
small fine serrations 



b) dextral anus 
i) oral papillae complete across upper labium 



z) no more than 3 rows of 
papillae below mouth 



zz) 4-7 rows of papillae below 
mouth 



50 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



*) body depressed, wider 
than deep; 2-3 rows of 
papillae above mouth; 
body and tail with 
dark spots 
Hyla debilis — mouth 
large, as wide as 
body 
Hyla lancasteri — 
mouth small, 
about % width of 
body 

*) body ovoid, depth 
equal to width; 1 row 
of papillae above 
mouth; body uniform 
tan, tail tan with flecks 
and dashes of darker 
pigment that tends to 
form cross-bars on 
dorsal surface of tail 
musculature 
Smilisca sordida 

ii) oral papillae incomplete across upper labium 
x) oral papillae incomplete across lower 
labium; short lowermost denticle row 
mounted on a vertically moveable 
extension 
Hyla boulengeri 
xx) oral papillae complete across lower labium 



body and tail spotted 
with dark 
Hyla tica — tail fin 
not extending 
onto body 
Hyla rivularis — tail 
fin extending onto 
body 
body black with golden 
lichenous markings, 
tail tan with clear fins 
Hyla pictipes 



y) spiracle ventrolateral 

*) tail fin extending onto 
body 
Phyllomedusa lemur 
**) tail fin not extending 
onto body 

body slightly deeper 
than wide 
Agalychnis sal- 

tator 
Agalychnis calli- 

dryas 
Agalychnis spur- 
relli 
body deep, at least 
15% deeper than 
wide 



yy) spiracle lateral 

*) body ovoid, depth and 
width approximately 
equal 

tail fin not extending 
onto body; mouth 
ventral 
Hyla pseudo- 
puma — mouth 
small 
Hyla angusti- 
lineata — mouth 
small 
Smilisca si la — 
mouth medium 
tail fin extending 
onto body; mouth 



LARVAE OF COSTA RICAN FROGS AND TOADS 



51 



Agalychnis annae 



anteroventral 
body dark brown 
or black 
Smilisca 
b audi nii — 
body not 
tuberculate 
Hyla fimbri- 
membra — 
body tuber- 
culate 
body pale brown 
to tan 
Hyla loquax 
Smilisca phae- 

ota 
Smilisca puma 
**) body deep, much deep- 
er than wide; tail fin 
extending onto body; 
mouth anteroventral, 
small 
Hyla elaeochroa 
Hyla staufferi 



Denticle Rows: 2/5 
oral disk complete; 
dextral anus 
Hyla legleri 



3/4-6 
oral disk complete; 
median anus 
Phrynohyas venulosa 



3/5-4 
oral disk indented; 
dextral anus 
Rana palmipes 
Rana vibicaria 



Denticle Rows: 6/4 

oral disk complete; dextral anus 
Rana warschewitschii 



6/9 

oral disk complete; dextral anus 
Hxla colxmba 



An Index to Illustrations 

Agalychnis 

annae Duellman 1963; 1970 
calcarifer (unknown) 
callidryas as helenae Starrett 1960 
callidryas Duellman 1970 
saltator Duellman 1970 
spurrelli Duellman 1970 
Anotheca 

spinosa Duellman 1970 
spinosa as corona ta Taylor 1954; 
Robinson 1961 



of Costa Rican Tadpoles 

Atelopus 

chiriquiensis (unknown) 

senex (unknown) 

varius Starrett 1967 
Bufo 

coccifer (unknown) 

coniferus (unknown) 

fastidiosus (unknown) 

haematiticus (unknown) 

holdridgei Novak and Robinson 1975 

luetkenii (unknown) 



52 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



marinus Breder, 1946; Savage 1960 

melanochloris (unknown) 

periglenes Savage 1967 

valliceps Limbaugh and Volpe 1957 
Crepidophryne 

epioticus (unknown) 
Centrolenella 

albomoculata (unknown) 

chirripoi (unknown) 

colymbiphyllum (unknown) 

euknemos (unknown) 

fleischmanni Starrett 1960 

granulosa Starrett 1960 

ilex (unknown) 

prosoblepon Starrett 1960 

pulverata (unknown) 

spinosa Starrett 1960 

talamancae (unknown) 

valerioi as reticulata Starrett 1960 

vireovittata (unknown) 
Colostethus 

nubicola Dunn 1924; Savage 1968 

talamancae Savage 1968 
Dendrobates 

auratus Breder 1946; Savage 1968; 
Silverstone 1975 

granuliferus (unknown) 

pumilio Starrett 1960; Savage 1968; 
Silverstone 1975 
Gastrophryne 

pictiventris (unknown) 
Glossostoma 

aterrimum (unknown) 
Hypopachus 

variolosus as caprimimusl Taylor 
1942 
Hyla 

angustilineata Duellman 1970 

boulengeri Duellman 1970 

colymba as albomarginata Dunn 
1924 

colymba Duellman 1970 

debilis Duellman 1970 

ebraccata Duellman 1970 

elaeochroa Starrett 1960; Duellman 
1970 

fimbrimembra Savage 1980 



lancasteri as moraviensis Starrett 
1960 

lancasteri Duellman 1970 

legleri Duellman 1970 

loquax Duellman 1970 

lythrodes (unknown) 

microcephala Duellman 1970 

miliaria (unknown) 

phlebodes Duellman 1970 

picadoi Robinson 1977 

pictipes Starrett 1966; Duellman 1970 

pseudopuma Starrett 1960; Duellman 
1970 

rivularis Starrett 1960; Duellman 1970 

rosenbergi Breder 1946; Duellman 
1970 

rufioculis Duellman 1970 

rufitela Duellman 1970 

staufferi Duellman 1970 

tic a Duellman 1970 

uranochroa Dunn 1924; Duellman 
1970 

xanthosticta (unknown) 

zeteki Dunn 1937; Starrett 1960; 
Duellman 1970 
Leptodactylus 

bolivianus Heyer 1970 

fragilis as labialis Heyer 1970 

melanonotus Heyer 1970 

pentadactylus Heyer 1970 

poecilochilus Heyer 1970 
Phrynohyas 

venulosa Zweifel 1964a; Duellman 
1970 
Phyllobates 

lugubris (unknown) 

vittatus as lugubris Savage 1968 

vittatus Silverstone 1976 
Phyllomedusa 

lemur Duellman 1970 
Physalaemus 

pustulosus Breder 1946 
Rana 

palmipes Volpe and Harvey 1958 

"pipiens" Orton 1952; Witschi 1956 

vibicaria Zweifel 1964b 

warschewitschii Starrett 1960 



LARVAE OF COSTA RICAN FROGS AND TOADS 53 

Rhinophrynus puma Duellman and Trueb 1966; 

dorsalis Orton 1943; Starrett 1960 Duellman 1970 

Smilisca sila Duellman and Trueb 1966; 
baudinii Duellman and Trueb 1966; Duellman 1970 

Duellman 1970 sordida Duellman and Trueb 1966; 
phaeota Duellman and Trueb 1966; Duellman 1970 

Duellman 1970 

Acknowledgments 

It is a pleasure to thank my long-time associates in the study of the Costa Rican 
frog fauna, Roy W. McDiarmid, Norman J. Scott and P. H. "Holly" Starrett, 
who have collected many of the tadpoles that have made this review possible. 
Over the years Dr. Starrett has provided insight into the many aspects of tadpole 
biology and her remarks on the manuscript are especially appreciated. 

The work could not have been completed without the continuous cooperation 
of the Universidad de Costa Rica and the Organization for Tropical Studies 
(OTS). Indirectly the National Science Foundation through grants G-6089 and 
GB-40747 has also aided the study. Finally, the opportunity to hold a John Simon 
Guggenheim Foundation Fellowship (1963-1964) provided the opportunity to ini- 
tiate the fieldwork essential to the completion of this work. 

Literature Cited 

Altig, R. 1970. A key to the tadpoles of the continental United States and Canada. Herp., 25(2): 180- 

207. 
, and R. A. Brandon. 1971. Generic key and synopses for free-living larvae and tadpoles of 

Mexican amphibians. Tulane Stud. Zool. Bot., 17(1): 10-15. 
Breder, C. M., Jr. 1946. Amphibians and reptiles of the Rio Chucunaque drainage, Darien, Panama, 

with notes on their life histories and habitats. Bull. Amer. Mus. Nat. Hist., 86(8):375-436. 
Duellman, W. E. 1963. A new species of tree frog, genus Phyllomedusa, from Costa Rica. Rev. 

Biol. Trop., 11(1): 1-23. 

. 1967. Courtship isolating mechanisms in Costa Rican hylid frogs. Herp., 23(3): 169-183. 

. 1970. The hylid frogs of Middle America. Mono. Mus. Nat. Hist. Univ. Kansas, 1(2 

vols.):xii-754. 
, and J. D. Lynch. 1969. Descriptions of Atelopus tadpoles and their relevance to atelopodid 

classification. Herp., 25(4):23 1-240. 
, and L. Trueb. 1966. Neotropical hylid frogs, genus Smilisca. Univ. Kansas Publ. Mus. Nat., 

17(7):281-375. 
Dunn, E. R. 1924. New amphibians from Panama. Occ. Paps. Boston Soc. Nat. Hist., 5:93-95. 
. 1937. The amphibian and reptilian fauna of bromeliads in Costa Rica and Panama. Copeia, 

1937(3): 163-167. 
Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae. Herp., 16(3): 183-190. 
Heyer, W. R. 1969. The adaptive ecology of the species groups of the genus Leptodactylus (Am- 
phibia, Leptodactylidae). Evol., 23(3):42 1-428. 
. 1970. Studies on the genus Leptodactylus (Amphibia: Leptodactylidae). II. Diagnosis and 

description of the Leptodactylus of Costa Rica. Rev. Bio. Trop., 16(2): 171-205. 
Limbaugh, B. A., and E. P. Volpe. 1957. Early development of the Gulf Coast toad Bufo valliceps 

Wiegmann. Amer. Mus. Nat. Hist. Nov., 1842:1-32. figs. 1-5. 
Lynch, J. D. 1976. The species groups of the South American frogs of the genus Eleutherodactylus 

(Leptodactylidae). Occ. Pap. Mus. Nat. Hist., Univ. Kansas, 61:1-24. 
Novak, R. M. and D. C. Robinson. 1975. Observations on the reproduction and ecology of the 

tropical montane toad, Bufo holdridgei Taylor in Costa Rica. Rev. Biol. Trop., 23(2):2 13—237. 



54 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Orton, G. L. 1943. The tadpole of Rhinophrynus dorsalis. Occ. Paps. Univ. Mich. Mus. Zool., 

472:1-7. 
. 1952. Key to the genera of tadpoles in the United States and Canada. Amer. Midi. Nat., 

47(2):383-395. 
Robinson, D. C. 1961. The identity of the tadpole of Anotheca coronata (Stejneger). Copeia, 1961 

(4): 495. 
. 1977. Herpetofauna bromelicola Costarricense y renacuajos de Hyla picadoi. Hist. Nat. 

Costa Rica, Mus. Nac. Costa Rica, 1:31-43. 
Savage, J. M. 1960. Geographic variation in the tadpole of the toad Bufo marinus. Copeia, 

1960(3):233-236, figs. 1-3. 

. 1967. An extraordinary new toad {Bufo) from Costa Rica. Rev. Biol. Trop., 14(2): 153-167. 

. 1968. The dendrobatid frogs of Central America. Copeia, 1968 (4):745-776. 

. 1976. A preliminary handlist of the herpetofauna of Costa Rica 2nd ed. Univ. Costa Rica: 

1-19. 
. 1980. The tadpole of the Costa Rican fringe-limbed tree-frog, Hyla fimbrimembra . Proc. Biol. 

Soc. Wash. In press. 
Silverstone, P. A. 1975. A revision of the poison-arrow frogs of the genus Dendrobates Wagler. Sci. 

Bull. Nat. Hist. Mus. Los Angeles Co., 21:1-55. 
. 1976. A revision of the poison-arrow frogs of the genus Phyllobates Bibron in Sagra (Family 

Dendrobatidae). Sci. Bull. Nat. Hist. Mus. Los Angeles Co., 27:1-53. 
Starrett, P. H. 1960. Descriptions of tadpoles of Middle American frogs. Misc. Publ. Mus. Zool. 

Univ. Michigan, 110:1-37, 1 pi. 33 text figs. 
. 1966. Rediscovery of Hyla pictipes Cope, with description of a new montane stream Hyla 

from Costa Rica. Bull. So. Calif. Acad. Sci., 65(1): 17-28. 
. 1967. Observations on the life history of frogs of the family Atelopodidae. Herp., 23(3): 195- 

203. 
. 1973. Evolutionary patterns in larval morphology. In Symp. Evol. Biol. Anurans. J. L. Vial, 

ed. Univ. Missouri, Kansas City Press. 
Taylor, E. H. 1942. Tadpoles of Mexican Anura. Univ. Kansas Sci. Bull., 28(l):37-55. 
. 1954. Frog-egg-eating tadpoles of Anotheca coronata (Stejneger) (Salientia, Hylidae). Univ. 

Kansas Sci. Bull., 36(8):589-596. 
Volpe, E. P. and S. M. Harvey. 1958. Hybridization and larval development in Rana palmipes Spix. 

Copeia, 1958(3): 197-207. 
Wager, V. A. 1965. The frogs of South Africa. Purnell and Sons. Capetown. 
Wake, D. B., and J. F. Lynch. 1976. The distribution, ecology and evolutionary history of pletho- 

dontid salamanders in tropical America. Nat. Hist. Mus. Los Angeles Co. Sci. Bull., 25:1-65. 
Wake, M. H. 1977. The reproductive biology of caecilians: an evolutionary perspective. In The 

reproductive biology of amphibians. D. H. Taylor and S. I. Guttman, eds. Plenum, N.Y. 
Witschi, E. 1956. Development of vertebrates. W. B. Saunders, N.Y. 
Zweifel, R. G. 1964a. Life history of Phrynohyas venulosa (Salientia: Hylidae) in Panama. Copeia, 

1964(l):201-208. 
. 1964b. Distribution and life history of the Central American frog, Rana vibicaria. Copeia, 

1964(2): 300-308. 

Accepted for publication April 10, 1980. 



Bull. Southern California Acad. Sci. 
79(2), 1980, pp. 55-64 

Ecology of a Coastal Salt Marsh after Long-Term 
Absence of Tidal Fluctuation 

H. Peter Eilers 

Abstract. — Ecology of a coastal salt marsh after long-term absence of tidal 
fluctuation by H. Peter Eilers, Bull. Southern California Acad. Sci., 79(2):55-64, 
1980. Tidal access to Bolsa Bay, California, was removed in 1899 then partially 
returned in late 1978. Investigation in 1977-1978 revealed that salt marsh vege- 
tation had adjusted to lowered water levels and subsidence by establishment of 
zonal patterns at anomalously low elevations. Net production in macrophytes 
decreased with elevation (4380 to 164 g rrr 2 yr _1 ), soil nitrogen (NH 3 ) remained 
high at all levels, and soil salinity was seasonally variable (7 to 54 ppt). Low 
redox potentials indicated persistent anaerobiosis. With the recent removal of 
tide gates and return of tidal fluctuation, much of the pre-existing marsh has been 
drowned. 

Department of Geography, California State University, Fullerton, California 
92634. Present address: United States Environmental Protection Agency, Cor- 
vallis, Oregon 97330. 



Introduction 



In the fall of 1978, limited tidal fluctuation was restored after nearly 80 years 
to a 52 ha section of Bolsa Bay, California. Because tidal marsh in southern 
California has been declining in area since the establishment of the State (USDI 
1972), this reclamation project is an important reversal of a long-standing trend. 
The research presented here was designed to consider the effects of such long- 
term tidal removal on the ecology of a salt marsh, especially macrophyte pro- 
duction, substrata characteristics and the relation of plants to potential tide levels. 

Bolsa Bay 

Bolsa Bay or Bolsa Chica Bay (the Spanish "little purse 11 ) occupies the seaward 
end of an alluvial-filled valley originally excavated at lower sea level in the Bolsa 
Chica-Huntington Mesa of western Orange County (Fig. 1). The present bay 
comprises two subdivisions. Outer Bolsa, directly to the west of Bolsa Chica 
Mesa, is tidally controlled through a narrow channel leading to adjacent Sunset 
Bay. A closure dam containing culverts with flap valves to prevent incursion of 
flood tides and extending from the southern edge of the mesa to the beach sep- 
arates Outer from Inner Bolsa. 

The history of human use and modifications of Bolsa Bay are considered in 
detail by Talbert (1952), Dillingham (1971), Speth et al. (1976), and EDAW (1978). 
First mentioned in the late 1790 1 s as within Rancho Bolsa Chica, Mexican title 
was established in 1840 and reconfirmed in the treaty of Guadalupe-Hidalgo 1948. 
The earliest detailed survey of Bolsa Bay, published in 1874 (Fig. 2), reveals an 
undisturbed wetland system of over 800 ha supplied with some terrestrial drainage 



56 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




PACIFIC OCEAN 



Fig. 1. Bolsa Bay, California. Dashed lines represent remnants of degraded marsh, old diked 
ponds, and filled areas. Roads interconnect oil well sites in Inner Bolsa. Square pattern denotes salt 
marsh. Dike is the landward boundary of Bolsa Chica Ecological Reserve and PCH is Pacific Coast 
Highway (State Highway 1). Note position of sampling transect. 



through the Freeman River and open directly to the Pacific Ocean. Alteration of 
the natural system began in 1899 with ownership of 211 ha granted to the Bolsa 
Chica Gun Club by the State of California. Dam emplacement in that year re- 
stricted tidal fluctuation to Outer Bolsa and presumably enhanced waterfowl hunt- 
ing. Loss of the tidal prism led to rapid siltation and permanent closure of the 
bay mouth within six months necessitating a new channel between Outer Bolsa 
and the present site of Huntington Harbor in adjacent Sunset Bay (Moffatt and 
Nichol 1971). Later the hunting club constructed a series of diked ponds in the 
more elevated marsh. Oil was discovered in 1926 with leases to the Signal Oil 
Company granted by the duck club decendants in 1940. Oil and gas exploitation 
followed extensive diking and well pad construction in 1943, and extraction con- 
tinues to the present. Signal Oil became sole landowner of Outer and Inner Bolsa 
in 1970 and initiated plans to convert the bay to a marina when soil resources are 
exhausted in about 15 years. 

Against this backdrop, however, existed pressure from local residents and the 
State of California to restore the wetlands and public access to them. In addition, 
study of tidally controlled wetlands suggested the importance of marshes to es- 
tuarine and coastal food chains (Teal 1962). Through a complex agreement be- 
tween the Signal Oil Company and the State of California in 1973, Signal retained 
title to all but 140 ha of wetland. An additional 90 ha will revert to the State in 
1987 if a new ocean channel is established. Following this agreement, the State 
of California through the Department of Fish and Game initiated the Bolsa Chica 
Ecological Reserve and began planning for the restoration of tidal fluctuation to 
Inner Bolsa and the construction of ocean access. 



ECOLOGY OF COASTAL SALT MARSH 



57 




Fig. 2. Bolsa Bay, 1874. Note ocean inlet. 



Methods 

To understand the effect of long-term absence of tidal fluctuation on the salt 
marsh system at Bolsa Bay, study was initiated in October 1977 and continued 
through March 1979. Concurrent investigations of other less altered southern 
California salt marshes (Sweetwater River, Los Penasquitos Lagoon, and Upper 
Newport Bay) were underway by me to provide comparative data (Eilers 1980). 

Vegetation 

A transect of five 2.0 x 2.5 m macroplots (BB-1 to BB-5) was located within 
the salt marsh perpendicular to the elevation gradient along the main channel in 
Inner Bolsa (Fig. 1). At intervals of six to eight weeks for a period of 13 months 
(October 1977 through October 1978), a 20 x 50 cm quadrat frame was located 
within each macroplot (at a different position each session) and all above-ground 
vascular plant material in the frame was harvested, placed in plastic bags, then 
refrigerated within two hours to minimize fermentation losses (Milner and Hughes 
1968). In the laboratory each sample was divided into component species. Species 
fractions were further divided into living and dead material. Subfractions were 
then dried to constant weight in a gravity convection oven at 85°C and weighed 
to the nearest 0.1 g. 

Annual net production estimates for macroplots were calculated by the method 
of Smalley (1959) applied to component species as follows: 1) If the net change 
between sampling periods was positive for both the live biomass (L) and dead 
biomass (D), then species production in the interval was equal to the sum (AL + 
AD); 2) if AL and AD were negative, production was assumed to be zero; 3) if 
AL and AD were + and -, respectively, then species production was equal to 
AL; and 4) if AL and AD were - and +, respectively, species production was 
assumed to be equal to the sum (AL + AD) if the sum was greater than zero, and 
equal to zero if the sum was negative. Net production for each species between 



58 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

harvests was summed to yield species annual net production, and macroplot 
annual net production was determined as the sum of component species produc- 
tion. 

Physical Parameters 

At the time of each vegetation harvest, soil water was analyzed for salt content 
with a Goldberg refractometer and oxidation-reduction potential (redox, as Eh in 
millivolts), ammonia, and pH was measured with an Orion specific ion meter 
(Behrens 1965; Banwart et al. 1972). Elevation of each macroplot with reference 
to National Geodetic Vertical Datum (NGVD) was established by survey and 
converted to potential mean high water (MHW) based on MHW for the nearest 
tidal bench mark (Warner Avenue bridge) and the work of Moffatt and Nichol 
(1971). All levels were double run with a closing error less than 0.2 cm. 

Results 

Vegetation 

Four vascular plant species were recorded in quadrat samples. They were, in 
order of occurrence from high to low sites, Frankenia grandiflora, Distichlis 
spicata, Salicornia virginica, and Spartina foliosa (Fig. 3). 

Net above-ground production varied from 4380 g m 2 yr * for a nearly pure 
stand of Frankenia (BB-1) to 164 g m~ 2 yr -1 for low-density Spartina (BB-5). 
More diverse stands (BB-2,3,4) were intermediate (2214, 3522, and 1561 g m~ 2 
yr 1 , respectively). A sharp decrease in net production accompanied a decrease 
in elevation (Fig. 3), and the low-elevation Spartina appeared to be dying out as 
the study progressed despite the fact that this species is characterized by loss of 
aerials in the winter. 

Physical Parameters 

The sampling transect was 30 m long and extended from -0.57 to —1.12 m 
MHW (Fig. 3). Mean annual soil salinity for sample sites varied from 20 to 41 
ppt with an increase toward lower sites (Fig. 4). The greatest annual range of soil 
salinity was recorded for BB-1 (7 to 45 ppt); for all sites minimum values occurred 
in the winter and early spring, while maximum values appeared in the summer. 

Oxidation-reduction potential (redox) is a convenient measure of soil aeration 
(Brereton 1971). Values less than 200 mV indicate poor drainage and anaerobic 
conditions (Stolzy and Fluhler 1978). Even though all macroplots recorded redox 
potentials greater than 200 mV in the early spring, annual means revealed per- 
sistent and increased anaerobiosis toward lower elevations along the transect 
(Fig. 4). Sampling sites were perpetually waterlogged, and disturbance of the 
peaty soil resulted in release of hydrogen sulfide. 

Mean annual soil nitrogen (as NH 3 ) varied greatly along the transect with no 
clear relationship to elevation, although there was a tendency toward increased 
annual range for lower macroplots (Fig. 4). Minimum and maximum nitrogen 
occurred in the spring and summer, respectively. 

Measurement of soil pH provides a general assessment of nutrient ion concen- 
tration, and variations in pH over time may indicate fluctuations in soil moisture 
(Neely 1962). Greater annual variation in pH for more elevated macroplots (Fig. 



ECOLOGY OF COASTAL SALT MARSH 



59 



5000-. 
4000- 
3000- 
2000- 

1000- 



NET PRODUCTION 
(g m- 2 yr- 1 ) 



FRANKENIA 
1 
DISTICHLIS 



SALICORNIA 



MHW-i 

-0.5- 
-1.0- 
-1.5- 



ELEVATION 
(METERS) 



1 1 

10 20 

DISTANCE ALONG TRANSECT (METERS) 



1 
30 



Fig. 3. Net production, species composition and macroplot elevation along sampling transect in 
Inner Bolsa. 



4) suggested that some drying takes place near the summit of the transect. Annual 
mean pH reflected increased alkalinity at lower elevations. 

Discussion 

Vegetation and environmental measurements together suggested that the ab- 
sence of tidal fluctuation for 80 years strongly influenced but did not totally alter 
the salt marsh system of Inner Bolsa Bay. Net production of vascular plants, 
except for Spartina at BB-5 was comparable to that observed elsewhere in south- 
ern California marshes (Winfield and Zedler 1976; Eilers 1980). 

The presence of Spartina in Inner Bolsa was curious. This grass dominates 
low marsh environments from northern California (Humboldt Bay) to the south- 
ern most extent of salt marsh at Laguna Ojo de Liebre, Baja California (Phleger 
1965; Macdonald 1969). A tall (1 m) rhizomatous grass with prominent aerenchy- 
ma tissue, Spartina foliosa is capable of withstanding long submergence (Purer 
1942). Macdonald (1977) recognizes two distinct groups of estuaries and coastal 
lagoons in southern California based in part of the presence of this species. The 
first of these is characterized by having deep channels, large tidal prism, perennial 



60 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



60-. 
40- 
20- 
0- 
400 
200H 

-200 
100-1 
75 
50 
25 
J 
8.0-. 
7.5- 
7.0 
6.5- 1 



SOIL SALINITY 
(PPT. ) 





SOIL REDOX POTENTIAL 
(Eh-mV) 



SOIL AMMONIA 
(10" 5 M) 




MHW- 
-0.5- 
-1.0- 

-1.5- 




1 1 1 

10 20 30 

DISTANCE ALONG TRANSECT (METERS) 



Fig. 4. Soil salinity, redox potential, ammonia and pH along sampling transect in Inner Bolsa. 
Annual means for sample macroplots shown as filled circles; vertical lines denote annual range. 



ocean contact and well-developed, pure stands of S. foliosa directly above mud- 
flat. The second group includes those with insufficient tidal prism to prevent 
seasonal or long-term closure of their ocean inlets. Invariably, according to Mac- 
donald (1977), S. foliosa is absent from the second group. Vigorous stands of S. 
foliosa are present at open tidal sites such as Upper Newport Bay (Vogl 1966), 
Tijuana Estuary (Zedler 1977) and Anaheim Bay (Speth et al. 1976); S. foliosa 
is absent at Los Penasquitos Lagoon where tidal access is restricted to winter 
months (Mudie et al. 1974). The restricted distribution of S. foliosa at Mugu 
Lagoon where tidal fluctuation is reduced (Warme 1971), and its absence from 
diked marsh at La Ballona Creek (Clark 1979), lends further support to the Mac- 
donald (1977) model. The occurrence of S. foliosa at Bolsa where tidal fluctuation 
was not permitted for 80 years is, therefore, difficult to explain. Perhaps the 



ECOLOGY OF COASTAL SALT MARSH 61 

Table 1 . Mean redox potential of substrata in four southern California salt marshes obtained at 
harvest sessions in 1977-1978 (Eh-mV). After Eilers (1980). 

1977 1978 





Oct/ 


Dec/ 












Marsh 


Nov 


Jan 


Feb 


Apr 


June 


July 


Aug 


Bolsa Bay 


65 


307 


-1 


27 


55 


1 


103 


Upper Newport Bay 


304 


407 


289 


337 


— 


289 


261 


Los Penasquitos Lagoon 


382 


381 


301 


384 


242 


276 


322 


Sweetwater River Estuary 


369 


— 


369 


311 


428 


354 


354 



unique environment created by artificial removal of tides is a factor. Yet, because 
the salinity regime and soil alkalinity at BB-3,4,5 where this grass was sampled 
were characteristic of sea water, and because elevations there were well below 
MHW, it is likely that some sea water inflow through the permeable beach sands 
occurred and permitted Spartina, which was probably present in the bay before 
1899, to persist. 

Low redox potentials and attendant anaerobiosis were among the more signif- 
icant effects of diking and removal of the tides. Redox measurements obtained 
in undiked southern California marshes were consistently higher than those for 
Bolsa Bay (Table 1). Peat accumulations 15 to 20 cm thick in each macroplot at 
Bolsa indicated poor nutrient turnover and restricted nutrient contribution to the 
adjacent mudflat and channel. 

The relationship between plants and tide levels has been a central theme in salt 
marsh research since the pioneer investigations of Ganong (1903) and Johnson 
and York (1915), through the work of Hinde (1954) and Chapman (1960), and 
continuing to the present with studies, for example, by Zedler (1977), Eilers (1979) 
and Frenkel et al. (In press). Salt marsh macrophytes have been shown to occupy 
relatively narrow elevational limits and the vertical boundaries of salt marshes 
tend to be somewhat uniform from one coastal location to the next. Chapman 
(1934 in Hinde 1954) observed that none of the higher plant associations or species 
in the British marshes he investigated had a lower limit below mean sea level 
(MSL). Mahall and Park (1976) recorded Spartina folios a in San Francisco Bay 
salt marshes from about MSL to MHW and Salicornia virginica from MHW to 
the upper limit of tidal fluctuation. Warme (1971) and Barbour et al. (1973) like- 
wise noted the presence of salt marsh above MSL, and Macdonald (1969) gen- 
eralized that most Pacific coast marshes become established at about mean lower 
high water (MLHW) regardless of local tidal range. The upper limit of salt marsh 
is often more difficult to define because a gradual transition between marsh and 
upland is frequently present (Harvey et al. 1978). The National Ocean Survey 
(NO A A 1980) surveyed 17 Pacific coast salt marshes and found a mean upper 
boundary (defined as the center of the transition zone) of 0.9 m above MHW for 
central and southern California locations. 

Most if not all of the work to date concerning salt marsh plants and tide levels 
has been completed in estuaries, bays and lagoons where tidal fluctuation is 
unrestricted or is at least a seasonal event occurring over several months. Com- 
parison of the vertical range of salt marsh vegetation at Inner Bolsa with that of 



62 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



-0.5' 



Fig. 5. Vertical range of salt marsh at Bolsa Bay compared to that of other Pacific Coast salt 
marshes. Abbreviations as follows: TJE, Tijuana River Estuary (Zedler 1977); SRE, Sweetwater River 
Estuary (Eilers 1980); MB, Mission Bay (Macdonald 1969); LPL, Los Penasquitos Lagoon (Eilers 
1980); UNB, Upper Newport Bay (Eilers 1980); BBa, Inner Bolsa Bay; BBb, Outer Bolsa Bay; ML, 
Mugu Lagoon (Warme 1971); SFB, San Francisco Bay (Hinde 1954); NRE, Nehalem River Estuary, 
Oregon (Eilers 1979). 



Outer Bolsa and other tidally influenced southern California marshes, therefore, 
provides some insight into the vertical distribution of plant species and marsh 
after long-term removal of tidal access (Fig. 5). Apparently, in the 80 year absence 
of periodic inundation, salt marsh plants abandoned positions at or above MHW 
and re-established at anomalously low elevations. Frankenia, for example, was 
located 0.3 to 0.9 m below its normal elevation and equal to or below the lowest 
Spartina or Salicornia found elsewhere as reported by Zedler (1977) and Hinde 
(1954). 

This anomalous condition may not be due to the removal of tidal fluctuation 
alone but to a combination of tidal removal and subsidence (resulting from oil or 
ground water withdrawals or both). Moffatt and Nichol (1971) studied bench mark 
levels for the time period 1933 to 1964 and estimated subsidence at between 23 
and 46 cm. Orange County bench marks along the west side of Inner Bolsa have 
been repeatedly surveyed and show continued loss of elevation. For example, 
bench mark IJ-29-68 near the sampling transect descended by 8.4 cm between 
1968 and 1976. From the available data, a mean subsidence rate for Inner Bolsa 
may be estimated at about 1 cm per year. Subsidence could account for a marsh- 
land shift downward of approximately 80 cm since the removal of tidal fluctuation 
and, together with some re-establishment or vegetation extention of marsh plants 
to lower levels, may be responsible for the anomaly. Whatever the cause, how- 



ECOLOGY OF COASTAL SALT MARSH 63 

ever, it is obvious that restoration of unrestricted tidal access to Inner Bolsa will 
lead to prolonged inundation and disruption of the existing salt marsh. 

Following the construction of a parking lot for visitors and a dike to prevent 
inundation of land owned by Signal (Fig. 1), partial tidal contact was returned to 
the Bolsa Chica Ecological Reserve in Inner Bolsa by removing flap valves on 
culverts through the old dike on 15 November 1978. The lower marsh at the time 
of writing (August 1979) was in an advanced state of decay and, due to the 
restricted nature of the culverts, water fluctuation (as recorded from a temporary 
staff gauge near the parking lot) was limited to 0.5 m above and below -0.75 m 
MHW. 

Acknowledgments 

I wish to acknowledge the contribution of Kathleen Kunz who served as field 
assistant and Inda Taylor and Michael Sweesy for their help in field work and 
laboratory analysis. I thank Keith Macdonald for critical review of the manu- 
script. This research was supported by U.S. Environmental Protection Agency 
grant number R805438-01-1 to California State University, Fullerton. 

Literature Cited 

Banwart, W. L., M. A. Tabatabai, and J. M. Bremner. 1972. Determination of ammonium in soil 

extracts and water samples by an ammonia electrode. Comm. in Soil Sci. and Plant Anal., 

3:449. 
Barbour, M. G., R. B. Craig, F. R. Drysdale, and M. T. Ghiselin. 1973. Coastal Ecology, Bodega 

Head. University of California Press, Berkeley. 
Behrens, E. W. 1965. Use of the Goldburg Refractometer as a salinometer for biological and geo- 
logical field work. Journal of Marine Research, 23:165-171. 
Brereton, A. J. 1971. The structure of the species populations in the initial stages of salt-marsh 

succession. J. Ecology, 59:321-338. 
Chapman, V. J. 1960. Salt marshes and salt deserts of the world. Interscience Publishers, Inc., New 

York. 
Clark, John. 1979. Ballona wetlands study. Part I. Report to School of Agriculture and Urban 

Planning. University of California, Los Angeles. 
Dillingham Environmental Co. 1971. An environmental evaluation of the Bolsa Chica area. I— III. La 

Jolla, California. 
Eilers, H. Peter. 1979. Production ecology in an Oregon coastal salt marsh. Estuarine and Coastal 

Marine Science, 8:399-410. 
. 1980. Production in coastal salt marshes of southern California. Report to Environmental 

Protection Agency, Corvallis, Oregon. 
Frenkel, R. E., H. P. Eilers, and C. A. Jefferson. In press. Oregon coastal salt marsh upper limits 

and tidal datums. Estuaries. 
Ganong, W. F. 1903. The vegetation of the Bay of Fundy salt and diked marshes: an ecological 

study. Botanical Gazette, 36:161-186, 280-302, 349-367, 429-455. 
Harvey, H. T., M. J. Kutilek, and K. M. DiVittorio. 1978. Determination of transition zone limits 

in coastal California wetlands. Report to United States Environmental Protection Agency, 

Corvallis, Oregon. 
Hinde, H. P. 1954. The vertical distribution of salt marsh phanerogams in relation to tide levels. 

Ecology, 24:209-225. 
Johnson, D. S., and H. H. York. 1915. The relation of plants to tide-levels. Carnegie Institute of 

Washington Publication 206. 
Macdonald, K. B. 1969. Quantitative studies of salt marsh mollusc faunas from the North American 

Pacific Coast. Ecology, 39:33-58. 



64 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



-. 1977. Coastal salt marsh. Pages 263-294 in M. G. Barbour, and J. Major (eds). Terrestrial 
Vegetation of California. University of California Press. 



Mahall, B. E., and R. B. Park. 1976. The ecotone between Spartina foliosa Trin. and Salicornia 

virginica L. in salt marshes of northern San Francisco Bay: I. Biomass and production. J. 

Ecology, 64:421-433. 
Milner, C, and R. E. Hughes. 1968. Methods for the measurement of the primary production of 

grassland. I.B.P. Handbook No. 6. Blackwell Scientific Publications, Oxford. 
Moffatt and Nichol, Engineers. 1971. Historic tideland investigation of Bolsa Chica and Anaheim 

Bays, Orange County, California. 
Mudie, P. J., B. Browning, and J. Speth. 1974. The natural resources of Los Penasquitos Lagoon 

and recommendations for use and development. State of California Department of Fish and 

Game. Coastal Wetland Series No. 7. 
National Oceanic and Atmospheric Administration, National Ocean Survey. 1980. The relationship 

between the upper limit of coastal wetlands and tidal datums along the Pacific Coast. Rockville, 

Maryland. 
Neely, W. W. 1962. Saline soils and brackish waters in management of wildlife fish and shrimp. 

Trans. N. Amer. Wildl. Conf., 27:321-334. 
Phleger, F. B. 1965. Sedimentology of Guerrero Negro Lagoon, Baja California, Mexico. Papers 

Colston Research Society, 17:205-237. 
Purer, E. A. 1942. Plant ecology of the coastal salt marshes of San Diego County, California. 

Ecological Monographs, 12:81-111. 
Smalley, A. E. 1959. The role of two invertebrate populations Littorina irrorata and Orchelinum 

fidicinium in the energy flow of a salt marsh ecosystem. Ph.D. Thesis, University of Georgia, 

Athens. 
Speth, J. W., B. M. Browning, and K. S. Smith. 1976. The natural resources of Anaheim Bay- 
Huntington Harbor. California Department of Fish and Game and U.S. Fish and Wildl. Service. 

Coastal Wetland Series No. 18. 
Stolzy, L. H., and H. Fluhler. 1978. Measurement and prediction of anaerobiosis in soils. Pages 

363-425 in D. R. Nielsen and J. G. MacDonald (eds). Nitrogen in the Environment. Vol. 1. 

Nitrogen Behavior in Field Soil. Academic Press, New York. 
Talbert, T. B. 1952. My sixty years in California. Huntington Beach News Press. 
Teal, J. M. 1962. Energy flow in the salt marsh ecosystem of Georgia. Ecology, 43:614-624. 
U.S. Department of Interior. 1972. Southern California estuaries and coastal wetlands — endangered 

environments. U.S. Department of Interior, Portland, Oregon. 
Winfield, T. P., and J. B. Zedler. 1976. Comparative productivity of salt marsh vegetation in the 

Tijuana Estuary (Southern California). Abstract of paper presented at 57th annual meeting of 

Western Society of Naturalists, Fullerton, California. 
Vogl, R. J. 1966. Salt-marsh vegetation of Upper Newport Bay, California. Ecology, 47:80-87. 
Warme, J. E. 1971. Paleoecological aspects of a modern coastal lagoon. University of California 

Publ. Geol. Sci., 87:131. 
Zedler, J. B. 1977. Salt marsh community structure in the Tijuana Estuary, California. Estuarine 

and Coastal Marine Science, 5:39-53. 

Accepted for publication April 8, 1980. 



Bull. Southern California Acad. Sci. 
79(2), 1980, pp. 65-77 

A General "Exact Test" for N x M Contingency Tables 

Harrington Wells and Jack Lester King 

Abstract. — A general "Exact Test" for n x m contingency tables by Harring- 
ton Wells and Jack Lester King, Bull. Southern California Acad. Sci., 79(2):65- 
77, 1980. A generalized Exact Test for n x m contingency tables based upon 
methodology of the Fisher- Yates Exact Test for 2 x 2 tables and the chi-square 
statistic is presented. The General Exact Test is a multi-tailed statistic with no 
minimum expected matrix cell values required. A computer program for the Gen- 
eral Exact Test is described and presented in the computer language BASIC. The 
number of possible tables that need to be considered for an Exact Test can be 
very large. However, when expected cell sizes are large the chi-square distri- 
bution can be used for an approximation to the probability. The generalized Exact 
Test should be very useful for n x m contingency tables where categories are 
nominally measured and some expected values are small (e.g. taxonomic prob- 
lems involving type specimens, or protein electrophoretic data), since other avail- 
able statistical tests require arbitrary lumping of categories and/or populations. 

Department of Biology, University of Tulsa, Tulsa, Oklahoma 74104, and De- 
partment of Biological Sciences, University of California, Santa Barbara, Cali- 
fornia 93106. 



Introduction 



Experimental situations often occur in which only small samples may be ob- 
tained, or where some types of events are rare and others are not. For example, 
medical studies on human subjects are often limited to a few individuals. Simi- 
larly, an environmental impact study of a rare or endangered organism might be 
limited to small samples. In other cases the high cost of acquiring data may result 
in few observations being made. On the other hand, genetic and other types of 
research often involve investigation of rare events (e.g. rare allele occurrence). 
Although the sample size in this latter type of study may be large, the number of 
observations of a particular type of event is small. If the data are of nominal 
measure, and there are low expected numbers of observation for some event 
type, then the data are essentially untestable unless nonarbitrary lumping of sam- 
ple populations or categories can be made, or unless the data form a 2 x 2 table 
and directional predictions are made a priori. 

This article describes a generalized Exact Test for n x m contingency tables. 
Two variations of the Exact Test are presented; one based on the multinomial 
distribution and the other on the multivariate hypergeometric distribution. The 
method combines the technique of the Fisher- Yates Exact Test for 2 x 2 matrices 
(Fisher 1934; Yates 1934), and the chi-square statistic (Pearson 1900, 1911) to 
produce a multi-tailed statistical test with no minimum expected frequency re- 
quirements for any size matrix. The technique is presented as a computer pro- 
gram. 



66 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

The investigator often obtains biological data from several independent samples 
and wishes to know whether they should be regarded as having come from the 
same statistical population. The problem is to determine whether the observed 
variation between samples is due to chance difference in random samples from 
the same population, or whether the variability signifies true differences among 
populations. If the researcher can assume that the samples are drawn indepen- 
dently from normally distributed populations of equal variance then the standard 
parametric F statistic can be used, which is the statistic for analysis of variance. 
However, if the assumptions of the parametric statistic cannot be made, a non- 
parametric alternative is required. For data which are at least ordinal there are 
several appropriate nonparametric statistics. However, there is basically only one 
general nonparametric statistic for nominal data in contingency tables larger than 
2x2: the chi-square (Lehmann 1975). 

The classic statistical test of the hypothesis that the sample populations have 
distributions which are homogeneous for data in a n x m contingency table has 
been the chi-square with (n - l)(m - 1) degrees of freedom. The chi-square test, 
however, requires that the expected values in each cell not be too small. When 
this requirement is violated the results of the test are meaningless since the chi- 
square distribution no longer is a good approximation to the sum of relative 
squared differences. Siegel (1956) recommends that fewer than 20 percent of the 
expected values for a x 2 with df larger than 1 be less than 5, and that no cell 
should have expected values less than 1. More recent studies involving known 
distributions suggest that these criteria may be conservative and adequate criteria 
for the x 2 test may be only that all expected values are greater than 1.333 (Yarnold 
1970; Larntz 1978). In either case, if data do not meet the criteria, columns and/ 
or rows must be lumped until the criteria are met. If there are only 2 categories 
of result and 2 populations in the problem, then the data are testable by the 
Fisher- Yates Exact Test regardless of expected cell values if the experimenter 
can use a one tail test (Siegel 1956). Frequently, studies involve more than two 
populations, or more than two categories of results, or are two tailed, and in 
addition deal with rare events. Genetic data often are untestable without lumping, 
since usually there are 1, 2, or 3 common alleles and an additional number of rare 
alleles. In other types of experiments it may be expensive to obtain large samples, 
or the data may have been collected by other researchers prior to conception of 
the current experiment. Taxonomic studies often can have this latter type of 
problem. Arbitrary lumping of data has been the only solution. 

Methods and Examples 
The data to be analyzed are assumed to consist of observations which can be 
placed into discrete mutually exclusive categories, and to be of either nominal 
or ordinal measure. Furthermore, the data are assumed to have come from two 
or more independent samples. Thus, random samples are drawn from n differ- 
ent sampling populations, and each observation in each sample can be classified 
as one of m different types. Then, the data can be expressed as the n by m 
contingency table (1); where k y represents the number of observations of the 
]th type in the random sample from the \th population for i = 1, . . . , n and 

m n 

j = 1, . . . , m, n = ^ ky for i = 1, . . . , n are the row sums, Cj = V k y for 

j=l i=l 



EXACT TEST FOR N x M CONTINGENCY TABLES 67 

n m 

j = 1, . . . , m are the column sums, and N = ^ ^ k u is the sum of all elements 

i=l j=l 

of the data matrix. 



(1) 





CATEGORY 


ROW SUM 




Ku K 12 . . . K lm 


i"i 


POPULATION 


K 2 i K 2 2 • • • k 2m 


r 2 




kni k n 2 • . . K nm 


r n 


COLUMN SUM 


c l c 2 • • • C m 


N 



The null hypothesis to be tested is that the proportion of individuals which 
can be classified as belonging to a given category is equal in all populations 
sampled, and that this supposition is true for each observation category. How- 
ever, proportionately equal numbers of individuals of different types is not 
hypothesized to necessarily occur in the sampled populations. Let p y denote 
the true probability that an observation chosen at random from the ith sample 
population will be of type j for i = 1, . . . , n and j = 1, . . . , m. Of course, 
the true probabilities {py} are not known. Since every observation can be 

n 

categorized as one of the given possible types, ^ p y = 1 for i = 1, . . . , n. 

j=i 
Then, the hypothesis to be tested may be algebraicly expressed as (2). 



H : Pij = p 2 j = • • • = Pnj for j = 1, . . . , m 
H^ The hypothesis H is not true. 



(2) 



As equation (2) states, the null hypothesis H asserts that all the distributions 
from the n different samples are actually alike, which is to say they are 
homogeneous. When the null hypothesis H is true the n random samples are 
drawn from the same distribution. Therefore, given that H is true, the maximum 
likelihood estimator of p y is the same for all values of i, and this estimator is 
pu = Cj/N for j = 1, . . . , n. It follows directly that under H the maximum 
likelihood estimator of the expected number of individuals of type j to be ex- 
perimentally observed from the ith sample population is ey = npy = r^c/N) 
for i = 1 , . . . , n and j = 1 , . . . , m. 

The crucial question in defining a general Exact Test is, what criteria does 
one choose to determine how deviant a given table is from the results expected 
if the null hypothesis is true? The difference between the actual number of 
observations recorded of type j from population i, and the expected number 
under H will tend to be smaller when H is true than when H is not. It seems 
reasonable, therefore, to base a test of the hypothesis (2) on values of the 
difference A = (k y - ey). However, it is obvious that the importance of a given 
deviation from expected is dependent on the expected value, and that since 
deviation can be either positive or negative from the expected value, a simple 
summing of the relative differences will result in cancellation of differences. 
We have chosen to use the statistic Q in (3) to correct for the summation 
and weighting difficulties because it has been shown to be a good test of dif- 
ference by Pearson (1900, 1911), and others, irrespective of the fact that the 



68 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

chi-square distribution is a good approximation to (3) when {e u } are large. 
For our purposes, the fact that the chi-square test gives a good approximation 
of the probability of obtaining a value of Q as large or larger than that ob- 
served when expected cell frequencies are large, is an additional reason for 
choosing (3). 

n m 

Q = 2 2 (k " ~ ^ )2/d " (3) 

i=l j=l 

The probability of obtaining a known table by random samples, given that 
the null hypothesis is true, can be found by using equation (4) since the sampling 
distribution is multinomially distributed under H ; where x! is x factorial. 



Tin 



11 'i- m / c .\£ 

Probability =1 ^--n (4 H 

nnw 1 



ik u 



(4) 



The generalized Exact Test finds the probability of observing a table with 
Q value greater than or equal to Q of the table actually obtained by the in- 
vestigator. The fact that Q is the divergence measure of the widely used x 2 
statistical test, attests to its appropriateness as a measure of deviation from 
results expected under H . The exact test is performed by creating all possible 
tables that have the same row (population) sums as the observed data matrix. 

If both the row (population) and column (category) marginals can be fixed 
then the probability of obtaining a known table by random samples, given that 
the null hypothesis is true, can be found by using equation (5) since the sampling 
distribution is multivariate hypergeometrically distributed under H ; where 
x! is x factorial. 



n m 



n ■-.! n *>'■ 

Probability = -!=! bl (5) 

Ni n n k .. ! 

i = l j = l 

Methodologically, the Exact Test using the multivariate hypergeometric dis- 
tribution is performed by creating all possible tables that have the same row and 
column marginal sums as the observed data matrix. The technique, then, is anal- 
ogous to the procedure of the Fisher- Yates Exact Test for 2 x 2 tables, though 
the criteria for determining what is a more deviant table differ. 

Using either distribution, for each table the Q statistic and probability of ob- 
taining the table when the null hypothesis is true are found. The probability of 
obtaining a table with value of Q as large or larger than Q of the observed data 
matrix, then, is the sum of the probabilities of those tables which have Q values 
greater than or equal to Q of the table actually obtained in the investigation. The 
General Exact Test is non-directional, so no a priori directional assumptions are 
required. Table 1 gives an example for a small matrix, and Table 2 for a large 
matrix. 

Exact Test problems are best solved by use of a computer (TRS-80 in our 
case), since the larger n, m, and N become the more tables need be considered 



EXACT TEST FOR N x M CONTINGENCY TABLES 



69 



Table 1. Relation between occurrence of individuals of morphological type defined by the 3rd 
principal coordinate analysis axis and grassland habitat location (from Wells 1979). 





Observed Data 








Principal Coordinate Axis III 




Grassland Location 


+ Cluster 


Cluster 


Total 


Main Group 

Pine Forest Isolated 

Total 


14 

1 

15 


83 
10 
93 


97 

11 

108 



Multinomial Exact Test P = .8964 
Hypergeometric Exact Test P = .7072 
X 2 = .2357 DF = 1 P = .803 

1 176 Tables were Created for the Multinomial Test 
12 Tables were Created for the Hypergeometric Test 



and the more difficult it becomes to calculate each table's probability. Unfortu- 
nately, even with the fastest computers it is not practical to test some problems 
with the Exact Test statistic method. There simply are too many tables to be 
considered. 

Use of the multinomial distribution for the Exact Test requires generation of 
many more tables than when the multivariate hypergeometric distribution is used, 
since the latter has only row marginals fixed. For example, Table 1 analyzed by 
use of the multinomial Exact Test requires generation of 1176 tables while the 
multivariate hypergeometric Exact Test requires generation of only 12 tables. 
Thus, the multinomial variation of the General Exact Test is more often impracti- 
cal to perform than is the hypergeometric version of the Exact Test. However, 
the criteria for the hypergeometric distribution are rarely truly met. That is, 
resampling the same set of populations, with population sample sizes equal to 
those of the first sample will, due to sampling error, often give results differing 
from those of the original sample. Even so, for the reasons shown by Fisher 
(1956) for the Fisher- Yates 2x2 Exact Test, the multivariate hypergeometric 
distribution will give a good approximation. Enumeration techniques (Wilson 



Table 2. Relation between occurrence of individuals of electrophoresis alkaline phosphatase phe- 
notypes and habitat type (from Wells 1979). 

Alkaline Phosphatase 



Habitat 



Individual Phenotype 



FS 



Total 



Oak Forest 


52 


4 










56 


Grassland 


88 


5 


3 




6 


102 


Chaparral 


100 


3 


1 







104 


Total 


240 


12 


4 




6 


262 




Hypergeometric 


Exact Test P = 


.02343 







70 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Table 3. Relation between occurrence of individuals of electrophoresis acid phosphatase pheno- 
types and habitat type (from Wells 1979). 

Acid Phosphatase 



Individual Phenotype 



Habitat S F Total 

Oak Forest 15 24 39 

Grassland 51 105 156 

Chaparral 29 77 106 

Total 95 206 301 

Hypergeometric Exact Test/ 5 = .412 

X 1 Estimated P = .416 



1941; Barnard 1945) seem only to enhance the apparent significance of the data. 
Therefore, use of the multivariate hypergeometric distribution for the Exact Test 
when the multinomial distribution may be impractical is conservative and closely 
approximates the multinomial probability. 

Finally, many cases in which tables are too large to be efficiently tested via 
either Exact Test method will have large expected matrix cell values. When the 
expected numbers of all cells are large the Q statistic is approximately distributed 
as x 2 , and a chi-square table can be used to obtain a close estimate of the true 
probability. For example, consider Table 3. The Exact Test probability (hyper- 
geometric) of an as likely or less likely table (value of Q) is .412 while the prob- 
ability obtained by using the x 2 distribution is .416 (by interpolation). Remember 
though that the Exact Test does not require large expected values whereas the 
chi-square approximation does. 

An exact test for 3x2 contingency tables has previously been proposed in a 
book by Pierce (1970). However, Pierce simply defined his Exact Test probability 
as the sum of the probabilities of the tables which have a probability of occurrence 
less than or equal to that of the observed data table. His method has no quick 
approximation when expected cell values are large, is only for 2 x 2 or 3 x 2 
tables, the power and efficiency have not been calculated as has been done for 
the x'\ an d the probability obtained by Pierce's method is not necessarily equal 
to that obtained with the x 2 method. Using the actually observed data of Table 
1 as an example, Pierce's method gives a probability of 1.000 which is larger than 
either Exact Test probability (multinomial P = .8963, hypergeometric P = 
.7072), or the chi-square ( x 2 = .2357, df = 1, P - .803). 

Finally, consider a one tail modification of the hypergeometric Exact Test for 
2x2 tables, and how the results compare to those of the Fisher- Yates Exact 
Test for 2 x 2 tables. For 2x2 tables the value of Q is equal to zero and is the 
minimum when the observed values are equal to the expected numbers. Fur- 
thermore, the value of Q is strictly increasing monotonically in either direction 
from the expected value. The proof is as follows. Let {e n } be the expected values, 
and x and y be some deviation from expected; where x = y + z for any y s= 
and z > 0. Q x and Q v are then given by equation (6). Since the rows may be 
interchanged without changing the answer, and since reversing the rows reverses 
the sign on y and z, the proof for any y =s and z < is also given by (6). 



EXACT TEST FOR N x M CONTINGENCY TABLES 71 



Table 4. Hypothetical example demonstrating how program initiates matrix and how all possible 
tables are formed. 



Hypergeometric ( 
Categories 


;4a) 






Observed Table 
Categories 








Multinomial (4b) 
Categories 




A 


BCD 


Total 




A 


B 


C 


D 


Total 




A 


BCD Total 


1 X_1 

- Y 5 

3 

P. 

£ Z 5 


7 4 2 
0< — 0< — 2 
0^0<— 


12 

5 
5 


| X 

3 

a 
cu Z 


5 
3 
1 


4 

1 

2 


2 


2 


1 
1 




12 

5 
5 


JS v 

3 J 

a 


12 

5 

5 


0< — 0< — 

ot^ol— 

0C^_ 


12 

5 
5 


Total 9 


7 4 2 


22 


Total 


9 


7 


4 


2 


22 


Total 22 





22 



Qx = [(en - x)(e 22 - x) - (e 21 + x)(e 12 + x)] 2 /S 

> [(en - y)(e 22 - y) - (e 21 + y)(e 12 + y)] 2 /S = Q y 
because [•] > [•] can be reduced to zN(N + 2y) > 0, which is (6) 
greater than since z > 0; where {e n , e 12 , e 21 , e 22 } are the expected 
frequencies, and S = product of all row and column sums. 

Now, if a sign is given to Q corresponding to the direction which the table 
deviates from expected, then Q changes strictly monotonically with the corre- 
sponding deviation of the Fisher- Yates test. Q is more extreme than the value of 
Q for the observed table if: 1) it is greater than the observed Q, given that a 
positive a priori prediction was made; or 2) it is less than the observed Q, given 
that a negative a priori prediction was made. Therefore, for 2x2 tables, by 
assigning a sign to Q values based on an a priori directional prediction the results 
obtained via the multivariate hypergeometric Exact Test are identical to those 
obtained via the Fisher- Yates Exact Test. 

Exact Test Computer Program 

We have designed a program to perform the Exact Test using either the mul- 
tinomial or multivariate hypergeometric distributions on any table of n popula- 
tions and m categories (see the appendix for the actual program). The program 
begins by entering the data table. Next a table of Log factorials is created. Prob- 
abilities are calculated using logarithms so that the numbers do not get too large 
due to the factorials. Creating the factorial table at the beginning of the program 
allows the problem to be solved much more rapidly. The third step is to calculate 
the value of Q (subroutine 400) and the probability of the table (subroutines 200 
and 250 for the multivariate hypergeometric distribution, subroutines 300 and 350 
for the multinomial distribution). Finally, all possible tables must be created. To 
do this, start by initializing the matrix. For the multivariate hypergeometric Exact 
Test this is done by setting all elements k u = for i > 1 and j > 1, set k u = jth 
column sum for j > 1 , set k n = \th row sum for i > 1, and set k,, = 1st column 
sum + \st row sum - total sum. An example is illustrated in Table 4a. All ele- 
ments k u , i > 1 and j > 1, are then systematically changed. When some element 
k u is changed to k u + a, also change k u to k,, - a, change k n to k u - a, and 
change k,, to k u + a. One only needs to test if k n s= to see if the table is a 
legitimate matrix of the Exact Test. For the multinomial Exact Test matrix ini- 



72 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

tialization is done by setting all elements k u = for i > 1, and setting k u = ith 
row sum for i = 1, . . . , n. An example is illustrated in Table 4b. All elements 
k u , i > 1, are then systematically changed. When some element k u is changed to 
k u + a, also change k n to k n - a. One needs only to test if k n ^ to see if the 
table is a legitimate matrix of the Exact Test. For Exact Tests based on either 
distribution, the statistic Q and the probability of observing each legitimate table 
are calculated. If Q is greater than or equal to Q of the observed data matrix the 
probability is summed. A version of this program written in the computer lan- 
guage BASIC for the TRS-80 Level II micro-computer (Radio Shack) is given as 
the Appendix. 

Conclusions 

The generalized Exact Test for n x m contingency tables is applicable to data 
of either nominal or ordinal measure. However, we expect that its primary use 
will be for nominal data where some cells have expected frequencies too small 
for traditional statistical testing of the data without lumping categories and/or 
populations (e.g. Wells and Wells 1980). As table size and row and column sums 
increase so do the number of tables that must be considered for the exact test. 
Thus, only the advent of widespread computer use has allowed this technique 
to be a realistic statistical method for researchers. Even so, very large contingency 
tables cannot be tested via either multinomial or multivariate hypergeometric 
Exact Tests in a reasonable length of time on even the fastest of contemporary 
computers. However, on large tables that have large expected frequencies the 
chi-square distribution gives a good approximation to the Exact Test probability. 

Acknowledgments 

We thank Dr. Dale M. Smith, Dr. Patrick H. Wells, and the reviewers for 
critically reading this manuscript, and for making valuable suggestions for its 
improvement. 

Literature Cited 

Bernard, G. A. 1945. A new test for 2 x 2 tables. Nature, 156:177. 

Cochran, W. G. 1952. The x 2 test of goodness of fit. Ann. Math. Stat., 23:315-345. 

Fisher, R. A. 1934. Statistical Methods for Research Workers. Oliver and Boyd. Edinburgh, Great 

Britain. 
. 1956. Statistical Methods and Scientific Inference. Hafner Publishing Co. New York, New 

York. 
Larntz, K. 1978. Small sample comparisons of exact levels for chi-square goodness of fit statistics. 

J. Am. Stat. Ass., 73:253-263. 
Lehmann, E. L. 1975. Nonparametrics. Holden-Day, Inc. San Francisco, California. 
Pierce, A. 1970. Fundamentals of Nonparametric Statistics. Dickenson Publishing Company, Inc. 

Belmont, California. 
Pearson, K. 1900. On the criterion that a given system of deviations from the probable in the case 

of a correlated system of variables is such that it can be reasonably supposed to have arison 

from random sampling. Phil. Mag., Series 5; 50:157-175. 
. 191 1. On the probability that two independent distributions of frequency are really samples 

from the same population. Biometrika, 8:250-259. 
Siegel, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw-Hill Book Company, 

Inc. New York, New York. 



EXACT TEST FOR N x M CONTINGENCY TABLES 



73 



Wells, H. 1979. Ph.D. Thesis. University of California, Santa Barbara, California. 

, and P. H. Wells. 1980. Are geographic populations equivalent to genetic populations in 

biennial species? A study using Verbascum virgatum (Scrophulariaceae). Genet. Research. 

36:17-28. 
Wilson, E. B. 1941. The controlled experiment and the four-fold table. Science, 93:557-560. 
Yarnold, J. K. 1970. The minimum expectation in the v/ goodness of fit test. J. Am. Stat. Ass., 

65:864-886. 
Yates, F. 1934. Contingency tables involving small numbers and the x 2 test. J. Royal Stat. Soc, 

1:217-235. 

Accepted for publication September 1, 1980. 



Appendix 

A computer program for the generalized Exact Test which is written in the computer language 
BASIC. The program will perform both the multinomial and multivariate hypergeometric variations 
of the Exact Test. The program will run on the Radio Shack TRS-80 Level II computer without 
modification. Odd number lines may be deleted without affecting the program. Lines are skipped to 
isolate sections of the program so that interpretation is easier. 

The program is interactive with the user. Requested are the data matrix and a decision as to the 
distribution on which to base the Exact Test (multinomial or multivariate hypergeometric). The user 
need only respond to the commands as they appear to use the program. P, P4, and P5 (see program 
comments 1-21) are printed for each legitimate table. Ql, Tl, T2, P3, P4, and P5 (see 1-21) are printed 
when the Exact Test is completed. 



I REM EXACT TEST FOR ANY SIZE MATRIX. 

3 REM VARIABLES BEGINNING WITH I,J,K,L,M, AND N ARE INTEGER 

VARIABLES BEGINNING WITH P ARE DOUBLE PRECISION TYPE. 
5 REM WE FIND MOST PROBLEMS DO NOT REQUIRE THAT THE FUNCTIONS 

LOG AND EXP BE DOUBLE PRECISION. 
7 REM RESERVED VARIABLE LIST. 

Tl = # OF TABLES WITH SAME ROW AND COLUMN SUMS. 

T2 = # OF TABLES WITH VALUE OF Q > OR = Ql. 
9 REM Q = "CHI-SQUARE" VALUE OF A TABLE. 

Ql = ••CHI-SQUARE , • VALUE OF THE OBSERVED DATA MATRIX. 

Ml = # OF ROWS IN THE DATA MATRIX = # OF POPULATIONS. 

II REM M2 = # OF COLUMNS IN THE DATA MATRIX = # OF CLASSES. 

N = MATRIX OF DATA WITH DIMENSIONS Ml BY M2. 

Nl = COLUMN MATRIX OF ROW SUMS WITH DIMENSION Ml. 
13 REM N2 = ROW MATRIX OF COLUMN SUMS WITH DIMENSION M2. 

N3 = TOTAL SUM. 

E = TABLE OF EXPECTED VALUES WITH DIMENSIONS Ml BY M2. 
15 REM F = MATRIX OF LOG FACTORIALS, F(J)=LOG(J!). 

P = PROBABILITY OF A GIVEN MATRIX OCCURRING. 

PI = LOG OF VARIABLE PORTION OF PROBABILITY FUNCTION. 
17 REM P2 = LOG OF CONSTANT PORTION OF PROBABILITY FUNCTION. 

P3 = OBSERVED TABLE PROBABILITY. 

P4 = PROBABILITY OF TABLES WITH Q > OR = TO Ql. 
19 REM P5 = TOTAL SUM OF PROBABILITIES = 1.0 AT END OF RUN. 

J AND K ARE RESERVED POINTERS IN MATRIX OPERATION. 

J1,J2,K1,K2, AND K3 ARE LOOP VARIABLES. 



74 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Appendix 
Continued 

21 REM VARIABLES TO BE ENTERED ARE Ml, M2, AND THE MATRIX N. 

23 REM CLEAR SCREEN AND DEFINE VARIABLE TYPES. 

24 CLS 

26 DEFINT I,J,K,L,M,N 

28 DEFDBL P 

29 REM ENTER MATRIX SIZE AND DIMENSION MATRICES. 

30 PRINT "ENTER NUMBER OF POPULATIONS = ROWS" : INPUT Ml 
32 PRINT "ENTER NUMBER OF CLASSES = COLUMNS" : INPUT M2 

34 DIM N(M1,M2), N1(M1), N2(M2), E(M1,M2) 

35 REM INITIATE VARIABLES BY SETTING THEM EQUAL TO ZERO. 

36 T1 = : T2=0 : P4=0 : P5=0 : N3=0 
38 FOR J=l TO Ml : N1(J) = : NEXT J 

40 FOR K= 1 TO M2 : N2(K)=0 : NEXT K 

41 REM ENTER DATA MATRIX AND CALCULATE ROW SUMS, COLUMN SUMS, 

AND THE TOTAL SUM. 

42 FORJ=lTOMl 
44 FORK=lTOM2 

46 PRINT "ENTER ELEMENT"; J; K : INPUT N(J,K) 

48 N1(J)=N1(J)+N(J,K) : N2(K)=N2(K) + N(J,K) : N3 = N3 + N(J,K) 

50 NEXT K 

52 NEXT J 

53 REM CALCULATE EXPECTED FREQUENCIES. 

54 FORJ=lTOMl 
56 FOR K= 1 TO M2 

58 E(J,K)=N1(J)*N2(K)/N3 

60 NEXT K 

62 NEXT J 

63 REM DIMENSIONS MATRIX OF LOG(FACTORIAL) AND CALCULATE ITS 

ELEMENTS. 

64 DIM F(N3) : F(0)=0 

66 FOR J=l TO N3 : F(J) = F(J-l) + LOG(J) : NEXT J 

67 REM DECIDE WHICH DISTRIBUTION TO USE - MULTINOMIAL OR 

MULTIVARIATE HYPERGEOMETRIC. 

68 PRINT "ENTER 1 TO USE THE MULTINOMIAL DISTRIBUTION" 

70 PRINT "ENTER 2 TO USE THE HYPERGEOMETRIC DISTRIBUTION" 

72 INPUT J : IF J= 1 THEN 134 

74 IF J=2 THEN 76 ELSE PRINT "INVALID ENTRY" : GOTO 68 

75 REM STATISTIC WILL BE BASED ON MULTIVARIATE HYPERGEOMETRIC 

DISTRIBUTION. 

76 CLS : PRINT "EXACT TEST USING HYPERGEOMETRIC DISTRIBUTION" 

77 REM CALCULATE Q AND PROBABILITY FOR OBSERVED TABLE. 

78 GOSUB 200 : GOSUB 250 : P3=EXP(P2-P1) : GOSUB 400 : Q1 = Q 

79 REM INITIATE MATRICES FOR THE PROCESS OF CREATING ALL POSSIBLE 

TABLES. 

80 FOR J=2TO Ml : N(J,1) = N1(J) : NEXT J 

82 FOR K=2TO M2 : N(1,K) = N2(K) : NEXT K 

84 FOR J=2 TO Ml : FOR K=2 TO M2 : N(J,K)=0 : NEXT K : NEXT J 

86 N(1,1) = N1(1) + N2(1)-N3 

87 REM INITIATE POINTERS AND BEGIN PROCESS OF TABLE GENERATION. 

88 J = M1 : K=M2 : GOTO 102 



EXACT TEST FOR N x M CONTINGENCY TABLES 75 

Appendix 
Continued 

89 REM FORM A NEW MATRIX BY INCREASING ONE ELEMENT OF THE 

MATRIX BY 1. 

90 N(J,K)=N(J,K)+1 

92 N(1,K) = N(1,K)-1 : N(J,1)=(J,1)-1 

94 N(1,1)=N(1,1)+1 

95 REM TEST MATRIX TO SEE IF IT IS LEGITIMATE, IF IT IS CALCULATE Q 

AND THE PROBABILITY ASSOCIATED WITH IT. 
IFN(J,1)<0THEN 112 
IFN(1,K)<0THEN 112 
J=M1 : K=M2 
IFN(1,1)<0THEN 90 

GOSUB 250 : P=EXP(P2-P1) : P5 = P5 + P : T1=T1 + 1 
GOSUB 400 : IF Q<Q1 THEN 110 
P4=P4+P : T2=T2+1 
PRINT TAB(63), "P = "; P, "P4 = "; P4; TAB(63), "P5 = "; P5 : GOTO 90 

REM POINTERS J AND K NEED TO BE RESET, AND ELEMENTS OF THE 

MATRIX NEED TO BE RESET. 
K=K-1 : If K>1 THEN 116 
J=J-1 ; K=M2 : IF J<2 THEN 178 
FOR Jl=JTO Ml 

IF J1=J THEN K2=K+ 1 ELSE K2=2 
IF K2>M2 THEN 130 
FOR Kl = K2TO M2 

N(J1,1)=N(J1,1) + N(J1,K1) : N(1,K1) = N(1,K1) + N(J1,K1) 
N(1,1)=N(1,1)-N(J1,K1) : N(J1,K1)=0 
NEXT Kl 
NEXT Jl 
GOTO 90 

REM STATISTIC WILL BE BASED ON MULTINOMIAL DISTRIBUTION. 
CLS : PRINT "EXACT TEST USING MULTINOMIAL DISTRIBUTION" 

REM CALCULATE Q AND PROBABILITY FOR OBSERVED TABLE. 
GOSUB 300 : GOSUB 350 : P3 = EXP(P2+P1) : GOSUB 400 : Q1 = Q 

REM INITIATE MATRICES FOR THE PROCESS OF CREATING ALL POSSIBLE 
TABLES 

FOR J=l TO Ml : N(J,1) = N1(J) : NEXT J 

FOR J=l TO Ml : FOR K=2 TO M2 : N(J,K)=0 : NEXT K : NEXT J 

REM INITIATE POINTERS, BEGIN PROCESS OF TABLE GENERATION. 
J = M1 : K=M2 : GOTO 150 

REM FORM A NEW MATRIX BY INCREASING ONE ELEMENT OF THE 

MATRIX BY 1. 
N(J,K) = N(J,K)+1 : N(J,1) = N(J,1)-1 
REM TEST MATRIX TO SEE IF IT IS LEGITIMATE, IF IT IS CALCULATE Q 

AND THE PROBABILITY ASSOCIATED WITH IT. 
IF N(J, 1X0 THEN 158 
J=M1 : K=M2 

GOSUB 350 : P=EXP(P2 + P1) : P5 = P5 + P : T1=T1+1 
GOSUB 400 : IF Q<Q1 THEN 156 
P4=P4+P : T2=T2+1 
PRINT TAB(63), "P = "; P. "P4 = "; P4; TAB(63), "P5 = "; P5 : GOTO 144 

157 REM POINTERS J AND K NEED TO BE RESET, AND ELEMENTS OF THE 
MATRIX NEED TO BE RESET. 



76 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Appendix 
Continued 

158 K = K-1 : IF K>1 THEN 162 

160 J=J-1 : K=M2 : IF J<1 THEN 178 

162 FORJl=JTOMl 

164 IF J1=J THEN K2=K+1 ELSE K2=2 

166 IF K2>M2 THEN 174 

168 FOR Kl = K2TO M2 

170 N(J1,1) = N(J1,1) + N(J1,K1) : N(J1,K1) = 

172 NEXT Kl 

174 NEXT Jl 

176 GOTO 144 

177 REM PRINT FINAL SOLUTION. 

178 CLS 



180 PRINT "Ql = 

182 PRINT "T2 = 

184 PRINT "Tl = 

186 END 



Ql, "P3 = "; P3 
T2, "P4 = "; P4 
Tl, "P5 = "; P5 



199 REM SUBROUTINE TO CALCULATE CONSTANT PORTION OF 

HYPERGEOMETRIC PROBABILITY FUNCTION. 

200 P2=0 

202 FOR J2=l TO Ml : P2=P2 + F(N1(J2)) : NEXT J2 

204 FOR K3=l TO M2 : P2=P2+F(N2(K3)) : NEXT K3 

206 P2=P2-F(N3) 

208 RETURN 

210 END 

249 REM SUBROUTINE TO CALCULATE VARIABLE PORTION OF 

HYPERGEOMETRIC PROBABILITY FUNCTION. 

250 P1=0 

252 FORJ2=lTOMl 

254 FOR K3= 1 TO M2 

256 P1 = P1 + F(N(J2,K3)) 

258 NEXT K3 

260 NEXT J2 

262 RETURN 

264 END 

299 REM SUBROUTINE TO CALCULATE CONSTANT PORTION OF 

MULTINOMIAL PROBABILITY FUNCTION. 

300 P2=0 

302 FOR J2=l TO Ml : P2=P2 + F(N1(J2)) : NEXT J2 

304 RETURN 

306 END 

349 REM SUBROUTINE TO CALCULATE VARIABLE PORTION OF 

MULTINOMIAL PROBABILITY FUNCTION. 

350 P1 = 

352 F0RK3=1T0M2 

354 P=0 

356 FORJ2=lTOMl 

358 P1 = P1-F(N(J2,K3)) : P=P+N(J2,K3) 

360 NEXT J2 

362 Pl = Pl + P*LOG(N2(K3)/N3) 

364 NEXT K3 

366 RETURN 

368 END 



EXACT TEST FOR N x M CONTINGENCY TABLES 




77 


Appendix 






Continued 






399 REM SUBROUTINE TO CALCULATE Q = 


"CHI-SQUARE. 




400 Q=0 






402 FOR J2=l TO Ml 






404 FORK3=lTOM2 







406 Q=Q+(N(J2,K3)-E(J2,K3))|2/E(J2,K3) 

408 NEXT K3 

410 NEXT J2 

412 RETURN 

414 END 



Bull. Southern California Acad. Sci. 
79(2), 1980, pp. 78-81 



Research Notes 

Morphological Variation of the Carinal Plate of the Stalked 
Barnacle Pollicipes polymerus Sowerby 



The intertidal area of Southeast Farallon Island, San Francisco County, Cali- 
fornia (37°42'N, 123°00'W) is riddled with numerous caves at sea level. Although 
the majority of these penetrate less than 8 m, a few reach lengths of over 60 m. 
The biota of these caves differ markedly from that occurring at corresponding 
tidal levels outside. While many organisms are thought to occur in caves because 
of reduced illumination, others appear to occupy caves because of the benefits 
derived from the cave's topography. The decreasing dimensions of a cave with 
length, combined with acute and oblique side channels, deep fractures and wall 
protuberances, tend to intensify and concentrate the waves that enter the cave. 
In these areas of considerable wave action the stalked barnacle Pollicipes polym- 
erus Sowerby (1833) occurs in large aggregations. This barnacle is generally as- 
sociated with, and is considered indicative of high energy intertidal areas along 
the coast of California (Ricketts & Calvin, 1968). 

During a biological survey of Southeast Farallon Island, several aggregations 
of P. polymerus were examined. These aggregations occurred in a moderately 
exposed, dimly illuminated side channel which leaves the main chamber of the 
cave at an acute angle. The side channel also opens seaward, but the position 
and level of the opening is such that neither large scale water movement nor 
illumination is enhanced. Two aggregations, each of approximately 20 typical P. 
polymerus, contained two and three individuals respectively, having an elongate 
carinal plate (Fig. 1). 

P. polymerus has been figured by Darwin (1851), Pilsbry (1907), Newman 
(1975), and others as having a carina that terminates below the apex of the tergum, 
a rostrum which extends above the accessory platelets, triangular accessory 
platelets and peduncular scales (Fig. 2). The membrane that surrounds the ap- 
erture has been described as being yellow to brown or red in color, the latter 
being associated with low illumination. 

In the Southeast Farallon Island variants (Fig. 3) the carinae extend as much 
as one centimeter beyond the apex of the tergum; the rostrum is reduced; the 
terga and scuta show signs of erosion (exfoliation) and both accessory platelets 
and peduncular scales are sub-rectangular in shape. The membrane that surrounds 
the aperture is red. 

Arnold Ross and William A. Newman (Scripps Institute of Oceanography) 
suggested that the elongate carina may only be apparent; the terga being reduced 
due to exfoliation of the laminae. Exfoliation of both terga and scuta is not un- 
common, as can be seen in Figure 3. 

To test the above suggestion we did a morphometric study of four variant and 
59 typical specimens of P. polymerus to determine (1) if the elongate carinae 
result from plate exfoliation or allometric growth, and (2) if the specimens with 
elongate carinae are allometrically different from typical specimens. As a bio- 



RESEARCH NOTES 



79 




Fig. 1. In situ Pollicipes polymerus with elongated carina (arrow). Southeast Farallon Island, 
California. T. Niesen photographer. 




Fig. 2. Typical Pollicipes polymerus showing measurements used for capitular biometric index: 
R-C DIA = rostral-carinal diameter, TH = tergal height. 



80 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 3. Variant Pollicipes polymerus with elongated carinae. Southeast Farallon Island, California. 
Drawing by E. Hamburg. 



metric index we used the relationship between the rostral-carinal (r-c) diameter 
and the tergal height (Fig. 2); specimens were measured to the nearest 0.01 mm 
with vernier calipers. 

The scatter diagram of tergal height on r-c diameter for the 59 typical specimens 
indicates a linear relationship for the two variables (Fig. 4). An analysis of co- 
variance showed no significant difference in the relationship of r-c diameter to 
tergal height between the variant and typical specimens. Thus the elongated ca- 
rinae of the variant specimens are not a result of tergal exfoliation. 

Fitted regression lines were then calculated for the four specimens with elon- 
gated carinae and the 59 typical specimens. Although the variants fall within the 
cluster of points defining the typical specimens, they do not appear to correspond 
to the fitted regression line for the typical specimens (Fig. 4). A Student's t-test 
of regression coefficients shows that the variant specimens are significantly dif- 
ferent (P < 0.01) from the typical specimens. Therefore, the capitular growth of 
the variants is allometrically different from the typical barnacles. Allometric 
growth of these barnacles may result from competition between individuals in the 
aggregation. All four variant barnacles were found in the center of their respective 
aggregations and were surrounded by typical specimens. 



RESEARCH NOTES 



81 



20- 










18- 








/. - 


16- 

s 

14- 

►- 

z 

2 12- 
in 

Z 




• 


• 
• 


• • 

• 


< 10 " 
O 

at 
ui 


...-%" 


* , y 






*- 8- 


• 


/ 




• TYPICAL 


6- 




/ 




OVA RIANT 




1 1 


1 1 1 1 


1 


1 1 1 1 



12 14 16 18 20 22 24 26 28 30 
ROSTRAL- CARINAL DIAMETER MM 



32 34 



Fig. 4. Scatter diagram and fitted regression lines of tergal height on rostral-carinal diameter for 
typical and variant specimens of Pollicipes poly merits. Correlation coefficient for typical specimens = 
.8479; for variants = .9929. 



Acknowledgments 

We thank William A. Newman and Arnold Ross (Scripps Institute of Ocean- 
ography) for their helpful suggestions and critical reading of the manuscript. 
Voucher specimens are deposited in the Department of Invertebrate Zoology, 
California Academy of Sciences (CASIZ Cat. No.: 004702). 

Literature Cited 

Darwin, C. 1851. A monograph of the sub-class Cirripedia. The Lepadidae. Ray Society, London, 

400 pp. 
Newman, W. A. 1975. Phylum Arthropoda: Crustacea, Cirripedia. Pp. 259-269 in Light's Manual: 

Intertidal Invertebrates of the central California coast, 3rd ed. (R. I. Smith and J. T. Carlton 

eds.), Univ. California Press, xviii + 716 pp. 
Pilsbry, H. A. 1907. The barnacles (Cirripedia) contained in the U.S. National Museum. Bull. U.S. 

Nat. Mus., 93:1-366. 
Ricketts, E. F., and J. Calvin. 1968. Between Pacific Tides, 4th ed. (Revised by J. W. Hedgpeth). 

Stanford Univ. Press, xiv + 614 pp. 

Accepted for publication July 7, 1980. 

Chet Chaffee and David R. Lindberg, Department of Invertebrate Zoology, Cal- 
ifornia Academy of Sciences, San Francisco, California 941 18. Present address: 
Center for Coastal Marine Studies, University of California, Santa Cruz, Cali- 
fornia 95064. 



Bull. Southern California Acad. Sci. 
79<2), 1980, pp. 82-86 



Observations of Feral Populations of Xenopus laevis 
(Pipidae) in Southern California 



Xenopus laevis, a clawed frog native to Africa, is known to colonize areas 
outside its natural range. Extralimital populations have been reported in the 
United Kingdom, Ascension Island, and the United States (Wyoming, Wisconsin, 
Arizona, New Mexico, Utah, California, Florida, Nevada, and Colorado). 

The characteristics that may have aided in the establishment of temporary or 
permanent populations outside of Africa appear to be: (1) distribution for scien- 
tific, medical, and pet uses; (2) ability to reproduce in a broad variety of conditions 
including temporary bodies of water, man-made ponds, and irrigation/drainage 
channels; (3) ability to avoid adverse conditions by short overland migrations, 
aestivation, and habitat modifications; (4) high reproductive potential; and (5) 
opportunistic feeding on a broad variety of food sources. 

The precise date of introduction in California in unknown. The first official 
report of feral frogs in California is 1968 (St. Amant and Hoover 1969). However, 
it is likely that Xenopus existed in California waters before this time. Clawed 
frogs were widely used in the 1930' s and 1940' s for human pregnancy diagnosis 
and subsequently for medical research. A few years later, clawed frogs became 
popular as aquarium pets and were readily available to the general public through 
pet stores. The presence of Xenopus in Orange County near metropolitan Los 
Angeles before 1968 undoubtedly stems from accidental or intentional release of 
captive frogs by humans. Subsequent reports of feral clawed frogs in Los Angeles, 
San Diego, Yolo, and Riverside counties generated an often sensationalized in- 
terest within California that may have prompted further human dispersal of 
clawed frogs. 

The San Diego Natural History Museum was often contacted by people seeking 
information on the frogs and the areas where they could be collected. However, 
few studies were conducted to gain insight into the potential impact of this exotic 
on the established fauna (see Munsey 1972; Zacuto 1975; Fritts and McCoid 
1976). 

From summer 1974 through spring 1977, we conducted a series of studies on 
the feeding habits, growth, and reproduction of Xenopus in California. During 
this period we made a number of incidental observations on population densities, 
dispersal, and aestivation. Because such data are important to understanding feral 
clawed frogs and to any eradication, control, or management of the species, the 
observations are summarized in this report. 

Generally, when X. laevis was encountered in San Diego and Riverside coun- 
ties, it was found in large numbers in restricted areas such as ponds and pools. 
In contrast to the broad distribution in the Sweetwater River drainage described 
on the basis of metamorphosing young by Mahrdt and Knefler (1973), we found 
clawed frogs to be conspicuous in only a few sites. We investigated several of 
the sites noted by Mahrdt and Knefler, but often found clawed frogs to be absent 
or present only as transients. The two sites we studied most intensively were: 
Site A — a pond at the junction of Avocado and Valencia streets, Spring Valley, 



RESEARCH NOTES 83 

San Diego County, California; and Site B — a pond 1.6 km southwest of Vail 
Lake, near Temecula, Riverside County, California. Site A was about 17 x 42 
m with a maximum depth of 1.5 m. Site B was about 6 x 18 m with depths of 
10-70 cm. Using a Peterson mark-recapture method, we computed population 
estimates of 602 and 494 adult frogs for Sites A and B respectively. Additionally, 
a far larger body of water in the flood plain of the Tijuana River in San Ysidro, 
San Diego County, California, was determined to have a large population of 
clawed frogs, but it was not studied. 

That each of these major populations was in a different river system and not 
adjacent to similar aggregations suggests that human-aided dispersal is more im- 
portant to colonization of clawed frogs in California than is natural dispersal of 
the frogs. The metropolitan San Diego populations possibly stem from a pet dealer 
who raised clawed frogs in outdoor tanks adjacent to the Sweetwater River in 
the mid-1960's (Douglas Ruth, pers. com.). The population at Site B was first 
reported to us in January 1975, shortly after the opening of a recreational vehicle 
park which encompasses Site B. Most of the campers and visitors to the camp- 
ground and park come from the metropolitan Los Angeles area where established 
populations of clawed frogs have existed for several years. Campers and visitors 
may have brought pets which were introduced into the waters in the campground. 

Within a stream system, clawed frogs are capable of dispersing over short 
distances (Mahrdt and Knefler 1973), perhaps in response to high population 
density and flooding conditions. After metamorphosis (generally in May but quite 
variable), large numbers of juveniles move downstream out of the source pond. 
We observed the colonization of a shallow pond 0.8 km downstream from Site 
B in May 1975 by 200-300 juveniles. Only a few juveniles were found in the 
intervening stream habitat despite our active search. Subsequently, we collected 
downstream from the second pond in a marsh area adjacent to Vail Lake but only 
one juvenile clawed frog was found on 4 May 1975. The majority of the frogs 
apparently stopped at the first available habitat. No adults were found in the 
downstream pond until several months after colonization. 

Juvenile clawed frogs also used sheet flooding situations to disperse. In San 
Diego County near Site A, young frogs moved across fields and school play- 
grounds with runoff after a heavy rain. Only recently metamorphosed frogs were 
observed in actively running water. All sites occupied by Xenopus that we ex- 
amined were noticeably disturbed habitats (i.e., directly altered by man in being 
artificially impounded, channelized, or excavated). 

It is commonly thought that adult Xenopus are unpalatable to predatory fishes 
(James St. Amant, pers. com.). In our experience, X. laevis is conspicuously 
absent from waters with predatory fishes (centrarachids) in Riverside and San 
Diego counties. We failed to find clawed frogs in much of the Sweetwater River, 
or in the San Diego and Santa Margarita rivers. Frogs were also absent from a 
number of large ponds seasonally continuous with the Sweetwater River and from 
the margins of Vail Lake. Because of the continuity of the drainage and proximity 
to established populations of frogs, Xenopus is presumed to have had opportu- 
nities to colonize these areas occupied by fish species. 

It is possible that Xenopus can be eliminated from a body of water by contin- 
uous predatory pressure on the larvae or smallest young. We have some circum- 
stantial evidence to suggest that black crappie (Pomoxis nigromaculatus) and 



84 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

green sunfish (Lepomis cyanellus) were responsible for the extirpation of all 
Xenopus larvae from the spring reproduction of 1976 at Site B. Calling male 
frogs, amplexing pairs, egg masses, and young larvae, between the stages of 5 to 
47, were observed during April and May 1976. However, no larvae older than 
stage 47, which coincides with the initiation of midwater schooling of clawed frog 
larvae, were found despite continued searches. Black crappie and green sunfish 
were present throughout the 1976 reproductive period. Clawed frogs did repro- 
duce successfully in the same area in 1975, but to our knowledge no predatory 
fishes were present at that time. 

In California (like Africa) X. laevis was subjected to occasional drought. During 
late summer and early fall, water levels dropped significantly at Site A. In 1974 
and 1975 this further concentrated an already dense population and caused a 
marked behavioral shift in the frogs. Surface water temperatures exceeded 30°C 
and as the pond reached depths of 10-15 cm, bottom temperatures approximated 
those at the surface. During this period, clawed frogs were concentrated in the 
deepest water. 

Upon examination of the actual areas occupied, we found the pond substrate 
had been excavated by the frogs, thereby increasing the vertical water column. 
Presumably in response to rising water temperatures, the frogs excavated pits in 
the soft mud. The pits were 30-40 cm deep and 10-45 cm wide. Each pit was 
occupied by one to two frogs but adjacent pits sometimes were connected in a 
reticulated pattern. The temperature at the bottom of the pits was much lower 
than nonexcavated areas and fluctuated around 20°C. Frogs often surfaced to 
breathe, but at other times remained in the pits. 

During 1974 the pond did not completely dry, but if it had, the pits would have 
concentrated the available water for the frogs, delaying the need for aestivation 
or other evasive behavior. Under extreme drought conditions the pits would 
presumably be the site of further excavations for aestivation chambers (Balinsky 
et al. 1967). 

To test this hypothesis, 11 frogs were removed from Site A on 10 September 
1974, transported to the laboratory and placed in one-gallon jars (3.78 1) filled 
with mud from the substrate of Site A. The jars were covered with wire mesh 
and the contents allowed to dry slowly. The frogs immediately burrowed into the 
mud, constructing a small space at the bottom of the jar with a vertical tube 
opening to the surface. Over a 3 month period, all 11 frogs were sequentially 
removed, examined, and preserved. In each instance, the frog was found in ver- 
tical position with the head oriented upwards. The frog occupied a chamber not 
greatly exceeding the dimensions of the body. The chamber had a vertical con- 
nection with the surface of the now hardened soil, which presumably served as 
a source of air for the aestivating frog. During the confinement in jars, no food or 
water was added and only two animals died before they were removed from the 
experiment. It should be noted that the frogs were not dormant during their 
confinement and occasionally were seen with the tip of the snout projecting from 
the vertical opening. On 13 December 1974, the last frog gained 31.6 gm in weight, 
presumably due to water uptake. 

Perhaps the greatest concern about the impact of X. laevis in California in- 
volves its feeding habits. The frogs have been characterized as voracious, non- 
specific predators of vertebrates (St. Amant et al. 1973). However, only indirect 



RESEARCH NOTES 85 

evidence exists to support this belief. There have been only two feeding studies 
on California populations (Lenaker 1972; McCoid and Fritts 1980). The latter 
study refuted the conclusions of the former by demonstrating that Lenaker' s 
method of collecting frogs inadvertently produced a bias in the stomach contents 
of the frogs. McCoid and Fritts concluded that Xenopus preyed almost exclu- 
sively on slow-moving invertebrates in southern California, a conclusion sup- 
ported by laboratory studies (Avila and Frye 1978). Mahrdt and Knefler (1972) 
cited the absence of presumed vertebrate prey species in the areas occupied by 
Xenopus, implying the possibility of local extinctions due to predation by Xen- 
opus. However, these authors gave no consideration to other factors that might 
affect the distribution of the prey species (i.e., habitat quality or seasonal vari- 
ation in distribution). 

During our investigations at Site A, Gambusia affinus, which was widely in- 
troduced in California for mosquito control, became extinct during a period of 
low water and high temperatures in 1974. There is little doubt that clawed frogs 
consumed some of the Gambusia as the water volume decreased. Xenopus is 
capable of capturing active vertebrates in confined situations (McCoid and Fritts 
1980). An unknown percentage of the Gambusia at Site A died possibly as a 
result of high water temperatures in those shallow areas not occupied by clawed 
frogs. On the basis of our observations, the Gambusia were faced with two 
alternatives — to remain in hot shallow water or invade the small pits occupied by 
clawed frogs. The confinement of pits constructed by clawed frogs presented an 
opportunity for the frogs to exploit a food source otherwise unavailable. There- 
fore, the extinction of Gambusia was a result of unusual conditions associated 
with drought and the presence of clawed frogs. Whether Gambusia would have 
become extinct in the absence of Xenopus cannot be determined. 

In view of our studies and a review of other data, we see the impact of X. 
laevis in a different light than other investigators. Our analysis of stomach con- 
tents (McCoid and Fritts 1980), distributions, and other aspects of the biology of 
feral clawed frogs, suggests that X. laevis is a nuisance species, but not a major 
threat to the established fauna (introduced or native) of California. In San Diego 
and Riverside counties, the frogs occupy marginal habitats and prey infrequently 
on aquatic vertebrates. The fact that reproduction of Xenopus is not always 
successful suggests that populations can be controlled and, perhaps with time, 
eradicated by game fishes. Certainly the failure of high concentrations of rotenone 
to remove a Los Angeles County population should not be used as testimony to 
the durability of an exotic frog that lacks gills and breathes air at the surface of 
the water (see St. Amant 1975; Zacuto 1975). 

Because the major factor in dispersal of clawed frogs in California appears to 
be human transport, populations will likely continue to appear and spread. An 
alternative way to curb the spread of Xenopus in California is through education 
of the general public on the implications of exotic introductions. 

Xenopus laevis is used widely in research laboratories and classroom instruc- 
tion (Deuchar 1975). As an animal capable of reproducing and forming dense 
populations under proper conditions, a potential exists for the clawed frog to be 
reared for commercial distribution to research institutions that currently depend 
upon imported Xenopus and native Rana sp. harvested from natural habitats. A 
commercial biological supply company in Wisconsin is currently rearing Xenopus 



86 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

in their laboratory. On the basis of the success of feral populations in California, 
it may be possible in special situations to harvest feral clawed frogs without 
detrimental environmental effects. 

Acknowledgments 

We thank the Riverside County Fish and Game Commission for partial support 
of field work at Site B. We are indebted to D. Ruth, C. Crumly, and H. Snell for 
assistance with field work. R. Matheson, J. Sites, Jr., and J. Dixon provided 
editorial comments on an early draft of this paper. 

Literature Cited 

Avila, V., and P. Frye. 1978. Feeding behavior of the African clawed frog {Xenopus laevis Daudin): 

(Amphibia, Anura, Pipidae): Effect of prey type. J. Herpetol., 12:391-396. 
Balinsky, J., E. Choritz, C. Coe, and G. Van der Schans. 1967. Amino acid metabolism and urea 

synthesis in naturally aestivating Xenopus laevis. Comp. Biochem. Physiol., 22:59-68. 
Deuchar, E. 1975. Xenopus: The South African clawed frog. John Wiley and Sons, London. 246 pp. 
Fritts, T., and M. McCoid. 1976. Colonization and biology of the African clawed frog {Xenopus 

laevis) in southern Riverside County, unpub. rep. Riverside Co. Fish Game Comm. 28 pp. 
Lenaker, R. 1972. Xenopus in Orange County, unpub. rep. California State Poly. Univ. 12 pp. 
Mahrdt, C, and F. Knefler. 1972. Pet or pest? Environ. Southwest, 446:2-5. 

. 1973. The clawed frog — again. Environ. Southwest, 450:1-3. 

McCoid, M., and T. Fritts. 1980. Notes on the diet of a feral population of Xenopus laevis (Pipidae) 

in California. Southwest. Nat., 25:272-275. 
Munsey, L. 1972. Salinity tolerance of the African pipid frog, Xenopus laevis. Copeia, 1972:584- 

586. 
St. Amant, J. 1975. Exotic visitor becomes permanent resident. Terra, 13(4):22-23. 
, and F. Hoover. 1969. Addition of Misgurnus anguillicaudatus (Cantor) to the Californian 

fauna. California Fish Game, 55:330-331. 
, , and G. Stewart. 1973. African clawed frog, Xenopus laevis (Daudin), established in 

California. California Fish Game, 59:151-153. 
Zacuto, B. 1975. The status of the African clawed frog (Xenopus laevis) in Agua Dulce and Soledad 

Canyons, unpub. rep. California Dept. Fish Game. 36 pp. 

Accepted for publication July 15, 1980. 

Michael J. McCoid, Department of Wildlife and Fisheries Sciences, Texas A&M 
University, College Station, Texas 77843, and Thomas H. Fritts, U.S. Fish and 
Wildlife Service, National Fish and Wildlife Laboratory, Tulane University Mu- 
seum of Natural History , Belle Chasse, Louisiana 70037. 



Bull. Southern California Acad. Sci. 
79(2), 1980, pp. 87-88 



Extension of the Range of Holothuria zacae, 
Deichmann 1937 



In October 1970, 20 unusual holothurians were found off Ship Rock, Santa 
Catalina Island, California (33°27'48"N; 118°29'26"W) on a shelly debris bottom, 
at a depth of 30 meters. One specimen was sent to the Smithsonian Institution 
and it was identified as Holothuria zacae (Deichmann 1937), a species previously 
known to range from Cedros and Guadalupe Islands off Baja California south to 
the Galapagos Islands, and into the Gulf of California. 

The Panamic region of the west coast of North America was explored exten- 
sively for shallow water holothurians during the Hancock Expeditions of 1932- 
1939 and 1944-1954 (Deichmann 1958). A search of these reports and other avail- 
able literature and collections leads us to believe that the Ship Rock population 
is the only one known from north of Guadalupe Island. The locality data from 
the Hancock Expeditions suggest a rather disjunct distribution of//, zacae, hence 
the occurrence of this species so far north of its known range may be unusual. 

Additionally, Pawson (pers. comm.) has suggested that since the genus is 
known to have a pelagic larva, this is probably an isolated population at the 
northern extreme of the range and that it has merely escaped detection until 
recently. 

The physical characteristics of the Ship Rock forms comply with the "forma 



nHonaHM 


9»w • 








JBI 


f*~ 






•■*^I*#"* '» 


_mr 




















fr 4 ■****"* ' f^H 




"' ' ^rttttPk 




K- *"" ■ 


cfff k * V~***s 










4*HhI 








-^1 


'Wffi 






?-.'- 


■+ -- w ^ 




K§5£ 




~fc 


" *■ 




BW^^*'^'- ' 










- 






** VKB 










" '•'■" • 


















" ' jm — ■- 






" .^' 




•f-T" 








"^^^ 






' • M " ^ 










* 4r!£ 




• v ir 



Holothuria zacae, adult. 90' of water, base of Ship Rock. Santa Catalina Island. 



88 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

iota" of H. zacae typical of the Baja California area (Pawson, pers. comm.). A 
representative individual (Fig. 1) was 40 cm long and 18 cm in circumference. Its 
basic coloration is a creamy background with two widely separated rows of dark 
brown papillae, each with a green tip. The rest of the dorsal surface is covered 
with closely spaced, lighter brown, warty projections. The numerous podia are 
green and distributed abundantly along the ventral surface. When disturbed, only 
a slight contraction of the body wall is exhibited and there is no evidence of 
evisceration. 

Another large holothurian that occurs at Catalina, Parastichopus parvimensis 
(Clark 1913), is generally smaller, caramel-brown in color, has fewer dorsal pa- 
pillae and fewer ventral podia than H. zacae. P. parvimensis has a great capacity 
for contraction and readily eviscerates when handled roughly. 

More recent observations (March-May 1978) of the Ship Rock H. zacae pop- 
ulation revealed only five individuals found exclusively on a shelly debris, around 
rocks, at depths of 25 to 35 meters. Some cursory investigations have been made 
for other populations in the southern California bight without success. 

We would like to thank D. L. Pawson of the Smithsonian Institution for his 
assistance in identifying our material and for searching the available literature. 
Further, we thank the Institute for Marine and Coastal Studies of the University 
of Southern California for logistical support. 

Literature Cited 

Clark, H. L. 1913. Echinoderms from Lower California, with Descriptions of New Species. Bull. 
Amer. Mus. Nat. Hist., 32:234-235. 

Deichmann, E. 1937. The Templeton Crocker Expedition IX. Holothurians from the Gulf of Cali- 
fornia, the west coast of lower California and Clarion Island. Zoologica (N.Y.), 22:161-176. 

. 1958. The holothurioidea collected by the Velero III and IV during the years 1932 and 1954. 

Part II, Aspidochirota. Allan Hancock Pacific Exped., 1 1(2):307— 3 10. 

Accepted for publication July 10, 1980. 

Nancy E. Muleady and Robert R. Given, Catalina Marine Science Center, P.O. 
Box 398, Avalon, California 90704. 

Contribution number 39 of the Catalina Marine Science Center. 



INSTRUCTIONS FOR AUTHORS 

The BULLETIN is published three times each year (April, August, and November) and includes articles in English 
n any field of science with an emphasis on the southern California area. Manuscripts submitted for publication 
should contain results of original research, embrace sound principles of scientific investigation, and present data 
n a clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide 
or contributors. Consult also recent issues of the BULLETIN. Authors should strive for directness and lucidity, 
ichieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous 
-eference of pronouns, and logically placed modifiers. 

MANUSCRIPT PREPARATION 

It is strongly recommended that, before submitting a paper, the author ask qualified persons to review it. The 
luthor is requested to submit at least two additional copies with the original, on 8Vi x 11 opaque, nonerasable 
Daper, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. 
Footnotes should be avoided. Manuscripts which do not conform to the style of the BULLETIN will be returned to 
he author. 

An abstract summarizing in concise terms the methods, findings, and implications discussed in the paper must 
iccompany & feature article. Abstract should not exceed 100 words. 

A feature article comprises approximately five to thirty typewritten pages. Papers should usually be divided into 
he following sections: abstract, introduction, methods, results, discussion and conclusions, acknowledgments, and 
iterature cited. Avoid using more than two levels of subheadings. 

A research note is usually one to six typewritten pages and rarely utilizes subheadings. Consult a recent issue 
of the BULLETIN for the format of notes. Abstracts are not used for notes. 

Abbreviations: Use of abbreviations and symbols can be determined by inspection of a recent issue of the 
BULLETIN. Omit periods after standard abbreviations: 1.2 mm, 2 km, 30 cm, but Figs. 1-2. Use numerals before 
anits of measurements: 5 ml, but nine spines (10 or numbers above, such as 13 spines). The metric system of 
weights and measurements should be used wherever possible. 

Taxonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International 
Zode of Botanical Nomenclature (Lawjouw et al. 1956), the International Code of Nomenclature of Bacteria and 
Viruses (Buchanan et al. 1958), and the International Code of Zoological Nomenclature (Stoll et al. 1961). Special 
ittention should be given to the description of new taxa, designation of holotype, etc. Reference to new taxa in 
itles and abstract should be avoided. 

The literature cited: Entries for books and articles should take these forms. 

McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. 

Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54:452-458. 

Brattstrom, B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), 
Univ. California Press, xii + 635 pp. 

Tables and figures (line drawings, graphs, or black and white photographs) should not repeat data contained in 
(he text. The author must provide numbers and short legends for tables and figures and place reference to each 
jf them in the text. Legends should be typed on a separate sheet of paper and placed at the end of the manuscript. 
Illustrations and lettering thereon should be of sufficient size and clarity to permit reduction to standard page size; 
ordinarily they should be no more than twice the size of intended reduction and should not exceed SVi by 1 1 inches 
in size. Photographs must be printed on glossy paper. Submit one photoduplicated copy of each illustration. All 
Illustrations accompanying Research Notes will be reduced to one column width. All half-tone illustrations will have 
light screen (grey) backgrounds. Special handling such as dropout half-tones, special screens, etc., must be requested 
by and will be charged to authors. 

A cover illustration pertaining to an article in the issue or one of general scientific interest will be printed on the 
:over of each issue. Such illustrations along with a brief caption should be sent to the Editor for review. 

PROCEDURE 

All manuscripts should be submitted to the Editor, Robert J. Lavenberg, Los Angeles County Natural History 
Museum, 900 Exposition Blvd., Los Angeles, California 90007. Evaluation of a paper submitted to the BULLETIN 
oegins with a critical reading by the Editor; several referees also check the paper for scientific content, originality, 
ind clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent 
':o the author at which time he or she may be requested to rework portions of the paper considering these rec- 
ommendations. The paper then is resubmitted and may be re-evaluated before final acceptance. 

Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He or she 
should promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations, legends, 
ind bibliographical references. He or she marks corrections on the galley (copy editing and proof procedures in 
Style Manual) and promptly returns both galley and manuscript to the Editor. Manuscripts and original illustrations 
will not be returned unless requested at this time. All changes in galley proof attributable to the author (misspellings, 
inconsistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with 
[he printer, not the Editor. 



CONTENTS 

A Synopsis of the Larvae of Costa Rican Frogs and Toads. By Jay M. 

Savage 45 

Ecology of a Coastal Salt Marsh after Long-Term Absence of Tidal Fluctu- 
ation. By H. Peter Eilers 55 

A General "Exact Test" for N x M Contingency Tables. By Harrington 

Wells and Jack Lester King 65 

Research Notes 

Morphological Variation of the Carinal Plate of the Stalked Barnacle Pollicipes polymerus 

Sowerby. By Chet Chaffee and David R. Lindberg 78 

Observations of Feral Populations of Xenopus laevis (Pipidae) in Southern California. By 

Michael J. McCoid and Thomas H. Fritts 82 

Extension of the Range of Holothuria zacae, Deichmann 1937. By Nancy E. Muleady and 

Robert R. Given 87 



COVER: Aerial oblique view of Inner Bolsa Bay, California, June 15, 1979. Pacific Ocean at right; 
oil field operated by Signal, Inc. at upper left. Diked area at center is part of Bolsa Chica Ecological 
Reserve to which tidal fluctuation was partially restored in 1978. 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Volume 79 



ULLETIN 



MAY 2 6 1981 



t-w YOKK 



Number 3 




BCAS-A79(3) 89-136 (1980) 



DECEMBER 1980 



Southern California Academy of Sciences 

Founded 6 November 1891, incorporated 17 May 1907 



OFFICERS 

Takashi Hoshizaki, President 

Fred G. Hochberg, Vice President 

Richard E. Pieper, Secretary 

Joseph E. Haring, Treasurer 

Robert J. Lavenberg, Editor 

Gretchen Sibley, Assistant Editor 



1978-1980 

Kristin H. Berry 

Robert A. Georges 

Joseph E. Haring 

Jerry D. Kudenov 

Donald J. Reish 



BOARD OF DIRECTORS 

1979-1981 

John Baird 

Jules Crane 
Fred G. Hochberg 
Richard E. Pieper 
Gloria Takahashi 



1980-1982 

Takashi Hoshizaki 
Alan J. Mearns 

Steven N. Murray 
Camm C. Swift 

Robert G. Zahary 



Membership is open to scholars in the fields of natural and social sciences, and to any person 
interested in the advancement of science. Dues for membership, changes of address, and requests for 
missing numbers lost in shipment should be addressed to: Southern California Academy of Sciences, 
the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 90007. 

Annual Members $ 12.00 

Life Members 150.00 

Fellows: Elected by the Board of Directors for meritorious services. 



The Bulletin is published three times each year by the Academy. Manuscripts for publication should 
be sent to the appropriate editor as explained in "Instructions for Authors" on the inside back cover 
of each number. All other communications should be addressed to the Southern California Academy 
of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, Los 
Angeles, California 90007. 



Date of this issue 7 May 1981 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 89-96 

Notes on a Collection of Centrolenid Frogs from the 
Colombian Choco 

Marc P. Hayes and Priscilla H. Starrett 

Abstract. — Notes on a collection of centrolenid frogs from the Colombian Cho- 
co by Marc P. Hayes and Priscilla H. Starrett, Bull. Southern California Acad. 
Sci., 79(3):89-96, 1980. A collection of 30 Chocoan centrolenids comprising 7 
species is reported. Centrolenella albomaculata, C. chirripoi, C. euknemos, C. 
fleischmanni, and C. ilex are first reports from Colombia. C. chirripoi is reported 
for the first time outside the type locality in Costa Rica. Prepollical spines found 
in C. spinosa are shown to lack sexually dimorphic character. A distinctive pro- 
tuberance of the third metacarpal in centrolenids is advanced as an apparently 
unique familial character. 

Department of Biological Sciences and Allan Hancock Foundation, University 
of Southern California, Los Angeles, California 90007. 



In 1968 and 1971, Philip A. Silverstone visited the Colombian Choco in the 
course of work on dendrobatid frogs. Silverstone (1973) gives a thorough descrip- 
tion of the 1971 study site. During these visits, a small, but significant collection 
of centrolenid frogs was made. This assemblage of 31 frogs was subsequently 
deposited in the herpetological collection of the Los Angeles County Museum of 
Natural History (LACM). 

It is the purpose of this paper to report on this collection, as several of the 
species are new to Colombia, and further, to comment on significant characters, 
augmenting existing knowledge of selected species and the family Centrolenidae. 
A single specimen is omitted from the following accounts, as it is being reserved 
for possible description as a new species in a separate account. Localities of 
individual specimens are summarized in the appendix following the discussion. 

Centrolenella albomaculata Taylor 

Specimens.— LACM 72909. 

Remarks. — This is the only specimen referable to this species and the first 
report from Colombia. We were struck by the superficial resemblance of this 
specimen to the Amazonian lowland slope form C. medemi (see Lynch and Duell- 
man 1973). This specimen suggested a new species, were it not for some large- 
spotted variants in the Costa Rican collections at the University of Southern 
California. Specimens of C. albomaculata show an increasing size progression 
of spots from head to vent, the largest spots located on the upper surface of the 
thigh. Further, Costa Rican individuals examined exhibit considerable variation 
in spot size among individuals. However, thigh spots of this Colombian example 
attain maximum diameters of 2.5-3.0 mm, a condition found only in the Costa 
Rican specimens with the largest spots. In this respect, the Colombian specimen 
represents the extreme of pattern variants examined. 



90 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Centrolenella chirripoi (Taylor) 

Specimens.— h ACM 72929. 

Remarks. — C. chirripoi was reviewed by Starrett and Savage (1973), who re- 
garded it as a valid species. The only collections of this species are the holotype 
and paratype series from Rio Cocolis, Provincia de Limon, Costa Rica (Taylor 
1958). This single collection represents the first reported collection outside the 
type locality. We take this opportunity to emphasize the characters which sep- 
arate C. chirripoi from its closest congeners, since centrolenids of the fleisch- 
manni group (sensu Savage 1967) are notoriously difficult to separate in a pre- 
served state. Starrett and Savage (1973) listed 8 characters, all of which are not 
equally important in characterizing this species. The characters most important 
are: 1) the distinctive webbing between fingers II and III, 2) the indentation 
between the nostrils when the head is viewed from above, 3) the only slightly 
protuberant eyes when viewed from above, 4) the slightly rounded nature of the 
snout when viewed in profile, 5) the long loreal region and 6) the weak canthus 
rostralis. The webbing character, perhaps the most distinctive, is superficially 
shared with C. pulverata. Figure 1 demonstrates the distinctive webbing in both 
species. The remarkable feature of this character is that it exhibits very little 
variation in all specimens examined. Starrett and Savage (1973) also noted that 
the color of the parietal pericardium was not known. Examination of the guanine 
distribution on this specimen and one of the paratype series (KU 36868) suggested 
that C. chirripoi has a transparent parietal pericardium. This was confirmed by 
Silverstone's field notes, taken on the specimen in life, which stated (quote):"bare 
heart." In this character and the indentation of the snout, C. chirripoi allies itself 
with C. colymbiphyllum. 

Centrolenella euknemos Savage and Starrett 

Specimens.— LACM 47066. 

Remarks. — Savage and Starrett (1967) report this species from Costa Rica and 
Panama. This specimen represents the first published record from Colombia and 
extends the known range of the species over 400 km to the south and east. 

Centrolenella fleischmanni (Boettger) 

Specimens.— LACM 47067-9, 47071^, 72930. 

Remarks. — This is the most wide ranging of centrolenid species (Lynch and 
Duellman 1973), although some (Goin 1964; Starrett and Savage 1973) suggested 
that several cryptic species may be involved. Duellman (1973) reports the range 
as from Guerrero and Veracruz, Mexico to Surinam and Ecuador. Occurrence 
in Colombia is based on a statement of Goin (1964), where he postulates its 
presence based on geographic encirclement by documented localities. These col- 
lections, therefore, represent the first definitive records from Colombia. 

Centrolenella ilex Savage 

Specimens.— LACM 47063, 47070, 72910, 72914. 

Remarks. — Starrett and Savage (1973) reported specimens from Costa Rica and 
Panama. Charles W. Myers (pers. comm.) recently informed us that the unlisted 
specimens of C. ilex reported by Lynch and Duellman (1973) are not referable 
to this species. These collections, therefore, represent the first definitive records 



CENTROLENID FROGS FROM COLOMBIA 



Ml 




Fig. 1. A) Volar aspect of left manus of Centrolenella chirripoi. Drawn from specimen KU 36863. 
B) Volar aspect of left manus of Centrolenella pulverata. Drawn from specimen CRE 7155 (USC 
collections). Scale mark = 2 mm. 



from Colombia. Radiographs of the males in this series confirm a previously 
unverified suspicion that the males of this species have distinctive humeral hooks 
(Fig. 2). Myers informs us that this is also the case in Panamanian C. ilex. The 
humeral hook in C. ilex is long, thin, lying parallel to the humerus and terminating 
in a sharp point (Fig. 3). It lies well-concealed by the arm musculature. This is 
in sharp contrast to the rather spatulate, flairing humeral hooks found in C. 
prosoblepon. Females have at most a small humeral boss, but no hook. We have 
also examined a specimen from Jaime Villa that was taken in the drainage of the 
Rio Indio, Departamento Zelaya, Nicaragua, extending the known range north 
to this site. 

Centrolenella prosoblepon (Boettger) 

Specimens.— LACM 47065, 72923, 72925-6. 

Remarks. — The three male specimens (LACM 72923,72925-6) lack the dorsal 
and tibial spots characteristic of most Costa Rican C. prosoblepon. Lynch and 
Duellman (1973) comment on the apparent continuum of variation between spot- 
ted and unspotted dorsal patterns which forced them to synonymize C. para- 
bambae with C. prosoblepon. Furthermore, snout shape in these specimens is 
the extreme of variants observed in Central American C. prosoblepon, the canthal 
platform being less pronounced. This variability in pigmentation, snout shape and 
our own observations on the shape and degree of flairing in humeral hooks did 
not permit separation of these specimens as a new species. As with C. fleisch- 
manni, cryptic species may be involved, but this is not determinable at this time. 
Lynch and Duellman (1973) cite specimens referable to this species from Depar- 
tamento Cauca, Colombia, but these collections are the first reported from De- 
partamento del Choco. 



92 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. Radiograph of a male Centrolenella ilex (LACM 72910), venter up. Dark arrow denotes 
metacarpal bulge and light arrow points to humeral hook. Specimen length = 29.5 mm. 

Centrolenella spinosa Taylor 

Specimens.— LACM 47064, 72911, 72913, 72917-8, 72920-2, 72924, 72927-8. 

Remarks. — Taylor (1949) described this species from an all male series and 
speculated on the presence of reduced prepollical spines in females. In a later 
paper (Taylor 1951), he notes that the spine is entirely concealed and restates 
that females have a less well developed spine, although he cites no new speci- 
mens. Savage (1967) mentions that C. spinosa males have both a free prepollex 
and prepollical spine and adds that females have a prepollex while implying they 
lack a prepollical spine. This Colombian series of C. spinosa includes 11 adults, 
3 females and 8 males. Radiographs of both this series and Costa Rican C. spinosa 




Fig. 3. Left humerus, ventral aspect of a male Centrolenella ilex (LACM 72910). Drawn from a 
radiograph. Scale mark = 2 mm. 



CENTROLENID FROGS FROM COLOMBIA 93 




Fig. 4. Radiograph of a female Centrolenella spinosa (LACM 72920), ventral aspect. Dark arrow 
denotes metacarpal bulge and light arrow points to the prepollical spine. Specimen length = 21.0 mm. 

show that both males and females have a prepollex and prepollical spine (Figs. 
4-5). Furthermore, the spine found in females is not significantly smaller than 
that found in males. Starrett and Savage (1973) report localities for this species 
from Costa Rica and Panama. Their addition that specimens examined from Co- 
lombia and Ecuador are referable to this species refers in part to the series being 
reported here. We concur with their determinations and confirm this series as the 
first definitive records of C. spinosa from Colombia. 

Discussion 

The species reported herein raise the total number of known Colombian cen- 
trolenids west of the Andean crest to 13, the others being Centrolene geckoideum, 
Centrolenella antioquiensis , C. buckleyi, C. grandisonae , C. grifftthsi and C. 
johnelsi (Cochran and Goin 1970; Duellman and Lynch 1973). There is no reason 
to believe that the wet lowland tropical forest species, C. pulverata and C. val- 
erioi, found in Panama (Starrett and Savage 1973), will not eventually be found 
in the Chocoan lowlands of Colombia. Similar predictions can be made for the 
mid-elevation Pacific slope species, C. peristicta and C. ocellifera, reported by 
Lynch and Duellman (1973) from Ecuador on biogeographic grounds. These 
species are expected in similar habitats on the Pacific slope of the Colombian 
Andes. 



94 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 5. Hard elements of left manus and forearm of a female Centrolenella spinosa (LACM 72920), 
ventral aspect. Drawn from a radiograph. Cartilaginous intercalated phalangeal elements are omitted. 
Scale mark = 2 mm. 



Taylor (1951) defined the family Centrolenidae. His familial definition is based 
primarily on the nearly complete fusion of the astragalus and calcaneus, a fusion 
pattern shared only by the genus Pelodytes (Pelobatidae). Other characters used 
by Taylor are the presence of terminal T-shaped (or Y-shaped) phalanges, present 
in certain hyperoliids, leptodactylids, microhylids, rhacophorids and possibly bu- 
fonids (Trueb 1973; Liem 1970) and the digital intercalated cartilage, a character 
shared by the Hylidae, the Hyperoliidae and the African microhylid Phrynome- 
rus. To this group, we add a character we believe will be useful in defining the 
family Centrolenidae. This is the presence of a distinct protuberance on the medial 
side of the third metacarpal (Figs. 2, 4 or 5). This protuberance was noted in all 
radiographed centrolenid material to date (both in Centrolene and 26 species of 
Centrolenella). This bulge, which varies in position from between two-thirds to 
one-third the distance from the distal end of the metacarpal, varies in size among 
species, protruding from Vi to 2 mm from its shaft. It was illustrated, but not 
discussed, in Eaton's anatomical study of Centrolenella prosoblepon (see Eaton 
1958:466). Presumably, this feature was regarded as anomalous, as limited com- 
parative material was then available. Radiographs of available dendrobatid, hylid, 



CENTROLENID FROGS FROM COLOMBIA 95 

leptodactylid, pelobatid and ranid material show that, at least in the material 
examined, the metacarpal protuberance is absent. Liem (1970) discusses a bony 
knob on the third metacarpal of some rhacophorids and hyperoliids. His illustra- 
tion (see Liem 1970:39) clearly shows this structure an extended expansion of the 
dorsolateral tip of the metacarpal, analogous to, but differing from the protuber- 
ance of centrolenids located on the midlateral metacarpal. Notably, the bony 
knob in rhacophorids and hyperoliids is associated with a slip of the humerodor- 
salis muscle in what Liem suggests as being the most derived condition of this 
muscle. In this condition, the muscle has two metacarpal slips inserting on the 
third and fourth digits. Liem suggests this condition is associated with arboreal 
life, noting the modification probably both increases extension efficiency of these 
digits and enables independent extension. A muscle dissection of Centrolenella 
prosoblepon shows the humerodorsalis with insertions with patterns paralleling 
those found in Liem's derived rhacophorids and hyperoliids. Further, the slip on 
metacarpal three apparently inserts on the protuberance previously described. 
Realizing the very arboreal nature of all members of the family Centrolenidae, 
we suggest that the condition of the humerodorsalis found in C. prosoblepon is 
an independently acquired adaptation to an arboreal life mode. The presence of 
the metacarpal protuberance suggests that the pattern of insertion of the humer- 
odorsalis may be a character which is uniform throughout the family. We reserve 
judgment, however, upon examination of the balance of the family. We believe 
careful examination of anuran families, particularily those with arboreal members, 
will support the apparent unique state of the metacarpal protuberance found in 
the Centrolenidae. 

Acknowledgments 

We thank John W. Wright and Robert L. Bezy (LACM) for the loan of the 
Silverstone collection. Further, we would like to thank William E. Duellman for 
the loan of the paratype series of C. chirripoi and for allowing us to open one of 
the paratypes to examine guanine distribution. Thanks are due Charles W. Myers, 
whose comments on C. ilex and generous loan of an unpublished manuscript 
benefited this paper. We thank Jaime Villa for allowing us to examine and report 
on the Nicaraguan C. ilex. We also thank Jay M. Savage for allowing us to dissect 
a specimen of Centrolenella prosoblepon, and Roy W. McDiarmid, Craig Guyer 
and Michael M. Miyamoto for taking of their time to review and enhance the 
manuscript. Special thanks are due Philip A. Silverstone, who took the time to 
go over the details of his field notes with us and clarify our questions. Finally, 
we dedicate this paper to the late Charles F. Walker, who long ago pointed out 
to one of us (Starrett) that available centrolenid radiographs had an interesting 
little protuberance on the third metacarpal. 

Appendix of Specimens Examined 

Large series of Costa Rican material from the collections at USC were used as 
comparative material. This material is not listed here since Starrett and Savage 
(1973) give a thorough listing. All material is listed by country first and then 
province, both in alphabetical order. 

Centrolenella albonuiculata. — COLOMBIA: Choco: Camino de Yupe in the 
drainage of the Rio Opogado, 420 m, LACM 72909. 



96 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Centrolenella chirripoi. — COLOMBIA: Choco: upper Rio Opogado between 
Rio Merendo and Rio Yupe, 75 m, LACM 72929. COSTA RICA: Limon: Rio 
Cocolis, near Suretka, KU 36862-4, 36866-70. 

Centrolenella euknemos. — COLOMBIA: Antioquia: Rio Arquia at Finca Los 
Llanos (above Vegaez), 100-200 m, LACM 47066. 

Centrolenella fleischmanni. — COLOMBIA: Antioquia: Rio Arquia, Finca 5 km 
w of Finca Chibiqui (40 km wsw of Urrao), 350-400 m, LACM 47067-9. Choco: 
upper Rio Opogado above Rios Yoto and Angostura, 60 m, LACM 47071^; 
upper Rio Opogado, 75-120 m, LACM 72930. 

Centrolenella ilex. — COLOMBIA: Antioquia: Belen, Rio Arquia upstream 
from Vegaez, 100 m, LACM 47063. Choco: trail between upper Rio Napipi (near 
its juncture with Rio Merendo) and upper Rio Opogado at 60-130 m, LACM 
47070; Camino de Yupe, 420 m, LACM 72910, 72914. NICARAGUA: Departa- 
mento Zelaya: Rio Indio, 75 m. 

Centrolenella prosoblepon. — COLOMBIA: Antioquia: Belen, Rio Arquia up- 
stream from Puerto Palacios (also above Vegaez), 100 m, LACM 47065. Choco: 
Camino de Yupe, 420 m, LACM 72923, 72925-6. 

Centrolenella spinosa. — COLOMBIA: Antioquia: Rio Arquia at Belen (up- 
stream from Vegaez), 100 m, LACM 47064. Choco: Camino de Yupe, 420 m, 
LACM 72911, 72913, 72917-8, 72920-2, 72924, 72927-8. 

Literature Cited 

Cochran, D. M., and C.J. Goin. 1970. Frogs of Colombia. U.S. Nat. Mus. Bull., pp. 288:xii + 654 pp. 
Duellman, W. E. 1973. Liste der rezenten Amphibien und Reptilien: Hylidae, Centrolenidae, Pseu- 

didae. Das Tierreich, 95:xix + 225 pp. 
Eaton, T. H., Jr. 1958. An anatomical study of a neotropical tree frog, Centrolene prosoblepon 

(Salientia: Centrolenidae). Univ. Kansas Sci. Bull., 39(10):459-472. 
Goin, C. J. 1964. Distribution and synonymy of Centrolenella fleischmanni in northern South Amer- 
ica. Herpetol., 20:1-8. 
Liem, S. S. 1970. The morphology, systematics and evolution of the Old World treefrogs (Rhaco- 

phoridae and Hyperoliidae). Fieldiana: Zool., 57:vii + 145 pp. 
Lynch, J. D., and W. E. Duellman. 1973. A review of the centrolenid frogs of Ecuador, with 

descriptions of new species. Occ. Pap. Mus. Nat. Hist. Univ. Kansas, 16:1-66. 
Savage, J. M. 1967. A new tree-frog (Centrolenidae) from Costa Rica. Copeia, 1967(2):325-331. 
. and P. H. Starrett. 1967. A new fringe-limbed treefrog (Family Centrolenidae) from lower 

Central America. Copeia, 1967(3): 604-609. 
Silverstone, P. A. 1973. Observations on the behavior and ecology of a Colombian poison-arrow 

frog, the Kokoe-pa (Dendrobates histrionicus Berthold). Herpetol., 29(4):295-301. 
Starrett. P. H., and J. M. Savage. 1973. The systematic status and distribution of Costa Rican glass- 
frogs, genus Centrolenella (Family Centrolenidae), with description of a new species. Bull. 

So. Cal. Acad. Sci., 72(2):57-78. 
Taylor, E. H. 1949. Costa Rican frogs of the genera Centrolene and Centrolenella. Univ. Kansas 

Sci. Bull.. 33(l):257-270. 
. 1951. Two new genera and a new family of tropical American frogs. Proc. Biol. Soc. Wash., 

64:33^10. 
. 1958. Notes on Costa Rican Centrolenidae with descriptions of new forms. Univ. Kansas 

Sci. Bull., 39(2):41-68. 
Trueb, L. 1973. Bones, Frogs, and Evolution. In Vial, J. L. ed. Evolutionary Biology of the Anurans: 

Contemporary Research on Major Problems. Columbia, Missouri: Univ. Missouri Press. 

pp. 65-132. 

Accepted for publication April 17, 1980. 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 97-108 

Five Insects Believed to be Newly Established or Recolonized 

on Santa Cruz Island, California 

(Dermaptera, Lepidoptera) 

Jerry A. Powell 

Abstract. — Five insects believed to be newly established or recolonized on 
Santa Cruz Island, California (Dermaptera, Lepidoptera) by Jerry A. Powell, 
Bull. Southern California Acad. Sci., 79(3):97-108, 1980. An earwig, 2 moths, and 
2 butterflies not recorded in 1939^1 and 1966-69 surveys of S.C.I, insects, be- 
came established during 1969-78. Evidence suggests that 3 were introduced by 
man. Sudden appearance of the butterflies, however, is not easily interpreted. 
Either might have immigrated often during 50-100 years S.C.I, has had their 
weedy hostplants. It is hypothesized that such species periodically colonize, are 
eliminated during stress such as overgrazing by feral sheep in drought years, then 
recolonize. An undersaturated nature of the insect fauna of offshore islands, 
especially badly perturbed ones, is proposed. Extinction presumably is higher 
than natural, immigration low, and/or colonization improbable owing to reduced 
patch sizes of native hostplants. 

University of California, 201 Wellman Hall, Berkeley, California 94720. 



Santa Cruz Island (S.C.I.) is the largest of the northern tier of California's 
Channel Islands and is the most diverse geographically and faunistically. Lying 
30 km off the coast of the mainland, about 40 km south of Santa Barbara, the 
island is approximately 34 km long, ranging from 3 to 10 km wide, with an area 
of ca. 249 km 2 . 

There is no record of the native insect fauna prior to extensive disturbance of 
the natural flora by feral sheep and pigs. Sheep and hogs may have been intro- 
duced in the early 1800s. Cattle were grazed on the island as early as 1865 (Hil- 
linger 1958), and according to Holder (1910) the Caire ranch was managing more 
than 30,000 head of sheep around the turn of the century. During the early era 
of insect exploration in California (1860-1930) inaccessibility of the Channel Is- 
lands and ownership visitation policies discouraged collecting visits. Sporadic 
survey by entomologists and sporadic collections by other biologists were made 
during the late 1800s and after the turn of the century, but most of these were on 
Santa Catalina and San Clemente Islands (see Miller and Menke 1980, for review 
of entomological investigations). No comprehensive systematic list of insects was 
produced for S.C.I, from these early visits. Even on the one major, organized 
Channel Islands survey, that of the Los Angeles County Museum in 1939^4 1 , 
Santa Cruz Island was relatively neglected, considering its size, with only a one- 
week visit in August (with 3 entomologists), and a 5 day visit in March, mainly 
at Pelican Bay by one entomologist, with some collecting at other sites by two 
persons (Comstock 1939, 1946). Therefore extensive overgrazing had taken place 
for 80 years or more before any appreciable insect survey occurred. 

A cooperative effort, beginning in 1940, between the Stanton family, owners 



98 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

of the island, and the Department of Biological Control, University of California, 
Riverside, to control prickly pear cactus, preceded the establishment of the U.C. 
Santa Cruz Island Reserve and field station on the Stanton property. According 
to Goeden et al. (1967), Mr. Stanton estimated that approximately 40% of 
the rangeland of the island had been rendered useless for grazing by dense growth 
of native prickly prears. After 24 years of efforts involving releases of several 
native southwestern Nearctic species of Hemiptera, Homopteraand Lepidoptera, 
a considerable degree of control was effected, primarily by cochineal scales, 
Dactylopius (Goeden et al. 1967). 

Presence of the field station, beginning in 1966, opened the door for much more 
diverse and intensive entomological survey work, so that a picture of the island's 
insect fauna has begun to emerge during the past 15 years. In general the insect 
fauna has been found to be depauperate relative to that of the mainland (Powell 
1967; Miller 1971; Opler 1974; Weissman and Rentz 1976). For example, only 33 
species of butterflies have been discovered in rather intensive search during all 
months of the year (Langston 1980; C. L. Remington pers. coram.; Cal- 
ifornia Insect Survey unpubl. data), while more than twice that number occur in 
a comparable area of adjacent mainland around Santa Barbara (Emmel and Em- 
mel 1973; S. Miller in litt.). The fauna contains only a few endemic species 
(Alexander 1973; Gordon 1976; Powell 1967; Opler 1971, 1977; Rentz and Weiss- 
man 1973). Probably those mostly are relics of past extensive mainland distri- 
butions, as is believed to be true of endemic plants (Thorne 1969). However, 
Rentz and Weissman (1973) give evidence for island speciation in one genus of 
sand-burrowing crickets. 

The establishment of a U.S. Naval operation on the island in the 1940s — and 
later the field station — has caused a considerable increase in the traffic of per- 
sonnel, foodstuffs, and equipment in recent decades. As a result, the chance of 
introduction and establishment of mainland species has increased. Five species 
appear to have become established on the island during the past 15 years, based 
on survey work carried out in 1966-1969 compared with more recent collections. 

No doubt the insects of Santa Cruz Island are incompletely surveyed, but 
comparison of collections from different institutions indicates that there has been 
sufficient seasonal coverage to enable reasonable confidence in statements about 
easily obtained species. Additions to the faunal list are expected in taxa where 
the insects are small or require specialized collection techniques (e.g., Berlese 
samples of litter dwellers, rearing of leaf miners, etc.), but the appearance of 
previously unrecorded, yet easily observed species can be taken as evidence of 
recent establishment or expansion of populations on the island. 

During 1966 and 1969, personnel of the California Insect Survey, University of 
California, Berkeley, made three general collecting trips to the island. The first, 
25 April to 2 May, involved 6 staff and graduate students, who devoted approx- 
imately 36 Full Time Equivalent (FTE) collector days. The second, 7-10 June 
1966, was made by two participants, an equivalent of 8 FTE days. The third trip 
took place 14-17 March 1969, with 5 experienced collectors, about 15 FTE days. 
All three visits were blessed with good weather and adequate transportation fa- 
cilities so that diverse habitats were surveyed, from Christi Beach near the west- 
ern end to the vicinity of Chinese Harbor in the northeastern portion of the island. 
Subsequent collecting has shown that many species were overlooked, especially 



SANTA CRUZ ISLAND INSECTS 99 

those active as adults only in late summer, fall, and winter. However, five species 
encountered during the 1970s are insects that likely would have been observed 
during our 1960s trips had they been present, as documented below. 

DERMAPTERA 

Forficula auricularia (Linnaeus) 

Although the European earwig was not discovered in California before 1923, 
it rapidly spread throughout the northern half of the state at low to moderate 
elevations during the 1930s. The earliest record in southern California was 1931, 
but the species was not considered established there until 1940 or later (Langston 
and Powell 1975). It was collected around Santa Maria during 1945-1947 and at 
Santa Barbara as early as 1952, but we were unable to document its established 
occurrence in the 1960s for most of southern California (Langston and Powell 
1975). Only one record was known from the Channel Islands at that time, a 
collection in 1972 on San Clemente Island. 

Langston and I made special efforts to look for earwigs on both of our 1966 
trips to Santa Cruz Island, as this was during the data-gathering years for the 
C.I.S. bulletin. Searches were made around the ranch buildings, the vegetable 
garden, and other likely sites, but we found no Dermaptera. 

In September 1978, M. E. Buegler and I found adults and immatures of For- 
ficula common around the field station. Therefore it appears that the European 
earwig was introduced to S.C.I, during the 1970s. Similarly, this earwig has re- 
cently turned up on Santa Rosa Island, the earliest specimen having been col- 
lected in 1971 (Langston and Miller 1977). That it was not introduced earlier to 
either island probably can be attributed to the fact that F. auricularia is better 
adapted to more mesic, northern habitats in California than arid ones. As a result 
this insect is not as abundant in southern California as is the ring-legged earwig, 
Euborellia annulipes (Gerstaecker), and probably it was not established in main- 
land areas adjacent to the islands before the 1950s or later. 

Euborellia annulipes, a wingless species, has been widely established in south- 
ern California since the 1880s and is known from several of the Channel Islands, 
but still has not been discovered on S.C.I. (Langston and Powell 1975; Langston 
and Miller 1977). Although earwig colonies may be overlooked owing to seasonal 
dormancy or other causes of restricted occurrence, the flightless E. annulipes is 
more likely to remain localized and therefore undiscovered than is Forficula. 

LEPIDOPTERA 

Pieris rapae (Linnaeus) 

The cabbage white butterfly occurs abundantly throughout California, except 
at the highest elevations and in the deserts, and is particularly common in coastal 
urban and agricultural situations. It is an Old World insect that was introduced 
into eastern North America about 1860 and is believed to have spread rapidly 
across the country (Holland 1898; Wright 1906). There is a possibility that it was 
introduced separately into California. In any event, P. rapae has been established 
in southern California coastal areas since before the turn of the century, the first 
specimen having been taken in 1883 (Wright 1906). Thus it was surprising when 
we did not find this species on Santa Cruz Island in 1966. Each trip included two 



100 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

lepidopterists, and other collectors were alerted to take samples of all butterflies 
seen. Moreover, P. rapae flies near the coast in southern California in every 
month of the year (Emmel and Emmel 1973). Thus, it is inconceivable that we 
could have overlooked the cabbage white even at a low population density. 

Known from Santa Barbara as early as 1916 (Coolidge 1923), it seems unlikely 
that this butterfly has never immigrated to S.C.I. , yet there appear to be no 
records prior to 1969. There are no specimens in the Los Angeles County Museum 
of Natural History from the 1939-41 faunal survey (S. Miller in lift.), and none 
could be located in the U.C. Riverside collection, which has some general insect 
survey material from the fieldwork associated with the biological control of prick- 
ly pear cactus on the island. Of course it is possible that P. rapae is such a weedy 
and ubiquitous insect that no voucher specimens were retained by early collectors 
who may have observed it. 

Subsequently Pieris rapae has become common in weedy habitats along the 
main water courses in the Central Valley and Prisoners Harbor Creek. Specimens 
were collected in early August 1969, by D. B. Weissman; C. L. Remington (in 
lift.) encountered the species in the early 1970s; and we found it numerous in 
September 1978. If the species had been unable to cross the channel on its own 
for nearly 90 years following its establishment in California, it is possible that it 
was inadvertently introduced, perhaps as larvae on garden nursery stock, during 
the late 1960s. Why this event did not occur during a much earlier ranching era 
is a mystery. 

Pyrgus communis albescens Ploetz 

The western checkered skipper is a homodynamic insect that occurs throughout 
southern California, especially in weedy areas where Malva is adventive. Thus 
it should not have been surprising when we found this species at scattered places 
on S.C.I. — Canada Cervada near the western end, Willow Cove on the south 
shore, and Prisoners Harbor on the north shore — during 26-29 September 1978. 
Those collections, however, evidently were the first records of P. communis on 
the island. 

We did not encounter this species during our 1966-69 trips, when 26 species 
of butterflies were taken; D. B. Weissman did not collect it among 16 species 
taken in early August 1969; and G. A. Gorelick failed to record it among 22 
species logged in June 1978 (Langston 1980 and in lift.). In addition, S. E. Miller 
(in litt.) did not find S.C.I, specimens in the LACM from the 1939-41 Channel 
Islands Biological survey. 

Spring and summer collections cannot be ruled out as seasonally asynchronous 
unless a markedly differing voltinism has been fixed genetically in the island 
population, limiting it to a fall flight. On the mainland, P. communis has been 
recorded from every month and commonly flies from February to October (Em- 
mel and Emmel 1973). Individuals are often seen by mid-March even in the San 
Francisco Bay area. 

Moreover, data from other Channel Islands suggest a restricted flight period is 
unlikely. Pyrgus communis has been known on Santa Catalina Island at least 
since 1932 (LACM) and has been taken during the winter (Meadows 1936). There 
are specimens from Santa Rosa Island taken in 1939 and 1941 according to Miller 
(in litt.) who encountered this species on his earliest visits to other northern 



SANTA CRUZ ISLAND INSECTS 101 

Channel Islands, on tiny, remote Santa Barbara I. in June 1978, and on East 
Anacapa in August 1978. Therefore failure of P. communis to appear in S.C.I. 
collections prior to September 1978 is perplexing. The widespread distribution on 
the island at that time suggests either that populations expand in numbers and in 
range as the season progresses, or that the species had quite recently (since June 
1978) reached the island, rapidly populating various parts of it. Neither hypothesis 
seems plausible, considering the overall distribution and voltinism of this insect. 

Laetilia coccidivora (Comstock) 

This pyralid moth was described from Washington, D.C. and is widespread 
through the southern and southwestern states, including southern California 
(Heinrich 1956). The larvae feed on various Coccidae, especially cochineal scales 
on cactus, and occasionally on the flowers of prickly pear. 

Cochineal insects, species of Dactylopius, were introduced to Santa Cruz Is- 
land from the mainland between 1940 and 1951 for control of prickly pear cactus 
(Goeden et al. 1967). Successful introductions apparently occurred in 1951 with 
colonies from Hawaii, via Riverside, of cochineal insects descended from a series 
of introductions from Mexico to Australia beginning in 1927. Distributions of 
Dactylopius -infested Opuntia pads from the ranch headquarters area of the orig- 
inal introduction were carried out in 1955-1960, and observations from 1961 to 
1966 indicated that Dactylopius had spread throughout the island and had effected 
substantial control of the cactus (Goeden et al. 1967). Goeden and his coworkers 
attributed the success of this biological control agent in part to the apparent 
exclusion of two of its natural predators, a coccinellid beetle (Hyperaspis) and 
Laetilia coccidivora. The latter was encountered at all mainland stations where 
Dactylopius was surveyed, but it was absent from their collections on Santa Cruz 
Island, although Goeden et al. admit the inadvertent introduction of these pred- 
ators during the course of the biological control work could not be ruled out. 

Laetilia coccidivora was first collected on Santa Cruz Island in October 1972, 
when C. L. Remington took a series of adults at lights at the U.C. field station. 
He collected another series in August 1974 at the same station, and I took them 
in blacklight traps at all sites sampled in late September 1978. Larvae collected 
by D. S. Green in June 1977 produced adults in mid-July. One Dactylopius- 
infested Opuntia pad I collected in late September 1978, at Willows Creek, 
produced adult Laetilia in October and November and again the following May 
and June (JAP 78J7). Heinrich (1956) cites collection dates throughout the year 
in Texas and Arizona and for March, October and November in southern Cali- 
fornia. The life cycle does not seem to be carefully documented, but the records 
suggest either a bivoltine pattern or continuous generations are possible in warmer 
areas. 

In addition to our negative evidence from blacklight trapping at the field station 
in May and June 1966, Remington sampled a diversity of microlepidoptera at the 
station in late July 1967 and 1968, and mid- August 1968, but did not find L. 
coccidivora. Presuming that population numbers build up during the season and 
that adults appear at lights only in exceptionally warm weather, our spring and 
June work could be ruled out as adequate for discovery of this species. However, 
the moth sampling by Remington in July and August 1967-1968, should have 
revealed Laetilia, had it become as abundant in the area as collections during 
1972-1978 indicated. 



102 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

It appears that Laetilia coccidivora was introduced to Santa Cruz Island during 
the biological control program in 1951-1961, or by some other means subsequent- 
ly. Probably the moth did not build up in population numbers or was localized 
on the island in areas remote from the vicinity of the ranch headquarters and field 
station, until the early 1970s. 

The deleterious effect on Dactylopius as a biological control agent of prickly 
pear cactus by appreciable levels of this predaceous moth have yet to be assessed. 
My rearings produced 15 adult Laetilia and one Chalcidae, presumably a para- 
sitoid of Laetilia, from a single Opuntia stem. 

Platynota stultana Walsingham 

The "omnivorous leaf roller" was described in 1884 from Sonora, Mexico, and 
its native range probably included adjacent parts of the southwestern U.S., as it 
was collected in Cochise County, Arizona in the 1890s by F. H. Snow (specimens, 
AMNH). As the common name implies, a vast array of larval hosts has been 
recorded (e.g., Atkins et al. 1957), most of which are in nursery and agricultural 
situations, whence the insect has expanded its geographical range during the past 
80 years. Platynota stultana seems not to be native in cismontane southern Cal- 
ifornia because it was not encountered by Coquillett and others in the early years 
of citrus investigations, yet it had become a pest of citrus by 1913 (Woglum 1920). 
The earliest record in California I have seen is at La Mirada, Los Angeles County, 
where it was reared from tomato in 1898 (specimen, NMNH). By 1913-1915, P. 
stultana had become an economic problem in various citrus and cut flower grow- 
ing areas of southern California (Woglum 1920; Bohart 1942). 

The species reached coastal Santa Barbara County by 1940, although the date 
of initial establishment and continuous residency there is unknown. There is one 
record for Carpinteria (40 km NW of S.C.I.) (NMNH), and P. stultana was reared 
from larvae collected by Henne on Anacapa Island during the LACM Channel 
Islands Biological Survey in 1940. About 1960 this species apparently underwent 
a change in physiological tolerance and during the following several years greatly 
expanded its geographic and ecological range in California (Powell 1981). 

Adults of Platynota stultana readily come to lights, so presence of a population 
is easily detected. I collected the moths at Goleta (45 air km N of S.C.I.) in June 
1965 and on both Santa Catalina and San Clemente Islands on my first visits to 
them, in April 1968 and March 1972, yet no collections were made on Santa 
Cruz Island prior to 1975. The adults fly virtually throughout the year in southern 
California, so that seasonal exclusion cannot be the reason. 

The first record for S.C.I, was a female taken at UV light at the field station 
in late July 1975, by C. L. Remington and R. Priestaf. We found the adults 
common at lights around the station, and one was taken in a blacklight trap at 
Prisoners Harbor in late September 1978. Therefore it seems probable that Pla- 
tynota stultana was introduced in the early 1970s, possibly via cut flowers or 
potted nursery plants, and is established on the island. 

This species has been reared from Eriogonum grande, a native endemic plant, 
at Isthmus Station on Santa Catalina Island, by G. A. Gorelick, so it will be 
interesting to monitor its invasion of the native flora on Santa Cruz. There are no 
undoubted records of the occurrence of P. stultana in native plant communities 
on mainland California, except in desert areas (Goeden and Ricker 1976a). The 
same authors (1976b, and earlier references) encountered the omnivorous leaf- 



SANTA CRUZ ISLAND INSECTS 103 

roller on several native ragweed species (Ambrosia) in cismontane southern Cal- 
ifornia, but it appears that most or all these records are from roadsides, aban- 
doned cropland and similar artificial expansions of Ambrosia habitat. It was not 
discovered on Ambrosia chamissonis in native situations by Goeden and Ricker 
or by me on Santa Cruz Island or at many other coastal sand dune localities. 

Discussion 

Probability of introduction by man. — There is reasonable evidence to indicate 
the recent introduction by man of 3 of the species recorded here, Forficula au- 
ricularia, Laetilia coccidivora, and Platynota stultana. The sudden, recent ap- 
pearance of the remaining two butterflies, however, is not easily interpreted. No 
data are available to explain the long period during which they apparently failed 
to colonize. 

Pieris rapae could have been imported on cabbage or other garden crucifers 
during the late 1960s, but an immigration via this route much earlier, when pro- 
duction and transport of agricultural products were less controlled, seems more 
plausible. Pyrgus communis is an unlikely candidate for accidental transport by 
man. Both species seem to be sufficiently strong fliers to have made the trip on 
their own many times during the 80-100 years their weedy hostplants surely must 
have been available on the island. Both have been resident for more than 50 years 
on Santa Catalina, a comparable sized island situated the same distance off the 
southern California coast (Meadows 1936). 

Depauperate nature of the fauna: — One of the most fascinating aspects of the 
Channel Islands fauna is its depauperate nature. While reasonable hypotheses 
can be proposed to explain absence of many species that occur on the adjacent 
mainland, others seem missing by chance. In the best documented insect group, 
the butterflies, Miller (/'// litt.) has recorded about 70 species in an area the size 
of S.C.I, in the Santa Ynez Mountains-Santa Barbara coastal shelf, but of these, 
only 33 have been discovered on S.C.I. , only 40 km away (Langston 1980; present 
data). Among the absentees, some can be theorized as either lacking from the 
original community at the time of separation from the mainland or subsequently 
eliminated and too sedentary to have immigrated (e.g., the lycaenids Euphilotes 
battoides and Philotes sonorensis, which have been demonstrated to possess low 
vagility rates, (Arnold 1980; Keller et al. 1966). Others might be missing because 
their restricted larval food preferences are for plants not found or too limited on 
the island to maintain insect populations (e.g., Colias eurydice on Amorpha cal- 
if ornica, Habrodais grunus on Quercus chrysolepis). Overgrazing by feral sheep 
in dry years might account for the paucity of grass feeding Lepidoptera, explain- 
ing the absence of 6 or 7 Santa Barbara Hesperiinae and the Satyrid Coeno- 
nympha California, which occurs in weedy areas throughout most of California. 

On the other hand, non-residents include several widespread, apparently vagile 
butterflies whose larval hosts are common on S.C.I. For example, Papilio rutulus, 
one of California's largest butterflies, and Limenitis lorquini, seem imponderably 
absent from suitable appearing willow-lined creek habitats; Phyciodes mylitta, a 
homodynamic species which feeds on weedy and native Asteraceae, occurs abun- 
dantly in disturbed situations at low elevations throughout the state north of Santa 
Barbara but not on S.C.I. ; Incisalia iroides is polyphagous on native plants, both 
monocot and dicot, and would seem to be as likely a colonist as another lycaenid. 



104 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

the resident Celastrina argiolus, which has similar larval food preferences. Four 
Eriogonum (Polygonaceae) specialists, the lycaenids Lycaena gorgon, Euphilotes 
battoides, E. enoptes, and Apodemia mormo, do not occur on S.C.I. , yet all live 
along the coast of the mainland, often in sympatry. Among Cercocarpus (Rosa- 
ceae) feeding Lepidoptera, two moths, Acleris folianus (Tortricidae) and Ethmia 
discostrigella (Oecophoridae) are common on S.C.I. , but a butterfly, Strymon 
tetra (Lycaenidae) is absent. All 3 are sympatric in scattered parts of California, 
such as at San Diego. Many such examples could be listed. 

Compliance to island biogeographic theory. — Orthoptera on the Channel Is- 
lands conform fairly well with the species numbers/area aspects of island biogeo- 
graphic theory (Weissman and Rentz 1976), and species numbers of butterflies 
are remarkably similar (S. Miller in lift.). Preliminary sampling of S.C.I. , San 
Nicolas, Santa Catalina, and San Clemente islands indicates that diversity for 
Lepidoptera as a whole will follow this pattern. Thus it is tempting to suggest 
that the Channel Islands possess dynamic equilibrium levels in insect species, 
that an extinction curve-immigration curve intersect has been reached. According 
to this hypothesis, the low species numbers compared to the mainland are a 
function of island area and ecological diversity. Theoretically, aspects of com- 
munity complexity, including competitive displacement, interact to maintain the 
insect fauna near its present diversity. In birds, area by itself is a relatively poor 
predictor of numbers of species on the Channel Islands compared to ecological 
parameters such as numbers of native plant species (Power 1972). For insects, 
a list of plant species provides a crude index to potential niche diversity in larval 
foods in some phytophagous groups but not necessarily in Orthoptera, many of 
which are scavengers or general plant feeders. The insular fauna of Orthoptera 
may be at or near equilibrium, with numbers of species related to ecological 
diversity, expressed by complexity of vegetation types, which is roughly corre- 
lated with area. By contrast, in Lepidoptera and other phytophagous insects 
having many species that depend upon particular plants, extinction and coloni- 
zation rates presumably are much more sensitive to disturbance, particularly 
overgrazing and invasion of weeds. Such perturbance favors hemodynamic, es- 
pecially polyphagous species, which are often alien, and these become dominant 
in population numbers per unit area. Native, host specific species are selected 
against by elimination or restriction of host plants to small habitat patches. As 
a result, although insect species numbers may be generally correlated with area 
and environmental complexity, in most taxa they are likely to be low compared 
to the original and potential equilibrium levels. 

The best documented studies of extinctions and turnover rates of insular ani- 
mals have concentrated on birds, primarily because they can be censused with 
confidence, and often long term data are available (e.g., Diamond 1969; Diamond 
and May 1977; Power 1976). Coincidentally, birds are vagile and presumably 
more easily capable of immigration than most animals. Therefore the supposition 
that such turnover rates represent a dynamic equilibrium of the MacArthur-Wil- 
son (1967) interpretation may be valid. With most insects, however, recent ex- 
tinction rates likely are higher than natural due to decimation of the plant com- 
munity or to fragmentation of larval host populations. Colonization rates may be 
low owing to low immigration rates and/or reduced patch sizes of suitable habitat. 
Thus, the extinction rate in large part is independent of the colonization rate. 



SANTA CRUZ ISLAND INSECTS 105 

Population lifetime under natural conditions is a fundamental problem in pop- 
ulation biology, and extinctions and recolonization of local populations due to 
natural causes have been recorded in butterflies (e.g., Ford 1945; Ehrlich et al. 
1975). A species can survive such periodic fluctuations at a colony site provided 
that it is established nearby in sufficiently large numbers to ensure recolonization. 
Thus normal periodic cycles in abundance resulting in local extinctions may not 
have lasting effects in continental populations but may be disastrous to populations 
in isolated localities such as islands. 

Problem of census in insect communities. — The primary problem in drawing 
inferences about extinction or other faunal changes in insects is a practical one, 
that of accurate census. It is essentially impossible to inventory the insects of a 
complex island such as S.C.I, with confidence. Year around effort would be 
needed to cope with seasonality, an array of collectors with differing specialized 
experience would be required to sample adequately the variety of habitats, and 
the expertise of an army of taxonomists would have to be recruited to provide 
specific discrimination, irrespective of nomenclatural decisions. Many of the 
smaller forms are undescribed, whether or not they are known to occur on the 
mainland. Moreover, prolonged diapause, the ability of individuals to postpone 
emergence for more than one season, is much more widespread in temperate 
zone insects than has been appreciated (Powell 1974). 

Therefore, except in the simplest possible communities, such as the mangrove 
islands manipulated by Simberloff and Wilson (1969), several years are needed 
to census with any degree of completeness. Based on sporadic surveys repre- 
senting nearly all months, I have recorded about 400 species of Lepidoptera on 
S.C.I. Comparing that figure to those given for the North American fauna (Daly, 
Doyen, and Ehrlich 1978), one can extrapolate an estimate of at least 3500 species 
of insects on the island, and probably the real count is much higher. As a result, 
efforts to assess extinction or turnover rates in insects must be confined to well 
known taxa with conspicuous, relatively easily sampled species, such as butter- 
flies. When an animal such as Pieris rapae appears at several spots on the island, 
it can be inferred that recent colonization or recolonization has occurred. By 
contrast, the apparently sudden appearance of a native insect species more often 
is likely the result of inadequate past field survey. For example, when I discovered 
the univoltine, autumnal tortricid moth, Decodes fragarianus, on S.C.I, in Sep- 
tember 1978, a plausible explanation is that we had overlooked the larvae during 
spring surveys of Quercus in 1966-1969, negating the statement by Opler 
(1974:22) that this species does not occur on the Channel Islands. 

Undersaturated state of the insect fauna. — While it is true that the number of 
species on S.C.I, is the product of colonization and extinction rates superimposed 
upon the original community at the time of separation from the mainland, the 
number does not appear to be at equilibrium in phytophagous insects. It seems 
probable that a suppression of the original number has occurred through restric- 
tion of the islands' sizes during the Pleistocene (Johnson 1978) and through man's 
influence, especially grazing of domestic and ultimately feral animals, during the 
past 150 years. Most native insect species are not highly vagile and/or are de- 
pendent upon a narrow larval food preference. As a result, in contrast to birds, 
colonization rates almost certainly are too low to balance extinction rates, and 
a dynamic equilibrium is not attained. Severe disturbance of the flora has frag- 



106 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

merited and reduced the size of many phytophagous insect populations on the 
island, enhancing chances of local extinction due to natural fluctuations in abun- 
dance. 

Therefore it is probable that insect diversity has declined since the advent of 
introduced mammals and weedy plants, and the island's insect fauna is under- 
saturated. The situation is comparable to defaunation experiments with mangrove 
islands except the relaxation time required to reach equilibrium (or 90% of equi- 
librium ^ 9 o , of MacArthur and Wilson) is much longer. Feral sheep were removed 
and excluded from a large portion of S.C.I, during the 1950s, which can be 
envisioned as the end of the defaunation process, with gradual establishment and 
expansion of formerly resident and new colonist plants and phytophagous insects 
having progressed since that time. 

The relaxation time for the fauna to return to equilibrium following perturbation 
(including immigration of alien species), may be very long (examples cited by 
Simberloff 1974). Thus the insect fauna of S.C.I. , and probably most islands 
inhabited by Western man, may be interpreted as undersaturated and out of 
equilibrium, a state that will continue indefinitely. Cessation of overt disturbance 
(grazing, grading, farming, etc.) may mark the beginning of the return to an 
equilibrium, but immigration effects of the disturbance (colonized weedy plants 
and animals) prevent return to the original equilibrium and together with differ- 
ential rates of immigration and abilities to colonize by mainland species that have 
not yet colonized, preclude return to the original community composition. 

Insects that need specific plants as larval food, such as native bunch grasses 
or Eriogonum, may have dropped out of the fauna, and immigration from con- 
specific populations on the mainland or on nearby islands has not occurred and/ 
or remaining patches of suitable habitat are too limited to have enabled coloni- 
zation by chance. Homodynamic species that are more vagile and often weed- 
feeding, such as Pieris rapae and Pyrgus communis, also may have undergone 
extinction in times of climatic or grazing stress, but if so, have been able to 
recolonize. 

Thus the insect community of S.C.I, is transitional in species membership, 
consisting primarily of two cohorts: a) relics of the original California mainland 
community, and b) species that have been able to colonize in spite of or even 
because of the disturbance by man. The latter group consists mostly of homo- 
dynamic, weed-feeding or polyphagous species, some of which may colonize and 
disappear periodically. Included are both alien (e.g., Forficula auricularia and 
Pieris rapae) and native species (e.g., Platynota stultana and Pyrgus communis). 

Acknowledgments 

Thanks are extended to the Santa Cruz Island Company and Dr. Carey Stanton, 
whose cooperation has enabled use of the U.C. field station for the past 15 years. 
Dr. Stanton was most helpful during our early visits, as has been Dr. Lyndal 
Laughrin, Santa Cruz Island Reserve, U.C. Santa Barbara, on recent visits. Dr. 
C. L. Remington, Peabody Museum, Yale University, provided facilities and 
hospitality enabling study of his 1967-1976 collections of small moths from S.C.I. 
A draft of the manuscript was reviewed by Dr. Stanton, Dr. J. M. Diamond, and 
S. E. Miller, each of whom provided comments aiding its revision. R. L. Langs- 
ton, S. E. Miller, and S. Frommer provided unpublished data on butterfly col- 



SANTA CRUZ ISLAND INSECTS 107 

lections from the island. The efforts of numerous other collectors who worked 
with me or independently are also greatly appreciated. N.S.F. grants GB-4014 
and GB-6813X provided support for our fieldwork and surveillance of rearing 
material in 1966-1969. 

Literature Cited 

Alexander, C. P. 1973. Undescribed species of Nearctic Tipulidae (Diptera), XI. Great Basin Nat- 
uralist, 33(3): 189-196. 
Arnold, R. A. 1980. Ecological studies of six endangered butterflies (Lepidoptera: Lycaenidae); the 

role of island biogeography and patch dynamics in the design of nature preserves. Ph.D. Thesis, 

U. Calif., Berkeley. 
Atkins, E. L., M. H. Frost, L. D. Anderson, and A. S. Deal. 1957. The "omnivorous leafroller," 

Platynota stultana Wlshm., on cotton in California: nomenclature, life history, and bionomics 

(Lepidoptera: Tortricidae). Ann. Entomol. Soc. Amer., 50(3):25 1-259. 
Bohart, R. M. 1942. Platynota stultana as a pest of field-grown carnations. J. Econ. Entomol., 

35:399^103. 
Comstock, J. A. 1939. Channel Islands biological survey. Introductory note. Bull. So. Calif. Acad. 

Sci., 38(3): 133-134. 
. 1946. Channel Islands biological survey. 33, Brief notes on the expeditions conducted be- 
tween March 16, 1940 and December 14, 1941. Bull. So. Calif. Acad. Sci., 45(2):94-107. 
Coolidge, K. R. 1923. California butterfly notes— I. Bull. Brooklyn Entomol. Soc, 18:159-161. 
Daly, H. V., J. T. Doyen, and P. R. Ehrlich. 1978. Introduction to insect biology and diversity. 

McGraw-Hill, Inc., x + 564 pp. 
Diamond, J. M. 1969. Avifaunal equilibria and species turnover rates on the Channel Islands of 

California. Proc. Natl. Acad. Sci., 64:57-63. 
, and R. M. May. 1977. Species turnover rates on islands: dependence on census interval. 

Science, 197:266-270. 
Ehrlich, P. R., R. R. White, M. C. Singer, S. W. McKechnie, and L. E. Gilbert. 1975. Checkerspot 

butterflies: a historical perspective. Science, 188:221-228. 
Emmel, T. C, and J. F. Emmel. 1973. The Butterflies of Southern California. Nat. Hist. Mus. 

L. A. Co., Sci. Ser. 26, 148 pp. 
Ford, E. B., 1945. Butterflies. Collins, London, xiv + 368 pp. 
Goeden, R. D., C. A. Fleschner, and D. W. Ricker. 1967. Biological control of prickly pear cacti on 

Santa Cruz Island, California. Hilgardia, 38(16):579-606. 
, and D. W. Ricker. 1976a. The phytophagous insect fauna of the ragweed Ambrosia duinosa 

in southern California. Environ. Entomol., 5(l):45-50. 
. 1976b. The phytophagous insect fauna of the ragweed Ambrosia psilostachya in southern 

California, U.S.A. Environ. Entomol., 5(6): 1169-1 177. 
Gordon, R. D. 1976. The Scymnini (Coleoptera: Coccinellidae) of the United States and Canada: 

key to genera and revision of Scymnus, Nephus and Diomus. Bull. Buffalo Soc. Natural 

Sciences, 28:1-362. 
Heinrich, C. 1956. American moths of the subfamily Phycitinae. U.S. Natl. Mus., Bull. 207, 581 pp. 
Hillinger, C. 1958. The California Islands. Academy Publications, Los Angeles, Calif., 158 pp. 
Holder, C. F. 1910. Channel Islands of California. A. C. McClurg & Co., Chicago, xvi + 397 pp. 
Holland, W. J. 1931. 1898. The Butterfly Book. Doubleday, Page, & Co., Garden City, N.Y., 382 pp. 
Johnson, D. L. 1978. The origin of island mammoths and the Quaternary land bridge history of the 

northern Channel Islands, California. Quat. Res., 10:204-255. 
Keller, E. C, R. H. T. Mattoni, and M. S. B. Sieger. 1966. Preferential return of artificially displaced 

butterflies. Anim. Behav.. 14:197-200. 
Langston, R. L. 1980. The Rhopalocera of Santa Cruz Island, California. J. Research Lepid. (in 

press). 
, and S. E. Miller. 1977. Expanded distributions of earwigs in California (Dermaptera). Pan- 
Pacific Entomol., 53(2): 1 14-1 17. 
, and J. A. Powell. 1975. The earwigs of California. Bull. Calif. Insect Survey. 20:1-25. 



108 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

MacArthur, R. H., and E. O. Wilson. 1967. The theory of island biogeography. Monogr. in Popul. 

Biol. 1. Princeton U. Press, 203 pp. 
Meadows, D. 1936. An annotated list of the Lepidoptera of Santa Catalina Island, California. Part 

I. Rhopalocera. Bull. So. Calif. Acad. Sci., 35:175-180. 
Miller, D. R. 1971. Mealybugs of Santa Cruz Island, California (Homoptera: Coccoidea: Pseudo- 

coccidae). Pan-Pacific Entomol., 47(4):293-303. 
Miller, S. E., and A. S. Menke. 1980. Entomological bibliography of the California islands. Santa 

Barbara Nat. Hist. Mus. (in press). 
Opler, P. A. 1971. Seven new leaf miners associated with Quercus agrifolia (Heliozelidae, Graci- 

lariidae). J. Lepid. Soc, 25(3): 194-211. 
. 1974. Biology, ecology, and host specificity of Microlepidoptera associated with Quercus 

agrifolia (Fagaceae). U. Calif. Publ. Entomol., 75:1-83. 
. 1977. A review of North American Rhodophaea (Phycitinae: Pyralidae), with description of 

six new species. J. Lepid. Soc. 31:41-56. 
Powell, J. A. 1967. A previously undescribed moth reared from Catalina ironwood on Santa Cruz 

Island, California (Lepidoptera: Plutellidae). Pan-Pacific Entomol., 43(3):220-227. 
. 1974. Occurrence of prolonged diapausae in ethmiid moths (Lepidoptera: Gelechioidea). 

Pan-Pacific Entomol., 50(3):220-225. 
. 1981. Expanding geographical and ecological range of Platy not a stultana in California (Lep- 
idoptera: Tortricidae). Environ. Entomol. (in review). 
Power, D. M. 1972. Numbers of bird species on the California islands. Evolution, 26:451^163. 

. 1976. Avifauna richness of the California Channel Islands. Condor, 78:394-398. 

Rentz, D. C, and D. B. Weissman. 1973. The origins and affinities of the Orthoptera of the Channel 

Islands and adjacent mainland California. Part 1. The Genus Cnemotettix. Proc. Acad. Nat. 

Sci., Phila., 125(6):89-120. 
Simberloff, D. S. 1974. Equilibrium theory of island biogeography and ecology. Ann. Rev. Ecol. and 

Syst., 5:161-182. 
, and E. O. Wilson. 1969. Experimental zoogeography of islands. The colonization of empty 

islands. Ecology, 50:278-296. 
Thorne, R. F. 1969. The California Islands. Ann. Missouri Bot. Gard., 56:391^108. 
Weissman, D. B., and D. C. Rentz. 1976. Zoogeography of the grasshoppers and their relatives 

(Orthoptera) on the California Channel Islands. J. Biogeogr., 3:105-114. 
Woglum, R. S. 1920. A recently discovered citrus pest, Platynota tinctana (Walk.) in California. 

Calif. Dept. Agric, Mo. Bull., 19:341-343. [misidentification by A. Busck off. stultana]. 
Wright, W. G. 1906. The Butterflies of the West Coast of the United States. 2nd ed. Publ. by author, 

San Bernardino, Calif., 257 + vii pp. 

Accepted for publication November 6, 1980. 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 109-111 

Crustaceans in the Shrimp By-catch from off the Coasts of Si- 
naloa and Nayarit, Mexico 

R. K. G. Paul and M. E. Hendrickx 

Abstract. — Crustaceans in the shrimp by-catch from off the coasts of Sinaloa 
and Nayarit, Mexico by R. K. G. Paul and M. E. Hendrickx, Bull. Southern 
California Acad. Sci., 79(3): 109-1 1 1, 1980. Samples of the shrimp by-catch were 
obtained from boats operating off the northwest Pacific coast of Mexico. The 
crustacean component of the by-catch was separated and examined. Twenty-four 
species of crustacean were found in the by-catch and several, including 5 species 
of large portunid crabs, have considerable commercial value but are presently 
unexploited in this region of Mexico. 

Centro de Ciencias del Mar y Limnologia, Estacion Mazatlan, UNAM. Apdo. 
Postal 811, Mazatlan, Sinaloa, Mexico. 

Introduction 

For many years a large commercial fishing fleet of shrimp trawlers has operated 
in the Gulf of California and off the Pacific coast of Mexico. Shrimps of the genus 
Penaeus form the basis of this fishery (Snyder-Conn and Brusca 1977; Edwards 
1978) and constitute more than 10% of the total catch (Chavez and Arvizu 1972; 
Rosales 1976). The rest of the total catch consists mainly offish, crustaceans and 
mollusks and is often referred to as the shrimp by-catch. Very little published 
information exists concerning the composition of the by-catch and, of the studies 
that have been made, many are limited to fishes. 

Research on the shrimp by-catch is important since, apart from the ecological 
information that can be obtained, the by-catch represents an important potential 
food source which is being vastly underutilized at present. 

In this paper the crustacean component of the by-catch is considered and qual- 
itative data are presented. 

Materials and Methods 

Two trips were made on commercial shrimp trawlers during November 1978 
and February 1979 between Mazatlan (Sinaloa) and San Bias (Nayarit). In the 
first trip samples were obtained from three trawls, and in the second trip from 6 
trawls. All the trawls were made within five miles of the coast at depths of 14 to 
48 metres. The average duration of each trawl was 4 hours. The nets used by 
commercial shrimp trawls are benthic otter trawls with a 15 metre wide mouth 
and a 6.5 centimetre stretched mesh size. 

Results 

Table 1 gives a list of the crustaceans which occurred in the trawl samples. For 
purposes of comparison, the presence or absence of these organisms is also in- 
dicated for Mazatlan Bay (based on monthly samples taken for one year with a 
small benthic trawl), and from the study of Rosales (1976) on the by-catch off the 
whole coast of Sinaloa. 



10 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 1. Crustaceans occurring in the shrimp by-catch from the present study. For comparison, 
the presence (+) or absence (-) of these organisms is indicated for Mazatlan Bay samples and the 
by-catch samples of Ro sales (1976). 











Bay By- 










of catch 










Maza- Rosales 


Order 


Section 


Family 


Species 


tlan (1976) 



Stomatopoda 



Decapoda 



Squillidae 



Macrura Scyllaridae 
Palinuridae 
Penaeidea Penaeidae 



Brachyura Portunidae 



Majidae 
Calappidae 



Anomura Diogenidae 

Porcellanidae 



Squilla panamensis Bigelow 
Squilla parva Bigelow 
Squilla mantoidea Bigelow 

Evibacus princeps Smith 
Palinurus gracilis Streets 
Sicyonia disdorsalis (Burkenroad) 
Trachypenaeus similis pacificus 

Burkenroad 
Solenocera florea Burkenroad 
Callinectes arcuatus Ordway 
Callinectes toxotes Ordway 
Callinectes bellicosus (Stimpson) 
Portunus asper (A. Milne Edwards) 
Portunus acuminatus (Stimpson) 
Portunus xantusii affinis (Faxon) 
Euphylax robustus Milne Edwards 
Euphylax dovii Stimpson 
Leiolambrus punctatissimus (Owen) 
Stenocionops ovata (Bell) 
Stenorhynchus debilis (Smith) 
Hepatus kossmanni Neumann 
Persephona townsendi (Rathbun) 
Calappa convexa de Saussure 
Petrochirus californiensis Bouvier 
Porcellana cancrisocialis Glassell 



+ 


+ 


+ 


- 


+ 


- 


- 


+ 


+ 


+ 


+ 


+ 



- 


+ 


- 


+ 


+ 


+ 


- 


+ 


+ 


+ 


+ 


- 


- 


+ 


+ 


+ 


+ 


+ 


- 


+ 



Discussion and Conclusions 

A total of 24 species of crustaceans representing 2 orders, 4 sections and 9 
families were identified from the by-catch samples. Rosales (1976) reported 34 
species of crustaceans from the by-catch, but it should be noted that the samples 
in this study were taken over a two year period (1964-66) and from a much larger 
area. Many of the species reported by Rosales, and not in the present study, 
occurred in one trawl only and from outside the area covered by the present 
study. 

Several species of commercial value occurred in large numbers in samples, 
including five species of swimming crabs: Callinectes arcuatus Ordway; Calli- 
nectes toxotes Ordway; Callinectes bellicosus (Stimpson); Euphylax robustus A. 
Milne Edwards; and, Euphylax dovii Stimpson. At present no fishery exists for 
crabs on the Pacific coast of mainland Mexico and large quantities are wasted 
during shrimp-fishing operations, since when the catch is sorted the crabs are 
often killed and thrown overboard. Fishermen believe them to be important pred- 
ators of penaeid shrimp. Other species with possible commercial value include 
the stomatopods (Squilla spp.) and species of Penaeidae such as Sicyonia dis- 



SINALOA AND NAYARIT SHRIMP 1 1 1 

dorsalis (Burkenroad), Trachypenaeus similis pacificus Burkenroad and Solen- 
ocera florea (Burkenroad), none of which are commercially exploited at present. 

Acknowledgments 

We would like to thank the staff of the Crustacean Section of the Allan Hancock 
Foundation for confirming the identification of specimens. 

Literature Cited 

Chavez, H., and J. Arvizu. 1972. Estudio de los recursos pesqueros demersales del Golfo de Cali- 
fornia, 1968-1969. III. Fauna de acompanamiento del camaron. Mem. IV Congreso Nac. 
Ocean. (Mexico):361-378. 

Edwards, R. R. C. 1978. The fishery and fisheries biology of penaeid shrimp on the Pacific coast of 
Mexico. Oceanogr. Mar. Biol. Ann. Rev., 16:145-180. 

Rosales, F. J. 1976. Contribution al conocimiento de la fauna de acompanamiento del camaron en 
alta mar, frente a la costa de Sinaloa, Mexico. Mem. Reunion sobre los recursos de pesca 
costera de Mexico, Veracruz, Nov. 1976:25-80. 

Snyder-Conn, E., and R. C. Brusca. 1977. Shrimp population dynamics and fishery impact in the 
northern Gulf of California. Ciencias Marinas, 1(3):54— 68. 

Accepted for publication October 7, 1980. 



Research Notes 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 112-118 



Notes on Seasonal Collections of Sharks 
Near Point Conception, California 



This note describes the results of eleven quarterly, week-long gill net collec- 
tions of sharks between June 1974 and December 1976 at three stations near Point 
Conception, California. These collections were part of an extensive marine en- 
vironmental survey (Dames and Moore 1977) encompassing quantitative sampling 
offish, benthos, plankton, and intertidal organisms. 

The Cojo Anchorage study area is approximately 25 km west of Santa Barbara 
and 5 km southeast of Point Conception, California. Three stations, designated 
PC-1, PC-2, and PC-3, were sampled at depths of approximately 17, 12 and 8 m 
MLLW, respectively. The three station locations corresponded approximately to 
the outer, middle, and inner boundaries of an extensive kelp bed dominated by 
Macrocystis sp. The dominant substratum at all stations was a shale pavement 
overlain with scattered shale or a veneer of sandy silt, with some higher rocky 
areas present at the inshore stations. 

In June and September 1974, sampling consisted of one to three 24 hr sets of 
single mesh 3.8 cm bar mesh surface and bottom gill nets at Station PC-2. These 
nets measured 30.5 x 3.1 m, and were constructed of braided nylon line. In 
December 1974 and March 1975, the sampling effort was expanded to include 
three stations with night and day sets of approximately 12 hr each. 

Since diver observations in the area indicated a more varied community of 
fishes than was represented in the collections from the single mesh gill nets, 
surface and bottom multimesh (1.3 to 6.4 cm bar mesh in 1.3 cm increments) 
monofilament nets were deployed for two 12 hr day and night sets at each of the 
three stations in June 1975. The multimesh nets measured 30.5 x 2.4 m; con- 
sisting of two 3.1 m wide panels of each mesh, and were constructed of monofila- 
ment line of varying thickness. Panels were in progressive size sequence. For 
comparative analysis, the single mesh nets were deployed coincidentally with 
multimesh nets during the June 1975 survey. From September 1975 through De- 
cember 1976, multimesh nets were deployed with surface and bottom nets fished 
concurrently at each station. 

Upon retrieval of the nets, specimens of sharks were recorded as to species, 
total length, and sex. Additional notes included the recording of the mesh size 
and location (surface or bottom) in which each specimen was caught. 

A total of 2190 hr of gill net fishing time resulted in the capture of 673 sharks, 
representing eight species (Table 1). Due to the concentration of effort for the 
first two 1974 surveys at Station PC-2, almost 50% of the total fishing time was 
at that station. Surface and bottom net fishing times were evenly distributed, with 
1075 hr for surface nets, and 1116 for bottom nets. 

Spiny dogfish 1 {Squalus acanthias) and swell sharks (Cephaloscyllium ven- 



1 Common and scientific names in accordance with American Fisheries Society (1970). 



RESEARCH NOTES 



113 



700 


— (636) 








600 










500' 


£-lSD 


5T (447) 






400 




£-2SD 






300 


1(279) 


S-1SD 


w {273) 




200 






|-1SD 




100 


p- 1SD 
£-(46) 


1(118) 


|(110) 
■jjjr- 1 SD 


— (141) 
*-2SD 

i-iSD 

|(46) 

iiLlSD 



All spp. 



S.a 



C.v. 



Other spp. 



LEGEND 

S.a. - Squal us acanthias 
C-v. - Cephaloscyll ium ventriosum 
1SD - ONE STANDARD DEVIATION 
2SD - TWO STANDARD DEVIATIONS 
xWS; RANGE 
• MEAN 

Fig. 1. Mean and range of number of individuals caught/1000 hours fished by species (all stations/ 
depths and seasons combined). 



114 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Table 1. Catch per 1000 hours fished by season (all stations combined). 









Season 






Species 


Spring 


Summer 


Fall 


Winter 


Mean 1 


Squalus acanthias 


7.6 (3) 2 


349.4(181) 


127.6(81) 


54.6 (35) 


137 


Cephaloscyllium ventriosum 


63.1 (25) 


148.7 (77) 


192.1 (122) 


70.2 (45) 


123 


Triakis semifasciata 


— 


17.4 (9) 


6.3 (4) 


51.5 (33) 


21 


Galeorhinus zyopterus 


— 


5.8(3) 


29.9(19) 


4.7 (3) 


11 


Squatina californica 


2.5(1) 


30.9 (16) 


11.0(7) 


— 


11 


Mustelus californicus 


— 


— 


3.2 (2) 


7.8(5) 


3 


Prionace glauca 


— 


1.9(1) 


— 


— 


<1 


Heterodontus francisci 


— 


— 


1.6(1) 


— 


<1 


Total catch/ 1000 hr 


73.2 (29) 


554.1 (287) 


371.7(236) 


188.8 (121) 


307 (673) 


Total hours actually 












fished 


396 


518 


635 


641 


2190 



1 Sum of actuals divided by 2190 multiplied by 1000. 

2 Actuals. 



triosum) were nearly equal in abundance and accounted for about 85% of the 
total catch (Fig. 1). Leopard sharks (Triakis semifasciata), Pacific angel sharks 
(Squatina californica) and soupfin sharks (Galeorhinus zyopterus) accounted for 
a combined total of 14% of the catch, with T. semifasciata being twice as abun- 
dant as the other two. A blue shark (Prionace glauca), a horn shark (Hetero- 
dontus francisci), and seven grey smoothhounds (Mustelus californicus) account- 
ed for the remainder of the sample. Combining the data on all species, females 
outnumbered males during all but the spring (March) surveys by as much as 2.3:1. 

Due to the nonrandom grid sampling and variations in methodology, the col- 
lected data are not suitable to rigorous statistical analysis. However, the data 
have been standardized to catch per unit effort (here equalling number of indi- 
viduals per 1000 hr fishing), and the significance of seasonal, spatial (station 
preference) and compositional changes in the shark populations has been inferred 
assuming a normal data distribution. 

When the 11 sample periods are considered separately, the mean catch per 
1000 hr of all species combined was 279 individuals with a range of 46 to 636 (Fig. 
1). Although there was no significant difference in the means of catch per unit 
effort between C. ventriosum and S. acanthias (110 and 118/1000 hr, respec- 
tively), these two species dominated the catches and the combined mean of these 
two species was significantly greater (a s= 0.001, one tailed) than all other species 
combined (x, all other species = 46). 

The seasonal mean shark density ranged from 73 to 554 per 1000 hr of fishing 
(Table 1) over the year with the summer catch the highest. C. ventriosum catch/ 
effort in the spring, fall and winter was significantly (a — 0.05) greater than the 
mean of any other species for those seasons as was the S. acanthias catch/effort 
in the summer. 

By station, catch/effort of all sharks ranged from 151 to 414, but was not 
significantly different between stations. C. ventriosum mean catch/effort was sig- 
nificantly (a — 0.05) greater at Station PC-1 (offshore) than the mean for all other 
species at that station, and the S. acanthias mean catch/effort was significantly 



RESEARCH NOTES 



115 



1150- 






T^r— (ma) 


1000- 






;>X; 


800- 






vX; 

B (587) 


500- 








350- 








250- 






•:•:•: 


200- 








150- 






;X\ 


100- 






Sfi£ -(136) 


50- 


Sg-(65) 
§jj(23) 






-i>: 


(0) 





SURFACE 



BOTTOM 



LEGEND 



Mtt RANGE 
• MEAN 

Fig. 2. Number o individuals caught/ 1000 hours fished by depth (all stations, species and seasons 
combined). 



116 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

(a — 0.05) greater at PC-3 (inshore) station. Both species were abundant (means 
greater than 1 SD above mean of all other species) at the mid-depth station. 

Mean catch/effort of male sharks was not significantly different from the mean 
for females; however, males were least abundant in the spring and females most 
abundant in the summer. Mean catch/effort of male and female C. ventriosum 
and S. acanthias showed no significant increase during any season, but female 
S. acanthias mean catch/effort showed a substantial increase (>1 SD above the 
mean for all species) in the summer season. 

The mean of seasonal surface (depth of to 3.1 m) caught sharks per 1000 hr 
of fishing was 23, compared with 587 for bottom (to within 3.1 m of bottom) 
caught sharks. Since the ranges do not overlap, the estimated 95% confidence 
limits would also not overlap (Fig. 2), so that it is clear that the bottom associated 
sharks dominated. 

Seasonal length frequency distribution for S. acanthias is shown in Figure 3. 
These data indicate (1) a substantial increase in the number of females in the 
summer months, (2) the majority of the females and males are beyond the length 
at which 98% of the individuals of this species are mature, and (3) the majority 
of the females are between 90 and 104 cm TL irrespective of season. 

The data collected during this study indicate that the shark populations of this 
site are predominantly demersal, with two species (S. acanthias and C. ventrio- 
sum) dominating the overall catch. Seasonal data indicate that the summer catch 
consists of more species and individuals with increases in the number of S. 
acanthias. C. ventriosum is relatively constant in catch/unit effort at the study 
area, with a higher abundance at the mid-depth (12 m) and outer (17 m) stations 
than nearshore. S. acanthias dominates the inshore catch. All other species show 
a more varied seasonal and station abundance. 

Feder et al. (1974) state that P. glauca, C. ventriosum and T. semifasciata 
have been observed to be associated (to varying degrees) with southern California 
kelp beds. They also report that T. semifasciata is gregarious and may appear in 
large aggregations for only short periods of time. The data in this report tend to 
support the documented gregariousness of T. semifasciata (24 of the 53 individ- 
uals were collected during one bottom net setting at PC-3) and the association of 
C. ventriosum with the kelp bed habitat. The relatively large numbers of S. 
acanthias reported here are greater than what would be expected based on other 
reports (Feder et al. 1974; Quast 1968). Multiple sightings of S. acanthias have 
been made, however, within the kelp beds near Big Fisherman's Cove, Catalina 
Island (Ted Hobson, pers. comm.) and this species historically supported a lim- 
ited local gill net fishery in a nearshore sedimentary bottom area offshore Ven- 
tura, California (John Richards, pers. comm.). Large scale migrations and pref- 
erence for inshore areas by S. acanthias have been documented by Templeman 
(1944) and Alverson and Stansby (1963); and increases in numbers of S. acanthias 
in San Francisco Bay during winter months have been reported by de Wit (1975). 
Although no fecundity data were collected during this study, it is possible that 
Cojo Anchorage is conducive to breeding and pupping activities for this species 
during the summer months. The relatively large numbers of this species appear, 
however, to be unreported for southern California kelp beds. 

The lack of substantial suitable habitat for S. calif omica (sandy bottom near 
rock areas; see Feder et al. 1974) may be the primary reason for the relative 



RESEARCH NOTES 



117 



Spring 



77777/ 



90 


Sur 


nmer 






























80 


































70 


































60 


































50 


































40 


































30 


































20 


































10 










m 


W/< 


: 




W/A 

















Fall 



W777?rT7777 



r~i 



30 
20 


Wint 


er 






























in 




























Vrrrri 


Iv.v.'r 






SO-W 


55-59,, 


h 60-M 


65-69 


70-74 


75-79 


80-84 


85-69 , 


90-94 


95-99 


100-104 


105-109 


110-114 


115-119 


I20-12* 


1J5-1Z9 



Total Length (cm] 



LEGEND 



^ 98% maturation Y//////X 

I length for males vyy///, males 

t 



98% maturation 
length for females 



femal es 



Fig. 3. Number of male and female Squalus acanthias caught/1000 hours fished by season (all 
stations combined). 



118 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

paucity of this species. This species was most common at the offshore station 
where kelp cover is minimal and expanses of sedimentary bottom are present. 

The study documents shark population distribution similar to those cited in 
Feder et al. (1974), but our data indicate the presence of some species that are 
more characteristic of bays and other protected waters. The paucity of P. glauca 
in these samples may have been due to the location of all the stations within the 
bounds of the kelp bed, as P. glauca is usually considered typical of open ocean 
habitats (Feder et al. 1974; Tricas 1979). 

Literature Cited 

Alverson, D. L.. and M. Stansby. 1963. The spiny dogfish (Squalus acanthias) in the northeastern 

Pacific. U.S. Fish and Wildlife Service Special Scientific Report — Fisheries No. 447, 25 pp. 
American Fisheries Society. 1970. A list of common and scientific names of fishes from the United 

States and Canada. AFS Special Publication No. 6, 150 pp. 
Dames and Moore. 1977. Supplemental data report, marine biology environment for proposed LNG 

facilities and associated gas transmission pipeline, Point Conception, California. D&M Job No. 

00011-168-02, January, 91 pp. 
de Wit, L. A. 1975. Changes in the species composition of sharks in south San Francisco Bay. 

California Dept. of Fish and Game, 61(2): 106-1 11. 
Feder, H. M., C. Turner, and C. Limbaugh. 1974. Observations on fishes associated with kelp beds 

in southern California. California Dept. of Fish and Game Fish Bulletin No. 160, 144 pp. 
Hobson, T., personal communication. National Marine Fisheries Service, Tiburon Laboratory, 3150 

Paradise Drive, Tiburon, California 93920. 
Richards, J., personal communication. University of California Cooperative Extension, 260 North 

San Antonio Road, Santa Barbara, California 93111. 
Quast, J. C. 1968. Fish fauna of the rocky inshore zone. Pp. 35-56 in Utilization of kelp bed resources 

in southern California (W. J. North and C. L. Hubbs, eds.), California Dept. of Fish and Game 

Fish Bulletin 139. 
Templeman, W. 1944. The life history of the spiny dogfish (Squalus acanthias) and the vitamin A 

values of the dogfish oil. Newfoundland Dept. of Natural Resources Fish Bulletin No. 15, 76 

pp. 
Tricas. T. C. 1979. Relationships of the blue shark (Prionace glauca) and its prey species near Santa 

Catalina Island. California. Fishery Bulletin, 77(1): 175-182. 

Accepted for publication September 1, 1980. 

Leray A. de Wit and Thomas B. Scanland, Dames and Moore, Consultants in 
the Environmental and Applied Earth Sciences, Suite WOO, 1 100 Glendon Ave- 
nue, Los Angeles, California 90024. 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 119-121 



A Remora, Remilegia australis. Attached to an Atlantic 
Spinner Dolphin, Stenella longirostris 



On 8 November 1979, in the eastern Caribbean Sea, off Venezuela (10°25'N, 
64°30'W), in about 180 m of water, a herd of approximately 250 dolphins was 
observed. They were identified as Atlantic spinner dolphins {Stenella cf. S. lon- 
girostris Gray), on the basis of their cape, elongated rostrum, triangular dorsal 
fin, and other pigmentation evidence (Perrin 1972). For about 30 minutes groups 
of dolphins rode the bow of the boat, a "Morgan 41," which was moving at about 
9 km/hr. Individuals were easily observable. 

One individual, whose behavior differed from the others in erratic changes of 
direction, caught our eye during a spinning leap, and was subsequently photo- 
graphed. The animal approached the boat to swim alongside, dashed ahead criss- 
crossing in front of the bow, and finally moved away with several leaps. Through- 
out this period there was a remora firmly attached to the dolphin's side. Figure 
1 shows the location of the attached fish. 

The remora was a pale sky-blue, about 40 cm long and 10 cm wide. We ten- 
tatively identified it as a whalesucker, Remilegia australis Bennett (Echeneidi- 
dae), the only remora reported to utilize cetaceans as hosts (Rice and Caldwell 
1961). This species appears to have a world-wide distribution in temperate and 
tropical waters. It has been reported as far north as Vancouver Island, British 
Columbia (49°N), and as far south as Cape of Good Hope (34°S) (Follett and 
Dempster 1960). 

Whalesuckers have been reported on the following cetacean species: Balaenop- 
tera musculus (Follett and Dempster 1960; Rice and Caldwell 1961; Rice 1978); 
Balaenoptera borealis (Rice and Caldwell 1961); unidentified rorqual (Nicklin 
1963, later identified as Balaenoptera edeni, Morejohn and Rice 1973); Eubalae- 
na sp. (Gudger 1922); Physeter macrocephalus (Gudger 1922; Follett and Demp- 
ster 1960); Delphinus delphis (Follett and Dempster 1960); Delphinus bairdii ( = 
D. delphis) (Radford and Klawe 1965); Stenella plagiodon (Mahnken and Gilmore 
1960); Stenella attenuata (Linehan 1979); Tursiops truncatus (Wallace 1977; 
Shane 1978); and an unidentified pelagic dolphin (Townsend 1916; Caldwell 1961). 

We are aware of additional unpublished observations of whalesuckers attached 
to Globicephala sp. and Steno bredanensis (W. E. Evans, personal communi- 
cation), and to Lagenorhynchus obliquidens (S. Leatherwood, personal com- 
munication). 

Stenella cf. S. longirostris is now added to this list of cetacean hosts for remo- 
ras. 

In most accounts, remoras have been reported to move about on a cetacean 
host, changing position frequently. In contrast, during the period of our obser- 
vation, this one seemed to cling tenaciously to one spot. Close to the attachment 
site, we observed several oval pink spots about the size of the remora* s sucker 
disc. The aerial acrobatics of Stenella longirostris, its characteristic high-speed 
spinning leaps, rapid turns and fast swimming would seem to require adhesive 



120 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 1. Atlantic spinner dolphin, Stenella longirosths, with a suckerfish attached to its left side. 
Photo by William A. Watkins. 



strength sufficient to account for both the unusual tenacity of the specimen ob- 
served and the oval abrasions of the dolphin's skin. Scars from suckerfish at- 
tachment on cetaceans have been mentioned by Krefft (1953) and Radford and 
Klawe (1965). 

The observation of remora on the spinner dolphin were corroborated by Karen 
E. Moore and Romaine Maiefski, on board during a radio tracking experiment 
on Balaenoptera edeni. 

Funding for the cruise was from the Hubbs-Sea World Research Institute (San 
Diego), the Woods Hole Oceanographic Institution, and the Oceanic Biology 
Program of the Office of Naval Research (Contract N00014-74-C-0071 NR 083- 
004). W. E. Evans, J. R. Jehl, Jr., S. Leatherwood, R. H. Rosenblatt and F. H. 
Wolfson made useful comments and suggestions. This is contribution number 
4585 from the Woods Hole Oceanographic Institution. 



Literature Cited 

Caldwell, D. K. 1961. Observations on an unidentified dolphin of the family Delphinidae in Jamaican 

waters. Caribbean Journ. of Sci., 1:133-134. 
Follett, W. I., and L. J. Dempster. 1960. First record of the echeneidid fish Remilegia australis 

(Bennett) from California, with meristic data. Proc. California Acad, of Sci., 31:169-184. 
Gudger, E. W. 1922. An odd place for refuge. Nat. Hist. New York, 22:243-249. 
Krefft, G. 1953. Ichtiologische Mitteilungen aus dem Institut fur Seefischerei der Bundenstalt fur 

Fischerei. Zool. Anz., 1953:150-151. 
Linehan, E. J. 1979. The trouble with dolphins. National Geographic Magazine, 155(4):540. 



RESEARCH NOTES 121 



Mahnken, T., and R. M. Gilmore. 1960. Suckerfish on a porpoise. Journ. of Mammal., 41(1): 134. 
Morejohn, G. V., and D. W. Rice. 1973. First record of Bryde's whale {Balaenoptera edeni) off 

California. California Fish and Game, 59(4):3 13-315. 
Nicklin, C. 1963. Whale tale. Skin Diver, November, 1963:12-15. 
Perrin, W. F. 1972. Color Patterns of Spinner Porpoises {Stenella cf. S. longirostris) on the Eastern 

Pacific and Hawaii, with Comments on Delphinid Pigmentation. Fish. Bull., 70(3):983-1003. 
Radford, K. W., and W. L. Klawe. 1965. Biological observations on the whalesucker, Remilegia 

australis — Echeneiformes: Echeneidae. Trans. San Diego Nat. Hist. Soc, 14(6):65-72. 
Rice, D. W. 1978. Blue Whale. Pp. 30-35. In: Marine Mammals of Eastern North Pacific and Arctic 

Waters (D. Haley, ed.), Pacific Search Press, 256 pp. 
, and D. K. Caldwell. 1961. Observations on the habits of the whalesucker (Remilegia 

australis). Norsk Hvalfangsttid., 5:181-189. 
Shane, S. 1978. Suckerfish attached to a bottlenose dolphin in Texas. Journ. of Mammal., 59(2):439- 

440. 
Townsend, C. H. 1916. Porpoises at sea. Bull. New York Zool. Soc, 19:1427-1428. 
Wallace, B. 1977. Cover photo. Sea Frontiers, 23(3). 

Accepted for publication September 30, 1980. 

Giuseppe Notarbartolo di Sciara, Hubbs-Sea World Research Institute, San Die- 
go, California 92109 and William A. Watkins, Woods Hole Oceanographic In- 
stitution, Woods Hole, Massachusetts 02543. 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 121-124 



Stereotyped Motor Patterns in Two Captive 
Bull Sharks, Carcharhinus leucas 



Various motor patterns and the associated social activities have been described 
for the bonnethead shark, Sphryna tiburo (Myrberg and Gruber 1974) and the 
gray reef shark, Carcharhinus amblyrhynchos (Johnson and Nelson 1973). The 
sterotyped nature of these body movements and postures, as well as their de- 
parture from normal swimming behavior, were similar to those the authors ob- 
served in two captive bull sharks, Carcharhinus leucas. This report describes the 
behavior of these two bull sharks when the proper releasing situation occurs. 

Both bull sharks were males, and were caught in the area of the Florida Keys. 
At capture in April 1978, the smaller was 180 cm total length (TL), and the larger 
was 218 cm TL upon his capture in October of the same year. Each shark was 
transported approximately one and one-half months after capture to Sea World 
San Diego, where they were placed in a 1,500,000 liter, filtered sea-water aquar- 
ium. This facility was designed and built specifically for the maintenance and 
study of sharks (Keyes 1979). The general dimensions of this aquarium are 30.4 
m long, 10.7 m wide and 4.6 m deep. The observations were carried out daily, 
both from top side and through the underwater tank windows, from March 
through June 1979. 



122 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 1. Comparison of the postures exhibited by the bull sharks: A. angle-up; B. sustained lean; 
C. roll-arc and D. the common vertical orientation. 



In addition to the bull sharks, this facility housed the following: 1) seven nurse 
sharks, Ginglymostoma cirratum, three males and four females (—1.5 to 3.0 m 
TL), 2) five Atlantic lemon sharks, Negaprion brevirostris, two males and three 
females (=2.0 to 3.0 m TL), 3) four sandbar sharks, Carcharhinus plumbeus, all 
females (=2.0 to 2.5 m TL), 5) six cleaner wrasse, Labroides dimidiatus. All of 
the above-mentioned sharks, excluding the two Pacific lemon sharks, had lived 
in this aquarium for at least seven months. 

During regular locomotion, both bull sharks swam with relatively smooth, 
slightly stiff body movements. When closely encountering another shark under 
specific circumstances, explained later in this paper, the bull sharks involved 
veered to one side and performed one or more of the following postures: 

Angle-up (Fig. 1A). — a sharp and sudden upward change of attitude, ranging 
from 35° to 90° from the horizontal swimming path. The swimming strokes of the 
tail did not stop. 

Sustained-lean (Fig. IB). — a prolonged roll (less than 90° from horizontal) along 
the longitudinal body axis. This lean was usually held for approximately three 
seconds while the shark continued swimming. 

Roll-arc (Fig. 1C). — most abrupt change from normal swimming, this posture 
began with a lean of the body to one side. The head and tail were then simulta- 
neously thrown laterally toward the opposite side of the body lean. This resulted 
in a posture with the head and tail directed upward in relation to the shark's 
original horizontal orientation. In most cases the dorsal surface of the bull shark 



RESEARCH NOTES 



123 




180 



C. Leucas 



„_Sj«44 _:^*>w_ ; tx_i£u<^j 



Fig. 2. Typical releasing situation, in which the approaching shark passes over the C. leucas at 
an angle of 45°, at point of intersection. 



faced the other shark. Near vertical orientation of the bull shark's body in the 
water column was common (Fig. ID). Many times the head of the bull shark 
broke the surface of the water. All swimming motion was halted during this 
posture. 

Each of these postures was culminated by the bull shark either swimming or 
gliding away, and eventually returning to normal swimming behavior. The glides 
varied in descending angles, sometimes actually becoming a nose dive, straight 
down to just above the tank floor. 

The bull shark postures were released when another shark's approach direction 
and body orientation satisfied all of the following three criteria: l) traveled ap- 
proximately on the same level or above the bull shark's body in the water column, 

2) swam close to the bull shark's body (—0 to 2.4 m distance, averaging .7 m), 

3) achieved either a sustained parallel, a close trailing, or an angular intersecting 
travel with reference to the bull shark's body. When the bull shark reacted to 
instances of intersecting angle, the angle between the two sharks' bodies was less 
than approximately 90°, as shown in Figure 2. 

All of the shark species in the aquarium released postures in our bull sharks. 
Out of 158 recorded cases, 95 (60%) were released by the five Atlantic lemon 



124 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

sharks. The two bull sharks stimulated each other in 16 (10%) instances. The 
sandbar, Pacific lemon and nurse sharks were involved in 33 (21%) instances, 
while in 14 (9%) cases the observer could interpret no influence by any individual. 

This disproportionate number of postures exhibited toward the Atlantic lemon 
sharks can be explained largely by the existence of a general depth stratification 
of shark species at the Sea World facility. The bull sharks, along with the sandbar 
sharks, tended to occupy the mid depths of the aquarium, although the latter 
species will usually swim below a nearby bull shark. The nurse and Pacific lemon 
sharks most often rested on, or swam along the floor. Except when being cleaned 
by the cleaner wrasse (Keyes in prep.), or rummaging along the bottom after 
feeding, the Atlantic lemon sharks usually circled in the upper quarter of the 
aquarium water column. This preference for the area above the bull sharks placed 
the Atlantic lemon sharks in more posture inducing situations. 

The sharks at which the bull sharks directed their postures sometimes made 
obvious responses, such as change of course and/or speed. Quantified data con- 
cerning these responses are presently insufficient to allow the authors to present 
a conclusion as to the effect these postures have on the other sharks' behavior. 
Without this conclusion, an assignment of function to the bull shark postures is 
impossible. 

The bull shark is native to inshore and estuary areas of the Atlantic Ocean and 
Gulf of Mexico (Bohlke and Chaplin 1968), and occurs world wide in both fresh 
and salt water. The postures exhibited by the two Sea World bull sharks may be 
a result of such conditions as the mixture and high concentration of shark species 
in the aquarium. Yet, the occurrence of similarly exaggerated and repetitive be- 
haviors of Sphyrna tiburo and Carcharhinus amblyrhynchos in their natural en- 
vironment (Myrberg and Gruber 1974; Johnson and Nelson 1973) promotes the 
possibility that the bull shark postures may also occur in the wild. 

Acknowledgments 

We thank Hubbs-Sea World Research Institute and Sea World San Diego for 
the opportunity to carry out this research. Also, we thank Drs. Donald R. Nelson, 
William E. Evans and Richard Bray for reviewing the manuscript. Special grat- 
itude is given to the Sea World aquarium staff for their patience and observations 
and to Ann Dawes for the illustration. 

Literature Cited 

Bohlke, J. E., and Chaplin, C. C. G. 1968. Fishes of the Bahamas and Adjacent Tropical Waters. 

Livingston Publishing Co., Wynnewood, Pennsylvannia. 
Johnson, R. H., and Nelson, D. R. 1973. Agonistic display in the gray reef shark, Carcharhinus 

menisorrah, and its relationship to attacks on man. Copeia, 1973:76-84. 
Keyes, R. S. 1979. Sea World shark center report: 79-1. 
Myrberg, A. A., Jr., and Gruber, S. H. 1974. The behavior of the bonnethead shark, Sphyrna tiburo. 

Copeia, 1974:358-374. 

Accepted for publication August 12, 1980. 

David M. Stalls, Department of Biology, California State University, Long 
Beach, California 90840 and Raymond S. Keyes, Curator of Fishes, Sea World 
San Diego, San Diego, California 92109. 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 125-126 



Possible Cleaning Behavior by a Juvenile California 
Sheephead, Semico ssyphus pulcher (Labridae) 



Cleaning behavior has been observed in a number of marine fishes inhabiting 
the nearshore waters of California. This group includes six species of embioto- 
cids, two species of labrids, two species of kyphosids, one pomacentrid, and one 
scorpidid (Limbaugh 1955, 1961; Feder 1966; Gotshall 1967; Turner et al. 1969; 
Hobson 1969, 1971, 1976; Hixon 1979; DeMartini and Coyer 1981). These fishes 
possess a small mouth and are generalized (sensu Hobson 1971) "substrate-pick- 
ers" or "plankton-pickers" as juveniles or adults. For some of these species, 
cleaning behavior is confined to the juvenile phase and/or to relatively few, spe- 
cialized individuals (Hobson 1971; DeMartini and Coyer 1981). This note reports 
possible cleaning by a juvenile California sheephead, Semicossyphus pulcher 
(formerly Pimelometopon pulchrum), a behavior previously unreported for this 
species. 

During an early afternoon dive on 2 December 1976, I observed a juvenile 
sheephead apparently cleaning a blacksmith, Chromis punctipinnis . The event 
occurred above a small algae-encrusted boulder at a depth of 4-5 m in a dense 
kelp (Macrocystis) forest at Starlight Beach, near the west end of Santa Catalina 
Island, California. Two blacksmiths (approximately 60 and 120 mm total length) 
with no obvious discolorations or wounds, hovered 0.5 m above the small boul- 
der. The juvenile sheephead was approximately 90 mm total length and possessed 
the normal jm .mile coloration: a brilliant red orange body with a yellow stripe 
along the lateral line, two black spots on the dorsal fin, a single black spot at the 
base of the caudal fin, and black pelvic and anal fins trimmed with white. 

When first noticed, the larger blacksmith was 10-15 cm from the juvenile sheep- 
head and had assumed a conspicuous head-up, tail-down posture with fins mo- 
tionless and erect. During the subsequent observation period, the sheephead 
briefly inspected the posing blacksmith, then picked once at the region just pos- 
terior to the right pectoral fin. The blacksmith reacted positively to the contact 
by continuing to pose for an additional 15-20 s. During this period, the sheephead 
continued to inspect, but did not approach the posing blacksmith. The event was 
terminated when the sheephead slowly swam away (perhaps because of my pres- 
ence). This individual was not captured and could not be located on later dives 
within the area. The smaller blacksmith hovered near the larger blacksmith during 
the apparent cleaning event, but did not pose and was not approached by the 
sheephead. Neither blacksmith followed the retreating sheephead. 

I believe this represents cleaning behavior for the following reasons: 

1) blacksmith vigorously solicit and accept cleaning by cleaning fishes (Lim- 
baugh 1961; Hobson 1971) 

2) the observed blacksmith assumed a pose typical of cleaning solicitation in 
this species (Limbaugh 1961; Hobson 1971) 

3) the pose resulted in a brief inspection and subsequent oral contact by the 



126 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

juvenile sheephead, behaviors consistent with other reported instances of 
cleaning (Hobson 1971) 
4) the blacksmith continued to pose after interspecific contact, a positive re- 
sponse frequently observed in cleaning events (Limbaugh 1961; Hobson 
1971) 

During numerous subsequent observations of sheephead, I did not notice such 
apparent cleaning behavior among either juveniles or adults. Therefore, the be- 
havior seems restricted to juveniles and is either an occasional activity for a very 
few individuals, or is a rare event among all juveniles. 

It is reasonable to expect some degree of cleaning behavior in juvenile sheep- 
head. Cleaning is widespread among labrids throughout the world (Randall 1958; 
Feder 1966; Ayling and Grace 1971) and is restricted to the juvenile phase in the 
temperate Pseudolabrus miles and Coris sandageh (Ayling and Grace 1971) and 
the tropical Bodianus pulchellus, B. rufus, and B. diplotaenia (Randall 1968; 
Hobson 1969). Furthermore, juvenile sheephead are small-mouthed, substrate- 
pickers, and cleaning behavior appears to be widespread among small-mouthed 
fishes that pick food items from a substratum or the water column (Hobson 1971). 

Acknowledgments 

This work was partially supported by Sea Grant USDC 04-6-158-44118 and is 
contribution No. 48 from the Catalina Marine Science Center. I thank R. R. 
Given for use of facilities and R. F. Ambrose, E. E. DeMartini, and R. J. Schmitt 
for helpful comments on an earlier draft. 

Literature Cited 

Ayling, A. M., and R. V. Grace. 1971. Cleaning symbiosis among New Zealand fishes. N. Z. J. Mar. 

Freshwat. Res., 5:205-218. 
DeMartini, E. E., and J. A. Coyer. 1981. Cleaning and scale-eating in juveniles of the kyphosid 

fishes Hermosilla azurea and Girella nigricans. Copeia 4, in press. 
Feder, H. M. 1966. Cleaning symbiosis in the marine environment. Pp. 327-380 in Symbiosis, Vol. 

1. (S. M. Henry, ed.). Academic Press, 635 pp. 
Gotshall. D. W. 1967. Cleaning symbiosis in Monterey Bay, California. Calif. Fish Game, 53:125- 

126. 
Hixon, M. A. 1979. The halfmoon, Medialuna californiensis, as a cleaner fish. Calif. Fish Game, 

65:117-118. 
Hobson, E. S. 1969. Comments on certain recent generalizations regarding cleaning symbioses in 

fishes. Pac. Sci.. 23:35-39. 

. 1971. Cleaning symbioses among California inshore fishes. Fish. Bull., U.S., 69:491-523. 

. 1976. The rock wrasse, Halichoeres semicinctus, as a cleaner fish. Calif. Fish Game, 62:73- 

78. 
Limbaugh, C. 1955. Fish life in the kelp beds and the effects of kelp harvesting. IMR Ref. 55-9. 

Univ. Calif. Inst. Mar. Res. 158 pp. 

. 1961. Cleaning symbiosis. Sci. Am., 205:42-49. 

Randall, J. E. 1958. A review of the labrid fish genus Labroides, with descriptions of two new 

species and notes on ecology. Pac. Sci., 12:327-347. 

. 1968. Caribbean reef fishes. T.F.H. Pub., 318 pp. 

Turner. C. H., E. E. Ebert, and R. R. Given. 1969. Man-made reef ecology. Calif. Dept. Fish Game, 

Fish Bull., 146:1-221. 

Accepted for publication November 13, 1980. 

James A. Coyer, Catalina Marine Science Center (University of Southern Cali- 
fornia), Box 398, Avalon, California 90704. Present Address: Interstate Electron- 
ics Corp., Box 3117, Anaheim, California 92803. 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 127-129 



Use of Incisors to Identify Rodent Genera 
in Owl Pellets 



Analysis of owl pellets has been used for predator food studies (e.g. Cunning- 
ham 1960), determining species distributions (e.g. Twente and Baker 1951), in- 
terpreting paleontological finds (e.g. Brain and Brain 1977), and for estimating 
mammalian populations (e.g. Cabon-Raczynska and Ruprecht 1977). The senior 
author has used owl pellets in environmental impact studies, reducing the cost 
of the rodent species list and the impact of the assessment itself on the living 
fauna of the area. Owls may preferentially select prey species (Voight and Glenn- 
Lewin 1978); thus, pellet analysis is useful for verifying what species are in an 
area, not as proof of their absence. Owl pellet analysis is also a simple but 
rewarding laboratory exercise in ecology classes. 

Presently, Ingles (1965) provides the best known key for identifying the mam- 
mals of California; it is predominantly based on cranial characteristics. Great 
Horned Owls (Bubo virginianus) and Barn Owls (Tyto alba) are known to re- 
gurgitate 95-100% of the crania of their prey (Dodson and Wexlar 1979). The 
mandible, however, is seen as often as the cranium in the pellets but no concise 
key appears to be available for this bone. 

We asked six college biology majors with no previous experience to key out 
nine rodent crania (from owl pellets) to genera using Ingles' key which included 
only likely local species; they took an average of 13.6 minutes per item and were 
correct 75% of the time. The identification of these species had been verified 
against a reference collection of skulls kept at California State University, Ful- 
lerton. In this paper we present a new key (with an emphasis on incisor arcs) for 
rodent species from coastal sage scrub and disturbed (urban) habitats of Southern 
California. We think our method is adaptable to other localities; it is designed for 
individuals with limited experience in the use of mammal keys. Our method 
reduced the students' handling time to below two minutes per item and their 
identification accuracy rose to 85 and 89% on crania and on mandibles. 

Our method is based primarily on both upper and lower incisor characteristics 
with a limited number of cranial and mandibular characters. A basic piece of 
equipment needed is a multiple-circle plastic template (circles in V32 inch se- 
quences; templates may not be in metric), readily available from art and graphic 
supply stores. We assume that the incisor is an arc of a circle. As Landry (1957) 
pointed out, this assumption is not accurate since incisor curvature occurs in two 
planes "so that the curve produced is not a flat circle but a shallow spiral or, 
more properly, a helix." One only needs to place an incisor on a flat surface and 
try to match its outer curvature to the inner curvature of an appropriate circle 
on the template. The best match occurs when the entire outer surface of the 
incisor is in contact with the inner curvature of the proper diameter circle; the 
diameter of the incisor arc can then be recorded from the template and used on 
our key. Upper incisors approximated an 180° arc of a circle while lower incisors 
were less than 135° of a circle for our species. Upper incisor morphology and 



128 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Upper Incisor 



THOMOMYS t 



DIPODOMYS 



% in each 
size group 

• 1-10 

• 11-30 

• 31-50 
#51-70 

#71+ 



NEOTOMA #~<>-4-0 



RATTUS #H>-n 



PEROGNATHUS 



PEROMYSCUS f-# 



X 



MIllllHl 

17 18 19 20 21 22 23 24 26 28 

f-f grooved inc. (16) 



(51) 



first molar— 3 roots (19) 
first molar— 5 roots (5) 



14 15 18 

MICROTUS* f f f molars— serrated shaft (55) 
grooved inc. (35) 



MUS 
REITHRODONTOMYS 



5 6 7 8 




notched inc. 1 
grooved inc. (23) 



DIAMETER OF INCISOR ARC in 1/32 

Fig. 1. Key to upper incisors, with cranial characters as needed, of the rodents found in owl 
pellets from coastal sage scrub and urban/disturbed habitats of Southern California. Taxa represent 
Reithrodontomys megalotis, Mus musculus, Peromyscus sp., Perognathus sp., Microtus californi- 
cus, Rattus sp., Neotoma sp., Dipodomys agilis, and Thomomys bottae. Sample sizes are in paren- 
theses. Inc. = incisor. 



Lower Incisor 



THOMOMYS * T ? f ? ? T complex ang. near en. (42) 



NEOTOMA 
RATTUS 



«»-«h4-«»-i» 



first molar— 2 roots (21) 
first molar- 5 roots (5) 



MICROTUS 
DIPODOMYS 



T T ? 

m 



28 3032 36 4044 

♦-4 molars— serrated shaft (52) 



(10) 



20 21 22 23 24 26 

PEROMYSCUS 



MUS 



PEROGNATHUS f~t 
REITHRODONTOMYS 



n 



m 




'£7 (25) 

"(20) 
"u"cusp. molars (33) 

/ ^0 (20) 



condyloid 

'(en.) 

angular 
(ang.) 



11 12 13 14 15 16 

DIAMETER OF INCISOR ARC in 1/32" 

Fig. 2. Key to lower incisors, with mandibular characters added as needed. 



RESEARCH NOTES 129 

diameter of arc can be used to separate out six of nine rodent genera found in the 
owl pellet samples. We used the right incisor for our measurements whenever 
possible. Most owl pellets were from Barn Owls in western Riverside, Orange, 
and southeastern Los Angeles counties of California in coastal-sage scrub and 
urban/disturbed habitats. 

Figures 1 and 2 show the key for upper and lower incisors with cranial or 
mandibular characters added if needed. The user first finds the column corre- 
sponding to the diameter of the incisor arc which narrows the possible rodent 
species; a second incisor character or a mandible/cranial character allows the 
identification of the specific genus. Our method cannot separate out closely re- 
lated species which are the same size (i.e. Rattus rattus versus R. norvegicus, 
Neotoma fuscipes versus N. lepida). Thus for our sampling area the following 
taxa can be identified; Reithrodontomys megalotis, Perognathus sp., Dipodomys 
agilis, Thomomys bottae, Microtus califomicus, Neotoma sp., Rattus sp., Mus 
musculus, and Peromyscus sp. Another difficulty with this method is removing 
incisors from Reithrodontomys and Perognathus; it may be necessary to remove 
some of the cranial bones in the process. 

Acknowledgments 

We thank the fall, 1979 Animal Ecology class of California State University, 
Fullerton for their help, especially Greg Jakust and Gordon Owens. Dr. Bayard 
Brattstrom provided some of the owl pellet material. Dr. Gene Jones provided 
helpful criticism on the figures. 

Literature Cited 

Brain, C. K., and V. Brain. 1977. Microfaunal remains from Mirabib: Some evidence of palaeo- 

ecological changes in the Namib. Madoqua, 10:285-294. 
Cabon-Raczynska, K., and A. L. Ruprecht. 1977. Estimation of population density of the common 

vole in Poland: An analysis of owl pellets. Acta Theriol., 22:349-354. 
Cunningham, J. D. 1960. Food habits of the horned and barn owls. Condor, 62:222. 
Dodson, P., and D. Wexlar. 1979. Taphonomic investigations of owl pellets. Paleobiology, 5:275- 

284. 
Ingles, L. G. 1965. Mammals of the Pacific States. Stanford University Press, x + 506 pp. 
Landry, S. O., Jr. 1957. Factors affecting the procumbency of rodent upper incisors. J. Mamm., 

38:223-234. 
Twente, C. K., and R. H. Baker. 1951. New records of mammals from Jalisco, Mexico, from barn 

owl pellets. J. Mamm., 32:120-121. 
Voight, J., and D. C. Glenn-Lewin. 1978. Prey availability and prey taken by long-eared owls in 

Iowa. Amer. Midi. Nat., 99:162-171. 

Accepted for publication November 4, 1980. 

Joel D. Weintraub and Gregory Shockley, Department of Biological Science, 
California State University, Fullerton, California 92634. 



130 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 130-132 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



On the Status of the Eastern Pacific Cymothoid Fish Parasite 

Braga occidentalis Boone, and Its Synonymy with 

B. patagonica Schioedte and Meinert (Crustacea: 

Isopoda: Cymothoidae) 



Amid the various collections examined by the naturalist Pearl Lee Boone in 
her lifetime, is a single specimen of cymothoid isopod she chose to name Braga 
occidentalis. The specimen was allegedly collected from an unspecified locality 
"off the west coast of California" by James D. Dana and John L. Le Conte in 
1866 (Boone 1918). This species has not been reported since and its validity has 
been seriously questioned by Lemos de Castro (1959), who suggested the pos- 
sibility that it is in reality a synonym of Braga patagonica Schioedte and Meinert, 
1884, a South American freshwater species. Lemos de Castro apparently did not 
examine the type of B. occidentalis. Our examination of this type specimen 
(deposited as a holotype in the Peabody Museum of Natural History, Yale Uni- 
versity, YPM No. 302) has confirmed Lemos de Castro's suspicions and we 
herein synonymize Braga occidentalis Boone, 1918 with B. patagonica. 

There is no original collection label associated with the type specimen, and 
considerable doubt is cast upon the data associated with it by Boone. First, there 
is no record of Dana and/or Le Conte having participated on any oceanic or 
coastal expeditions in the alleged year of the collection (1866). Secondly, the five 
other known species of the genus Braga are restricted in their distribution to 
freshwater habitats within the eastern drainage of South America (Trilles 1973). 
Lastly, no other specimens of B. occidentalis, or any congeners, have been 
reported from the Pacific or any other marine habitat. 

During the year 1866, when the specimen was reported by Boone to have been 
obtained, neither Dana nor Le Conte were in a position to make such a collection. 
Just a year after the Civil War, Le Conte (ending an appointment as Surgeon of 
Volunteers) was acting geologist on an overland survey for the extension of the 
Union Pacific Railroad. Dana was gradually recovering from a severe physical 
breakdown following his completion of the United States Exploring Expedition 
Report on the Crustacea and Zoophytes, and was never again capable of extended 
travel. 

The possibility still exists that either Dana or Le Conte collected the specimen 
Boone designated as B. occidentalis, but not in the year she reported. Le Conte 
traveled on an expedition to California in 1850-1851 collecting a considerable 
variety of material in many marine taxa, including crustaceans. The list of Crus- 
tacea was subsequently published by Dana (1854), but contained only intertidal 
and terrestrial species, suggesting that offshore collections were not made. There 
appears to be no record of Le Conte ever accompanying an expedition to the east 
coast of South America (although he did travel as far south as Honduras). 

Dana, on the other hand, traveled extensively around the world as a geologist 
and naturalist for the U.S. Exploring Expeditions of 1838-1842, collecting along 
both the Atlantic and Pacific coasts of South America. Dana might have collected 



RESEARCH NOTES 131 

the specimen in question from a freshwater or estuarine habitat along the east 
coast of South America, failing to describe it in his report on the Crustacea (Dana 
1853). The Crustacea listed in his report included five genera of cymothoid iso- 
pods: Cymothoa and Lironeca (3 species each), collected from Rio de Janeiro 
and the Hawaiian Islands; Nerocila (6 species) from Rio de Janeiro; Aegathoa 
(2 species), from the Bahamas, Rio de Janeiro, and Tierra del Fuego; and Cer- 
atothoa (2 species), from the Atlantic coast of North America and the Indo- 
Pacific. All 16 of these species are strictly marine in distribution, and none were 
reported from the eastern Pacific. Unfortunately, Dana did not include collection 
dates for any of the species he discussed. 

We believe there are two possible explanations regarding the collection data 
reported by Boone for Braga occidentalis. First, one of Le Conte's South Amer- 
ican colleagues may have sent the specimen to him, Le Conte subsequently for- 
warding it to Dana. Second, Dana may have collected the specimen along the 
Atlantic coast of South America while on the U.S. Exploring Expedition and 
later sent it to Le Conte for examination. Le Conte then would have returned the 
specimen (unidentified) years later, after the completion of Dana's report. The 
year presently associated with the specimen might then be the year Dana received 
the specimen back from Le Conte, and/or the year it was accessioned into the 
Yale Peabody Museum. Many years passed until Pearl Boone found the specimen 
on the shelves of the Yale Peabody Museum and described it, assuming it to be 
part of the Le Conte-Dana California collection. 

Ho (1975) reported the only other record of Braga (n. sp.) from the eastern 
Pacific. His specimen subsequently has been lost and recent communication with 
Ho suggests that it was probably a case of mistaken generic identification. Based 
on his description it appears to have been a juvenile Lironeca. 

We thank Thomas E. Bowman and Ernest W. Iverson for their critical review 
of this paper. 

This is contribution no. 387 of the Allan Hancock Foundation, University of 
Southern California. 

Appendix 

I. Chronology of J. L. Le Conte pertinent to the present paper (1825-1883) 

1850-1851 Expedition to California; stopped at Panama to make collections. 
Extended explorations through Colorado desert. 

1857 Accompanied the Honduras Inter-Oceanic Survey, under J. C. 

Trautwine. Visited Fuente de Sangu. 

1862-1865 Appointed Surgeon of Volunteers during Civil War. 

1867 Geologist, surveyed for extension of Union Pacific Railroad south- 

ward to Fort Craig, Colorado. 

1869-1872 Traveled in Europe, Algiers and Egypt. 

1874 President of American Association for the Advancement of Sci- 

ence. 

Comments: Although Le Conte worked extensively in the field of entomology 
(particularly the Coleoptera), he also contributed numerous scientific articles 
to the fields of vertebrate paleontology and mammalogy. 



132 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

II. Chronology of J. D. Dana pertinent to the present paper (1813-1895) 

1834 Upon graduation from Yale appointed as Instructor of Mathemat- 

ics to the midshipmen of the United States Navy. Traveled exten- 
sively. 

1836-1838 Chemical assistant at Yale College. 

1838-1842 Served as geologist and naturalist to the U.S. Exploring Expedi- 
tion to the southern and Pacific Oceans under Comm. Charles 
Wilkes. The next 13 years were devoted largely to the study of the 
material collected by the expedition and preparation of his report. 

1850 Appointed Professor of Geology and Natural History at Yale Col- 

lege. 

1859 Suffered a severe physical breakdown. Traveled abroad in Europe 

for a year to regain health. 

1862-1872 Published several books: Manual of Geology (1862), Text Book of 
Geology (1864), Systematics of Mineralogy, fifth edition (1868), 
Manual of Geology, new edition (1874), Corals and Coral Islands 
(1872). 

1887-1890 Revisited the Hawaiian Islands (the "Sandwich Islands") and the 
Volcano of Kilauea. 

Comments: Dana's principal publications dealt with hermatypic corals 
("zoophytes"), crustaceans and geological formations. He was the first to 
propose the concept of cephalization in animals (1852), "The domination of 
the brain in determining the development of an animal organism,'" and he 
proposed a subsidence theory of atolls independently of Darwin's. His zoo- 
geographical analysis and techniques (1853) represent a landmark in marine 
biogeographical studies. 

Literature Cited 

Boone. P. L. 1918. Description often new isopods. Proc. U.S. Natl. Mus., 54(2253):591-604. 
Dana. J. D. 1853. United States Exploring Expedition. Vol. XIII. Crustacea, Part II. 1618 pp. Printed 

by C. Sherman, Philadelphia, Pennsylvania. (Dated 1852). 
. 1854. Catalogue and descriptions of Crustacea collected in California by Dr. John L. Le 

Conte. Proc. Phil Acad. Natrl. Sci., 7:175-177. 
Ho, J.-S. 1975. Parasitic Crustacea, in Lane, E. D. and C. W. Hill (Eds.)., The marine resources of 

Anaheim Bay. Calif. State Fish and Game, Bull. No. 165:69-72. 
Lemos de Castro, A. 1959. Sobre as especies sul-Americanas do genero Braga Schioedte and Meinert 

(Isopoda, Cymothoidae). Arquiv. Mus. Nac. 49:69-95. 
Trilles, J. -P. 1973. Notes documentaries sur les Isopodes Cymothoadiens parasites de poissons d'eau 

douce de l'Amerique du Sud. Bull. Mus. Nat. d'Hist. Natur.. ser. 3. No. 114, Zool., 88:239- 

272. 

Accepted for publication December 2, 1980. 

Melinda A. Thun and Richard C. Brusca, Allan Hancock Foundation, University 
of Southern California, Los Angeles, California 90007. 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 133-134 



One Mass Stranding, Not Two, of Sperm Whales at La Paz, 
Baja California, 1954 



In January 1954, news arrived in San Diego of a mass stranding of sperm whales 
at La Paz, southern Baja California, gulf side. 

I visited La Paz on 6 February 1954, and found three sperm whales in advanced 
stages of decomposition, on an inner, shallow beach, just east of the city of La 
Paz. I also received details of the mass stranding from two informants in La Paz: 
Mrs. Margaret Waters, stringer for the New York Times, and teacher of English; 
and Sr. Luis Collins, manager of Hotel Mision. Collins also gave me photos. 

Waters and Collins agreed on all essential details: The date was 16 January; 22 
individuals had stranded at the same time; all were males, 35 to 40 feet in length; 
nineteen had been towed to an outer beach before my arrival. Photos showed 
partial flensing of many carcasses by local inhabitants to obtain blubber to render 
for the waxy oil, good for lubrication. 

In May 1956, Cockrum reported two mass strandings at La Paz, as follows: 
"An Associated Press dispatch datelined February 13, 1954, from San Diego, 
California . . . reported [that] Mr. Marvin Grisby, a pilot from San Diego, . . . 
saw 24 sperm whales stranded in the shallow cove at La Paz . . . and, two weeks 
later, at the same place, 34 more . . ." 

In early 1957, appeared my report on the stranding. It had been submitted prior 
to Cockrunfs publication. It had all the details and the photos that I gathered on 
6 February 1954. 

In 1959, I recorded all the known mass strandings in the world that I was able 
to obtain at that time. In it, I included the stranding at La Paz of 16 January 1954. 
I also stated that I had no information of a second stranding a short time after 
the first and at the same place; and that the dispatch of the Associated Press that 
Cockrum had used was in error. 

All this is important because: 1) I visited La Paz, 6 February 1954, while some 
sperm whales were still on the beach. I also gathered what I considered accurate 
information on the mass stranding. I continued correspondence and conversations 
with Mrs. Waters, and visited La Paz again in February 1955, 1956, 1957, and 
in December 1959; 2) Cockrum used a newspaper dispatch of doubtful accuracy — 
no accuracy, as it turned out; 3) the report of two strandings has now entered the 
popular literature (Nayman 1973); and 4) two recent mass strandings of sperm 
whales have raised much interest in the phenomenon, often with references to 
earlier cases. The two recent mass strandings were: 1 January 1979, 56 sperm 
whales, 9 of which were females, at a beach between Mulege and Santa Rosalia, 
eastern coast of mid Baja California (see Payne 1979); and 16 June 1979, 41 sperm 
whales — 16 males and 25 females — near Florence, Oregon (Rice 1979). 

I met Mr. Marvin Grisby some time in the early or mid 1950s. He operated a 
small plane out of San Diego for charter service to Baja California, and possibly 
was using his and rumored information to obtain business. 



134 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Cockrum in 1956 also briefly mentioned another mass stranding of sperm 
whales in Golfo California — this case accurately. In 1957, I gave full details with 
photos of this stranding: nine sperm whales, all males, at Cabo Tapoca, Sonora 
side of the upper Golfo, 12 April 1953. 

My visit to La Paz on 6 February 1954 was during air census of the Californian 
population of gray whales with Dr. Gifford C. Ewing, owner and operator of a 
plane at Scripps Institute of Oceanography, La Jolla, California. 

Incidentally, Nayman mentioned two dates for the alleged two mass strandings 
of 1954: 3 February for 24 whales, and 27 February for 34. These dates were not 
in Cockruufs report, and presumably, from the context of his report were not in 
the dispatch from the Associated Press. All this adds to the confusion of Grisby, 
Associated Press, Cockrum, and Nayman. 

I am indebted to Mr. Robbin B. Patten for the reference to and the quotation 
from Nayman. 

Literature Cited 

Cockrum, E. L. 1956. Sperm-whales stranded on the beaches of the Gulf of California. J. Mamm., 

37(2):288, May. 
Gilmore, R. M. 1957. Whales aground in Cortes' Sea. Pac. Disc, Calif. Acad. Sci., S.F., 10(1):22- 

27. 
. 1959. On the mass stranding of sperm-whales. Pac. Naturalist, Beaudette Found. Biol. 

Research, Solvang, 1( 10):9— 16. 
Nayman. J. 1973. Whales, dolphins and man. Hamlyn Publ. Group, Ltd., London, pp. 128. 
Payne, M. 1979. Burning whales. Oceans [Mag.], S.F., 12(2):61-63. 
Rice, D. 1979. Preliminary report on the mass stranding of sperm whales at Florence, Oregon, on 

16 June 1979. Natl. Marine Mammal Lab., NW & Alaska Fisheries Center, Natl. Marine 

Fisheries Service, Seattle, pp. 6, map. Mimeo. 

Accepted for publication November 20, 1980. 

Raymond M. Gilmore, Natural History Museum, P.O. Box 1390, San Diego, 
California 92112. 



Bull. Southern California Acad. Sci. 
79(3), 1980, pp. 135-136 

INDEX TO VOLUME 79 

Adenostroma fasciculatwn, 5 

Braga occidentalis Boone, 130 

Braga patagonia Schioedte and Meinert, 130 

Brusca, Richard, see Melinda A. Thun 

Carcharhinus leucas, 121 

Carothers, Linn E., see George F. Howe 

Chaffee, Chet and David R. Lindberg: Morphological Variations of the Carinal 
Plate of the stalked Barnacle Pollicipes polymerus Sowerby, 78 

Coyer, James A.: Possible Cleaning Behavior by a Juvenile California Sheep- 
head, Semicossyphus pulcher (Labridae), 125 

de Wit, Leray A. and Thomas B. Scanland: Notes on Seasonal Collections of 

Sharks Near Point Conception, California, 112 
di Sciara, Giuseppe Notarbartolo and William A. Watkins: A Remora, Remilegia 

australis, attached to an Atlantic Spinner Dolphin, Stenella longirostris, 119 

Eilers, H. Peter: Ecology of a Coastal Salt Marsh After Long-Term Absence of 

Tidal Fluctuation, 55 
Eleutherodactylus , 1 3 

Fiske, Shirley, J. and J. C. Weibel: Navajo Social Interactions in an Urban 

Environment: An Investigation of Cognition and Behavior, 19 
Fritts, Thomas H., see Michael J. McCoid 

Gilmore, Raymond M.: One Mass Stranding, not Two, of Sperm Whales at La 

Paz, Baja California, 1954, 133 
Given, Robert R., see Nancy E. Muleady 

Hayes, Marc P. and Priscilla H. Starrett: Notes on a Collection of Centrolenid 

Frogs from the Colombian Choco, 89 
Hendrickx, M. E., see R. K. G. Paul 
Hogue, Charles L.: Discovery of the Male Katydid Idiostatus viridis Rentz, 

with Descriptions and Biological Notes (Orthoptera: Tettigoniidae: Dectici- 

nae), 1 
Holothitria zacae, 87 
Howe, George F. and Linn E. Carothers: Postfire Seedling Reproduction of 

Adeno 'Stoma fasciculatwn H. and A., 5 

Idiostatus viridis, 1 
Index to Volume 78, 42 

Keyes, Raymond S., see David M. Stalls 
King, Jack Lester, see Harrington Wells 

Lindberg, David R., see Chet Chaffee 



136 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

McCoid, Michael J. and Thomas H. Fritts: Observations of Feral Populations 

of Xenopus laevis (Pipidae) in California, 82 
Muleady, Nancy E. and Robert R. Given: Extension of Range of Holothuria 

zacae, Deichmann 1937, 87 

Paul, R. K. G. and M. E. Hendrickx: Crustaceans in the Shrimp By-catch From 

off the Coasts of Sinaloa and Nayarit, Mexico, 109 
Pollicipes poly merits, 78 
Powell, Jerry A.: Five Insects to be Newly Established or Recolonized on Santa I 

Cruz Island, California (Dermaptera, Lepidoptera), 97 

Remilegia australis, 119 

Savage, Jay M.: A New Frog of the Genus Eleutherodactylus (Leptodactylidae) 

from the Monteverde Forest Preserve, Costa Rica, 13 
Savage, Jay M.: A Synopsis of the Larvae of Costa Rican Frogs and Toads, 45 
Scanland, Thomas B., see Leray A. de Wit 
Semicossyphus pulcher, 125 
Shockley, Gregory, see Joel D. Weintraub 
Stalls, David M. and Raymond S. Keyes: Stereotyped Motor Patterns in Two 

Captive Bull Sharks, Carcharhinus leucas, 121 
Starrett, Priscilla H., see Marc P. Hayes 
Stenella longirostris , 119 

Thun, Melinda A. and Richard Brusca: On the Status of the Eastern Pacific 
Cymothoid Fish Parasite Braga occidentalis Boone and Its Synonymy with 
B. patagonia Schioedte and Meinert (Crustacea: Isopoda: Cymothoidae), 130 

Watkins, William A., see Giuseppe Notarbartolo di Sciara 

Weibel, J. C, see Shirley J. Fiske 

Weintraub, Joel D. and Gregory Shockley: Use of Incisors to Identify Rodent 

Genera in Owl Pellets, 127 
Wells, Harrington and Jack Lester King: A General "Exact Test" for N x M 

Contingency Tables, 65 
Wicksten, Mary K.: Range Extensions of Four Species of Crangonid Shrimps 

in the Eastern Pacific Ocean (Decapoda: Crangonidae), 38 

Xenopus laevis, 82 



INSTRUCTIONS FOR AUTHORS 

tie BULLETIN is published three times each year (April, August, and November) and includes articles in English 
any field of science with an emphasis on the southern California area. Manuscripts submitted for publication 

lould contain results of original research, embrace sound principles of scientific investigation, and present data 
a clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide 

r contributors. Consult also recent issues of the BULLETIN. Authors should strive for directness and lucidity, 

;hieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous 

ference of pronouns, and logically placed modifiers. 

MANUSCRIPT PREPARATION 

It is strongly recommended that, before submitting a paper, the author ask qualified persons to review it. The 
jthor is requested to submit at least two additional copies with the original, on SV2 x 11 opaque, nonerasable 
iper, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. 
ootnotes should be avoided. Manuscripts which do not conform to the style of the BULLETIN will be returned to 
ie author. 

An abstract summarizing in concise terms the methods, findings, and implications discussed in the paper must 
^company a feature article. Abstract should not exceed 100 words. 

A feature article comprises approximately five to thirty typewritten pages. Papers should usually be divided into 
ie following sections: abstract, introduction, methods, results, discussion and conclusions, acknowledgments, and 
terature cited. Avoid using more than two levels of subheadings. 

A research note is usually one to six typewritten pages and rarely utilizes subheadings. Consult a recent issue 
f the BULLETIN for the format of notes. Abstracts are not used for notes. 

Abbreviations: Use of abbreviations and symbols can be determined by inspection of a recent issue of the 
ULLETIN. Omit periods after standard abbreviations: 1.2 mm, 2 km, 30 cm, but Figs. 1-2. Use numerals before 
aits of measurements: 5 ml, but nine spines (10 or numbers above, such as 13 spines). The metric system of 
eights and measurements should be used wherever possible. 

Taxonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International 
jOde of Botanical Nomenclature (Lawjouw et al. 1956), the International Code of Nomenclature of Bacteria and 
iruses (Buchanan et al. 1958), and the International Code of Zoological Nomenclature (Stoll et al. 1961). Special 
tention should be given to the description of new taxa, designation of holotype, etc. Reference to new taxa in 
ties and abstract should be avoided. 

The literature cited: Entries for books and articles should take these forms. 

McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. 

Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54:452-458. 

Brattstrom, B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), 
Univ. California Press, xii + 635 pp. 

Tables and figures (line drawings, graphs, or black and white photographs) should not repeat data contained in 
ie text. The author must provide numbers and short legends for tables and figures and place reference to each 
f them in the text. Legends should be typed on a separate sheet of paper and placed at the end of the manuscript. 
lustrations and lettering thereon should be of sufficient size and clarity to permit reduction to standard page size; 
rdinarily they should be no more than twice the size of intended reduction and should not exceed 8V2 by 1 1 inches 
l size. Photographs must be printed on glossy paper. Submit one photoduplicated copy of each illustration. All 
lustrations accompanying Research Notes will be reduced to one column width. All half-tone illustrations will have 
ght screen (grey) backgrounds. Special handling such as dropout half-tones, special screens, etc., must be requested 
y and will be charged to authors. 

A cover illustration pertaining to an article in the issue or one of general scientific interest will be printed on the 
over of each issue. Such illustrations along with a brief caption should be sent to the Editor for review. 

PROCEDURE 

All manuscripts should be submitted to the Editor, Robert J. Lavenberg, Los Angeles County Natural History 
luseum, 900 Exposition Blvd., Los Angeles, California 90007. Evaluation of a paper submitted to the BULLETIN 
egins with a critical reading by the Editor; several referees also check the paper for scientific content, originality. 
nd clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent 
5 the author at which time he or she may be requested to rework portions of the paper considering these rec- 
mmendations. The paper then is resubmitted and may be re-evaluated before final acceptance. 

Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He or she 
hould promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations, legends, 
nd bibliographical references. He or she marks corrections on the galley (copy editing and proof procedures in 
'tyle Manual) and promptly returns both galley and manuscript to the Editor. Manuscripts and original illustrations 
/ill not be returned unless requested at this time. All changes in galley proof attributable to the author (misspellings, 
iconsistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with 
ie printer, not the Editor. 



CONTENTS 

Notes on a Collection of Centrolenid Frogs from the Colombian Choco. 

By Marc P. Hayes and Priscilla H. Starrett 89 

Five Insects Believed to be Newly Established or Recolonized on Santa 

Cruz Island, California (Dermaptera, Lepidoptera). By Jerry A. 

Powell 97 

Crustaceans in the Shrimp By-catch from off the Coasts of Sinaloa and 

Nayarit, Mexico. By R. K. G. Paul and M. E. Hendrickx 109 

Research Notes 

Notes on Seasonal Collections of Sharks Near Point Conception, California. By Leray A. 

cie Wit and Thomas B. Scanland 112 

A Remora, Remilegia australis, Attached to an Atlantic Spinner Dolphin, Stenella longi- 

rostris. By Giuseppe Notarbartolo di Sciara and William A. Watkins 119 

Stereotyped Motor Patterns in Two Captive Bull Sharks, Carcharhinus leucas. By David 

M. Stalls and Raymond S. Keyes 121 

Possible Cleaning Behavior by a Juvenile California Sheephead, Semicossyphus pulcher 

(Labridae). By James A. Coyer 125 

Use of Incisors to Identify Rodent Genera in Owl Pellets. By Joel D. Weintraub and 

Gregory Shockley 127 

On the Status of the Eastern Pacific Cymothoid Fish Parasite Braga oecidentalis Boone, 

and Its Synonymy with B. patagonica Schioedte and Meinert (Crustacea: Isopoda: 

Cymothoidae). By Melinda A. Thun and Richard C. Brusca 130 

One Mass Stranding, Not Two, of Sperm Whales at La Paz, Baja California, 1954. By 

Raymond M. Gilmore 133 

Index to Volume 79 135 



COVER: Stereotyped predictable behavior of a bull shark, Carcharhinus leucas. The bull shark is 
seen beneath an Atlantic lemon shark, Negaprion brevirostris. These sharks were photo- 
graphed in captivity by David M. Stalls. 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




ULLETIN 




Number 1 



APRIL 1981 



Southern California Academy of Sciences 

Founded 6 November 1891, incorporated 17 May 1907 

© Southern California Academy of Sciences, 1981 



OFFICERS 

Fred G. Hochberg, President 

Richard E. Pieper, Vice President 

Camm C. Swift, Secretary 

John J. Baird, Treasurer 

Robert J. Lavenberg, Editor 

Gretchen Sibley, Assistant Editor 



1979-1981 

John Baird 

Jules Crane 

Fred G. Hochberg 

Richard E. Pieper 

Gloria Takahashi 



BOARD OF DIRECTORS 

1980-1982 

Takashi Hoshizaki 
John D. Soule 

Steven N. Murray 
Camm C. Swift 

Robert G. Zahary 



1981-1983 

Kristin H. Berry 
Peter Castro 

Peter L. Haaker 
Martin L. Morton 
Richard M. Straw 



Membership is open to scholars in the fields of natural and social sciences, and to any person 
interested in the advancement of science. Dues for membership, changes of address, and requests for 
missing numbers lost in shipment should be addressed to: Southern California Academy of Sciences, 
the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 90007. 

Annual Members $ 12.00 

Student Members 8.00 

Life Members 150.00 

Fellows: Elected by the Board of Directors for meritorious services. 



The Bulletin is published three times each year by the Academy. Manuscripts for publication should 
be sent to the appropriate editor as explained in "Instructions for Authors" on the inside back cover 
of each number. All other communications should be addressed to the Southern California Academy 
of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, Los 
Angeles, California 90007. 

Date of this issue 16 December 1981 



Bull. Southern California Acad. Sci. 

80(1), 1981, pp. 1-12 

© Southern California Academy of Sciences, 1981 

A New Species of Murre, Genus Uria, from the 
Late Miocene of California (Aves: Alcidae) 

Hildegarde Howard 

Abstract. — A new species of murre, genus Uria, from the late Miocene of 
California (Aves: Alcidae) by Hildegarde Howard, Bull. Southern California 
Acad. Sci., 80(1): 1 — 12, 1981. A new species of murre, Uria brodkorbi, is repre- 
sented by impressions of the skull and incomplete skeleton of one individual on 
two slabs of diatomite from the late Miocene Sisquoc Formation, exposed near 
Lompoc, California, U.S.A. This is the seventh species of fossil bird to be de- 
scribed from this formation. Uria brodkorbi is very similar morphologically to 
the Recent Uria aalge that is common along the California coast today, but differs 
in being of heavier build and having a shorter beak, a stronger sternum and more 
massive wings. 



Introduction 



A previously unrecorded fossil bird specimen from the diatomite deposits of 
the Sisquoc Formation near Lompoc, California, was recently made available to 
me for study and description through the generosity of Dr. Pierce Brodkorb, of 
the University of Florida. The new fossil consists of the impressions of the skull 
and the anterior elements of the skeleton on two contiguous slabs of diatomite; 
it represents a new species of murre, family Alcidae. 

Six species of marine birds were described from the same late Miocene deposits 
at Lompoc by Miller (1925). These include three species of the booby-gannet 
family (Pelecaniformes: Sulidae): Miosula media, Sula willetti and Morns lom- 
pocanus; a shearwater (Procellariiformes: Procellariidae), Puffinus diatomicus; 
a godwit (Charadriiformes: Scolopacidae), Limosa vanrossemi; and an auklet 
(Charadriiformes: Alcidae), Cerorhinca dubia, of smaller size than the alcid spec- 
imen now at hand and related to the puffins rather than to the murres. 

Although the number of fossil bird specimens collected from the Sisquoc For- 
mation has nearly doubled since the time of Miller's report (1925), no additional 
species have been described from the deposits. The holotypes and many of the 
referred specimens are in the collections of the Museum of Paleontology at the 
University of California, Berkeley. The others are in the Natural History Museum 
of Los Angeles County and the California Academy of Sciences. Most of the 
approximately 20 avian specimens now known from these deposits are skeletal 
imprints in diatomite with no bones remaining. One specimen, assigned to Cero- 
rhinca dubia (LACM 74068), retains badly fragmented wing bones. 

The specimen now at hand is in the collection of Dr. Pierce Brodkorb (PB 
7960). It was acquired by Dr. Brodkorb many years ago under a Cooperative 
Agreement with the Florida State Geological Survey. Dr. Brodkorb was unable 
to learn particulars regarding the collection of the specimen. The label data read, 
"Sisquoc Formation, Santa Barbara Co., Cal. Diatomaceous earth quarries in 
this area. Johns Manville Co. ,, 



2 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Photographs of PB7960 viewed in certain lights cause the image of the impres- 
sions to be reversed, so that the skeletal elements appear in relief (Figs. 1 and 2). 
In this aspect the skeleton so closely resembles that of the present-day murres, 
genus Una, that had the deposit been of Pleistocene age, the specimen might 
have been identified as the Common Murre, Uria aalge, that is found along the 
California coast today. More detailed study, however, reveals important differ- 
ences. Therefore, a new species is here described. 

Methods and Materials 

In order to observe more closely the details of the skeleton of this fossil murre, 
latex molds were made of the impressions in the diatomite. Two sets of molds 
were prepared, each made directly from the slabs in an effort to assure maximum 
accuracy of detail. One set remains with the holotype, the other is in the cast 
collection of the Natural History Museum of Los Angeles County. Although the 
skeletal impressions in the diatomite slabs were checked for measurements, the 
actual study of the specimen was based largely on the latex molds. 

Abbreviations. — The following acronyms are used for specimens cited in the 
text: ANSP, Academy of Natural Sciences, Philadelphia; CSULB, California 
State University, Long Beach; LACM, Natural History Museum of Los Angeles 
County; PB, Pierce Brodkorb collection; USNM, National Museum of Natural 
History, Smithsonian Institution. 

Bone terminology follows Howard (1929). 

Comparative material. — Recent: Complete skeletons of Uria aalge californica 
(17), U. a. inornata (6), U. lomvia lomvia (2, one lacking complete skull), U . I. 
arra (10), from the collections of CSULB, LACM and PB. Also disassociated 
elements of Uria spp. from Aleut middens in Amchitka, Alaska (CSULB): 48 
humeri, 45 coracoids and 36 carpometacarpi. With the exception of the genus 
Alle, specimens representing all other North American genera of Alcidae were 
also compared. 

Fossil: Uria antiqua (Marsh 1870), cast of holotype humerus (ANSP 13357); 
Uria affinis (Marsh 1872), cast of holotype humerus (ANSP 13358); Australca cf. 
grandis Brodkorb 1955, proximal and distal ends of humeri (USNM 192758 and 
178136) and ulnae (USNM 193326 and 215652, complete coracoid (USNM 215513) 
and proximal end of carpometacarpus (USNM 215906) from the Lower Pliocene 
Yorktown Formation, Lee Creek, North Carolina; Miocepphus cf. mcclungi 
Wetmore 1940, humerus (USNM 25668) from the Middle Miocene Calvert For- 
mation, Maryland; lUria sp., proximal end of humerus (LACM 52018) from the 
late Miocene Monterey Formation, Orange County, California. 

Systematics 

Class Aves Linnaeus 1758 

Order Charadriiformes (Huxley 1867) 

Family Alcidae Vigors 1825 

Subfamily Alcinae (Vigors 1825) 

Genus Uria Brisson 1760 

In the fossil specimen from Lompoc, as exposed on the diatomite slabs and 
shown in the latex molds, the following characters shared with Uria, Alca and 



NEW SPECIES OF LATE MIOCENE MURRE 3 

Pinguinus (genera that Storer (1960) groups together in the tribe Alcini) are ob- 
servable: sharply ridged temporal fossa of the cranium, carpometacarpus with 
long process of metacarpal I, and humerus with depressed, ovoid pectoral at- 
tachment and with external tuberosity projecting anconad. 

The straight beak and well developed wing elements immediately distinguish 
the fossil specimen from Pinguinus. The straight beak also distinguishes the fossil 
from the genus Alca. Other distinctions from Alca include the more rounded 
shaft of the humerus and the less acute bend in the anconal profile of its distal 
end, and a narrower coracosternal connection. In all of these characters, as well 
as in the general proportions of the skeleton, the fossil resembles the murres of 
the genus Uria. 

Uria brodkorbi new species 
Figures 1-4 

Holotype. — PB7960, consisting of impressions of skull and anterior portion of 
skeleton on two slabs of diatomite. Slab PB7960A contains the skull in lateral 
aspect, and seven cervical vertebrae: slab PB7960B contains the incomplete fur- 
cula, sternum, coracoids, scapula, ribs and wing bones. 

Plastotypes. — Latex molds made from the holotype PB7960A and PB7960B are 
stored with the holotype and at LACM. 

Formation and age. — Sisquoc Formation, late Miocene, Clarendonian land 
mammal age. 

Locality. — Johns Manville diatomite quarry near Lompoc, California. Collector 
and date of collection unknown. 

Etymology. — The new species is named in honor of Dr. Pierce Brodkorb. 

Diagnosis. — Pre maxillary symphysis shorter than in Uria aalge or U. lomvia 
arra, closer to U. lomvia lomvia. Height of mandible at angular greater than in 
either Recent species of Uria. Sternum with broadly curved anterior margin, in 
contrast to straight contour dorsal to forward thrust of the carinal apex typical 
of Recent Uria ; tip of carina more truncated and anterior carinal margin more 
protruded anteriorly. Coracoid with well developed procoracoid as in Recent 
Uria, but with the tip sharper and more upturned; scapular facet more deeply 
cup-shaped than in U. aalge, and rounder, less oval than in U. lomvia; glenoid 
facet broader than in either Recent species. Humerus with area below head nar- 
rowed by flange extending mediad from pectoral attachment below external edge 
of head, and, internally, by raised area marking medial extent of capital groove; 
line of M. latissimus dorsi anterioris not paralleling shaft as in Recent species of 
Uria, but slanting palmad from distal edge of pectoral scar along external side of 
shaft, nearly 20 mm in length. Distal metacarpal symphysis of carpometacarpus 
with metacarpal III sloping distad rather than forming distinct right angle as in 
U. aalge; closer to U. lomvia, but distal margin more raised. 

Measurements (in millimeters). — Because of the condition of the specimen, 
many of the measurements are based on the latex mold, with the skeletal impres- 
sions used as a check. At best all measurements are approximate. 

Skull: Greatest length 93.6; length of rostrum 53.0; length of premaxillary sym- 
physis 25; greatest height of mandible 12.5. 

Sternum: Length from carinal apex to tip of posterior lateral process 122; height 
from carinal apex to ventral lip of coracoidal sulcus 36. 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 1 . Una brodkorbi, holotype in diatomite slabs PB7960A (top) and PB7960B (bottom). Lighting 
of photograph makes impressions appear in relief. Length from tip of beak to posterior tip of sternum 
338 mm. 



NEW SPECIES OF LATE MIOCENE MURRE 




Fig. 2. Uria brodkorbi, holotype slab PB7960B showing impression on left side. Depending on 
the angle viewed, the skeletal elements appear raised or impressed. Length of complete carpo- 
metacarpus 47.5 mm. 



Coracoid: Length from foremost (anterior) edge of coracohumeral surface to 
external tip of sternal facet 42.5; depth of shaft to tip of procoracoid 11.5; breadth 
of glenoid facet 6.5; length of glenoid facet 8.0; distance from procoracoid to 
foramen 6.6. 

Humerus: Greatest length 90 approx.; breadth of proximal end across external 
and internal tuberosities 18; proximodistal height of head 7; length of pectoral 
attachment through external tuberosity 12; greatest breadth of pectoral attach- 
ment 4; greatest distance from distal end (externally) to point of contact of ect- 
epicondylar prominence with shaft 12.3; depth of external side of distal end 8.0; 
depth of shaft above distal end 7.5. 

Ulna: Depth of distal end, externally, 8.0. 

Radius: Length 66 approx. 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




V 

Fig. 3. Uria brodkorbi, latex molds of holotype showing left side. Length from tip of beak to 
posterior tip of sternum 338 mm. 



NEW SPECIES OF LATE MIOCENE MURRE 




Fig. 4. Uria brodkorbi, humerus on latex mold of holotype. Approximately natural size. 



Carpometacarpus: Greatest length, externally, 47.5; length of process of meta- 
carpal I, 9.3; depth of distal end from internal tuberosity of metacarpal II through 
distal metacarpal symphysis, measured on right carpometacarpus, 8.0. 

Wing phalanges: Lengths: digit I, phalanx 1, 21.2; digit II, phalanx 1, 20.8; 
digit II, phalanx 2, 21.5; digit III, phalanx 1, 9.3. 

Description. — As exposed on the diatomite slabs and shown in the latex molds, 
the left side of the skull and sternum and the elements of the left side of the 
skeleton are best preserved. 

Although the cranial part of the skull is incompletely preserved, the rounded 
posterior contour of the supraoccipital and the narrow, sharply ridged temporal 
fossa are distinguishable. The latex mold (Fig. 3) defines the extent of the short 
beak, which more closely resembles that of Uria I. lomvia than any of the adult 
specimens of either U. I. arm or U. aalge at hand. The mandible depth is even 
greater than in U. I. arra (see Table 1). 

The position of the furcula is indicated on holotype slab PB7960B by two holes 
anterior to the coracoids. Filled with latex in preparing the molds, the poorly 
defined tips appear, but reveal no dependable characters. 

The sternum is damaged posterior to the coracoidal sulcus, and the area of the 
costal ridges is pushed forward and folded onto the carina so that from this point 
to the posterior end the dorsal surface is exposed. Because of this damage, the 
total length as measured on the specimen (and given above) could be as much as 
10 mm less than the actual length of the sternum. The anterior contour of the 
carina shows clearly in lateral view (Fig. 2) as a broad arch, in contrast to the 
straight contour dorsal to the forward thrust of the apex typical of the Recent 
species of Uria. Further distinctions lie in the more truncated tip and the more 
forward-protruding flange along the anterior carinal margin. Only one Recent 
specimen of murre (U. aalge californica, LACM 674) has a suggestion of these 
characters. 



8 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

The brachial tuberosity is poorly shown on the right coracoid, and is not visible 
on the left, which is otherwise better preserved. The glenoid facet is relatively 
broader than in the modern species of Uria. The scapular facet is deeper than in 
U. aalge, being closer to U. lomvia in this respect, though more rounded, rather 
than oval. The length of the procoracoid resembles the condition in Recent Uria, 
but the projection is sharper and more upturned than in most of the specimens 
examined (see Fig. 2). The small foramen is well below the tip of the procoracoid; 
the size and position of the foramen is variable in the sample of specimens of the 
Recent species. Although the sternal facet of the coracoid is not exposed, the 
shape of the coracoidal sulcus of the sternum suggests that the sternal end of the 
coracoid is long and narrow as in Uria, with the facet possibly even more laterally 
extensive than in the Recent species. This condition is in contrast to that found 
in the puffins, or even in Alca, in which this articular surface is shorter and 
deeper. 

The left scapula is incompletely revealed in ventral view (Fig. 2). Neither the 
acromion nor the coracoidal articulation is visible, and the shape of the glenoid 
facet is not clear. 

The broken left humerus (Fig. 4) lies with the proximal end presenting an aspect 
slightly lateral of anconal, while the distal fragment is turned so as to expose 
more of the external side. The proximal end of this element shows the most 
notable differences from the Recent species of Uria. As observed in the latex 
mold (Fig. 4), the pectoral attachment is broad, with a distinct flange extending 
below the head, narrowing the area between the attachment and the median crest; 
the head appears to be less sharply undercut. A flange of variable extent occurs 
in a few specimens of both U. aalge and U. lomvia, but in none is it as strongly 
developed or as evenly contoured as in the fossil. The line of the anterior latis- 
simus dorsi muscle in U. brodkorbi is as long as in Recent Uria, but the slope 
is more markedly palmad and is emphasized by the more rounded anconal aspect 
of the shaft. Distally, the external tricipital groove is sharply rimmed; the ent- 
epicondyle is incomplete and the width of the internal groove is not clear. How- 
ever, the slight anconal flare toward the distal end, as shown in the latex mold 
(see Fig. 3) suggests that the internal groove is broader than the external, thus 
resembling the condition in Uria. This contrasts with Alca in which the anconal 
contour of the shaft bends palmad, and the two grooves are of equal size. The 
tip of the ectepicondylar process is more prominent than in most specimens of 
Recent Uria. There is, however, considerable variation in the Recent series, and 
a few specimens approach the same prominence, although only one (of maximum 
size) is as elevated above the distal end, and the ectepicondylar process is less 
vertically placed with respect to the shaft. 

The ulna and radius are crushed proximally, and provide little information other 
than approximate length of the radius. 

The external side of the complete left carpometacarpus, and the internal side 
of the distal end of the right carpometacarpus are exposed. In overall length, as 
well as in the length of the process of metacarpal I, the element falls within the 
size range of U. I. arra (see Table 1). The rounded distal contour of metacarpal 
III (see Fig. 2) is approached in some specimens of U. lomvia. 

Comparison with previously recorded fossil Alcidae. — Three extinct species of 
Uria have been previously described, each on the basis of the humerus: Uria 



NEW SPECIES OF LATE MIOCENE MURRE 9 

antiqua (Marsh 1870) from the Lower Pliocene of North Carolina; U. affinis 
(Marsh 1872) from the Pleistocene of Maine; and U. ausonia Portis 1887 from 
the Pliocene of Italy. U. ausonia is based only on the distal end of a humerus, 
which I have not examined. 

A cast of the holotype of each of the other two species is available. Both are 
longer than the humerus in the holotype of Uria brodkorbi (U. antiqua length 
96.2 mm, U. affinis length 95.0 mm as given by Marsh (1870 and 1872), and 
closely approximated on the casts), with greater breadth between the pectoral 
attachment and the median crest. Both also have the line of M. latissimus dorsi 
anterioris paralleling the shaft as in Recent species of Uria, in contrast to U. 
brodkorbi in which the muscle line slopes palmad. U. affinis further resembles 
Recent Uria, distally, in the unequal breadth of the tricipital grooves, the internal 
being wider, as appears also to be true of U. brodkorbi. In U. antiqua the grooves 
are of equal breadth. The humerus of U. antiqua resembles that of U. brodkorbi in 
the rounding of the shaft toward the proximal end, although the shaft is heavier 
and the apex more anconal in position in U. antiqua. 

There is one previous tentative record (Howard 1978) of Uria from the Miocene 
of California. The record is based on an incomplete proximal end of a humerus 
(LACM 52018) from locality LACM 6902 in the late Miocene Monterey Formation 
of Orange County. The specimen is slightly smaller than the humerus of the 
approximately contemporaneous U. brodkorbi, lacks the flange from the pectoral 
attachment to the head, and is more excavated below the head. It resembles the 
humerus of U. brodkorbi, however, in the rounding of the shaft and the palmad 
trend of the line of M. latissimus dorsi anterioris. 

Two extinct genera are worthy of consideration in comparison with U. brod- 
korbi: Australca Brodkorb 1955, genotype A. grandis from the Middle Pliocene 
Bone Valley Formation of Florida; and Miocepphus Wetmore 1940, genotype M. 
mcclungi from the Middle Miocene Calvert Formation of Maryland. The holotype 
of A. grandis is a coracoid, with humerus, radius, ulna, carpometacarpus and 
tibiotarsus referred (Brodkorb 1955). The holotype of M. mcclungi is a humerus. 
This and a subsequently referred humerus (Wetmore 1943) from the type locality 
are the only recorded specimens of Miocepphus. 

Through the kindness of Dr. Storrs L. Olson, of the National Museum of 
Natural History, specimens that he considers referable to Australca and Mio- 
cepphus have been made available (see Comparative Material above). The Aus- 
tralca material is similar in size to measurements given for A. grandis (Brodkorb 
1955), but is not from the type locality. The Miocepphus humerus, however, 
comes from the same formation as the holotype of M. mcclungi, though in Zone 
13 of the Calvert Formation (the type is from Zone 12). The specimen is inter- 
mediate in size between the holotype (Wetmore 1940) and the referred specimen 
(Wetmore 1943) of M. mcclungi, both of which are markedly smaller than the 
humerus of U. brodkorbi. As size is not a generic character, the specimens of 
both Australca and Miocepphus are analyzed on the basis of their qualitative 
characters. 

The major distinction of the Australca coracoid from that of U. brodkorbi lies 
in the more posterior position of the procoracoid (inset from the median edge of 
the coracoidal shaft). This character is shown clearly in both the referred speci- 
men at hand and in the illustration of the holotype (Brodkorb 1955:50, fig. 24). 



10 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



en <o — 

© >/"> On 



fNl r-; o 

© — ■ — 



t— nO 00 


— o ^r 


r^ \D — 


ON (N 


Tfr — 


oo — — 



* <D 

E E 



5i 



S u u 

i2 j2 j= 

C/3 b? C/5 

« E « 

P « « 



NO </"> 

tT O 
-<t — 



o< "5. 
o ? 



TJ- t-~ nC 

rn r~ — 

ON — — 



m v> m 



ri — 




(N 


NO 


ON 

On 


oc 


CM 




■* 


t-; 


■/"> 


r-- 


tt 


q 




^r 


on 


m 


oo 


sd 


Z 



«n 


_- 


,— , 


_ 


— 


v~» 


ON 


r- 


— . 


</~> 


o^ 


CM 




rf 






00 

x: 

o 

u 






NO 
>< 

o 

■_ 

a 


vC 


© 


w-i 


u-> 


>o 


>/"> 


rt 


q 


ro 


n3 


c*i 


>/-, 


c-i 


CM 


^-4 


NO 


O 


oc 


CM 


nC 


3N 


CS 


— 


Tf 


>— ' 




ON 


— 


— 


NO 



« s fl 



E ■§ 






a a, a g w q cq 
u 



C 1- C 



a a, w -o j 

TO 



oo oo 
no oo 



ON ON 



& B O 

rt u p 



U 



W J 



kj u 



■a E 



£ <4- 



E'-S 



E— en 
O 



u 


E 




u 


ed 




CJ 


K) 




>, 


"3 


on 



E § 
Z oo 



NEW SPECIES OF LATE MIOCENE MURRE 1 1 

With respect to this character comparison with U. brodkorbi is best observed on 
the right coracoid of the holotype in which the close proximity of the lower end 
of the procoracoid to the median edge of the shaft is discernible on the latex 
mold. In the humerus of Australca, the area below the head is broader and the 
pectoral scar longer and relatively flatter than in U. brodkorbi; distally the apex 
of the ectepicondylar process is more prominent and forms a more acute angle 
with the shaft. Although the Australca carpometacarpus resembles that of U. 
brodkorbi in the long process of metacarpal I, the angle between the process and 
the external trochlear crest is less acute. U. brodkorbi resembles the Recent 
species of Uria in this character. 

The humerus of Miocepphus is similar to that of U. brodkorbi in having a 
flange from the pectoral attachment narrowing the area below the head. The 
attachment itself, however, is flatter and less ovoid, and the external tuberosity 
is less prominent anconally. The line of the M. latissimus dorsi anterioris slopes 
slightly palmad, as in U. brodkorbi, but it is shorter. Distally, the tricipital 
grooves are equal in size as in Alca, as distinguished from the condition in Uria. 
The ectepicondylar process is even more prominent than in Australca, and ter- 
minates in a distinct papilla. 

Summary and Conclusions 

The holotype skeletal impression of Uria brodkorbi, transformed by latex 
molds into bones in relief, provides a rare opportunity to study the morphology 
of the associated elements of one individual fossil bird. 

Viewed as a whole, the skeleton resembles that of the murres, genus Uria. In 
relation to Recent species of the genus it appears to have been of sturdier build, 
with a stronger sternum and a short beak. Although in length the coracoid and 
humerus are within the size range of U. aalge calif ornica (see Table 1), the 
breadth of these elements equals or surpasses those of the larger U. lomvia arra. 
The carpometacarpus is similar to some specimens of U. lomvia in the rounding 
of the distal end of metacarpal III, and closely resembles U. I. arra in size. 

Some of the individual elements, if found isolated, might well suggest generic 
distinction from Recent Uria. This is particularly true of the sternum and hu- 
merus. However, the large series of modern skeletons studied reveals trends 
toward one or more of the apparently distinctive characters. Although no single 
Recent skeleton of Uria has a combination of characters matching those of the 
fossil, there is sufficient similarity within the comparative series to indicate that 
U. brodkorbi is entirely consistent with what would be expected in a Miocene 
ancestor within the genus. 

Therefore, the evidence weighs in favor of the new species being retained in 
the genus Uria, rather than in the erection of a new genus. 

Acknowledgments 

I am especially indebted to Pierce Brodkorb for the opportunity to study this 
important specimen from the Sisquoc Formation, and for providing the photo- 
graph taken by Robert D. Weigel (Fig. 2). I am also deeply grateful to my hus- 
band, Henry Anson Wylde, for preparing the latex molds of the specimen and 



12 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

for photographing Figures 1 , 3 and 4. Richard Meier (LACM) assisted by making 
the excellent print of the photograph for Figure 1. 

My thanks are also extended to Storrs L. Olson, Stuart L. Warter, Jean Geary 
and Diana Matthiesen for providing comparative material. My continuing appre- 
ciation is extended to members of the two science divisions of the Natural History 
Museum of Los Angeles County for their cooperation. 

The following persons critically reviewed the manuscript, providing helpful 
suggestions: Lawrence G. Barnes, Pierce Brodkorb, Kenneth E. Campbell, and 
Storrs L. Olson. 

Literature Cited 

Brodkorb, P. 1955. The avifauna of the Bone Valley Formation. Florida Geological Survey Report 

of Investigations, No. 14:i-iii, 1-57. 
Howard, H. 1929. The avifauna of Emeryville Shellmound. Univ. Calif. Publ. Zool., 32(2):301-394. 

Osteological terminology and illustrations reprinted 1980, in Papers in Avian Paleontology 

honoring Hildegarde Howard, K. E. Campbell, Jr., ed. Contrib. Sci. Nat. Hist. Mus. Los 

Angeles Co., 330:xxvii-xxxviii. 
. 1978. Late Miocene marine birds from Orange County, California. Natural History Museum 

of Los Angeles County Contrib. Sci., 290:1-26. 
Marsh. O. C. 1870. Notice of some fossil birds from the Cretaceous and Tertiary formations of the 

United States. Amer. Jour. Sci. ser. 2, 49(146): 205-2 17. 
. 1872. Notice of some new Tertiary and post-Tertiary birds. Amer. Jour. Sci., ser. 3, 

4(22): 256-262. 
Miller, L. 1925. Avian remains from the Miocene of Lompoc, California. Carnegie Inst. Wash. Publ., 

349:107-117. 
Storer, R. W. 1960. Evolution in the diving birds. Proc. Xllth Internat. Ornith. Cong., Helsinki, 

1958, pp. 694-707. 
Wetmore, A. 1940. Fossil bird remains from Tertiary deposits in the United States. Jour. Morphol., 

66(l):25-37. 
. 1943. A second specimen of the fossil Guillemot, Miocepphus. Auk, 60:604. 

Accepted for publication 19 February 1981. 

Chief Curator Emeritus, Natural History Museum of Los Angeles County, 900 
Exposition Boulevard, Los Angeles, California 90007. 



Bull. Southern California Acad. Sci. 

80(1), 1981, pp. 12-22 

© Southern California Academy of Sciences, 1981 

Reproduction of the Onespot Fringehead, Neoclinus uninotatus, 
in Monterey Harbor, California 

David G. Lindquist 

Abstract. — Reproduction of the onespot fringehead, Neoclinus uninotatus in 
Monterey Harbor, Monterey, California by David G. Lindquist, Bull. Southern 
California Acad. Sci., 80(1): 12-22, 1981. Males have greater development of the 
primary ocular cirrus. Nesting males were observed January to March and June 
to September. Males guarded large egg masses attached to the ceiling and sides 



NEOCLINUS UNINOTATUS 13 

of the inner refuge. Eggs were 1 to 1.3 mm in diameter with one large yellow- 
orange oil droplet. Larvae hatched at a total length (=notochord length) of 5 mm. 



Introduction 



Except for the pioneering studies of Carlisle, Turner and Ebert (1964) and 
Turner, Ebert and Given (1969) on man-made artificial reefs and the work of 
Stephens and his associates on King Harbor fishes (Terry and Stephens 1976; 
Stephens and Ellison 1977; Ellison, Terry and Stephens, 1979; Ehrlich et al. 1979; 
Stephens 1978), little attention has been paid the life histories of the fishes living 
in and around man-made structures in California's marine environment. Further- 
more, these previous studies have been designed primarily to define the effects 
of man-made alterations of the marine environment on the fish community as a 
whole. My report is a preliminary analysis of reproduction in a population of 
Neoclinus uninotatus using a man-made wharf habitat. 

The natural history of the three California species of the genus Neoclinus is 
poorly known. Only a few minor observations of these fishes in nature have been 
published (Stephens et al. 1970 on N. stephensae and Feder, Turner and Lim- 
baugh 1974 on N. uninotatus, blanchardi and stephensae). The most detailed 
studies of the species of this genus were published by Japanese workers (Shiogaki 
and Dotsu 1972; Fukao 1980). 

Methods and Materials 

A total of 26 SCUBA dives (30.6 hours) was made under the Monterey Wharf 
#2 to observe and record populations of the onespot fringehead between June 
1969 and June 1971. Most observations (17 hours) were conducted during the last 
eight months of the study. During various phases of the study, specimens were 
collected using quinaldine, an anesthetic, mixed with ten parts of 70% isopropanol 
(Gibson 1967; Stephens et al. 1970). 

The study was conducted at the east end of the wharf (Fig. 1). Haderlie and 
Donat (1978) have prepared a detailed description of the configuration and con- 
struction of the wharf. 

To facilitate behavioral observations, a square grid was constructed of lines 
and cinder blocks in the study area (Fig. 1). The grid measured 15 m on a side 
and was subdivided into nine 5 x 5 m squares. I placed the grid 15 m east of the 
wharf pilings on a flat, relatively barren sand bottom in 10 m of water during 
November 1970. The three subsquares closest to the wharf initially contained 30 
bottles, three of which housed one each N. uninotatus. These three subsquares 
served as the experimental habitat, whereas the six subsquares furthest from the 
wharf remained undisturbed during the study. From December 1970, to March 
1971, 32 adult (size range = 100-180 mm TL, x = 134 mm TL) .V. uninotatus, 
within their refuges, and 195 clean bottles and jars (orifice diameter range = 16- 
28 mm, x = 18 mm), were placed randomly within the three experimental sub- 
squares. Care was taken to place the orifices of the refuges level with the sub- 
stratum to encourage use of them by N. uninotatus. The locations of individuals 
were marked on a prepared map of the grid. 



14 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



- N - 



SAN 
FRANCISCO 



MONTEREY BAY 



36°40' - 





MONTEREY 



FISHERMAN S 
WHARF 




1 1 2°00 




MARINA 



MONTEREY 



300 



MUNICIPAL 
WHARF NO.2 



METERS 



Fig. 1. Configuration of Monterey Harbor in relation to the study area. 



Results and Discussion 

Sexual dimorphism and function of the ocular cirri. — Although Clark Hubbs 
(1953) presented six morphometric comparisons of the sexes of N. uninotatus 
that indicated dimorphism, I was unable to use these to determine the sex of 
specimens in nature. Jordan and Evermann (1896-1900), in their description of 
N. uninotatus (misidentified as N. blanchardi) , pointed out that males had an 
ocular cirrus that was twice as long as the female's. Hubbs (1953) did not treat 
the ocular cirri in his account. However, his illustrations (Figs. 13 and 14) clearly 
indicate sexual differences in the ocular cirri development. I found the length and 
amount of branching in the primary ocular cirrus to be a reliable method of sexing 
individuals in nature. 

The male's primary ocular cirrus is not only longer but is also wider with more 
branching at the cirrus tip (Fig. 2). I devised a cirrus index (product of cirrus 
length and number of primary and secondary distal cirrus branches) as a measure 



NEOCLINUS UNINOTATUS 



15 




MALE 



1MM 




FEMALE 

Fig. 2. Tips of the primary ocular cirri for both sexes of N. uninotatus. Both male and female 
measured 120 mm SL: length of male's cirrus is 11 mm and female's is 6 mm. Total branching for 
male is 27 and for female is 1 1. 



of cirrus development. A plot of the cirrus index on standard length indicates a 
greater and more rapid development of the male's cirri (Fig. 3). The cirrus index 
of males appears more closely related to standard length as suggested by the 
regression coefficient, r. However, additional data are needed for females. 

J. S. Stephens, Jr. and H. D. Hickman, Jr. (Occidental College, pers. comm.) 
found N. uninotatus males in southern California with a mean primary orbital 
cirrus length of 1.7 orbits, never shorter than one orbit and females with a mean 
of 0.6 orbits, never greater than one orbit. They also found the male's cirrus was 
stouter and the branching was more palmate, distally. Randall (1966) presented 
data indicating that the number of ocular cirri of Hypleurochilus aequipinnis also 
increase with size. 

The tropical blennioid families Labrisomidae, Clinidae, Blenniidae, and Chae- 
nopsidae, are notable for the various development of cirri over the head (nasal. 



16 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



450 



400 



350 



300 



250 



2£ 200 

C9 



150 



100 



50 







V = 3.37 x- 176.9 
r-0.64 

P< 0.01 



MALES 



FEMALES 



y=1.03x-67.8 

r=.58 

P< 0.10 



95 105 115 125 135 145 155 165 

STANDARD LENGTH (MM) 

Fig. 3. Least squares regression analysis of cirrus development and SL for both sexes of N. 
uninotatus. 



ocular and nape cirri). However, the functional significance of cirri in blennies 
has been unstudied, by and large, by previous workers. Springer and Gomon 
(1975) examined the total nape cirri of Malacoctenus triangulatus over its entire 
geographic range and reported an inverse relationship between nape cirri and 
lateral-line scales. Based on observations of Ford (1959), who reported the cirri 
of Malacoctenus hubbsi to be innervated by at least three cranial nerves, Springer 
and Gomon (1975) inferred a sensory function for the cirri and suggested that M. 
triangulatus compensates for the reduction in the lateral-line pores by an increase 
in the nape cirri. Evidence for a sensory function for the cirri is strengthened by 
the work of Schulte and Holl (1972), who found nerve bundles and sensory organs 
resembling taste buds in the ocular cirri of Blennius tentacularis. 

Shiogaki and Dotsu (1972) describe the ocular cirri of Neoclinus bryope as 
"cryptic organs." Their photograph (Fig. 1) of both sexes also suggests that males 
have a greater development of the ocular cirri. Since male blennies, in general, 
spend much of their time resting with the head at the entrance of the refuge, it 



NEOCLINUS UNINOTATUS 17 

seems advantageous for selection to have favored a greater degree of camouflage 
(i.e. cirri development) for the male. The male's cirri blend in well with refuge 
encrusting bryozoans, hydroids, and algae, thus allowing the male a greater de- 
gree of crypticity and, thus, protection against predators while stationed at the 
entrance of the refuge. An alternative but non-exclusive hypothesis has been put 
forth by Zander (1975). He observed that the male's ocular cirri in four Mediter- 
ranean Blennius are significantly enlarged, especially during the breeding season. 
Zander reasoned that the enlarged cirri serve as sign stimuli that aid in attracting 
females to the spawning refuge. Still another possible function of the cirri is 
suggested by the fact that the form of the cirri is frequently species specific, such 
that the shape of the cirri often serve as diagnostic characteristics. Blennies, thus, 
may also recognize conspecifics at least partially on the basis of the form of the 
cirri. 

Spawning season. — A total of 16 nests, each guarded by a single male, were 
found in the study area during 12 SCUBA dives as follows: one in January; two 
in February; one in March; eight in June; one in August; and three in September. 
Observations at the wharf were made during all months of the year except May. 
These preliminary observations suggest that the wharf population has either a 
protracted spawning season from January to September with peak activity in 
early summer or two separate spawning seasons: January to March; and June to 
September. At present, I am inclined to favor the latter hypothesis because my 
observations of 24 mature males in refuges during four SCUBA dives in April 
revealed none guarding eggs. 

Feder et al. (1974) report N. uninotatus spawning in April and May. However, 
the location(s) of these observations is (are) not stated. The observations were 
made by the late Conrad Limbaugh (J. S. Stephens, Jr., pers. comm.) and could 
have been made as far north as Monterey Bay but were more likely to have taken 
place in southern California (i.e. San Diego and Orange Counties), where most 
of his intensive observations were made (Feder et al. 1974). J. S. Stephens, Jr. 
and H. D. Hickman, Jr. (pers. comm.) have observed gravid females of N. uni- 
notatus from late January through at least September and with males guarding 
eggs in July off Redondo Beach in Los Angeles County. 

Neoclinus blanchardi, the closest relative of N. uninotatus, spawns from Jan- 
uary through August (Feder et al. 1974). Shiogaki and Dotsu (1972) report inter- 
tidal spawning for N. bryope during January to April, with peak activity in March 
and April in Nomozaki, Japan (32°35'N latitude). Feder et al. (1974) report the 
young of N. stephensae in August and September. 

Nesting behavior. — In all cases, the 16 nesting males observed in nature were 
guarding eggs located within glass bottles or, in one case, a glass jar. Although 
the volume of the containers varied from approximately 0.5 to 2.0 1, most had 
larger orifices (e.g. 25-30 mm). The outisde of the glass containers were variously 
encrusted with sessile invertebrates. The refuges were embedded one to two cm 
into the substratum such that the bottom of the orifice was level with the sub- 
stratum. Eggs were deposited in large thick masses within the center section of 
the refuge. The eggs adhered to the ceiling and side walls of the refuge by means 
of adhesive filaments. The male positioned his body among the eggs with only 
the head protruding from the refuge. If the refuge was lifted by the diver, the 
male would depart but would return soon after the refuge was replaced. 



18 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

In January 1971, I discovered a male and female within a one-half 1 bottle. 
Both fish were approximately 140 mm in total length. This was the first evidence 
of spawning since September 1970. Eggs were present in one small opaque mass 
and adhered to the ceiling and one side of the bottle. Close inspection revealed 
the eggs to be in a very early stage of development, indicating that the egg mass 
had been laid recently. The size of the egg mass was estimated at 60 by 30 mm 
and 15 mm thick. Shortly after I disturbed the pair, the female was forced from 
the refuge by the male. One month later I returned and removed the male and his 
refuge to the laboratory for study. The egg mass now covered the entire ceiling 
and portions of both sides of the refuge. Three distinct developmental stages were 
present within the egg mass. The most developed embryos had fully formed eyes 
and the tail overlapped the head. I estimated the total number of eggs present at 
15,350 by the volumetric method (Lagler 1956). The male remained with the egg 
mass in the aquarium and fanned the eggs with his pectoral fins at a variable rate 
of 34 to 71 beats per minute (x = 50, N = 10). Many of the eggs were later 
attacked by fungus and none hatched. 

Shiogaki and Dotsu (1972) reported that Neoclinus bryope spawned within the 
dead tubular vermetid gastropod shells of Serpulorbis imbricatus. These eggs 
were deposited in a single layer on the inner walls of the tube and were also 
guarded by the male. Egg clusters consisted of a maximum of five different de- 
velopmental stages and eggs ranged in number from 90 to 250. 

Within the Blennioidea, the habit of living and spawning within tubular refuges 
occurs in some of the blenniids (Abel 1962, 1964; Fishelson 1975; Losey 1968, 
1976; Wickler 1965), in many of the chaenopsid blennies (Robins, Phillips and 
Phillips 1959; Bohlke and Chaplin 1968; Kerstitch 1971; Lindquist 1971, 1975; 
Thomson, Findley and Kerstitch 1979; Longley and Hildebrand 1940, 1941; Ste- 
phens, Hobson and Johnson 1966), and in a few labrisomids, namely Paraclinus 
marmoratus (Breder 1939, 1941) and the species of Neoclinus (Feder et al. 1974; 
Fitch and Lavenberg 1975; Lindquist 1971; Shiogaki and Dotsu 1972; Fukao 
1980). Although spawning within a refuge of some sort (below a stone or in a 
cave, etc.) seems to be the general rule for the blenniids (Wirtz 1977), such is not 
the case for clinids and labrisomids. Most clinids and labrisomids studied are 
either live bearers or open substrata spawners (Wirtz 1977). The few tripterygiids 
studied are apparently divided between cavity spawning and open substrata 
spawning (Wirtz 1977, 1978; Thomson et al. 1979). Unfortunately, because the 
spawning behavior and general habits of too few species of the five aforemen- 
tioned blennioid families have been studied, it would be premature to attempt 
any broad implications of evolutionary relationships based on these characteris- 
tics. 

However, Neoclinus does occupy a unique position among the labrisomids 
because of its tubiculous habits. (The sponge dwelling habits of Paraclinus mar- 
moratus appear to be a special case.) The tube dwelling habits of Neoclinus 
would then seem to align this genus more closely with the blenniids and chae- 
nopsids than with the labrisomids and clinids. Indeed, Hubbs (1952, 1953) placed 
Neoclinus in the subfamily Chaenopsinae and family Blenniidae. Later taxonomic 
revisions removed Neoclinus to the Clinidae (Springer 1955; Bohlke 1957) and 
elevated the chaenopsins to family level (Stephens 1963). In doing so, Neoclinus 
was considered to be the closest relative of the clinid ancestor that gave rise to 
the chaenopsids (Stephens 1961, 1963; Stephens and Springer 1971). George and 



NEOCLINUS UNINOTATUS 



19 




1MM 




1MM 



Fig. 4. An intermediate stage egg and preflexion larva of N. uninotatus . 



Springer (1980) elevated the subfamily Labrisominae, to which Neoclinus pre- 
sumably belongs, to family rank. Although the higher classification of the tropical 
blennioid groups is still tentative and subject to controversy (Bohlke and Robins 
1974; Rosenblatt and Stephens 1978), I believe that the close position of Neocli- 
nus to the chaenopsids and blenniids is suggested by their similar tube dwelling 
habits. 

Egg and larval description. — The eggs of N. uninotatus range in size from 1 
to 1.32 mm in diameter (.v = 1.19, N = 60). The eggs are spherical and sometimes 
slightly indented when packed tightly together. Each egg has many adhesive 
filaments by which it attaches to other eggs or to the substrata. A single large 
yellow-orange oil droplet is present and ranges in size from 0.25 to 0.33 mm in 
diameter (.v = 0.28 mm, /V = 30). Other much smaller oil droplets are also pres- 
ent. An intermediate stage egg with a larva having 14 myomeres is illustrated in 
Figure 4. Melanophores begin to develop ventrally along the post-anal myomeres 
shortly after passing the 14-myomere stage. 

Eggs were hatched in the aquarium shortly after removal from nature. The 



20 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

length of the development period in nature was not determined. The larvae swam 
near the top of the aquarium, suggesting that they were photopositive. The larvae 
hatched at an advanced stage of development (Fig. 4). Little yolk remained in 
the yolk sac at hatching. Dense melanophores are located dorsal to the heart, air 
bladder, and rectum, as well as ventral to the liver and rectum. A series of 29- 
32 small melanophores is present along the ventral margin of the myomeres. The 
myomere count 11 + 34-36 = 45-47. 

The 14-myomere stage of N. uninotatus and bryope (Shiogaki and Dotsu 1972) 
differ in respect to: 1) the head shape of the embryo; 2) the number of smaller oil 
droplets in the egg; 3) the presence of Kupfer's vesicles in N. bryope; 4) the 
development of the eyes; and 5) the size of the egg. The dorsal head profile of 
N. bryope is slightly pointed compared to the flat profile of N. uninotatus. There 
are 18 smaller oil droplets in the yolk of N. bryope compared to two or three in 
N. uninotatus. Kupfer's vesicle is not developed in the 14-myomere stage of N. 
uninotatus. The eyes are more completely developed in N. uninotatus. The eggs 
of N. bryope are slightly larger, 1.28 to 1.49 mm, than those of N. uninotatus. 

The 5.5 mm larva (Fig. 4) of N. uninotatus compares with the 7.5 mm larva 
of N. bryope (Shiogaki and Dotsu 1972) but differs in the following respects: 1) 
size; 2) placement and number of melanophores; 3) myomere count; and 4) size 
of the pectoral fins. Neoclinus bryope has more myomeres (13 + 35-37 = 48- 
50), hatches at a larger size (6.6 mm compared to 5.0 mm), has a large melano- 
phore over the yolk in the stomach, has fewer post-anal melanophores (17-20), 
and has a smaller pectoral fin. Shiogaki and Dotsu (1972) suggested that N. bryope 
spends a longer time in the plankton than other intertidal fishes because the 
juveniles attained a size of 25 to 29 mm (about one-half the adult size) before 
entering the benthic mode. The smallest benthic juvenile of N. uninotatus was 
46 mm SL, suggesting that N. uninotatus may also have a lengthy planktonic 
stage. 

Acknowledgments 

This study is an expanded portion of my M.A. thesis conducted at the Moss 
Landing Marine Laboratories of the California State Colleges and Universities. 
I thank the members of my thesis committee (Nybakken, chairman, Tomlinson, 
and Yarberry) for their guidance and support. John S. Stephens, Jr., of Occidental 
College, shared his unpublished observations on Neoclinus and has been a source 
of encouragement and enthusiasm for this study. 

Literature Cited 

Abel, E. F. 1962. Freiwasserbeobachtungen an Fischen im Golf von Neapel als Beitrag zur Kenntnis 

ihrer Okologie und ihres Verhaltens. Int. Revue ges. Hydrobiol., 47:219-290. 
. 1964. Freiwasserstudien zur Fortpfianzungsethologie zweir Mittelmeerfische, Blennius ca- 

nevae Vine, und Blennius inaequalis C. V. Zeit. fur Tierpsychol., 21:205-222. 
Bohlke, J. E. 1957. A review of the blenny genus Chaenopsis and a description of a related genus 

from the Bahamas. Proc. Acad. Sci. Phil., 109:25-27. 
, and C. G. Chaplin. 1968. Fishes of the Bahamas and adjacent tropical waters. Livingston. 

771 pp. 
, and C. H. Robins. 1974. Description of a new genus and species of clinid fish from the 

Western Caribbean, with comments on the families of the Blennioidea. Proc. Acad. Nat. Sci. 

Phil., 126:1-8. 



NEOC LINUS UNINOTATUS 21 



Breder, C. M. 1939. On the life history and development of the sponge blenny, Paraclinus mar- 

moratus (Steindachner). Zoologica, 24:487^196. 
. 1941. On the reproductive behavior of the sponge blenny, Paraclinus marmoratus (Stein- 
dachner). Zoologica, 24:233-235. 
Carlisle, J. G., C. H. Turner, and E. E. Ebert. 1964. Artificial habitat in the marine environment. 

Calif. Fish and Game, Fish Bull., 124:1-93. 
Ehrlich, K. F., J. S. Stephens, G. Muszynski, and J. M. Hood. 1979. Thermal behavioral responses 

of the speckled sanddab, Citharichthys stigmaeus: laboratory and field investigations. Fish. 

Bull., 76:867-872. 
Ellison, J. P., C. Terry, and J. S. Stephens, Jr. 1979. Food resource utilization among five species 

of embiotocids at King Harbor, California, with preliminary estimates of caloric intake. Mar. 

Biol., 52:161-169. 
Feder, H. M., C. H. Turner, and C. Limbaugh. 1974. Observations on fishes associated with kelp 

beds in Southern Califronia. Calif. Fish and Game, Fish Bull., 160:1-138. 
Fishelson, L. 1975. Observations on behavior of the fish Meiacanthus nigrolineatus Smith- Vaniz 

(Blenniidae) in nature (Red Sea) and in captivity. Aust. J. mar. Freshwat. Res., 26:329-341. 
Fitch, J. E., and R. J. Lavenberg. 1975. Tidepool and nearshore fishes of California. Univ. Calif. 

Press. 156 pp. 
Ford, R. F. 1959. A study of the major classification of blennioid fishes based upon cranial nerve 

evidence. M.A. Thesis, Stanford Univ. 51 pp. 
Fukao, R. 1980. Review of the Japanese fishes of the genus Neoclinus with description of two new 

species and notes on habitat preference. Publ. Seto Mar. Biol. Lab., 25:175-209. 
George, A., and V. G. Springer. 1980. Revision of the clinid fish tribe Ophiclinini, including five 

new species, and definition of the family Clinidae. Smith. Contrib. Zool., 307:1-31. 
Gibson, R. N. 1967. The use of the anesthetic quinaldine in fish ecology. J. Anim. Ecol., 26:295- 

301. 
Haderlie, E. C, and W. Donat III. 1978. Wharf piling fauna and flora in Monterey Harbor, California. 

Veliger, 21:45-69. 
Hubbs, C. 1952. A contribution of the classification of the blennioid fishes of the family Clinidae, 

with a partial revision of Eastern Pacific forms. Stanford Ichthyol. Bull., 4:41-165. 
. 1953. Revision and systematic position of the blenniid fishes of the genus Neoclinus. Copeia, 

1953:11-23. 
Jordan, D. S., and B. W. Evermann. 1896-1900. The fishes of North and Middle America. Bull. 

U.S. Natl. Mus., 47:1-3313. 
Kerstitch, A. N. 1971. Sailfin blenny from Gulf of California. Salt Water Aquarium, 7:79-81. 
Lagler, K. F. 1956. Freshwater fishery biology. Wm. C. Brown Co. 417 pp. 
Lindquist, D. G. 1971. The comparative behavior of three species of blennioid fishes. M.A. Thesis, 

Calif. State Univ., Hayward. 117 pp. 
. 1975. Comparative behavior and ecology of Gulf of California chaenopsid blennies. Ph.D. 

Thesis, Univ. Arizona. 147 pp. 
Longley, W. H., and S. F. Hildebrand. 1940. New genera and species of fishes from Tortugas, 

Florida, Pub. Carnegie Inst. Wash., 517:225-285. 
, and . 1941. Systematic catalogue of the fishes of Tortugas, Florida with observations 

on color and local distribution. Papers from Tortugas Laboratory, 34:1-331. 
Losey, G. S. 1968. The comparative behavior of some Pacific fishes of the genus Hypsoblennius 

Gill (Blenniidae). Ph.D. Thesis, Univ. Calif., San Diego, 276 pp. 
. 1976. The significance of coloration in fishes of the genus Hypsoblennius Gill. Bull. So. 

Calif. Acad. Sci., 75:183-198. 
Randall, J. E. 1966. The West Indian blenniid fishes of the genus Hypleurochilus, with the description 

of a new species. Proc. Biol. Soc. Wash., 79:57-72. 
Robins, C. R., C. Phillips, and F. Phillips. 1959. Some aspects of the behavior of the blennioid fish 

Chaenopsis ocellata Poey. Zoologica, 44:77-84. 
Rosenblatt, R. H., and J. S. Stephens, Jr. 1978. Mccoskerichthys sandae, a new and unusual chae- 
nopsid blenny from the Pacific coast of Panama and Costa Rica. Cont. Sci. Nat. Hist. Mus. 

Los Angeles County, 293:1-22. 
Schulte, E., and A. Holl. 1972. Feinbau der Kopftentakel und ihrer Sinnesorgane bei Blennius 

tentacularis (Pisces, Blenniformes). Mar. Biol., 12:67-80. 



22 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Shiogaki, M., and Y. Dotsu. 1972. The life history of the blenniid fish, Neoclinus bryope. Bull. Fac. 

Fish., Nagasaki Univ., 34:1-8. 
Springer, V. G. 1955. The taxonomic status of the fishes of the genus Stathmonotus , including a 

review of the Atlantic species. Bull. Mar. Sci., 5:66-80. 
, and M. F. Gomon. 1975. Variation in the Western Atlantic clinid fish Malacoctenus trian- 

gulatus with a revised key to the Atlantic species of Malacoctenus. Smith. Cont. Zool., 200: 1- 

11. 
Stephens, J. S., Jr. 1961. Range extension of the temperate blennioid fish, Neoclinus bryope, into 

the tropical Western Pacific. Copeia, 1961:484^185. 
. 1963. A revised classification of the blennioid fishes of the American family Chaenopsidae. 

Univ. Calif. Pub. Zool., 68:1-133. 
. 1978. Breakwaters and harbors as productive habitats for fish populations — Why are fishes 

attracted to urban complexes? In J. Baskin (chairman). Symposium on Urban Harbor Envi- 
ronments. Ocean Studies Consortium, Calif. State Univ., Pomona. Pp. 49-60. 
and J. P. Ellison. 1977. A study offish food habitats as related to the biological enrichment 

of an area. Univ. Wash. Sea Grant Pub. WSG-WO 77-2:19-24. 
, E. S. Hobson, and R. K. Johnson. 1966. Notes on distribution, behavior, and morphological 

variation in some chaenopsid fishes from the Tropical Eastern Pacific, with descriptions of two 

new species, Acanthemblemaria castroi and Coralliozetus springeri. Copeia, 1966:424-438. 
, and V. G. Springer. 1971. Neoclinus nudus, new scaleless clinid fish from Taiwan with a 

key to Neoclinus. Proc. Biol. Soc. Wash., 84:65-72. 
, R. K. Johnson, G. S. Key, and J. E. McCosker. 1970. The comparative ecology of three 

sympatric species of California blennies of the genus Hypsoblennius Gill (Teleostomi, Blen- 

niidae). Ecol. Monog., 40:213-233. 
Terry, C. B., and J. S. Stephens. Jr. 1976. A study of the orientation of selected embiotocid fishes 

to depth and shifting seasonal vertical temperature gradients. Bull. So. Calif. Acad. Sci., 

75:170-183. 
Thomson, D. A., L. T. Findley, and A. N. Kerstitch. 1979. Reef fishes of the Sea of Cortez: The 

rocky-shore fishes of the Gulf of California. Wiley and Sons. 302 pp. 
Turner, C. H., E. E. Ebert, and R. R. Given. 1969. Man-made reef ecology. Calif. Fish and Game, 

Fish Bull., 146:1-221. 
Wickler, W. 1965. Zur Biologie und Ethologie von Ecsenius bicolor (Pisces Teleostei, Blenniidae). 

Zeit. fur Tierpsychol., 22:36-49. 
Wirtz, P. 1977. Zum Verhalten blennioider Fische, insbesondere der mediterranen Tripterygion- 

Arten. Ph.D. Thesis, Univ. Miinchen. 91 pp. 
. 1978. The behavior of the Mediterranean Tripterygion species (Pisces, Blennioidei). Zeit. fur 

Tierpsychol., 48:142-174. 
Zander, C. D. 1975. Secondary sex characteristics of blennioid fishes (Perciformes). Pubbl. Staz. 

Zool. Napoli, 39:717-727. 

Accepted for publication 12 March 1981. 

Department of Biology, University of North Carolina at Wilmington, Wilming- 
ton, North Carolina 28406. 



Bull. Southern California Acad. Sci. 

80(1), 1981, pp. 23-31 

© Southern California Academy of Sciences, 1981 



The Fish Population Associated with an 
Offshore Water Intake Structure 

Mark Helvey 1 and Philip Dorn 2 

Abstract. — The fish population associated with an offshore water intake struc- 
ture by Mark Helvey and Philip Dorn, Bull. Southern California Acad. Sci., 
80(1):23-31. This area off Southern California is described from underwater ob- 
servations. The intake structure was monitored for a year and found to support 
a diverse population of water column and benthic oriented species. Many fishes 
resided at the intake year round while others were seasonal visitors. Several 
water column species including blacksmith, senorita, and jack mackerel inter- 
acted with the inwardly flowing water current for short periods of time. The most 
common response was a positive rheotaxis as they maintained station within the 
flow. Although the intake water current may be attractive to certain species 
exhibiting strong rheotactic beahvior it does not influence the composition of the 
intake fish community. 



Introduction 

Man-made structures function as artificial reefs when located in areas of the 
marine environment that lack natural bottom relief (Turner et al. 1969). Artificial 
reefs provide new sources of food and shelter, thereby concentrating fish in pre- 
viously unsuitable areas. The majority of man-made structures have been inten- 
tionally established for enhancing local sport fisheries, although several studies 
have shown that industrial structures such as offshore platforms similarly serve 
this purpose (Carlisle et al. 1964; Treybig 1971; Hastings et al. 1976; Simpson 
1977). Other types of industrial structures that may serve as artificial reefs are 
the cooling water intakes of coastal power generating stations. These intakes 
normally consist of large vertical concrete conduits which rise several meters 
above the surrounding sand bottom and are topped with a concrete lid (velocity 
cap) situated above the conduit opening. Seawater is drawn through the opening 
between the top of the conduit and velocity cap creating primarily a horizontal 
water current. 

One distinction between cooling water intakes and other artificial reefs is that 
intakes continuously withdraw seawater which generates a constant water current 
moving towards the structure. The response to a water current or rheotaxis 
(Fraenkel and Gunn 1940) plays an important role in the ecology of several fishes 
(Arnold 1974) such as in the feeding behavior of planktivorous species (Hobson 
and Chess 1976, 1978; Stevenson 1972). The purpose of this investigation was to 
evaluate the fish population at an offshore cooling water intake structure and 
determine what kind of fishes responded to the water current of the intake. 



24 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




^3 

Fig. 1. The offshore cooling water intake structure with its adjacent rock boulder substrate. 



Materials and Methods 

Study Area 

The offshore cooling water intake structure is located 300 m offshore of Re- 
dondo Beach, California, bordered by King Harbor on one side and the Redondo 
Submarine Canyon on the other side. The intake rests in 15 m of water and 
consists of a concrete conduit or riser bowl that rises 3 m above the ocean floor 
(Fig. 1). Scattered quarry rock strewn non-uniformly around the base of the riser 
bowl forms a 15 m radius of rip-rap. A concrete velocity cap is supported by 
concrete pillars 1.2 m above the top of the riser bowl. Water drawn into the 
opening has entrance velocities ranging between 48.8-100.6 cm/sec, averaging 
73.7 cm/sec (Larson 1978). 

Procedures 

The fish population at the intake structure was sampled 33 times between Sep- 
tember 1976 and August 1977 by SCUBA divers using a Super 8 mm underwater 
motion picture camera (Kodak XL 360). Each sample (cinetransect) consisted of 
six 10 m transect stations. Four extended diagonally from each corner of the 
intake and two extended perpendicularly from its two longer sides. Filming was 
accomplished between 0830 and 1200 hours by SCUBA divers swimming along 
each transect course just over the substrate up to the base of the structure. 
Artificial lighting was not used. The camera was tilted to film individual fish in 
the transect. When fish schools were encountered on the transect course, the 
camera was panned to include the entire group. Because cryptic and secretive 
fishes were not easily photograhed, numbers of such species within the tran- 
sect course were recorded on a plastic slate by a second diver. 

Analysis of movie films included species identification and their abundance as 



FISH AT OFFSHORE WATER INTAKE 25 

Table 1. Summary of ANOVA (one-way) for seasonal and daily variation in species diversity. 









ANOVA 






Source of variation 


df 


SS 


MS 


F 


P 


Seasonal (among) 
Daily (within) 
Total 


3 
28 
31 


17.0 
236.8 
253.9 


5.7 
8.5 


0.7 


0.6 



determined by use of slow motion and stop-action projection. More than 95% of 
the fish population recorded on film were identifiable. The fish community was 
analyzed for species diversity using species number (Poole 1974). Community 
structure was analyzed by combining the log of the proportionate abundance of 
each species with its frequency of occurrence into a single measure (Mason and 
Bryant 1974; Alevizon and Brooks 1975). Species were grouped into one of three 
categories based on frequency of occurrence per season: 

A — species occurring in more than 67% of samples; 
B — species occurring between 33-67% of samples; 
C — species occurring in less than 33% of samples. 

The log proportionate abundances were grouped into four categories: 

1. relative abundance >100; 

2. relative abundance between 10-100; 

3. relative abundance between 1-10; 

4. relative abundance <1. 

The analysis classified each species into any one of 12 categories, ranging from 
A-l (dominant species seen frequently) to C-4 (rare species seen infrequently). 

The responses of fish to the intake water current were analyzed by divers 
during 245 minutes of additional observation during the study, between Septem- 
ber 1976-January 1977. Individual fish moving into the flow area between the 
bottom edge of the velocity cap and the top of the riser bowl and extending 1.0 
m horizontally out from the intake opening were observed and timed using an 
underwater watch. The amount of time an individual remained within the study 
zone was recorded on plastic slates. 

Results 

A total of 39 species was recorded at the intake structure either in the water 
column or exposed along the rock substrate surrounding the intake. The number 
of species encountered seasonally ranged from 26 species in the spring to 29 
species during the winter. Results of a one-way ANOVA showed no significant 
difference (P > .05) for species diversity between quarters (Table 1). 

Total fish densities varied between seasons with highest densities occurring 
during the winter and summer seasons (Table 2). This principally reflected the 
large recruitment of juvenile blacksmith, Chromis punctipinnis, in the winter 
months and the seasonal residence of shiner surf perch, Cymatogaster aggregata, 
during the summer months. 

The blacksmith, the senorita, Oxyjulis californica, and the blue rockfish, Se- 



26 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



E 

CO 


O 








— 


U 


o 


cO 


Oh 

o 






E c 
Si 3 



c 






1) 


■o 


o 


i 


c 


d 




CO 


r 




U 




4> 


2 


J3 


^ 






eO 


or 


u 


c 








u. 




u 


O 


u 


U 


611 


0/1 




oj 


ro 



£ CO C 

> O (U 

■£ O <u 

~ c & 

co o w 



o c u 

'5 * "O 

™ 11 C 



O o t. 
D. <h-5 



S3 & «5 

a 52- . 

<D U y 

■S "C <u 

~ O Q. 

C cS L) 

o o co 

■- CD 

o * t * H 

3 S o 



CD 



<u 



l. co 

CO "O 

a c 

I -a 



o vr- 



— — • CO 

« O C 

t— ' O 3 

cj X) 

CO CO 



a! ! 



OO SOi 



co 1: $ u C a. c 



»S^|i- 



lO a. to -J O c 



to U a. w O O 



Hi 



%a> gjr >. >,.c g <u o 



HI I 



O £ to Ot^Q $ to U 



(J < < 



O U U (J 



FISH AT OFFSHORE WATER INTAKE 27 

Table 3. Fish displaying rheotropic behavior during 245 minutes of observation (n = 21). Species 
are listed alphabetically. 









Mean time 




No. of 


% frequency 


within flow (sec) 


Species 


individuals 


occurrence 


± S.E. 


Chromis punctipinnis 


27 


76 


10.48 ± 1.98 


Damalichthys vacca 


9 


38 


6.78 ± 1.69 


Hypsurus caryi 


1 


5 


— 


Medialuna californiensis 


2 


9 


3.50 ± 0.5 


Oxyjulis californica 


23 


81 


18.82 ± 3.49 


Paralabrax clathratus 


19 


52 


9.18 ± 2.15 


Phanerodon furcatus 


13 


33 


13.69 ± 3.50 


Sebastes mvstinus 


2 


9 


20.00 ± 0.0 


Sebastes serranoides 


1 


5 


— 


Trachurus symmetricus 


4 


14 


69.25 ± 57.26 



bastes mystinus, were consistently encountered in the water column throughout 
the year, accounting for a large proportion of the intake population (Table 2). The 
blacksmith and senorita regularly swam within the area of high flow for intervals 
averaging 10 and 19 second intervals, respectively (Table 3). The most typical 
response of these two species was a positive rheotaxis (Fraenkel and Gunn 1940) 
against the water being drawn into the intake opening. These fishes occasionally 
turned upward to meet the curvilinear flow of water being drawn over the edge 
of the velocity cap (Fig. 2). While maintaining positive rheotaxis, it was not 
unusual for these species to "switch" back and forth between the rectilinear and 
curvilinear flows. 

Other common residents included the olive rockfish, Sebastes serranoides, the 
kelp bass, Paralabrax clathratus, the sheephead, Pimelometopon pulchrum, and 
the pile surfperch, Damalichthys vacca. The white perch, Phanerodon furcatus, 
was also a common member except during the spring quarter. The kelp bass, pile 
and white surfperches frequently responded to the flow (Table 3) normally dis- 
playing positive rheotaxis and the "switching" behavior. During late summer, 
olive rockfish aggregated in groups of up to 15 individuals in a stationary, tail-up 
attitude immediately adjacent to the riser bowl and just below the intake opening, 
a low flow area (L. Larson, pers. comm.). 

Numerous benthic oriented species including the barred sand bass, Paralabrax 
nebulifer; copper rockfish, Sebastes caurinus; brown rockfish, S. auriculatus; 
calico rockfish, S. dallii; sculpin, Scorpaena guttata; cabezon, Scorpaenichthys 
marmoratus; painted greenling, Oxylebius pictus; blackeye goby, Coryphopterus 
nicholsii; and bluebanded goby, Lythrypnus dalli, resided at the intake. These 
species were encountered along the rock boulder substrate that surrounds the 
intake structure. None of these species swam against the intake water current. 
Although barred sand bass were occasionally observed on top of the velocity cap, 
they also were never seen orienting to the intake water current. 

Jack mackerel, Trachurus symmetricus, were very common during the fall and 
winter quarters but infrequently seen during the spring and totally absent during 
the summer. Dense schools of up to 400 jack mackerel frequently maintained 
station within the horizontal flow zone of the intake for lengthy periods (Fig. 3). 
They remained within the flow for an average of 69 seconds per visit. 



28 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. Positive rheotactic response by a senorita, Oxyjulis californica, to the curvilinear water 
current drawn over the edge of the intake velocity cap. The edge of the velocity cap is in the 
foreground. An aggregation of jack mackerel, Trachurus symmetricus, is pictured in the background. 



Some members of the intake fish assemblage included seasonal migrators. Wall- 
eye surfperch, Hyperprosopon argenteum, moved into the area during late sum- 
mer and were continually sighted until mid-fall. They were easily disturbed by 
divers, which biased accurate appraisal of their numbers. They were never ob- 
served orienting to the flow. Shiner surfperch, Cymatogaster aggregata, initially 
appeared at the intake towards the end of spring, accounting for 6% of the pop- 
ulation, and increased their numbers during the summer to account for 73% of 
the total population. Shiner surfperch completely encircled the structure during 
the summer months, with many individuals also orienting to the intake water 
current. 



Discussion 

The number of species identified at the intake compares favorably with other 
artificial reefs in the area (Turner et al. 1969). The species comprising the intake 
assemblage occur at the adjacent King Harbor breakwater (Stephens and Zerba 
1981) and included such residents as blacksmith, senorita, olive and blue 
rockfish, barred and kelp bass as well as seasonally occurring jack mackerel, 
walleye and shiner surfperch. In spite of seasonal changeovers in community 
composition, the intake exhibited a diverse assemblage that remained consistently 
complex throughout the course of the study. 

Undoubtedly, the heterogeneous rock layer surrounding the intake influenced 
the presence of numerous species. The significance between fish diversity and 
habitat complexity is well established (Risk 1972; Luckhurst and Luckhurst 1978) 



FISH AT OFFSHORE WATER INTAKE 



29 




Fig. 3. Positive rheotactic response by a school of jack mackerel, Trachurus symmetricus , to the 
rectilinear water current being drawn to the intake structure shown on the right. 



and evolves from the need of many demersal, cryptic and secretive fishes to 
remain sheltered from predators. The intake rip-rap also permitted successful 
recruitment as demonstrated by the presence of juvenile blacksmith (<100 mm) 
close to the cover of rock boulder interspaces during the winter quarter. 

Many temperate reef species do not utilize shelter during their inactive pe- 
riods — perhaps a consequence of low predation pressures (Ebeling and Bray 1976; 
Stephens and Zerba 1981). Several of these species, including surfperch, are 
suprabenthic and forage almost exclusively upon epifaunal organisms (Feder et 
al. 1974; Bray and Ebeling 1975; Ellison et al. 1979). Stephens and Zerba 
(1981) concluded that food resources along the adjacent King Harbor breakwater 
were not limiting and actually played a major role in accommodating numerous 
species within the harbor. 

Several intake members were active throughout the day in the water column 
which indicates a feeding behavior independent of the substrate. Love and Ebel- 
ing (1978) found blue and olive rockfish as well as kelp bass to feed primarily on 
planktonic or nektonic organisms. Senorita are also facultative planktivores (Bray 



30 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

and Ebeling 1975; Hobson and Chess 1976) while blacksmith are obligatory plank- 
tivores (Limbaugh 1964; Hobson and Chess 1976; Bray in press). Naturally, fishes 
active in the water column will passively drift unless they obtain orientation cues 
such as those perceived by the optomotor response (Lyon 1904; Arnold 1974) 
which permits maintenance of station (Edmundson et al. 1968). Certainly the high 
relief of the structure allowed these species to extend their water column activities 
areally without losing visual contact of the high profiled intake structure. 

Some members of the intake community were observed to periodically enter 
the flow region and remain for short durations. It is noteworthy that many of 
these species are planktivorous and may utilize the flow as a feeding station. 
However, this behavior may not pertain to all species. The presence of substrate 
grazers such as pile and white surfperch may not be related to feeding in the flow 
but only represent a behavioral response as they contact the flow in transit be- 
tween the top of the velocity cap and surrounding rip-rap. Both areas were char- 
acterized by rich epifaunal invertebrate assemblages. In addition, the presence 
of jack mackerel schools within the flow suggests that the intake may serve as a 
schooling companion (Hunter and Mitchell 1967; Klima and Wickham 1971) allow- 
ing this species to maintain its normal schooling formation without losing sight 
of a stationary object. 

Conclusions 

The species recorded at the intake structure also occur at the adjacent break- 
water. This suggests that the intake structure, like other artificial reefs, promotes 
ocean productivity in an otherwise barren location by providing necessary re- 
source requirements for the associated fish assemblage. Although the additional 
environmental parameter of a constant water current may offer advantages for 
some species, it does not appear to influence the structure of the intake fish 
community. 

Acknowledgments 

We thank David G. Lindquist, John B. Palmer, John S. Stephens, Jr., and John 
Wintersteen for their helpful comments in an earlier version of this manuscript 
and to Michael H. Horn and an anonymous reviewer for their editorial assistance 
in a recent version. We are grateful for the assistance of Crescent Boccardi in 
preparing the illustration, Kim Zerba in computer analyses, Janet Helvey for 
typing earlier versions, and to Mike Curtis, Jenny Yuge, and Carol Ranken in 
collecting field data. Funding was provided by Southern California Edison Com- 
pany to Occidental College under contract #U0695903 to Dr. Stephens. 

Literature Cited 

Alevizon, W. S., and M. G. Brooks. 1975. The comparative structure of two western Atlantic reef- 
fish assemblages. Bull. Mar. Sci., 25:482^90. 

Arnold, G. P. 1974. Rheotropism in fishes. Biol. Rev., 49:515-576. 

Bray, R. N. 1981. Influence of water currents and zooplankton densities on the daily foraging 
movements of the blacksmith, Chromis punctipinnis , a planktivorous reef fish. Fish. Res. Bull., 
78:829-841. 

, A. W. Ebeling. 1975. Day and night surveys of fishes inhabiting the kelp beds off of Santa 

Barbara, California. Fish. Res. Bull., 74:372-390. 



1 



FISH AT OFFSHORE WATER INTAKE 31 



Carlisle, J. G. Jr., C. H. Turner, and E. Ebert. 1964. Artificial habitat in the marine environment. 

Calif. Dept. Fish Game, Fish Bull., 124:1-93. 
Ebeling, A. W., and R. N. Bray. 1976. Day versus night activity of reef fishes in a kelp forest off 

Santa Barbara, California. Fish. Res. Bull., 74:703-717. 
Edmundson, E., F. E. Everest, and D. W. Chapman. 1968. Permanence of station in juvenile Chinook 

salmon and steelhead trout. J. Fish. Res. Bd. Can., 25:1453-1464. 
Ellison, J. P., C. Terry, and J. S. Stephens, Jr. 1979. Food resource utilization among five species 

of embiotocids at King Harbor, California, with preliminary estimates of caloric intake. Mar. 

Biol., 52:161-169. 
Feder, H. M., C. H. Turner, and C. Limbaugh. 1974. Observations on fishes associated with kelp 

beds in Southern California. Calif. Dept. Fish Game, Fish Bull., 160:1-144. 
Fraenkel, G. S., and D. L. Gunn. 1940. The orientation of animals. Clarendon Press, Oxford, 

352 pp. 
Hastings, R. W., L. H. Ogren, and M. T. Mabry. 1976. Observations on the fish fauna associated 

with offshore platforms in the northeastern Gulf of Mexico. Fish Res. Bull., 74:387-402. 
Hobson, E. S., and J. R. Chess. 1976. Tropic interactions among fishes and zooplankters near shore 

at Santa Catalina Island, California. Fish. Res. Bull.. 74:567-598. 
, and . 1978. Tropic relationships among fishes and plankton in the lagoon at Enewetak 

Atoll, Marshall Islands. Fish. Res. Bull., 76:133-153. 
Hunter, J. R., and C. T. Mitchell. 1967. Association of fishes with flotsam in the offshore water of 

Central America. Fish. Res. Bull., 66:13-29. 
Klima, E. F., and D. A. Wickham. 1971. Attraction of coastal pelagic fishes with artificial structures. 

Trans. Amer. Fish. Soc, 100:86-99. 
Larson, L. E. 1978. Flow into offshore intake structures. M.Sc. Thesis, Dept. of Engin. Cal. State 

Univ. Long Beach, Calif., 45 pp. 
Limbaugh, C. 1964. Notes on the life history of two California pomacentrids: garibaldis, Hypsypops 

rubicunda (Girad), and blacksmiths, Chromis punctipinnis (Cooper). Pac. Sci., 18:41-50. 
Love, M. S., and A. W. Ebeling. 1978. Food and habitat of three switchfeeding fishes in the kelp 

forests off Santa Barbara, California. Fish. Res. Bull., 76:257-271. 
Luckhurst, B. E., and K. Luckhurst. 1978. Analysis of the influence of substrate variable on coral 

reef fish communities. Mar. Biol., 49:317-323. 
Lyon, E. P. 1904. On rheotropism. I. Rheotropism in fishes. Amer. J. Physiol., 12:149-161. 
Mason, C. F., and R. J. Bryant. 1974. The structure and diversity of the animal communities in a 

broadland reed-swamp. J. Zool., 172:289-302. 
Poole, R. W. 1974. An introduction to quantitative ecology. McGraw Hill, New York. 
Risk, M. J. 1972. Fish diversity on a coral reef in the Virgin Islands. Atoll Res. Bull., 153:1-6. 
Simpson, R. A. 1977. The biology of two offshore oil platforms. Inst. Mar. Res., Univ. Calif., IMR 

Reference 76-13, 16 pp. 
Stephens, J. S., Jr., and K. Zerba. 1981. Factors affecting fish diversity on a temperate reef. 

Envir. Biol. Fishes 1981; 6:111-121. 
Stevenson, R. A., Jr. 1972. Regulation of feeding beahvior of the bicolor damselfish (Eupomacentrus 

partitus Poey) by environmental factors. In Behavior of marine animals. Vol. 2: Vertebrates 

(H. E. Winn and B. L. Olla, eds.). Plenum Press, New York, pp. 278-302. 
Treybig, D. L. 1971. How offshore platforms help fishing. Ocean Ind., 6:64-65. 
Turner, C. H., E. Ebert, and R. R. Owen. 1969. Man-made reef ecology. Calif. Dept. Fish Game, 

Fish Bull., 146:1-221. 

Accepted for publication 18 December 1980. 



1 Occidental College, Los Angeles, California 90041. 

2 Present address: Shell Development Co., P.O. Box 1380, Houston, Texas 77001 



Bull. Southern California Acad. Sci. 

80(1), 1981, pp. 32-35 

© Southern California Academy of Sciences, 1981 

A New Coralline Boring Species of Poly dor a 
(Polychaeta: Spionidae) from Northern California 

James A. Blake 

Abstract. — A new coralline boring species of Poly dor a (Polychaeta: Spionidae) 
from Northern California by James A. Blake, Bull. Southern California Acad. 
Sci., 80(l):32-35. A new species of Polydora is described from Tomales Point in 
northern California. The species is a borer in coralline algae and related to the 
P. giardi group. It differs from its closest relatives in having posterior notopodial 
spines and a four lobed pygidium. 



Nine species of Polydora and Boccardia (Family Spionidae) were reported to 
be borers in coralline algae by Blake and Evans (1973). While examining samples 
of Lithophyllum pacificum Foslie collected from Tomales Point (Point Reyes 
National Seashore) in northern California, a new species of Polydora was dis- 
covered. This new species shares affinities with P. giardi Mesnil, which occurs 
in the same alga (Day and Blake 1979), but differs in several significant features. 
A description of this new species is presented herein, along with a discussion of 
its taxonomic affinities. 

The type collection is deposited in the National Museum of Natural History 
(USNM). 

Polydora bifurcata, new species 
Figures 1-2 

Material examined. — CALIFORNIA, Point Reyes National Seashore, Tomales 
Point, 38°14'N; 122°59'W, intertidal, boring into the coralline alga, Lithophyllum 
pacificum, March 1970, coll. J. A. Blake, holotype (USNM 58976) and one para- 
type (USNM 58977). 

Description. — Holotype incomplete, measuring 6.7 mm long and 0.5 mm wide 
for 63 segments; paratype complete, broken into two parts, measuring 4.7 mm 
long and 0.5 mm wide for 35 segments. Color: light tan in alcohol, no body 
pigment. 

Prostomium strongly bifurcate, forming two prominent lobes (Fig. 1A); carun- 
cle with folds, extending posteriorly into setiger 5; narrow field of nuchal ciliation 
surrounding caruncle; no occipital tentacle; no eyes. Peristomium narrow; palps 
short, possibly regenerating on both specimens. 

Setiger 1 well developed, with parapodia shifted dorsally; postsetal noto- and 
neuropodial lamellae prominent; capillary noto- and neurosetae present. Noto- 
setae of setigers 2-4, 6 and subsequent setigers with two-tiered fascicles of uni- 
limbate capillaries, setae of first tier being shorter and thicker than second tier; 
posterior setigers with five to six long, thin capillaries and three to four thicker 
curved, pointed acicular spines (Fig. 2D). Neurosetae of setigers 2-4, and 6 sim- 



NEW SPECIES OF POLYDORA 



33 





300,um 




Fig. 1. Polydora bifurcata (paratype, USNM 58977): A. Anterior end in dorsal view; B. Posterior 
end in dorsal view; C. Pygidium in posterior view. 



ilar in arrangement and form to notosetae; hooded hooks from setiger 7, num- 
bering three to four hooks per neuropodium throughout most of body, accom- 
panied by two to three inferior capillaries for about 20 segments; capillaries 
lacking thereafter; hooks lacking constriction on shaft (Fig. 2C), with reduced 
angle between teeth, but with wide angle between main fang and shaft; fringe of 
minute bristles present on hood opening. 

Setiger 5 strongly modified, overlapping setiger 6 with heavy dorsal muscula- 
ture (Fig. 1 A); setae including a superior dorsal fascicle of broad geniculate bris- 
tled setae (Fig. 2B), a curved row of major spines alternating with bristled com- 
panion setae (Fig. 2A) and a ventral fascicle of unilimbated capillaries. Major 
spines falcate, with one large accessory tooth on curved edge and a thin, narrowly 
adhering spur on convex side. 

Branchiae from setiger 8, short at first, reaching to full size by setiger 12, each 
gill extending maximally one-third of the distance across an individual segment. 



34 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




Fig. 2. Polydora bifurcata (paratype, USNM 58977): A. Three major spines and two companion 
setae from setiger 5; B. Geniculate dorsal seta from setiger 5; C. Hooded hook from an anterior 
neuropodium; D. Posterior notopodium in dorsal view, anterior edge toward the right, indicating 
position of acicular spines and capillaries. 



Pygidium four-lobed (Fig. 1B-C), with dorsal pair being slightly smaller than 
ventral pair; each pygidial lobe with longitudinal striations composed of separate 
bacillary glands. 

Remarks. — Among approximately 66 species of Polydora, P. bifurcata is most 
similar to the widespread P. giardi Mesnil and the central Pacific species, P. 
tridenticulata Woodwick (1964). Each of these species has an accessory tooth on 
the major spines of setiger 5, hooded hooks without a constriction or manubrium 
on the shaft and branchiae beginning from setiger 8 or more posteriorly. P. bi- 
furcata differs from both of those species in having instead of lacking posterior 
notopodial spines and in having a pygidium with four lobes instead of a complete 
disc or cuff. 

Polydora bifurcata was associated with two other polydorids, P. giardi and 
Boccardia columbiana Berkeley, in the Lithophyllum crusts. All three species 
bore directly into the alga. 

Distribution. — Northern California in the vicinity of Tomales Point. 



NEW SPECIES OF POLYDORA 35 



Literature Cited 

Blake, J. A., and J. D. Evans. 1973. Polydora and related genera as borers in mollusk shells and 

other calcareous substrates. The Veliger, 15(3):235— 249. 
Day, R. L., and J. A. Blake. 1979. Reproduction and larval development of Polydora giardi Mesnil 

(Polychaeta: Spionidae). Biol. Bull., 156:20-30. 
Woodwick, K. H. 1964. Polydora and related genera (Annelida, Polychaeta) from Eniwetok, Majuro, 

and Bikini Atolls, Marshall Islands. Pacific Science, 18:146-159. 

Accepted for publication 13 August 1980. 

Battelle's New England Marine Laboratory, P.O. Drawer AH, Duxbury, Mas- 
sachusetts 02332. 



Bull. Southern California Acad. Sci. 

80(1). 1981, pp. 36-41 

© Southern California Academy of Sciences. 1981 



Research Notes 

Observations on the Deep-water Chiton, Leptochiton rissoi 
(Nierstrasz 1905) in the Eastern Pacific 



Knowledge of the deep-sea chiton fauna is still fragmentary, even tentative; 
although some 20 species of chitons are known to be endemic to depths greater 
than 200 m (Ferreira 1980), in most cases observations have been confined to 
only a few specimens, often from a single locality. Accordingly, understanding 
of the anatomical and biological characteristics of such species has been limited. 

This paper reports on new data about an eastern Pacific deep-water chiton, 
Lepidopleurus rissoi Nierstrasz 1905, heretofore known only from Nierstrasz' 
(1905) account based on specimens obtained in the course of the Siboga expe- 
dition. These observations rest upon the examination of the lectotype in reposi- 
tory at the Zoologisk Museum of Amsterdam (ZMA), and material in the Benthic 
Collection of the Scripps Institution of Oceanography, La Jolla, California (SIO) 
and the Department of Invertebrate Zoology of the California Academy of Sci- 
ences, San Francisco, California (CASIZ). 

Polyplacophora de Blainville 1816 

Neoloricata Bergenhayn 1955 

Lepidopleuridae Pilsbry 1892 

Leptochiton Gray 1847 

Leptochiton rissoi (Nierstrasz 1905) 

Figures 1-2 

Lepidopleurus rissoi Nierstrasz 1905:6-7, figs. 5, 52-55 — Ferreira 1979:163, figs. 
30-32; 1980:59, tbl. 1. 

Type material. — Lectotype, designated by Ferreira (1979), and nine paralec- 
totypes at ZMA. 

Type locality.— Eastern Pacific, 3°27.1'N, 125°18.7'W, at 2053 m, Siboga sta- 
tion no. 126. 

Material examined. — In addition to the lectotype (Ferreira 1979:163, figs. 30- 
32), six other specimens were studied: 

1) SIO M559— 5°58.8'N, 81°38.2'W, criss cross dredge, 1190 m, leg. T. Chase, 
18 March 1963; 1 specimen, in alcohol, 11.8 mm long. 

2) SIO M1392— off Arica, Chile, 18°30.0'S, 70°34.5'W, rock dredge, 402-311 
m, R/V T. Washington, leg. A. Sontar & S. Luke, 7 May 1972; 2 specimens, in 
alcohol, 14.0 mm and ca. 7 mm long. 

3) SIO M1393— off Chile, 21°23.7'S, 70°18.2'W, rock dredge, 450-420 m, R/V 
T. Washington, leg. A. Sontar & S. Luke, 1 May 1972; 1 specimen, in alcohol, 
ca. 10 mm long. 

4) CASIZ 016695— off Oregon, 44°39.1'N, 125°11.0'VV, 1420 m, R/V Ancona, 
cruise no. 6304c, 27 April 1963; 1 specimen, in alcohol, ca. 10 mm long. 

5) CASIZ016696— Guide Seamount, California, 37°09'40"N, 123°04'40"W, 



RESEARCH NOTES 



37 




Fig. 1. Leptochiton rissoi (Nierstrasz 1905). Specimen 1 1.8 mm long (SIO M559). A) Dorsal view. 
B) Close-up of central and lateral areas of anterior valves. 



1098-732 m, R/V Mulberry, Sta. 45, 15 February 1950; 1 specimen, in alcohol, 
ca. 15 mm long [CASIZ Color Slides series nos. 727-728]. 

Description. — The specimens (Fig. 1) conform well to Nierstrasz' (1905) de- 
scription of Lepidopleurus rissoi, and the lectotype (Ferreira 1979). Uniformly 
white to light cream in color, they vary in length from 7 to 15 mm. A specimen 
(SIO M1392), 14.0 x 8.5 x 0.8 mm, was disarticulated for close study. The gir- 
dle, 0.8 mm wide at the level of valve iv, is covered with spiculoid scales (Figs. 
2A-a, and 2B), 60 x 10 /xm, sharply pointed, and deeply striated. The underside 



?8 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



^ 



<^ 








/00A- 



fW- 



50 Km 




B 

Fig. 2. Leptochiton rissoi (Nierstrasz 1905). Specimen 14.0 mm long (SIO M1302). A) Girdle 
elements of upper surface (a), and undersurface (b). [measured bar = 100 /xm]. B) Spiculoid scales 



of the girdle is similarly paved with translucid spiculoid scales (Fig. 2A-b), 80 x 15 
/Am, longitudinally striated. Gills, posterior, abanal, extending about one half of 
the foot's length, comprise some 15 plumes per side. The posterior valve is ab- 
solutely and relatively longer than the others (Table 1): the relative width of its 
sinus (width of sinus/width of sutural lamina) is 1.5 mm/215 mm = 0.6. 
The radula is 5.0 mm long (29% of the specimen's length), and comprises about 



RESEARCH NOTES 



39 




I 00 /A rv\ 




of upper surface of girdle [measured bar = 60 /xm]. C) Median and first lateral teeth of radula [mea- 
sured bar = 100 /xm]. D) Head of second (major) lateral tooth [measured bar = 100 fim]. 



35 rows of mature teeth. Median tooth (Fig. 2C), quadrangular, 100 /xm long, 75 
/xm wide at the anterior blade. First lateral teeth (Fig. 2C), somewhat rectangular, 
about 80 jxm long, 60 ^m wide. Second lateral teeth with a bicuspid head, the 
outer cusp much larger than the inner one (Fig. 2D). Spatulate teeth ("Seiten- 



40 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Table 1. Length (1) and width (w) of the valves of a 14 mm long specimen of Leptochiton rissoi 
(Nierstrasz 1905). 

1 w 

Valves (mm) (mm) 1/w 

i 1.5 5.0 .30 

ii 2.0 5.6 .36 

iii 1.6 6.0 .27 

iv 1.7 6.8 .25 

v 2.0 7.0 .29 

vi 1.9 6.8 .28 

vii 1.8 6.8 .26 

viii 3.3 5.8 .57 



plate") with a very simple spoonlike shape, about 125 /xm long. Outer marginal 
teeth, elongate, 100 x 40 /xm. 

Distribution. — The range of Leptochiton rissoi extends from 44°N to 21°S, and 
from depths of 216 m to 2053 m. 

Remarks. — The assignment of rissoi to Leptochiton Gray 1847 is in agreement 
with the current interpretation of the genus (Ferreira 1979). Compared to other 
congeneric Pacific species, Leptochiton rissoi differs in the shape of the valves, 
tegmental sculpture, girdle elements, and radula (see Ferreira 1979). But the 
differential diagnosis between L. rissoi and the recently described L. batialis 
Sirenko 1979 deserves extended consideration in view of similarities in tegmental 
sculpture. From the illustrations of L. batialis (Sirenko 1979, fig. 2), and mea- 
surements thereupon, the two species are seen to differ in significant features: 1) 
the valves are relatively longer in L. batialis than in L. rissoi (length/width ratio 
of valve v is 0.44 in L. batialis, 0.29 in L. rissoi), 2) the sutural laminae in valve 
viii are triangular in L. batialis, rectangular in L. rissoi, 3) the relative width of 
the sinus [width of sinus/width of sutural lamina] of valve viii is 1.4 in L. batialis, 
0.6 in L. rissoi, and 4) the median tooth of the radula is more elongated in L. 
batialis than in L. rissoi (width/length ratio of the tooth, 0.35 and 0.75, respec- 
tively). 

Acknowledgments 

I wish to express my appreciation to Dr. William A. Newman, and Spencer R. 
Luke, Scripps Institution of Oceanography, La Jolla, California, Dr. Welton L. 
Lee, Dustin D. Chivers, Dalene Drake, and Barbara Weitbrecht (credited for the 
camera lucida drawings), California Academy of Sciences, San Francisco, Cali- 
fornia, and Dr. H. E. Coomans, Zoologisk Museum of Amsterdam, for their 
contribution and assistance in several phases of this work. 

Literature Cited 

Ferreira, A. J. 1979. The family Lepidopleuridae (Mollusca: Polyplacophora) in the eastern Pacific. 
Veliger, 22(2): 145-165, 4 pits., 8 text figs. 

. 1980. A new species of Lepidopleurus Risso, 1826 (Mollusca: Polyplacophora) in the deep 

waters of the eastern Pacific. Veliger, 23(1):55-61, 1 pit., 5 text figs. 

Nierstrasz, H. F. 1905. Die Chitonen der Siboga-Expedition. Siboga-Expeditie, 48:1-112 (+ adden- 
dum), 8 pits. 



RESEARCH NOTES 41 



Sirenko, B. I. 1979. On the composition of the family Lepidochitonidae Dall, 1889 (=Lepidopleuridae 
Pilsbry, 1892) (Polyplacophora) with description of a new bathyal species. Proc. Zool. Inst. 
Acad. Sci. USSR, 80:116-121 [in Russian]. 

Accepted for publication 4 February 1981. 

Antonio J. Ferreira, Research Associate, California Academy of Sciences, Gold- 
en Gate Park, San Francisco, California 941 18. For reprints: 2060 Clarmar Way, 
San Jose, California 95128. 



Bull. Southern California Acad. Sci. 

80(1), 1981, pp. 41-45 

© Southern California Academy of Sciences, 1981 



Apparent Wind-oriented Behavior in the Gray Whale 
(Eschrichtius robustus) 



In recent years, intensified field observations have been made over large sec- 
tions of the migratory range of the gray whale (Eschrichtius robustus Lilljeborg). 
These observations have produced a much clearer picture of its behavior and 
natural history. Since the observations and writings of Captain Charles Scammon 
(1874), behavioral biologists have been trying to explain repetitive patterns of 
activities seen in the gray whale along a corridor from Beringia to the lagoons of 
Baja California, Mexico. 

Migration was one of the first behavior patterns to be identified and described. 
Scammon and others discovered that the gray whales calved and mated primarily 
in the lagoons of Baja California Sur. Gilmore (1960), Walker (1971), Sauer (1963), 
Rice and Wolman (1971), Samaras (1974), and others have described the repro- 
duction of these whales and have established some regionally-repetitive, funda- 
mental patterns of socially interactive behavior. Spyhopping, breaching, lobtail- 
ing, fluke-stands, and respiratory-swimming patterns may be involved in other 
activities: feeding, courtship, precopulative interaction, visual orientation and/or 
navigation, low-frequency audio communication over great distances, shaking off 
lice-loosened barnacles (Samaras and Durham, in press), and/or recreation. 

I would add yet another gray whale activity to the growing list of characteristic 
forms of behavior — sailing. Roger Payne (1976) has observed, photographed, and 
described sailing as a characteristic "recreational" behavior of the southern right 
whale, Eubalaena australis, in the region of the Valdez Peninsula of Patagonia 
(Payne 1976). Previously this activity had not been described for any other ce- 
tacean. Sailing, or more appropriately, fluke-sailing posture and physical condi- 
tions include: the caudal peduncle extended nearly vertically above the sea sur- 
face; the ventral surface of the flukes cupped over and facing into the wind (Figs, 
la and 2); and sailing done in an available wind force of at least 10-20 knots 
(Beaufort Scale 3-5). Because observers have been at low elevation on murky, 
choppy water, the subsurface posture of a fluke-sailing gray whale can only be 
conjectured (Fig. la). 

The apparent sailing activity of adult gray whales was observed in upper Mag- 



42 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




FLUKE SAILING 




Fig. 1. a. Gray whale fluke-sailing orientation. The subsurface posture of a gray whale involved 
in fluke-sailing is interpretive. Because of the observed physical orientation of the extended caudal 
peduncle and cupped flukes, in relation to the wind, the body of the whale is either perpendicular to 
the surface or bowed slightly with venter up as indicated. The vertical positioning, with flukes main- 
tained high out of the water for prolonged periods of time, is probably facilitated through a sculling 
action of the pectoral fins. b. Gray whale calf wind-oriented, lunging/porpoising posture as observed 
in Laguna Guerrero Negro during a moderate gale. 



RESEARCH NOTES 



43 




« WIND 



■< WAKE 







i m _ 

Hknum 


C 




J^^ 


^^^ 




^HH ***•»> 



44 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

dalena Bay, Baja California Del Sur on at least three separate occasions between 
13 and 16 February 1978. In all cases, large, mature whales engaged in this 
activity. Fluke-sailing occurred in the afternoon when the prevailing westerlies 
had freshened sufficiently. The flukes were observed deliberately turning into the 
wind as they slowly emerged from the water. This activity would last for at least 
a minute, accompanied by obvious movement or drift of the flukes with the wind 
(Fig. 2a). 

During this mid-February segment of the annual gray whale migration, the 
whales in upper Magdalena Bay are mostly, if not exclusively, females with 
calves. Consequently, the whales involved in the apparent fluke-sailing activity 
were, in all probability, sexually mature females. Similar wind-oriented behavior 
has not been reported from other traditional calving and mating lagoons, i.e., 
Laguna San Ignacio, Scammon Lagoon (Laguna Ojo de Liebre) and Laguna Guer- 
rero Negro. 

During February 1977, in Laguna Guerrero Negro, Baja California Del Sur, I 
encountered a different wind-oriented behavior among the lagoon's population of 
gray whale calves. During this time of year, the prevailing westerlies freshen 
briskly every afternoon from approximately 1300 hr to well after sunset. On the 
afternoon of 12 February 1977, the wind force rose to a moderate gale (28-33 
knots; Beaufort Scale 7). The water in Laguna Guerrero Negro heaped up, and 
white foam from the breaking wind-generated waves was blown in streaks along 
the direction of the wind. Coarse silica sand from Great Dune Island, at least a 
kilometer to the west, was blown across the lagoon onto our observing site on 
the abandoned salt-loading pier. The gray whale calves and cows swam around 
the perimeter of the lagoon in a counterclockwise movement, usually perpendic- 
ular to the prevailing wind direction. 

As the sea surface approached the spindrift stage, all the visible calves on both 
sides of the lagoon began lunging out of the water with a low-level porpoising 
maneuver (Fig. lb). The cows (female whales) continued to glide slowly through 
the water, close by their calves, without any apparent change in their normal 
lagoon swimming-respiratory activity. The calves did not appear to be under 
stress or in a state of panic, as the observed respiratory interludes were generally 
regular. But as they lunged or porpoised forward, they would surface head-first 
at an estimated 30° angle. The lunging motion produced a distinct bow or 
compression wave ahead and to each side. During each low-level porpoising 
maneuver, their heads and blowholes (nares) would rise above the spindrift stra- 



Fig. 2. a. A mature, female gray whale (Eschrichtius robustus) fluke-sailing during a gentle breeze 
with an estimated wind force of 7-10 knots (Beaufort Scale 3). The whale is sailing from right to left 
as indicated by the wind direction and wake in the photo. In b, the flukes are just emerging and 
turning into the wind. Sailing has already begun as indicated by the cupped over flukes and a dis- 
cernible wake stretching to the right of the caudal peduncle. This sailing episode took place in a 
moderate breeze with an estimated wind force of 11-16 knots (Beaufort Scale 4). In c, the peduncle 
and flukes of the whale have completely emerged and turned into the wind coming from the direction 
of the sand dunes in the background. The wind force during this observation was a strong breeze 
with a velocity of 22-27 knots (Beaufort Scale 6). These fluke-sailings took place in Upper Magdalena 
Bay, Baja California Del Sur on three separate occasions. The sailings occurred during the afternoon 
between 13 and 16 February 1978 and were three different whales. 



RESEARCH NOTES 45 

turn at the sea's surface (Fig. lb). This activity continued into dark. The wind 
did not moderate until after midnight. 

The apparent coordinative and continuous nature of the activity, with a definite 
relationship to severe wind and sea-surface conditions, suggested a learning and/ 
or conditioning experience. The exposure of the heads and blowholes above most 
of the spindrift during porpoising could serve to reduce the amount of salty spume 
inhaled during respiration. 

Acknowledgments 

I wish to thank Mary Wicksten, Allan Hancock Foundation U.S.C., for her 
valued criticism and technical advice. 

Literature Cited 

Gilmore, R. M. 1960. A census of the California gray whale. U.S. Fish Wildl. Serv., Spec. Sci. Rep., 

Fish, 342:1-30. 
Payne, Roger. 1976. At home with right whales. Natl. Geog. Mag., 149:3:322-339. 
Rice, D. W., and A. A. Wolman. 1971. The life history and ecology of the gray whale {Eschrichtius 

robustus). Amer. Soc. Mamm., Spec. Publ. No. 3, viii + 142 pp. 
Samaras, W. F. 1974. Reproductive behavior of the gray whale, Eschrichtius robustus, in Baja 

California. Bull. So. Calif. Acad. Sci., 73:2:57-64. 
, and F. E. Durham. The gray whale (Eschrichtius robustus) and its epizoite Crustacea. In 

Press. 
Sauer, E. G. F. 1963. Courtship and copulation of the gray whale in the Bering Sea at St. Lawrence 

Island, Alaska. Psychlogische Forschung, 27:157-174. 
Scammon, C. M. 1874. The marine mammals of the northwestern coast of North America. John H. 

Carmany and Co., San Francisco, 319 pp. 
Van Dorn, W. G. 1974. Oceanography and seamanship. Dodd, Mead & Co., New York, 481 pp. 
Walker, T. J. 1971. The California gray whale comes back. Natl. Geog. Mag., 193:394-415. 

Accepted for publication 22 May 1980. 

William F. Samaras Research Associate in Marine Mammals, Los Angeles Coun- 
ty Museum of Natural History , 900 Exposition Blvd., Los Angeles, California 90007 
and Research Associate in Marine Mammals and Geology, Cabrillo Marine Mu- 
seum, 3720 Stephen White Drive, San Pedro, California 90731 . 



Bull. Southern California Acad. Sci. 

80(1). 1981, pp. 45-48 

© Southern California Academy of Sciences, 1981 



A New Articulate Brachiopod from the Lower Cambrian 
Latham Shale, Southeastern California 



The lower Cambrian Latham Shale of Hazzard (1954) crops out at localities 
scattered widely throughout the Mojave Desert of southeastern California. Al- 
though the formation is highly fossiliferous, descriptive paleontologic documen- 
tation of its fauna is limited to a very few reports (Resser 1928; Crickmay 1933; 
Riccio 1952) dealing mainly with the abundant and diverse assemblage of trilo- 
bites. Other notes (Hazzard 1933, 1954; Mount 1974b, 1976) have touched on the 
stratigraphic allocation of some of the trilobite and non-trilobite taxa. 

The purpose of the present report is to provide the description of a new species 
of Nisusia, the only brachiopod of the Class Articulata known to occur in the 
Latham Shale. Nisusia Walcott 1905:247, is the earliest and most primitive ar- 
ticulate brachiopod genus and is recorded from the lower and middle Cambrian 
of North America, Asia, and Europe. The discovery of the new taxon from the 
Latham Shale provides one of the oldest known records of the genus from the 
Cordilleran area. 

References to the Department of Earth Sciences, University of California, Riv- 
erside, are hereafter abbreviated as UCR. 

Order Orthida 

Family Nisusiidae 

Genus Nisusia Walcott, 1905 

Nisusia fulleri new species 

Figures 1-2 

Nisusia n. sp.: Mount 1974c, p. 47, pi. 1, figs. 1, 2; Mount 1976, p. 176, fig. 4. 

Diagnosis. — The new species is distinguished from all other species of the 
genus by having the following combination of characteristics: shell medium in 
size, transversely subrectangular, greatest width at the hinge; sulcus shallow and 
wide; ornamentation of numerous, narrow, low costellae of three different 
strengths, with narrow interspaces. 

Description. — Shell medium in size for the genus, apparently thin; outline 
transversely subrectangular, wider than long with the greatest width at the hinge 
and maximum length at mid width; cardinal extremities nearly rectangular; lateral 
margins nearly straight or slightly convex; anterior margin broadly rounded. Sul- 
cus apparently very shallow and wide, originating in anterior third of shell and 
extending to anterior margin. Umbo short, widely convex, moderately swollen 
and protruded posterior to the posterior margin; apical angle 112°. Interarea nar- 
row and apparently apsacline. Ornamentation consists of numerous, somewhat 
regularly arranged, low, narrow, rounded costellae of three different strengths, 
gauging 4 per mm on the holotype; separated by very narrow interspaces with 
widths from V6 to 1 times the width of the costellae. Surface marked by numerous, 
fine, slightly raised concentric lines of growth. Interior characteristics unknown. 

Holotype.— UCR 10/2031, length 6.5 mm, width 8.0 mm, brachial valve? 



RESEARCH NOTES 



47 





Figs. 1-2. Nisusia fulleri Mount, new species, holotype UCR 10/2031, brachial? view, x5. 
(2) Nisusia fulleri Mount, new species, paratype UCR 10/2019, pedicle? view, x3. 



Paratypes.— UCR 10/2019, length 11.3 mm, width 15.0 mm, pedicle valve? 
UCR 10/2032, eight partial valves on one piece of shale. UCR 10/2037, length 
11.0 mm, width 12.3 mm, pedicle valve? 

Type locality.— UCR locality 10; at elevation of 1160 ft (305 m), 100 m west of 
old quarry at the southern end of the Marble Mountains, San Bernardino County; 
125 m west and 650 m south of northeast corner of Sec. 11, T.5N., R.14E., Danby 
15' quadrangle (1956 edition); same as locality M-5 of Hazzard (1933). 

Stratigraphic position. — The type locality is in the middle one-third of the 
Latham Shale which is 15 m thick at the site. 

Associated fauna. — Anthozoa: Bergaueria radiata Alpert 

Inarticulata: Paterina prospectensis (Walcott), Mickwitzia occidens Walcott 

Mollusca: Hyolithes whitei Resser 

Annelida: new genus and species (Mount 1976, fig. 20) 

Trilobita: Olenellus clarki (Resser), O. fremonti Walcott, O. gilberti Meek, 
O. mohavensis (Crickmay), O. nevadensis (Walcott), O. new species (Mount 
1976, fig. 11), Bhstolia anteros Palmer, B. bristolensis (Resser), B. insolens 
(Resser), B. new species (Mount 1976, fig. 17), Peachella iddingsi (Walcott), 
Onchocephalus new species (Mount 1976, fig. 18) 

Malacostraca: Anomalocaris canadensis Whiteaves 

Eocrinoidea: Gogia ojenai Durham 

Age. — Late Early Cambrian; Bhstolia Subzone of Mount (1974a, b), Bonnia- 
Olenellus Zone of Rasetti (1951), Waucoban Stage. 

Discussion. — Nisusia fulleri new species bears similarity in general form and 
outline to N. bivia (Walcott 1912:750) from the lower Cambrian portion of the 
Carrara Formation in the Resting Springs Range, Inyo County, California, but 
differs in details of surface sculpture. The latter species has coarser, wider spaced 
costellae with fine radial lines superimposed on the primary costellae. 

Nisusia montanaensis Bell 1941:238, from the middle Cambrian of Montana, 
has the fine sculpture of N. fulleri, however, it is larger, more quadrate, and the 
hinge line is slightly less than the width of shell at the middle. 

Nisusia festinata (Billings 1861:10), the type species for the genus, from the 
lower Cambrian of eastern North America is closely similar to the new form 
described here. The former taxon is spino-costellate and appears to have costellae 
of more even size and a narrower, deeper sulcus. 



48 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Nisusia kotujensis Andreeva 1962:87 from the lower Cambrian of the Siberian 
Platform of eastern Asia is distinguished from the new California species by 
having coarser costellae and narrower interspaces in proportion to the width of 
the costellae. Moreover, the hinges are extended into elongated cardinal extrem- 
ities. 

Nisusia mantouensis Resser and Endo 1937:141 and N. nasuta ramosa Nikitin 
1956:19 from the middle Cambrian of central Asia have some resemblance to N. 
fulled, however, these Asian species have coarser sculpture, a deeper sulcus and 
more rounded extremities. 

Etymology. — The new brachiopod is named for Mr. James E. Fuller of the 
University of California, Riverside, who collected the holotype and one of the 
paratypes. 

Acknowledgments 

I wish to thank Messrs. John Kniffen, James E. Fuller and David L. Willoughby 
who provided the specimens described in this report. 

Literature Cited 

Andreeva, O. N. 1962. Nekotorye Kembriyskie Brakhiopody Sibiri i Sredney Azii. Paleontologi- 
cheskiy Zhurnal, 1962:87-96. 

Bell. W. C. 1941. Cambrian Brachiopoda from Montana. J. Paleon., 15:193-255. 

Billings, E. 1861. New species of Silurian fossils. Can. Geol. Survey, Paleozoic Fossils, 1:1-24. 

Crickmay, C. H. 1933. Paleontology. Pp. 71-76 in Notes on the Cambrian rocks of the eastern 
Mohave Desert, California. Univ. Calif. Publ. Bull. Dept. Geol. Sci., 23:57-80. 

Hazzard, J. C. 1933. Notes on the Cambrian rocks of the eastern Mohave Desert, California. Univ. 
Calif. Publ. Bull. Dept. Geol. Sci., 23:57-80. 

. 1954. Rocks and structure of the northern Providence Mountains, San Bernardino County, 

California. Calif. Div. Mines Geol. Bull., 170(4):27-35. 

Mount, J. D. 1974a. Early Cambrian faunas from the Marble and Providence Mountains, San Ber- 
nardino County, California. Bull. So. Calif. Paleon. Soc, 6:1-5. 

. 1974b. Biostratigraphy of the lower Cambrian in southeastern California. Geol. Soc. Amer- 
ica, Abstracts With Programs, 6(3):224. 

. 1974c. Early Cambrian articulate brachiopods from the Marble Mountains, San Bernardino 

County, California. Bull. So. Calif. Paleon. Soc, 6:47-52. 

. 1976. Early Cambrian faunas from eastern San Bernardino County, California. Bull. So. 

Calif. Paleon. Soc, 8:173-182. 

Nikitin, I. F. 1956. Brakhiopody kembriya i nizhnego ordovika severo-vostoka Tsentral'nogo Ka- 
zakhstana. Alma-Ata. 141 pp. 

Rasetti, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smith- 
sonian Misc. Colin., 116(5): 1-277. 

Resser, C. E. 1928. Cambrian fossils from the Mohave Desert. Smithsonian Misc. Colin., 81(2): 
1-14. 

, and R. Endo. 1937. Description of the fossils. Pp. 103-301 in The Sinian and Cambrian 

formations and fossils of southern Manchoukuo. Manchurian Sci. Mus. Bull., 1:1-474. 

Riccio, J. F. 1952. The lower Cambrian Olenellidae of the Marble Mountains, California. Bull. So. 
Calif. Acad. Sci., 51:25-49. 

Walcott, C. D. 1905. Cambrian Brachiopoda, with descriptions of new genera and species. Proc 
U.S. Nat. Mus., 28:227-337. 

. 1912. Cambrian Brachiopoda. U.S. Geol. Survey Monogr., 51:1-872. 

Accepted for publication 30 October 1980. 

Jack D. Mount, Dept. of Earth Sciences, University of California, Riverside, 
California 92521. 



INSTRUCTIONS FOR AUTHORS 

The BULLETIN is published three times each year (April, August, and November) and includes articles in English 
n any field of science with an emphasis on the southern California area. Manuscripts submitted for publication 
hould contain results of original research, embrace sound principles of scientific investigation, and present data 
n a clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide 
or contributors. Consult also recent issues of the BULLETIN. Authors should strive for directness and lucidity, 
ichieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous 
eference of pronouns, and logically placed modifiers. 

MANUSCRIPT PREPARATION 

It is strongly recommended that, before submitting a paper, the author ask qualified persons to review it. The 
Author is requested to submit at least two additional copies with the original, on SV2 x 11 opaque, nonerasable 
liaper, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. 
footnotes should be avoided. Manuscripts which do not conform to the style of the BULLETIN will be returned to 

he author. 

i An abstract summarizing in concise terms the methods, findings, and implications discussed in the paper must 
accompany a feature article. Abstract should not exceed 100 words, 
t A feature article comprises approximately five to thirty typewritten pages. Papers should usually be divided into 

he following sections: abstract, introduction, methods, results, discussion and conclusions, acknowledgments, and 

iterature cited. Avoid using more than two levels of subheadings, 
i A research note is usually one to six typewritten pages and rarely utilizes subheadings. Consult a recent issue 

if the BULLETIN for the format of notes. Abstracts are not used for notes. 
1 Abbreviations: Use of abbreviations and symbols can be determined by inspection of a recent issue of the 

JULLETIN. Omit periods after standard abbreviations: 1.2 mm, 2 km, 30 cm, but Figs. 1-2. Use numerals before 
fcinits of measurements: 5 ml, but nine spines (10 or numbers above, such as 13 spines). The metric system of 
veights and measurements should be used wherever possible. 

Taxonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International 
Code of Botanical Nomenclature (Lawjouw et al. 1956), the International Code of Nomenclature of Bacteria and 
;/iruses (Buchanan et al. 1958), and the International Code of Zoological Nomenclature (Stoll et al. 1961). Special 

ttention should be given to the description of new taxa, designation of holotype, etc. Reference to new taxa in 

itles and abstract should be avoided. 

■ The literature cited: Entries for books and articles should take these forms. 

I McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. 

Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54:452-458. 
» Brattstrom, B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), 
Univ. California Press, xii + 635 pp. 

Tables and figures (line drawings, graphs, or black and white photographs) should not repeat data contained in 
|he text. The author must provide numbers and short legends for tables and figures and place reference to each 
>f them in the text. Legends should be typed on a separate sheet of paper and placed at the end of the manuscript. 
illustrations and lettering thereon should be of sufficient size and clarity to permit reduction to standard page size; 
.ordinarily they should be no more than twice the size of intended reduction and should not exceed SV2 by 11 inches 
In size. Photographs must be printed on glossy paper. Submit one photoduplicated copy of each illustration. All 
illustrations accompanying Research Notes will be reduced to one column width. All half-tone illustrations will have 
light screen (grey) backgrounds. Special handling such as dropout half-tones, special screens, etc., must be requested 
;>y and will be charged to authors. 

A cover illustration pertaining to an article in the issue or one of general scientific interest will be printed on the 
:over of each issue. Such illustrations along with a brief caption should be sent to the Editor for review. 

PROCEDURE 

All manuscripts should be submitted to the Editor, Robert J. Lavenberg, Los Angeles County Natural History 
vluseum, 900 Exposition Blvd., Los Angeles, California 90007. Evaluation of a paper submitted to the BULLETIN 
>egins with a critical reading by the Editor; several referees also check the paper for scientific content, originality, 
ind clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing i