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BULLETIN
Volume 93 Number 3
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BCAS-A93(3) 91-134 (1994) DECEMBER 1994
Southern California Academy of Sciences
Founded 6 November 1891. incorporated 17 May 1907 44
© Southern California Academy of Sciences, 1994
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Date of this issue | December 1994
&) This paper meets the requirements of ANS!I/NISO Z39.48-1992 (Permanence of Paper).
SOUTHERN CALIFORNIA
ACADEMY OF SCIENCES
1995Annual Meeting
May 5-6
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Telephone: 213/744-3384
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Sample Abstract
MICROBIAL ACTIVITY IN THE DIGESTIVE TRACT OF THE HALFMOON, Meadi-
aluna californiensis. J. S. Kandel’. J. R. Paterek> and M. H. Horn’. ‘California State Univ.
Fullerton, CA 92634 and *Agouron Institute, La Jolla, CA 92037.
We report the presence of a diverse microbial flora and of microbial fermentation products
in the hindgut region of the halfmoon. Medialuna californiensis, a seaweed-eating fish from
southern California coastal waters. Viable aerobic and anaerobic bacteria were found in all
sections of the gut. but were of highest concentration (10°—10*/ml) in the hindgut. Microscopy
revealed vibrios, spirilla. rod-shaped bacteria and flagellated protozoa in the midgut and
hindgut. but primarily vibrios and rods in the stomach and foregut. Acetic, isobutyric and
butyric acids, the volatile products of microbial breakdown of carbohydrates, were found
only in the hindgut. as was ethanol, a nonvolatile product. These results provide the first
evidence for microbial fermentation and its possible contribution to the energy supply in a
north-temperate herbivorous fish.
6" ——
west ith
Bull. Southern California Acad. Sci.
93(3), 1994, pp. 91-109
© Southern California Academy of Sciences, 1994
Cultural Change and Geographic Variation in the
Songs of the Belding’s Savannah Sparrow
(Passerculus sandwichensis beldingt)
Richard A. Bradley
Department of Zoology, Ohio State University,
1465 Mt. Vernon Ave., Marion, Ohio 43302
Abstract.—The Belding’s Sparrow is a subspecies of the Savannah Sparrow that
occurs only in isolated patches of Salicornia marsh habitat between Goleta, Cal-
ifornia and El Rosario Lagoon, Baja California Norte. Song samples were obtained
from 14 local populations during 1973 and 7 populations were resampled during
1987/88. Turnover in the element lexicon of each population was low. A few
elements changed in their proportional occurrence rate. During the initial survey
it was noted that certain song types were common among males at any one locality.
After 15 or 16 years the most common (popular) types in some populations remain
dominant. At other localities types have changed in popularity. Sequence com-
parison analysis of elements within songs was conducted to produce an objective
inter-song dissimilarity matrix. This analysis revealed that song variation within
local populations is much less extensive than among populations. The features
used to define the original dialects remain distinctive of the local populations.
Geographic variation in bird song often exhibits discontinuous variation with
local populations of birds singing distinctive local variants of the species-specific
vocalization. Past work on long-term temporal shifts in bird song have assessed
the extent to which population-specific “‘dialects’”» have remained stable (Mc-
Gregor and Thompson 1988). In the current study, I studied population-specific
geographic patterns of song structure in the Belding’s Savannah Sparrow (Pas-
serculus sandwichensis beldingi). 1 conducted an initial study of the singing be-
havior in 14 populations of the Belding’s Sparrow between 1972 and 1974 (Bradley
1977). Seven of the largest of these populations were re-sampled during the springs
of 1987 and 1988.
The main focus of the current study is the analysis of shifts in the patterns of
use of the specific song types within local dialect populations. At this scale song
variation reflects patterns of cultural drift within a population rather than potential
exchange among populations. Many of the specific song elements or complete
song types that were recorded in 1973 were still being used in populations 15 or
16 years later. This study evaluates the extent to which the proportion of males
using these song elements and complete song types have changed. Such changes
in song-type popularity reflect the cultural ‘fitness’ of learned song variants. Sam-
ples from 7 different populations represent replicate natural experiments in cul-
tural evolution.
In addition to analysis of cultural change within local populations, I used se-
quence comparison analysis to quantify song variation among populations. The
91
92 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
second goal of the study was to examine the hypothesis that clear discontinuities
occur within the range of geographic variation. As a corollary of this hypothesis
I aver that these discontinuities correspond to local populations and reveal a
history of behavioral isolation.
The subject of this work is the Belding’s Savannah Sparrow, a well-marked
subspecies (beldingi) of a widespread species (Passerculus sandwichensis). Each
population of the Belding’s Sparrow is geographically distinct, inhabiting small
patches of salt marsh along the coast of California and Baja California del Norte
(Grinnell and Miller 1944; Bradley 1973; Massey 1977; Fig. 1). Savannah Spar-
rows of this subspecies have been considered permanent residents (Grinnell and
Miller 1944; Garrett and Dunn 1981). There are very few reports of this distinctive
subspecies outside of these salt marshes (Grinnell and Miller 1944), and there
may even be a physiological restriction to dispersal beyond saline environments
(Cade and Bartholomew 1959; Poulson and Bartholomew 1962). Belding’s Spar-
rows exhibit better weight maintenance (Cade and Bartholomew 1959) and are
more active (Poulson and Bartholomew 1962) when kept in captivity with sea
water than with fresh water. If distinct discontinuities exist among the populations
of Belding’s Sparrows, they may be attributable to the isolation created by this
subspecies’ habitat specificity.
Methods
During 1973, recordings were made with a Uher 4000 Report S recorder and
a Uher M 512 microphone mounted ona 61 or 76 cm fiberglass parabolic reflector.
During 1987 and 1988 recordings were made with a Marantz CP221 cassette
recorder and a Sennheiser K3U microphone with an ME88 directional head.
Copies of the 1973 recordings are on file in the Bioacoustics Archive of the Florida
Museum of Natural History, Gainesville, Florida. Copies of the 1987-88 record-
ings are on file at the Borror Laboratory of Bioacoustics at Ohio State University,
Columbus, Ohio. Recording methodology was similar for both samples. I recorded
individual males by walking through the occupied habitat on one or two mornings
and recording singing territorial males as I entered their small display territories.
I did not retrace my path or record in the same area more than once. When
recordings were made on more than one morning, separate areas of the habitat
were sampled. This method sacrifices maximal sample size to insure that no
individual was sampled more than once (Bradley 1977). Population estimates for
these populations were between 16 pairs at Agua Hedionda Lagoon and over 2000
pairs at San Quintin Bay (Bradley 1973; Massey 1977). The song samples include
recordings from about 29% of the individuals in each of these small local pop-
ulations (median, Table 1). Long-term study (6 years) in conjunction with color
banding has established that individual male Belding’s Sparrows sing only one
song type (Bradley 1977; Massey 1979).
Audiospectrograms were made on either Kay Elemetrics 6061b, 7029a, or DSP
5500 audiospectrographs using both narrow and wide band filter settings and 160-
16000 Hz scale. A lexicon of all distinguishable song elements was made by
examining these audiospectrograms (Bradley 1977). A song element is any acous-
tical feature of a song that is separated from others by a silent gap. The sequence
for a particular song type was represented in the analysis (coded) by listing the
element codes as they occurred in the songs (Bradley 1977; Bradley and Bradley
SONG EVOLUTION IN BELDING’S SPARROWS 93
California
Baja California Norte
100km
N
W E
ER
S
Fig. 1. A map of the 7 localities sampled during this study. Localities for which only small sample
sizes were obtained are included on this map even through lexicon analyses could not be conducted
for these sites. Coastline distances between the sites are in km. AB = Anaheim Bay, NB = Newport
Bay, SM = Santa Margarita Estuary, AH = Agua Hedionda Lagoon, LP = Los Penasquitos Lagoon,
SQ = San Quintin Bay, ER = El Rosario Lagoon.
1983). Rapid buzzes (> 250 pulses/s) were classified as single elements. There was
relatively little variation among individuals with respect to buzz fine structure,
but buzzes differed in length. To include this variation in the analysis, the code
for a buzz was repeated once for each 0.2 seconds of duration (0.2 s was the modal
94 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Table 1. Summary of samples and song types for the 5 largest populations.
1973 1987/88
Number Domi- Number 74 Domi-
Locality (population)! N? of types nance? N of types nance
Anaheim Bay (267) 61 21 56% 70 31 34%
Newport Bay (83) 29 12 59% 24 15 38%
Santa Margarita (106) 29 13 41% 21 12 52%
Los Penasquitos (52) 21 12 48% 16 8 63%
San Quintin Bay (1000) 34 15 41% 31 20 29%
Total 174 73 50%+ 162 86 39%
! Estimated number of breeding pairs from Massey (1979), estimate for San Quintin from Bradley
(1973).
2? Number of males recorded.
3 Percent of males singing one of the three most common song types.
4 Based on the 3 most common song types from each locality.
length of the shortest single buzz phrase). Patterns of similarity among the song
elements were used to construct an inter-element dissimilarity matrix (Bradley
and Bradley 1983).
All of the elements taken together comprise a song type, sometimes referred to
as a theme or motif by other authors. The general song structure of Belding’s
Sparrows is described in detail elsewhere (Bradley 1977), but I will review the
terminology briefly. The song is composed of a number of similar introductory
elements followed by a series of short elements in a variety of patterns. The second
half of the song is composed of a buzz phrase composed of relatively long buzzes
as well as intervening short elements, and finally there is a terminal flourish. The
terminal flourish follows the last buzz in the song and is short (1 to 5 elements,
<0.3 s) and relatively invariant within a population. Chew (1981) employed a
slightly different nomenclature for the sequence of song segments in Canadian
populations of this species including an introductory (A) section that corresponds
to my introductory elements; the note complexes that follow are classified by
Chew as (B) and transition sections (C); the buzz section is referred to as the trill
section (D); and finally the terminal flourish (not always present in Canadian
songs) is referred to as the (E) section. The main difference between Chew’s and
my approach is that I focus on finer-scale individual element sequences while he
uses section sequences in his analysis.
For the present study each unique sequence-list corresponds to one unique song
type. Each male sang only one song type. The only exceptions were for the minor
variation which occurred within a song type in the number of repetitions of the
introductory note or the exact length of a buzz phrase. For males that sang songs
with different numbers of introductory notes, the longest song was used to rep-
resent that male. The song type of each male in the sample was coded as a sequence
of elements (based on the lexicon). The song sequences were compared directly
and with a series of computer programs (BELDINGS) developed for this purpose
(Bradley and Bradley 1983). These programs use an optimal-matching algorithm
for the sequence comparison that generates an inter-song dissimilarity measure
(Bradley and Bradley 1983). Briefly, this method compares the coded sequences
representing any two songs and calculates the minimum number of changes (ad-
SONG EVOLUTION IN BELDING’S SPARROWS 95
ditions, deletions and substitutions) necessary to produce an identical match. The
numerical value of a substitution is weighted by the acoustical similarity of the
two song elements involved. This method provides a sensitive and objective
measure of song similarity. For a detailed description of the method consult
Bradley and Bradley (1983).
A randomization test analogous to ANOVA was used to compare differences
among individuals within a population and between populations and sampling
years (Sokal and Rohlf 1981). For this randomization test, the test statistic was
the ratio of the mean within-group song dissimilarity to the mean between-group
‘song dissimilarity. Discriminant function analysis was used to produce a model
to assess the distinctiveness of the song variation among populations. Pearson
product-moment correlation analysis was used to compare mean song dissimi-
larity and geographic separation among populations.
Only the 5 localities represented by the largest sample sizes were analyzed for
variation in the occurrence of song type variants and shifts in the element lexicon.
The other two localities’ samples (Agua Hedionda Lagoon N = 10 (1973) and 3
(1988), El Rosario Lagoon N = 11 (1973) and 16 (1988)) were too small to warrant
such analyses. A Chi-square test of goodness-of-fit was used to assess the similarity
of occurrence patterns of individual elements between the sampling years for each
population. One-way ANOVA was used to assess variability among two song
pattern variables (number of elements per song type, number of element types
per song type). A simple index of similarity (1) was used to compare the lexicon
patterns among populations. The index I = 2c/(a + b) where a and b are the
number of different element types in each population and c is the number of
elements common to both populations.
Results
Occurrence of Song Types
Although patterns of variation within populations are less dramatic than be-
tween populations, clear differences exist among the individuals within a popu-
lation. These differences are primarily in the number and form of elements between
the introductory notes and the buzz sections of the songs. The number of different
song types represented by recordings from each sampling locality varied from 8
to 31 and generally reflects the number of males recorded (16 to 70, Table 1). In
each population there are certain song types that are shared by many males (Fig.
2). In the 1973 sample 50% of males sang one of the three dominant song types
at each locality (Bradley 1977). The 1987/88 data reveal that 39% of males sang
one of the three dominant types (Table 1). If the occurrence of song types merely
reflected random sampling one would expect 21% of the males from the 1973
sample and 17% of the males from the 1987/88 sample to sing three types (Table
1). Most song types have changed in popularity between samples (Table 2). A
sum of all individuals singing the most popular song type in 1973 across these 5
populations is 40 of 174 or 23% of the sample. Only 15 of 162 (9%) sang these
same types in the 1987/88 sample. In only 2 of the 7 populations did the rank 1
song remain the most popular song type. In 2 of the remaining 5 populations the
new first rank position was occupied by birds singing the previous rank 2 song.
At San Quintin Bay the third most popular type in 1973 became the most popular
SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
96
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Rank of Popularity
Fig. 2. A histogram of the number (vertical axis) of the individual Belding’s sparrows that sang
songs with the rank of popularity indicated (horizontal axis). The total sample is composed of 336
birds singing 159 song types. Each of the seven populations is evaluated twice, once for 1973 and
once for 1987 or 1988. Rank 1 songs from all populations are combined in the first bar, rank 2 in the
second bar, etc. If songs were distributed evenly and randomly one would predict that only 28
individuals would sing the most popular (rank 1) songs.
SONG EVOLUTION IN BELDING’S SPARROWS 97
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Fig. 3. Examples of popular songs that retained a similar form in subsequent sample. The rank 1
song at Anaheim Bay in 1973 (A), the same song type remained as rank | in 1987 (B). The songs A
& B have a dissimilarity value of 0.23. The rank 2 song at Anaheim Bay in 1973 (C) was used by
only two individuals in 1987 (D), dropping to rank 17. The songs C & D have a dissimilarity value
of 0.17. In contrast the average of the four dissimilarity measures among different songs in this figure
is 0.40.
type in 1988. For the remaining 2 populations a new song type rose to first rank.
Nevertheless, some of the dominant song types in 1973 were still popular in their
respective populations in the 1987/88 sample (Table 2, Fig. 3). For example the
single most common type (rank 1) recorded at Anaheim Bay in 1973 was also
the most common in 1987. In contrast the rank 2 song from 1973 at Anaheim
Bay became rare in 1987 and was sung by only two males (rank 17).
Despite changes in popularity, the fact that many song types remained un-
changed indicates that song types are often copied intact. Examining all song types
(not just the dominant ones) reveals that relatively few types were shared between
samples (Table 3). Nevertheless these 12 shared types represent 19% of the in-
dividual males in the 1987/88 sample. In addition to invariant song types, some
song types changed only slightly (Fig. 4A, B). In other populations very different
song types gained popularity (Fig. 4C, D).
98 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Table 2. Summary of the shifts in dominance of song types between samples.
1973 1987/88 ~
most popular 3 types use of same 3 types
Locality No. birds (rank) No. birds (rank)
Anaheim Bay 18 (1) 8 (2) 6 (3) 13 (1) 2 (17) 0
Newport Bay 7 (1) 6 (2) 4 (3) 0 6 (1) 0
Santa Margarita 5 (1) 4 (2) 3 (3.5)! 0 0) 1 (16)
Agua Hedionda 3 (1) 2 (2.5)! 2 (2.5)! 3 (1) 0 0
Los Penasquitos 4(1) 3 (2) 2 (3.5)! 0 0 0
San Quintin 6 (1) 5 (2) 4 (4.5)? 2 (22) 0 8 (1)
El Rosario 6 (1) 4 (2) 1 (3) 0 15 (1) 0)
1 The use of a fractional rank indicates a tie between two ranks.
2 Four song types tied for third rank.
Changes in Element Lexicon
Initial classification produced a lexicon of 46 distinct element types from the
5 populations studied in detail here. This lexicon is largely a subset of the one
illustrated in Figures | and 2, p. 62 and 64 in Bradley (1977). A total of 9 elements
present in the 1973 songs are absent among the 1987/88 songs. Five new elements
appear in the recent sample. This appears to represent a 30% degree of “turnover”
in song elements when expressed as the total number of changes divided by the
combined lexicon (Table 4). This figure is a distortion because most of the lost
or gained elements were sung by very few individual birds. Of the 9 element types
lost, 8 were sung by 3 or fewer individuals. Likewise, of the new 5 elements, 3
were sung by 3 or fewer individuals. When the estimate of turnover is “weighted”
by considering the frequency of occurrence of elements there appears to be a
relatively minor change in the overall lexicon of just 3% (Table 4). In other words,
the only changes in the lexicon lists for the two samples were among relatively
rare elements. Sampling error may contribute to this turnover estimate; if some
elements which were actually present were missed, the real turnover rate would
be even lower. What about element occurrence between years within each locality?
Chi-square analysis reveals that the proportional frequencies of the elements
Table 3. Song types shared in both 1973 and 1987/88 samples for the 5 largest populations sam-
pled.
Percent of birds in
Number of shared 1987/88 sample singing
Locality N! patterns? shared patterns?
Anaheim Bay 47 5 26
Newport Bay WS 2 8
Santa Margarita 24 1 19
Los Penasquitos 19 1 6
San Quintin Bay 32 3 19
Total 147 12 19
' Number of different song types in both samples combined.
* Number of birds in 1987/88 singing song types that are identical to song types recorded during
1973.
° The number of individual males that sang one of the shared types as a percentage of the 1987/88
sample.
SONG EVOLUTION IN BELDING’S SPARROWS 99
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SECONDS
Fig. 4. Examples of popular songs that were replaced in the population by new or previously
unpopular types. The most popular song type at Los Penasquitos in 1973 (A) is slightly different from
the most popular type recorded at that locality in 1988 (B). The songs A & B have a dissimilarity.
value of 0.61. The most popular song type at El Rosario in 1973 (C) is slightly different from the rank
1 song in 1988 (D). This new popular song (D) was the second most popular type at El Rosario in
1973. The songs C & D have a dissimilarity value of 0.61. The average of the four dissimilarity
measures among different songs in this figure is 0.71.
differed significantly between years in 4 of the 5 population samples analyzed (P
< 0.01 for each). Only the Anaheim Bay samples were indistinguishable. This
reflects the changing patterns of song-type popularity. The Anaheim Bay popu-
lation differs from the other four in that the single most popular song type at
Anaheim Bay in 1973 was still the most popular in 1987.
Geographic Variation
Variation among populations involves the form of song elements (lexicon) as
well as the sequence and general form of song arrangement. The terminal flourish
is the most distinctive feature of songs shared by individuals at each locality that
differs among localities. Song dissimilarity can vary between 0.0 (identical songs)
to 1.0 (no similarity). Mean values calculated from my sample vary from 0.047
(within Agua Hedionda Lagoon, 1988) to 0.854 (between El Rosario Lagoon in
100 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Table 4. Summary of changes to the element lexicons for the 5 largest samples.
1973 1987/88 Elements New Weighted
Locality lexicon lexicon lost elements Turnover’ turnover?
Anaheim Bay 23 21 3 1 9% 1%
Newport Bay 23 17 9 3 30% 6%
Santa Margarita 18 14 5) 1 33% 3%
Los Penasquitos 15 16 1 2 10% 4%
San Quintin Bay 17 15 4 2 19% 2%
All localities 41 35 9 5 30% 3%
' The number of changes divided by the total lexicon for both samples combined.
2 The number of occurrences of changed elements divided by the number of occurrences of unchanged
elements. This value effectively weights the change by frequency of element use.
1973 and Newport Bay in 1988). The sequence comparison analysis reveals that
variation among songs between each population of Belding’s Sparrows is signif-
icantly greater than song variation within the populations for both the original
(1973) and subsequent (1987/88) song samples (P < 0.001, Table 5).
Three separate discriminant function analysis models were created based on
the sequence comparison results. One model was created for each sample sepa-
rately, and a third model was built for the two samples combined. The model for
the 1973 data correctly assigned 95% of individuals to the correct group (the
population where they were recorded). The 1987/88 model correctly assigned
100% of individuals. The model based on songs from both samples combined
correctly assigned 99% of individuals to the a-priori groups. Thus variation in
song between years and between individuals is relatively minor compared to
variation among recording localities.
Mean song dissimilarity among populations that are relatively near each other
Table 5. Song similarity as assessed by sequence comparison analysis among individuals within
each locality and between localities. The values in the table are mean dissimilarity measures. This
measure approaches zero for identical songs.
1973 1987/88
Locality N! Within locality N Within locality
Anaheim Bay 1830 328 2415 338
Newport Bay 406 38 276 .286
Santa Margarita 406 .332 210 .290
Agua Hedionda 45 .256 3 .050
Los Penasquitos 210 .347 120 .348
San Quintin Bay 561 391 465 .296
E] Rosario 55 151 120 -086
Within Groups
All localities 3513 336 3609 316
Among Groups
All localities 15.402 .534* 12,681 MWR Z
** Indicates that these values are different from within groups values at P < 0.001.
* Number of comparisons or one half of a symmetrical matrix excluding diagonal elements (self
comparisons).
SONG EVOLUTION IN BELDING’S SPARROWS 101
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Distance between populations (km)
Fig.5. The relationship between the mean song dissimilarity (vertical axis) measured with sequence
comparison for each pair of populations with the map distance (horizontal axis) between these localities.
Data from both 1973 and 1987/88 samples are included. Dissimilarity measures can range from 0.0
(identical songs) to 1.0 (no similarity).
was either large or small, but distant populations nearly always have distinctly
different songs (Fig. 5). All of the comparisons where the localities were separated
by more that 160 km exhibited a mean song dissimilarity of 0.575 or greater (Fig.
5). I found no correlation between the mean song dissimilarity measure among
populations and the geographic distance separating them (r = 0.22, ns; Fig. 5,
Fig. 1).
Inspection of the audiospectrograms reveals that the distinctive terminal flour-
ish, characteristic of each population in 1973 (Bradley 1977), has remained un-
changed for most populations (Fig. 6). The most typical form of terminal flourish
in 1973 was still dominant in 1987/88 in 6 of the 7 populations (Fig. 6). The
sample from El Rosario Lagoon in 1973 contained two common terminal flour-
ishes. The less common form which was used by only 4 of 11 males recorded in
1973 was present in the songs of all 16 males recorded in 1988.
There were no significant differences among populations with respect to the
number of elements in the song types representative of each (F = 1.11, df 4,154,
ns). Internal song-type complexity as revealed by the number of different element
102 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
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= — =,| ™ = =) Ze] © 4
= = = = '
= = = z =a A i
ED Ty Te TNT TREN ET
as as as as
S- G H Q =e R
= @ Ss 8 = ww = =. 8 | s
= x = x —_ = _— = Sa
n — 4 =a al —
= ;
as o> as as
(SéDamSs) (seconds)
i J
Pa
= 3 z., ™
= z — = er —
Fig. 6. Examples of the typical terminal flourish for each population for both sampling periods.
The terminal flourish is a complex of song elements that concludes the song and occurs just after the
buzz phrase (Bradley 1977). A) Anaheim Bay 1973. B) Anaheim Bay 1987. C) Newport Bay 1973,
D) Newport Bay 1987, E) Santa Margarita Estuary 1973 (2-element form). F) Santa Margarita Estuary
1988 (2-element form), G) Santa Margarita Estuary 1973 (4-element form). H) Santa Margarita Estuary
1988 (4-element form). I) Agua Hedionda Lagoon 1973. J) Agua Hedionda Lagoon 1988, K) Los
Penasquitos Lagoon 1973 (2-element form), L) Los Penasquitos Lagoon 1988 (2-element form), M)
Los Penasquitos Lagoon 1973 (3-element form), N) Los Penasquitos Lagoon 1988 (3-element form).
O) San Quintin Bay 1973, P) San Quintin Bay 1988, Q) El Rosario Lagoon 1973, R) El Rosario
Lagoon 1988.
types per song did vary significantly between populations (F = 18.96, df 4,154,
P < 0.001). Anaheim Bay songs contain more variety (¥ = 10.7 element types/
song type), while Santa Margarita and Los Penasquitos contain less (x = 7.7, 7.9
element types/song type respectively). For each population the lexicon similarity
index was highest when the 1973 sample was compared to the later sample from
that locality, the mean index of similarity (I) for the 5 such comparisons equals
0.83. There was no relationship between the index of similarity computed between
the two samples from a locality and its population size (r = 0.00, ns), or sample
size (r = +0.07, ns). Thus shifts in the lexicon do not seem to be related directly
to population size. Comparisons among the populations reveal that neighboring
populations share somewhat similar element lexicons. The Spearman rank cor-
relation of the index of similarity with distance between populations was —0.76
(P < 0.01). This relationship does not appear to be linear (Fig. 7).
SONG EVOLUTION IN BELDING’S SPARROWS 103
0.670 e
0.617
ed @
+
-
.<
(qe) @
—
=
‘ & C)
oO @
W 0.565
S
(e) @
UO
om
x
ced)
—)
0.513
@
@
@
0.460 ) : @ )
T ]
(0) 125 250 375 500
Distance between populations (km)
Fig. 7. The relationship between the similarity of the lexicon lists (vertical axis) of a pair of
populations with the map distance (horizontal axis) between them. Data from the 5 largest populations
for both sampling periods are included. The line represents the regression line for this relationship (r?
= 0.40). The index of similarity can range from 0.0 (no similarity) to 1.0 (identical lexicon lists).
Discussion
The patterns of song variation among Belding’s Sparrow populations is some-
what different from that described for the eastern Canadian populations studied
by Chew (1981). Chew indicates that variation in the trills (=buzzes of this study)
characterized the regional populations. In Belding’s Sparrows variation in buzz
phrases was relatively minor. Chew found terminal song sections were absent in
20% of the Savannah Sparrows that he recorded (Chew 1981). The terminal
sections of Canadian Savannah Sparrows (Chew 1981) were much less complex
and shorter than the corresponding parts of Belding’s Sparrow songs. The se-
quences of song sections were consistent among Belding’s Sparrow populations
and varied significantly among the Canadian populations studied by Chew (1981).
Two factors may explain these differences; first, Chew used a different scale for
his grammatical analyses than I employ in the sequence comparison analysis.
Chew (1981) compares sequences of song sections while I compare sequence of
individual song elements. Second, the populations studied by Chew were separated
104 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
by much greater geographic distances than the populations of Belding’s Sparrows.
One clear similarity between these two studies is that a few songs were used by
the majority of males at any one locality. Among Canadian populations 61-80%
of the individuals sang one of three song types (Chew 1981, p. 710). Among
Belding’s Sparrow populations 34-63% shared the three most popular types (Ta-
ble 1).
Variation within populations of the Belding’s Sparrow is characterized by a few
popular song types used by many of the singing males. Most populations that I
studied experienced a shift in the proportional use of song types between 1973
and 1988. If juvenile males imitate their father’s song as is the case in a few
species (Immelmann 1969; Nicolai 1959; Gibbs 1990; Zann 1990) these popularity
shifts might represent an indirect measure of the reproductive success of their
fathers. Gibbs (1990) has demonstrated that song variation is related to repro-
ductive success in Geospiza fortis and suggests that cultural evolution and bio-
logical evolution reinforce each other. If young birds model their songs after other
(unrelated) individuals in the population, shifts in song-type use may simply reflect
shifts in the songs that were frequently copied as has been demonstrated in the
Saddleback, Philesturnus carunculatus rufusater (Jenkins 1977) and other species
(Petrinovich 1988; Payne and Payne 1993). Distinguishing between these two
alternatives (learning from father or neighbors) would be difficult in the Belding’s
Sparrow because large groups of neighboring males sing exactly the same song
type. In one case a wild Savannah Sparrow learned the song of a White-crowned
Sparrow (Cooper and Murphy 1985) so it is clear that they are capable of learning
songs different from that of their father. Laboratory work suggests that birds learn
songs from individuals with whom they interact regularly, or which are dominant
in a particular setting (Baptista and Petrinovich 1984; Petrinovich 1985; Slater
et al. 1988). In most of the Belding’s Sparrow populations that I studied, males
with similar songs are spatially clumped (Bradley 1977). In studies of color-banded
males there was no evidence of males changing their songs or imitating the songs
of their neighbors for periods up to 4 years (Bradley 1977; Massey 1979). The
type of songs which are common in local populations (Table 2) may reflect the
fact that males in some song groups exert a disproportionate influence on the
process of song learning. These song groups expand at the expense of others. Song
groups are not completely homogeneous, some individuals singing different song
types can be found within them. This pattern may reflect the settlement patterns
of individual males.
Local song groups and dialects in other species may be the result of both
philopatry and imitation of the song variants used by resident birds. Work on
Indigo Buntings, Passerina cyanea, has shown that young males arriving to breed
for the first time copy the songs of established territory holders with whom they
interact (Payne 1981b, 1983: Payne and Payne 1993). Because such imitation is
mediated by direct interactions among individuals it has been dubbed the “social
adaptation model’ (Payne 1981a). The pattern that results from this social in-
teraction is a series of small neighborhoods of 2-12 males singing similar songs
(Payne et al. 1981). Strong social interactions were influential in determining which
song young White-crowned Sparrows (Zonotrichia leucophrys) learned in captivity
(Baptista and Petrinovich 1986) and in the wild (Baptista and Petrinovich 1984;
Baptista 1985; Petrinovich and Baptista 1984). Similar learning patterns were
SONG EVOLUTION IN BELDING’S SPARROWS 105
Table 6. Comparison of geographic distance and sequence-comparison song dissimilarities among
the five large population samples of Belding’s Sparrows. The lower half of the table are the coastline
distances (km) and the upper half are the mean song dissimilarities for both sampling periods combined.
Dissimilarity measures can range from 0.0 (identical songs) to 1.0 (no similarity).
AB NB SM LP SQ
Anaheim Bay (AB) — .68 of tl .69 .67
Newport Bay (NB) 24 — .66 .65 .68
Santa Margarita (SM) 87 63 — .65 71
Los Penasquitos (LP) 124 100 37 — .66
San Quintin Bay (SQ) 432 428 365 328 —
observed in wild populations of this species (Baptista and Morton 1988; DeWolfe
et al. 1989). Saddlebacks acquire songs similar to the territorial neighbors with
which they have direct social contact, thus forming local song neighborhoods or
song groups (Jenkins 1977).
Song variation within Belding’s Sparrow populations is relatively minor when
compared to variation among the populations (Table 5). Beyond this simple
pattern, correlation analysis indicates that there is no cline in vocal similarity
among adjacent populations at the level of entire songs (Fig. 5). It is surprising
that the two pairs of close populations (Newport Bay— Anaheim bay 24 km and
Los Penasquitos— Santa Margarita 37 km) share song sequences that are no more
similar than some of the most widely separated populations (Table 6). The An-
aheim Bay and Newport Bay populations have been separated by extensive urban
development (except immediate beach area) for a relatively long period. This is
not the case between Santa Margarita Estuary and Los Penasquitos Lagoon where
coastal development is much less extensive, even today. This result reinforces my
interpretation that all of these populations have been isolated for many genera-
tions.
Mundinger (1982) suggests that analysis of the patterns of geographic distri-
bution of song elements (isogloss analysis) is an effective method of assessing the
evolution of microgeographic variation in vocal behavior. Comparisons of the
lexicon of elements among populations in Belding’s Sparrows yield a weak clinal
geographic pattern (Fig. 7). I consider the imitation of song elements between
birds from different source populations to be a form of behavioral exchange of
cultural memes (sensu Dawkins 1976) analogous to gene exchange. I conclude
that exchange of memes among populations of Belding’s Sparrows has probably
involved incorporation of new elements rather than entire songs.
The discovery that little meme exchange has occurred among the populations
of the Belding’s Sparrow implies that these populations have either been very
isolated from each other, or that subsequent behavioral divergence has been
relatively rapid. Estimates of annual survival and population turnover are im-
portant to an assessment of the significance of shifts in occurrence of song types
between these two samples. Massey (1979) in a six year color banding study
estimated that annual adult male survival was about 64%. This estimate is at the
high end of those published for this species (Wheelwright and Rising 1993). Using
the formula in Gill (1990) the mean life expectancy based on this estimate would
be 2.6 yr. Mortality may drop in individuals older than 5 years (Wheelwright and
106 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Rising 1993). Another factor indicates that the actual life expectancy is probably
lower; there is much higher mortality among first year male Savannah Sparrows
(Wheelwright and Rising 1993). I conclude that there have probably been three
to four population turnovers in the 15 years between the two samples in this
study. Re-sampling indicates that relatively few changes in songs have occurred
during this period. McGregor and Thompson (1988) reviewed the evidence for
stability in song dialects and concluded that stasis has been documented for periods
ranging from 6 to 28 years among 7 species of oscine passerines. Cultural shifts
over relatively short periods occurred among Indigo Buntings, yet a few song types
persisted for 15 years (Payne et al. 1981). The boundaries between song dialects
have shifted position in some populations of White-crowned Sparrows (Trainer
1983), indigobirds (Vidua) (Payne 1985) and European Starlings (Sturnus vulgaris)
(Adret-Hausberger 1986).
Bowman (1979) demonstrated that a relationship exists between song structure
and the physical acoustical environment in Galapagos finches. Gish and Morton
(1981) have argued that geographic variation in the songs of the Carolina Wren
are the result of selection for songs which possess physical properties appropriate
for transmission in the particular acoustical environment of the geographic lo-
cality. Handford (1988) attributed a significant portion of the geographic variation
in Zonotrichia capensis songs to adaptation to the acoustical propagation prop-
erties of the local vegetation. Thus it is possible that patterns of geographic vari-
ation in bird song reflect differences in the local habitat. Adaptation to local habitat
acoustics seems to be an unlikely explanation for dialects in the case of the
Belding’s Sparrow for two reasons. First, all populations of this sparrow occupy
coastal salt marsh dominated by the same 2 or 3 plant species (Bradley 1977)
with virtually identical topography, vegetation physiognomy and climate. Thus
these populations probably share the same selective environment with respect to
song degradation. Second, the features of the fine structure of the songs that vary
among Belding’s Sparrow populations do not differ in ways that would likely affect
their transmissibility. The differences are mainly in the arrangement of elements
and variants of elements which share the same frequency and amplitude envelopes.
The songs also share the same general acoustical pattern (Bradley 1977).
Some authors have considered geographic variants as selectively neutral epi-
phenomena that are a consequence of imperfect song learning (Andrew 1962;
Wiens 1982). Marked discontinuities in singing behavior have been implicated
as potential behavioral isolating mechanisms (Marler and Tamura 1962; Notte-
bohm 1969; Baker 1975; Baker and Cunningham 1985). The available genetic
evidence for such isolation has been challenged (Zink and Barrowclough 1984;
Hafner and Petersen 1985; Lougheed and Handford 1992). Others have shown
that song learning may occur after dispersal, which would presumably dilute the
effectiveness of local songs as markers of local origin (Kroodsma 1974; Jenkins
1977; Baptista and King 1980; Baptista and Morton 1982; Baptista and Petri-
novich 1984; McGregor et al. 1988). In separate reviews, Payne (198la) and
Baptista (1985) have concluded that geographic variation in song is a product of
the social milieu and that little evidence for an behavioral-isolation function exists.
It has also been shown that mate choice was unrelated to song type of potential
mates in a mixed-dialect population (Chilton et al. 1990). In the case of the
Belding’s Sparrow, isolation may result from geographic separation combined
SONG EVOLUTION IN BELDING’S SPARROWS 107
with habitat specificity and behavioral isolating mechanisms may be of relatively
minor importance.
Even if the songs themselves do not serve to enhance isolation of local popu-
lations the patterns may provide us with a record of historical contact (Baker and
Thompson 1985). Belding’s Sparrow populations exhibit geographic disconti-
nuities in song structure, as well as little evidence of similarity among adjacent
populations. The conclusion that I draw from the analysis of these song types is
that there has been relatively little meme exchange between the isolated popu-
lations of Belding’s Sparrows.
Rothstein and Fleischer (1987) provide an alternate hypothesis for the main-
tenance of song dialects that does not involve isolation. Their hypothesis avers
that males singing local song dialects are providing an “honest signal’’ of local
origin which may be used by females in selecting mates. Familiarity with local
song types may also enhance efficiency in assessment of territorial intrusion.
Morton (1982) has suggested a mechanism for this is comparison of a song to the
stored memory of a familiar undegraded song. Thus familiar local song variants
may be exploited by birds for their information value in both social and physical-
acoustical ways.
Acknowledgements
I thank Dave Bradley for countless hours of computer analysis and for his expert
statistical advice. Amy Tovar provided valuable help in the field and produced
many of the audiospectrograms. Lynda Barry and Terry Hermsen read the manu-
script and provided useful advice. I thank the Division of Natural Resources,
Camp Pendelton Marine Base, and the Seal Beach National Wildlife Refuge for
permission to study on lands under their care.
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Accepted for publication 27 September 1993.
Bull. Southern California Acad. Sci.
93(3), 1994, pp. 110-117
© Southern California Academy of Sciences, 1994
Helminth Parasites of Some Southern California Fishes with a
Redescription of Proctoeces magnorus Manter, 1940
(Digenea: Fellodistomidae) and Description of
Choanodera moseri sp. n.
(Digenea: Apocreadidae)
Patrick J. Frost and Murray M. Dailey
Ocean Studies Institute, California State University,
Long Beach, California 90840
Abstract. — Approximately 1400 marine fishes collected in and around Los Angeles
and Long Beach Harbors were examined for parasitic helminths between May
1979-—March 1992. Ten families of helminth parasites were collected from 18
species of southern California marine fish that represent new host-parasite records.
A new species is described, Choanodera moseri sp. n., aS well as a rediscription
of Proctoeces magnorus Manter, 1940.
Parasites of southern California marine fishes were surveyed from 1975-1979
(Dailey et al. 1981). During this study, 2268 fishes were examined, primarily for
nematodes of the two genera Anisakis and Pseudoterranova (Phocanema). In 1983,
Love and Moser published a check list of parasites of marine fishes for the western
United States that included southern California species.
During the present study, parasitic helminth were recovered from a total of
approximately 1400 marine fishes captured between May 1979—March 1992, in
and around the waters of Los Angeles and Long Beach harbors. The results of
this investigation, which includes new host-parasite records, a rediscription of
Proctoeces magnorus Manter, 1940 (Digenea: Fellodistomidae) and an original
description of a new species of Choanodera (Digenea: Apocreadidae) are presented
in this paper.
Methods
Fish were collected by trawl aboard the R/V Yellowfin for examination by
students enrolled in general parasitology at California State University, Long
Beach (CSULB). Fish were identified using Miller and Lea (1972), sexed, mea-
sured, eviscerated (gills and gut), and packed on ice immediately after capture.
Trematodes were fixed in alcohol-formalin-acetic acid (AFA), stained in Semi-
chon’s carmine, dehydrated in ethyl alcohol, cleared in xylene, and mounted in
balsam resin. Nematodes were killed in hot 70% ethyl alcohol, cleared in glycerin,
and mounted in glycerine jelly. Drawings were made with the aid of a drawing
tube. Measurements are in micrometers unless otherwise indicated, with ranges
followed by means in parentheses.
Results
Eleven families of helminth parasites were collected from 18 species of fishes
in southern California waters that represented new host-parasite records (Table
110
HELMINTH PARASITES OF SOME SOUTHERN CALIFORNIA MARINE FISHES 111
1). Six specimens of Proctoeces magnorus were recovered from two California
sheephead, Semicossyphus pulcher (Ayres) and 11 specimens of Choanodera mo-
seri sp. n. were found, four and seven respectively from two barred sand bass,
Paralabrax nubulifer (Girard) and two ocean whitefish, Caulolatilus princeps (Jen-
yns).
Description
Choanodera moseri sp. n.
(Gee, 1)
Choanodera moseri sp. n. Apocreadidae Skrjabin, 1942. The following descrip-
tion based on 11 specimens (2 adults, 9 immature).
Specific diagnosis. — Body elongate, thick, with widest part at level of acetab-
ulum. Anterior part of worm only slightly tapering, posterior end rounded. Length
0.601 to 2.052 mm (1.247 mm), width 0.281 to 0.595 mm (0.468 mm). Cuticle
not covered by spines or scales. Forebody folded in ventrally, forming scoop
shaped pocket, flaring anteriorly, coming together just posterior to the acetabulum.
Oral sucker subterminal, circular, 71.9 to 156.9 in diameter. Acetabulum circular,
sessile, 93.9 to 196.2, slightly wider than long. Pharynx unmodified, 45.3 to 130.8
long by 42.1 to 137.3 wide. Intestinal bifurcation midway between suckers. Cecae
end blindly near posterior of body. Genital pore median, just anterior to acetab-
ulum. Testes tandem, intercecal, wider than long. Anterior testes 48.6 to 340.0
wide by 48.6 to 294.3 long. Posterior testes 48.6 to 340.0 wide by 48.6 to 300.6
long. Cirrus and cirrus sac lacking. Seminal vesicle oval, median, extending to
just posterior to acetabulum, narrowing to a tubular pars prostitica with well
developed prostate gland.
Ovary circular, 22.7 to 163.5 in diameter, anterior to the anterior testis and
dextral of midline. Uterus pretesticular, mostly to left of ovary, emptying into
genital sinus. Eggs large, 64.8 to 100.44 by 48.6 to 81.0. Seminal receptacle round,
median, in pretesticular space, dorsal io, and slightly overlapping ovary and
anterior testis. Vitelline follicles large, extending from posterior edge of acetab-
ulum to posterior of body, confluent posterior to testes.
Type host.—Barred Sand Bass, Paralabrax nebulifer (Girard).
Location. — Intestine.
Locality. —Long Beach harbor, Long Beach, California.
Holotype. —USNM Helm. Coll. No. 82770.
Etymology. —This species is named in honor of Dr. Mike Moser, University
of California, Santa Cruz, for his many contributions to marine parasitology.
Remarks.—The new species is distinct from C. caulolotili Manter, 1940, the
only other species in this genus, by its smaller body size (0.601—2.052 mm) versus
2.5-3.3 mm for C. caulolotili, lack of tegumental spines and shape of ovary (round
of C. moseri sp. n. and lobed for C. caulolotili).
Proctoeces magnorus Manter, 1940
(Fig. 2)
Redescription based on 6 specimens. Body aspinose, cylindrical, equally wide
along most of length, 1.456 to 4.882 mm (3.356 mm) long by 0.297 to 0.446 mm
(0.350 mm) wide. Oral sucker subterminal, muscular, funnel shaped, 327 to 523
112 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Table 1. New host-parasite records for southern California fishes.
yy
USNM
Helm.
Family Genus/species Coll. + Host
Digenea
Acanthocolpidae Stephanostomum casum 82779 California halibut
(Paralichthyes californicus)
Allocreadidae Genitocotyle acirrus 82780 White croaker
(Genyonemus lineatus)
Helicometrina nimia 82773 Starry rockfish
(Sebastes constellatus)
Apocreadidae Choanodera moseri sp. nov. 82770 Barred sand bass
(Paralabrax nebulifer)
Ocean whitefish
(Caulolatilus princeps)
Fellodistomidae Procioeces magnorus 82778 Califormia sheephead
(Semicossyphus pulcher)
Gorgoderidae Probolitrema calijorniense 82777 Thornback ray
(Platyrhinoides triseriata)
Hemiuridae Parahemiurus merus 82772 Deepbody anchovy
(Anchoa compressa)
Opecoelidae Opecoelus lotellae 82774 California sheephead
(Semicossyphus pulcher)
Scorpion fish
(Scorpena guttata)
Opecoelina scorpaenaez* 82775 Ganbaldi
(Hypsypops rubicundus)
White croaker
(Genyonemus lineatus)
Opecoelina scorpaenae 82776 California sheephead
(Semicossyphus pulcher)
Monogenae
Microcotylidae Microcotyle sebasies 82771 Green rockfish
(Sebastes rastrellager)
Pile surf perch
(Paralichthys vacca)
Shortspine thomyhead
(Sebastolobus alascanus)
Speckled rockfish
(Sebastes ovalis)
Starry rockfish
(Sebastes constellatus)
Nematoda
Camallanidae Spirocamallanus pereirai 82781 Round hermng
(Etrumeus acuminatus)
Speckled fin midshipman
(Porichthys myriastes)
Queenfish
(Seriphus politus)
White croaker
(Genyonemus lineatus)
Spirocamallanus pereirai Topsmelt
(larvae) (Atherinops afjinis)
* New locality record.
HELMINTH PARASITES OF SOME SOUTHERN CALIFORNIA MARINE FISHES 113
Fig. 1. Line drawing of Choandera moseri from composite of immature and mature specimens.
A, acetabulum; GP, genital pore; O, ovary; OS, oral sucker; P, pharynx; SR; seminal receptacle; SV,
seminal vesicle; T, testis; U, uterus; V, vitellaria.
deep by 359 to 497 wide. Acetabulum pedunculate, 255 to 359 long by 261 to
327 wide, with muscular groove within cavity (not apparent in all specimens).
Prepharynx short; pharynx large, very muscular, 233 to 366 long by 117 to 281
wide. Cecae conspicuous, reaching posterior end of body. Intestinal bifurcation
114 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Fig. 2. Line drawing of mature Proctoeces magnorus. C, cirrus; CP, cirrus pouch; I, intestine; ISV,
internal seminal vesicle; GP, genital pore; O, ovary; OS, oral sucker; P, pharynx; PP, pars prostitica;
T, testis; U, uterus; V, vitellaria.
just posterior to pharynx. Genital pore median, anterior to acetabulum stalk,
posterior to intestinal bifurcation. Testes oval, diagonal, separated by a few folds
of uterus, in posterior half of body, 155 to 385 long by 119 to 281 wide. Cirrus
sac claviform, extending to half way between ovary and acetabulum in relaxed
HELMINTH PARASITES OF SOME SOUTHERN CALIFORNIA MARINE FISHES 115
specimens, just posterior to acetabulum in contracted specimens, 392 to 810 long
by 65 to 98 wide at greatest width (near base). Posterior portion of seminal vesicle
sac like, separated from anterior coiled tubular portion by a constriction. Pars
prostitica long, cirrus short, empties into anterior part of a large genital atrium.
Ovary oval, pretesticular, 61 to 379 long by 64 to 255 wide, separated from
anterior testes by a few coils of uterus. Seminal receptacle seen only in small
immature worms, at level of, or slightly posterior to ovary. Uterus extends to and
fills posterior portion of body, empties into genital atrium. Eggs small, 19 by 35.
Vitelline follicles few, in two lateral groups extending from anterior edge of ovary
to anterior testis.
Host. —California Sheephead, Semicossyphus pulcher (Ayres).
Location. — Intestine.
Locality. —Long Beach harbor, Long Beach, California.
Voucher specimen.—USNM Helm. Coll. No. 82778.
Remarks. — Freeman and Llewellyn (1958) synonymized P. eurythreaus Odhner,
1911 with P. subtenuis (Linton, 1907) Hanson, 1950 and concluded that P. mag-
norus was a synonym of P. subtenuis (=syn. P. eurythreaus). Gibson and Bray
(1980) placed P. magnorus as species inquerendi until more specimens were
described. Our specimens compare well with Manter’s (1940) original description
(USHM #9359), except in three areas: 1. The musculature of the oral sucker is
much more distinct in our specimens. 2. The internal seminal vesicle is longer
and separated into a sac like posterior section and a coiled tubular anterior portion,
and 3. The ovary and testis of our specimens are oval rather than spherical as in
Manter’s specimen. Since differences within species of Proctoeces can be very large
(Gibson and Bray 1980; Bray 1983), the variation seen between our specimens
and that of Manter, is probably due to variations within the species. Based on
this, we concur with Manter (1940) that P. magnorus is indeed a distinct species.
Discussion
Opecoelina scorpaenae Manter, 1934 (Table 1) has not been recorded from any
west coast fishes prior to this study (Love and Moser 1983; Gibson and Bray
1984). We found O. scorpaenae in three different hosts (Table 1), the California
sheephead, Semicossyphus pulcher (Ayres), garibaldi, Hypsypops rubicundus (Gi-
rard), and the white croaker, Genyonemus lineatus (Ayres), which suggests that
this parasite is common and wide spread.
Sekerak and Arai (1977) reported that specimens of Stephanostumum casum
(Linton, 1910) found from inshore species of rockfish from the north-eastern
Pacific were really S. californicum Manter and Van Cleave, 1951 and agreed with
Durio and Manter’s (1969) assumption that S. casum is restricted to tropical
waters. Stephanostumum casum has been found in fish from the Caribbean, Ber-
muda, the Mexican Pacific, Galapagos Islands, and the Red Sea (Durio and Manter
1969). Our specimen (Table 1), from the California halibut, Paralichthys califor-
nicus (Ayres) caught in southern California may represent the northern most extent
of S. casum’s range.
Parahemiurus merus (Linton, 1910) Manter, 1940 is widely distributed in the
Gulf of Mexico, Atlantic and Pacific Oceans, as well as Japanese waters, where
it is found mostly in carangid, salmonid, clupeid, and engraulid fishes (Bray 1990).
Parahemiurus merus was reported from 19 species of fish from the western Pacific
116 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Ocean (Love and Moser 1983). The deepbody anchovy. Anchoa compressa (Gi-
rard) is a new host record for this parasite (Table 1). One other host in the family
Engraulididae from southern California, the northern anchovy, E ngraulis mordax
(Girard), has also been found to be infected with P. merus (Woolcock 1935). Bray
(1990) suggested that P. merus could play an important role as a biological in-
dicator. Due to the economic and commercial importance of A. compressa and
E. mordax, and its wide spread prevalence, P. merus may make an important
biological marker for the management of these fisheries. Parahemiurus merus, for
example. is used as a marker for the migration of masu salmon, Oncorhynchus
masu in Japan (Awakuru and Nomura 1983) as well as a stock indicator for the
Pacific herring. C/upea harengus pullasi in central California (Moser and Hsieh
1992).
Another potentially important biological indicator may be the monogene, M7-
crocotyle sebastes Goto 1984, since it is found on the gills in a large number of
hosts (Love and Moser 1983). Five new hosts were found to be infected with this
parasite (Table 1). Two of the hosts, the starry rockfish. Sebastes constellatus
(Jordan and Gilbert) and the shortspine thornyhead. Sebastolobus alascanus Bean
are fished commercially in southern California (Eschmeyer et al. 1983). Micro-
cotyle sebastes reach very high levels of infection in the bocaccio. Sebastes pau-
cispinis Ayres (Jensen et al. 1982) and may present a potential health problem
for stocks of rockfish. a very important component of the sport and commercial
fishery in southern California (Eschmeyer et al. 1983).
Nobel and King (1959) listed five new host records for the nematode Spiro-
camallanus pereirai Annereaux, 1946 from southern California. Our study in-
cludes four new hosts from southern California waters (Table 1): the round herring.
Etrumeus acuminatus (Dekay). speckledfin midshipman. Porichthys myriaster
(Hubbs and Schultz), queenfish, Seriphus politus (Ayres), and the white croaker.,
Genyonemus lineatus. Third stage larvae of S. pereirai were also found in the
intestines of topsmelt, Atherinops affinis (Ayres) collected in Malibu Lagoon.
Malibu. California.
Acknowledgments
I would like to thank Carol Lyons for the drawings of Choanodera moseri and
Proctoeces magnorus.
Literature Cited
Annereaux, R.R. 1946. A new nematode, Procamallanus pereirai, L. with a key to the genus. Trans.
Amer. Micr. Soc., 65:299-303.
Awakura, T.. and T. Nomura. 1983. Studies of parasites of marine salmon, Oncorhynchus masu—
V1. Hemiurid trematodes found in alimentary tract. Sci. Reports of the Hokkaido Fish Hatchery,
38:39-46.
Bray. R. A. 1983. On the fellodistomid genus Proctoeces Odhner. 1911 (Digenea), with brief com-
ments on two other fellodistimid genera. J. Nat. Hist.. 17:321—339.
——. 1990. A review of the genus Parahemiurus Vas & Pereira, 1930 (Digenea: Hemiuridae).
Systematic Parasitology, 15:1—21.
Dailey, M. D.. L. A. Jensen. and B. W. Hill. 1981. Larval anisakine roundworms of marine fishes
from southern and central California. with comments on public health significance. Calif. Fish
and Game, 67(4):240—245.
Durio, W.O.,and H. W. Manter. 1969. Some digenetic trematodes of marine fishes of New Caledonia.
HELMINTH PARASITES OF SOME SOUTHERN CALIFORNIA MARINE FISHES 117
III. Acanthocolpidae, Haploporidae, Gyliauchenidae, and Cryptogonimidae. J. Parasit., 55(2):
293-300.
Eschmeyer, W. N., E. S. Herald and H. Hammann. 1983. Peterson field guides: Pacific coast fishes.
Houghton Mifflin Company, Boston, Massachusetts, 355 pp.
Freeman, R. F. H., and J. Llewellyn. 1958. An adult digenetic trematode from an invertebrate host:
Proctoeces subtenuis. J. Mar. Bio. Ass. U.K., 43:113-123.
Gibson, D. I., and R. A. Bray. 1980. The Fellodistomidae (Digenea) of fishes from the north east
Atlantic. Bull. British Mus. Nat. Hist., 37(4):259-293.
1984. On Anomalotrema Zhukov, 1957, Pellamyzon Montgomery, 1957, and Opecoelina
Manter, 1934 (Digenea: Opecoelidae), with a description of Anomalotrema koiae sp. nov. from
north Atlantic waters. J. Nat. Hist., 18:949-964.
Jensen, L. A., R. A. Heckmann, M. Moser, and M. M. Dailey. 1982. Parasites of Bocaccio, Sebastes
; paucispinis, from southern and central California. Proc. Helminthol. Soc. Wash., 49(2):314—
317.
Love, M. S., and M. Moser. 1983. A checklist of parasites of California, Oregon, and Washington
marine and estuarine fishes. NOAA Technical Report. NMFS SSRF-777. U.S. Dept. of Com-
merce.
Manter, N. W. 1940. Digenetic trematodes of fishes from the Galapagos Islands and the neighboring
Pacific. Allan Hancock Pacif. Exped., 2(14):329-496.
Miller, D. J., and R. N. Lea. 1972. Guide to the coastal marine fishes of California. Fish Bull. no.
157. Cal. Dept. Fish and Game, Sacramento, California.
Moser, M.,andJ. Hsieh. 1992. Biological tags for stock separation in Pacific herring, Clupea harengus
pullasi in California. J. Parasit., 78(1):54-60.
Nobel, R. R., and R. E. King. 1960. The ecology of the fish Gillichthys mirabillis and one of its
nematode parasites. J. Parasit., 46:679-685.
Sekerak, A. D., and H. P. Arai. 1977. Some metazoan parasites of rockfishes of the genus Sebastes
from the northeastern Pacific Ocean. Syesis., 10:139-144.
Woolcock, V. 1935. Digenetic trematodes from some Australian fishes. Parasitology, 27(3):309-311.
Accepted for publication 15 January 1993.
Bull. Southern California Acad. Sci.
93(3), 1994, pp. 118-126
© Southern California Academy of Sciences, 1994
On the Identity of Snapping Shrimp Described and
Identified by W. N. Lockington, 1878 4,
Mary K. Wicksten
Department of Biology, Texas A&M University,
College Station, Texas 77842
Abstract.—W.N. Lockington (1878) wrote the first key and guide to the snapping
shrimp of North America. Many records came from the eastern Pacific Ocean,
from California to Panama. His paper includes 10 species recognizable today,
one record that probably includes more than one species under the same name,
and reports and descriptions of another six species that are unrecognizable or
questionable. The five species described as new by Lockington probably are var-
iants of species that have been described since 1878; if so, Lockington’s species
names may be available as senior synonyms. The identities of some of these
species are likely to remain uncertain because the original descriptions were brief
and without illustrations, the types have been lost and the range of variation
within most alpheid species is unknown.
The first attempt to prepare a summary and key to the snapping shrimp (family
Alpheidae) of North America was that of W. N. Lockington of the California
Academy of Sciences. In 1878, he published ““‘Remarks on some new Alphei, with
a synopsis of North American Species,”’ which presented 18 species. Most of these
came from the Pacific Ocean, including the Gulf of California. The paper included
species described by Lockington as well as others previously described by Say
(1818) and Kingsley (1878a). Lockington considered all of the species to belong
either to Alpheus or Betaeus.
In comparing Lockington’s account with modern works on alpheid shrimp, one
is surprised by the small number of species. The modern reader should remember
that extensive collections had yet to be made in much of the United States and
Mexico. Like many curators of the time, Lockington relied upon amateur collec-
tors and fishermen to send specimens to him for examination. The majority of
the species known in 1878 were either intertidal species or shallow subtidal species
that could be collected in holdfasts or rocks cast ashore after storms.
As was customary at the time, Lockington presented brief descriptions of new
species without illustrations. New species were compared with known species,
with important morphological differences emphasized. The features presented by
Lockington (1878) in many cases only allow the modern reader to determine the
genus to which the animal belongs. Features such as presence or absence of
epipods, shape of the dactyls of the walking legs, and many of the ridges, grooves
and spines of the chelae were not included or were described in general terms.
Although locations for the species were given, no type locality was designated if
118
IDENTITY OF SNAPPING SHRIMP 119
specimens came from more than one collecting site, nor was a particular specimen
mentioned in the text as being a holotype.
Lockington’s type material was deposited at the California Academy of Sciences.
A few duplicate or exchange specimens seem to have been sent to the Smithsonian
Institution or European museums, but as yet, no alpheid specimens have been
located. In 1906, almost all of the specimens of the Academy as well as much of
the building housing them were destroyed by the fire following the major earth-
quake in San Francisco. (See Bronson 1959 for an account of how a few of the
Academy’s specimens were rescued.) No invertebrate specimens are known to
have been saved; indeed, being preserved in alcohol, the crustaceans probably
were among the first specimens to catch fire.
As part of an ongoing attempt to prepare a master species list of carideans of
the tropical eastern Pacific Ocean, M. E. Hendrickx (Estacion Mazatlan, Universi-
dad Autonoma de Mexico) and I have been examining historic records of shrimp
of the area. Despite the uncertainty of identification of some of the species, the
records given by Lockington are useful for comparison of modern and historic
distributions of alpheid species. Kim and Abele (1988) considered five of Lock-
ington’s species of A/pheus to be valid, but noted that three of the supposed species
were unreported since Lockington’s report. However, most of Lockington’s names
have been ignored for over 50 years, and, if found to be senior synonyms of other
names, could be suppressed (International Commission on Zoological Nomen-
clature Article 79c, 1985). This paper provides the modern identification, when
known, of Lockington’s species of 1878 and attempts to identify the species that
remain unrecognized.
Methods
In the account of the species, the name used by Lockington and the page on
which it appears is provided, along with the localities as given in the paper of
1878. Localities are given in quotations if they are vague or questionable in the
original text. However, Lockington himself was inconsistent in giving localities,
giving localities as ‘““Port Escondido, Gulf of California,’ which can be interpreted
either as Port Escondido in the Gulf of California or Port Escondido as well as
the rest of the Gulf of California. (Probably the former interpretation is correct.)
Species described prior to 1878 and their current taxonomic status and known
range are given first. Species described in the paper are discussed later, with clues
to their current identification.
Many of Lockington’s specimens came from “Port Escondido” on the Gulf of
California (now Puerto Escondido, Baja California Sur, Mexico; 25°50’N,
111°19’W). More recent collecting has been carried out there by the University
of Southern California. I examined published records of alpheids collected at
Puerto Escondido during the trips of the Velero III and later expeditions (Wicksten
1983; Kim and Abele 1988), as well as unpublished records in the card catalogue
of the Hancock collections at the Natural History Museum of Los Angeles County.
The alpheids from these collections have not been completely identified to species
and catalogued by station. However, the existing records give some indication of
what species might occur there and therefore might be the same as some of
Lockington’s species.
120 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Alpheid Species Reported by Lockington (1878)
1. Species Described Prior to 1878:
y,
Alpheus affinis Kingsley, 1878a: Panama (p. 476).
Status: Now Alpheus normanni Kingsley, 1878b. (Alpheus affinis Kingsley,
1878a is a homonym of A. affinis Guise, 1854; see Kim and Abele 1988.)
Western Atlantic Ocean from Virginia to Sao Paulo, Brazil and eastern Pacific
Ocean from Gulf of California to Galapagos Islands (Kim and Abele 1988).
Alpheus bellimanus Lockington, 1877a: San Diego (California) (p. 470).
Status: Valid species; Monterey Bay, California to Galapagos Islands (Wick-
sten 1983; Kim and Abele 1988).
Alpheus clamator Lockington, 1877b: Santa Barbara “Islands,” California and
San Bartolme Bay, Baja California (p. 469).
Status: Valid species; Dark Gulch, Mendocino County, California (T. Chess,
U.S. National Marine Fisheries, personal communication, range extension) to
San Bartholome Bay, Baja California Sur (Wicksten 1984).
Alpheus cylindricus Kingsley, 1878a: Pearl Island, Bay of Panama (p. 478).
Status: Valid species; eastern and western Atlantic Ocean from Islands of
Principe, Sao Tome and Annobon; Bimini, Bahamas, Bermudas and Florida
through Barbados to Bahia, Brazil; Flower Gardens Reefs off Texas; eastern
Pacific Ocean from Gulf of California to Galapagos Islands (Kim and Abele
1988).
Alpheus aequidactylus Lockington, 1877b: Monterey, California (p. 472).
Status: Now Alpheopsis equidactylus; Monterey Bay to Cortez Bank, Cali-
fornia (Wicksten 1984).
Alpheus floridanus Kingsley, 1878a: Fort Jefferson, Florida (p. 476).
Status: Valid species; eastern and western Atlantic Ocean from Guinea to
Congo and Principe Island; Gulf of Mexico to Bahia, Brazil; eastern Pacific
Ocean from Gulf of California to Ecuador (Kim and Abele 1988).
Alpheus heterochaelis Say, 1818: La Paz, San José Island, Amortiguado Bay,
Mulege Bay and Port Escondido, west side of Gulf of California; Magdalena
Bay, Baja California (p. 475).
Status: Although A. heterochaelis is a valid species, it is considered to inhabit
only the western Atlantic (Christoffersen 1984).
Remarks: Kim and Abele (1988) reported that records of A. heterochaelis
from the eastern Pacific Ocean by Kingsley (1878a) and Rathbun (1900) in-
cluded in part A. bouvieri A. Milne Edwards, 1878. However, it is unclear how
they came to this identification, for they did not indicate that they examined
the specimens on which the records of A. heterochaelis were based. Lockington’s
specimens may have included 4. californiensis Holmes, 1900 which has been
collected at Magdalena Bay, and other species of the ““Edwardsi” group of
Alpheus, which resemble A. heterochaelis. Lockington noted that his larger
specimens “‘showed traces of a varied coloration,” which suggests that he had
more than one species.
Alpheus minor Say, 1818: no locality given by Lockington (p. 472). Say reported
it from the “coasts of the southern states, and of East Florida.”
Status: Now Synalpheus minus (Say, 1818); Bermuda and North Carolina to
Alagoas, Brazil (Chace 1972).
IDENTITY OF SNAPPING SHRIMP 121
Alpheus panamensis Kingsley, 1878a: Acajutla, ““Central America”’ (now Pacific
coast of El Salvador) and Panama (p. 473).
Status: Valid species; eastern Pacific Ocean from Costa Rica to Peru (Kim
and Abele 1988).
Alpheus parvimanus Kingsley, 1878a: Panama (p. 477).
Status: Identity unknown.
Remarks: Kingsley’s description suggests that the species belonged to the
diverse ““Edwardsi” group, but, other than mentioning that the larger chela had
‘“‘a constriction of both margins posterior to the articulation of the dactylus,”
there are too few details to compare the shrimp with those described more
recently.
Alpheus sulcatus Kingsley, 1878a: Bay of Panama, Zorritas, Peru (p. 475). Kingsley
did not designate a type locality.
Status: Valid species; circumtropical except western Atlantic Ocean, eastern
Pacific Ocean from Gulf of California to Peru (Wicksten 1983; Kim and Abele
1988).
Betaeus aequalis (Kingsley, 1878a): Catalina Island, California (p. 478).
Status: Synonym of Betaeus harfordi (Kingsley, 1878a); Fort Bragg, California
to Magdalena Bay, Baja California Sur (Hart 1964).
Betaeus longidactylus Lockington, 1877a: San Diego, California (p. 480).
Status: Valid species; Elkhorn Slough, California to Baja California Norte;
also northern Gulf of California to Tepoca Bay, Sonora, Mexico (Hart 1964).
2. Species Described as New in Paper of 1878:
Alpheus barbara Lockington, 1878: Santa Barbara, California (p. 471).
Status: Probably a damaged or malformed specimen of Alpheus clamator
Lockington, 1877b (Wicksten 1990).
Remarks: Lockington differentiated the supposed new species from A. cla-
mator by the absence of a spine on the “basal joint of the antennae”’ (the
basicerite), ‘different proportions of the carpal joints of the second pair,” and
the ‘“‘want of meral spines on the posterior pairs” (of pereopods). However,
about 2% of specimens of A. clamator lack a spine on at least one basicerite.
The “different” proportions of the carpal joints are so close to those of a typical
A. clamator as to be nearly indistinguishable. Specimens of A. clamator usually
have a strong spine at the distal end of the merus of the third pereopod, but
this could be missing in an animal regenerating a limb. (Kingsley [1878a] and
Lockington [1878] both mentioned that the unique holotype of A. barbara was
““damaged)’”. Examination of over 1000 specimens collected throughout the
range of A. clamator failed to locate any animals matching Lockington’s de-
scription. The type locality of A. barbara, Santa Barbara, California, is inhabited
by A. clamator.
Alpheus fasciatus Lockington, 1878: “‘Port Escondido, Gulf of California” (p. 478).
Status: Kim and Abele (1988) treated A. fasciatus as a distinct and valid
species ‘“‘until more information about this species is available.”
Remarks: Kim and Abele (1988) believed that A. fasciatus was distinct from
A. paracrinitus Miers, 1881 on the basis of the length of the stylocerite, which,
according to Lockington’s description, was “longer than the first segment of the
peduncle.” Otherwise, Lockington’s description is almost identical to that of
122 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
A. paracrinitus. According to Lockington, A. fasciatus had “‘no ocular spines,
rostrum about equal to diameter of eye, surface between rostrum and eyes
slightly depressed but with no distinct sulcus, small spine on basai joint of
antennae below, larger hand smooth, nearly cylindrical, dactylus works verti-
cally, smaller hand small, smooth, cylindrical and slender.”
The name “fasciatus,” meaning banded, gives a clue to the species’ modern
identity. Lockington described the color as ““carapax (sic) and abdomen alter-
nately banded with bright red and white: larger hand red, with marblings of
white in some cases.” This color pattern is common in A. paracrinitus. Alpheus
paracrinitus has been collected in the southern Gulf of California, where A.
fasciatus also was reported. Alpheus paracrinitus is reported to be circumtrop-
ical, in the eastern and western Atlantic Oceans. the Gulf of Mexico, tropical
Indo-West Pacific to Hawaii, and eastern Pacific Ocean from the Gulf of Cal-
ifornia to the Galapagos Islands (Kim and Abele 1988).
Coutiére (1897) reported A. fasciatus as occurring at Djibouti in the Red Sea.
He mentioned seeing specimens of the species from Baja California, collected
by Leon Diguet. However, Banner and Banner (1981) noted that the specimens
from the Red Sea probably were A. alpheopsides Coutiére, 1905, which has
been collected there recently. Coutiére’s specimens of A. fasciatus have not been
located at the Muséum National d’Histoire Naturelle in Paris. but. should they
exist, they would be very useful for comparison with other specimens in de-
termining the modern identity of A. fasciatus.
Knowlton and Mills (1992) compared color patterns of specimens of A. par-
acrinitus from the Pacific coast of Panama with those of specimens from the
Caribbean coast, and found that three different color patterns existed. Specimens
from the Caribbean coast showed two discrete patterns that were different from
those of the Pacific population. At present. it is not known whether or not these
color patterns correspond to separate species or to within-species variation in
A. paracrinitus. If the color patterns are shown to be related to morphological
or enzymatic differences in the shrimp, it may be useful to distinguish the eastern
Pacific population by a different species name. Further study may indicate that
A. fasciatus Lockington is an available senior synonym for the eastern Pacific
species.
Alpheus laeviusculus Lockington, 1878: Port Escondido, Mulege Bay, and other
points on the “Californian” shore of the Gulf of California (p. 474).
Status: Probably a synonym of Synalpheus digueti Coutiére, 1909.
Remarks: Lockington’s choice of the trivial name was poor from the start.
for A. leviusculus Dana, 1852 isa homonym. However. the description indicates
that Lockington’s specimen probably belonged to the genus Synalpheus. The
description states that the front was “‘trispinose. the ocular spines triangular.
almost equal in length to the triangular rostrum, which is not divided from the
eye-shields by a rostrum,” the chelipeds were “very unequal” in size, and the
posterior pereopods had “‘dactyli bifid at tip, the upper spine longer than the
lower.”
Coutiére (1909) recognized that Lockington’s species was a Synalpheus and
attempted to redescribe and rename it. However, Coutiére’s new specimens
came from off San Nicolas Island in southern California. not the Gulf of Cal-
ifornia. The new specimens, named S. Jockingtoni, probably did not belong to
IDENTITY OF SNAPPING SHRIMP 123
Lockington’s original species. In S. /ockingtoni, the rostrum is longer than the
orbital spines, not almost equal to their length. The upper spine of the basicerite
is shorter than the lower, but is noticeable; Lockington described this spine as
““small’’ for his species. In S. /ockingtoni, the stylocerite is shorter than the first
segment of the antennular peduncle; in Lockington’s species, the stylocerite was
described as reaching the middle of the second segment of the antennular
peduncle.
Lockington’s description could belong to Synalpheus digueti Coutiére, 1909.
In the species, the rostrum is the same length as the orbital spines. The scapho-
cerite does not reach to the end to the antennular peduncle, the merus of the
larger cheliped bears a small spine, and the fingers of the chelae bear dark tips.
These features, mentioned by Lockington, occur in specimens of S. digueti.
Synalpheus digueti is a common intertidal species of the southern Gulf of
California, and has been collected at Puerto Escondido (Jens Knudsen station
K121, collections of Allan Hancock Foundation). It ranges from Guaymas, Gulf
of California to the Galapagos Islands (Wicksten 1983).
Alpheus spinicaudatus Lockington, 1878: Port Escondido (p. 477).
Status: Kim and Abele (1988) considered A. spinicaudatus to be a valid
species.
Remarks: Kim and Abele (1988) considered A. spinicaudatus to be a distinct
species on the basis of the movable finger of the major chela moving obliquely.
However, Lockington’s description otherwise resembles that of A. hebes Kim
and Abele, 1988 or a related species of the ““Edwardsi’”’ group. The rostrum is
described by Lockington as “‘very short, continued backward between the eye-
shields as a low carina, no ocular spine.” According to the description by
Lockington, the stylocerite was as long as the first segment of the antennular
peduncle, the spine of the scaphocerite was longer than the blade or antennular
peduncle. The major chela lacked a spine on the merus, the hand bore “‘con-
strictions” on the upper and lower surfaces and a sulcus running “backward”
longitudinally at right angles to the upper constriction. The smaller chela was
rounded and smooth, with the fingertips sharp and curved inwards, crossing
each other. The carpus of the second pereopod had 5 articles, the merus of the
third to fifth pereopods lacked a spine, the telson was elongate with tapering
sides and 2 pairs spinules on the upper surface as well as a pair of posterior
spines.
Alpheus hebes, which most closely resembles the description of A. spinicau-
datus, has a short rostrum and no noticeable eyeshields. It ranges from Ensenada
de San Francisco, Sonora, Gulf of California to the Galapagos Islands, and has
been collected at Puerto Escondido (Velero IIT sta. 670-37) (Kim and Abele
1988). Although A. hebes is described as having the movable finger of the major
chela closing vertically, Banner and Banner (1982, fig. 83) showed that the shape
and angle of the finger varied in 4. edwardsi within the species. Whether or not
similar variation occurs in A. hebes is unknown.
Alpheus tenuimanus Lockington, 1878: Port Escondido, Gulf of California (p.
473).
Status: Coutiére (1899) tentatively identified this as a species of Synalpheus.
Remarks: Lockington described this species as “arched in profile, much higher
in the centre (sic).”’ The front was “‘trispinose, rostrum much longer than ocular
124 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
spines, reaching middle of second joint of peduncle of antennulae,”’ the ocular
spines were “slender, projecting from the centre of the convex front of the
eyeshields.” The stylocerite reached beyond the first segment of the azitennular
peduncle, the basicerite bore a spine and the scaphocerite reached the end of
the antennular peduncle. The first pereopods bore a spine at the distal end of
the merus. The major chela was elongated and smooth, rounded above and
below, with a shallow sulcus and a shorter one “‘above”’ extending obliquely
upward from the carpal articulation on the inner side of the hand. The smaller
chela had a similar merus and an elongated, smooth, cylindrical chela. The
second pereopod had a 5-jointed carpus, the third to fifth pereopods were
without spines or spinules. The telson was elongate, slightly tapering, with 2
pairs of spinules on the dorsal surface and a spine on each posterolateral border.
Lockington’s mention of the “hands” (of the chelipeds) “‘equal in length, not
greatly differing size, dissimilar’’ casts some doubt on the identification of A.
tenuimanus as a species of Synalpheus, for in the latter genus, the hands differ
greatly in size. In species of the eastern Pacific Ocean, it is easy to distinguish
between a major and a minor chela; the major chela is not “elongated.”” The
ocular spines were described as “‘projecting from the centre of the convex front
of the eyeshields,” rather than “triangular,” which suggests that the species
belonged to Alpheus, not Synalpheus.
Two other “‘trispinose’’ genera of alpheids occur in the Gulf of California:
Alpheopsis and Salmoneus. However, Lockington himself commented on the
“triangular projection on each side of the base of the telson” in the description
of A. aequidactylus and yet made no mention of such a feature in the description
of Alpheus tenuimanus. Species of Salmoneus have peculiar chelipeds and a very
broad rostrum, easily distinguished from that of species of Alpheus. It seems most
likely that A. tenuimanus was a species of Alpheus.
Most of the description of A. teniumanus suggests a species of the “‘Sulcatus”’
group. Of the species known in the eastern Pacific Ocean, A. panamensis Kingsley,
A. felgenhaueri Kim and Abele and A. splendidus Coutiére are the most similar.
All have a rostrum at least as long as the first segment of the antennular peduncle;
the major chela neither has spinules on the merus or characteristic spines, notches
or bumps, and there are no large sulci adjacent to the rostrum. However, Lock-
ington stated that the posterior pairs (of pereopods) were “without spines or
spinules on any of the joints.”
Alpheus exilis Kim and Abele, 1988, described from specimens without che-
lipeds, may be found to be asynonym of A. tenuimanus. Both descriptions mention
the ocular spines; in both, the third pereopod lacks spines and spinules. However,
the rostrum and stylocerite are reported to be shorter than the first segment of
the antennular peduncle in A. exi/is. Length of the rostrum, however, has been
found to be variable in at least two species of A/pheus (Banner and Banner 1982,
figs. 20, 43). Alpheus exilis has been taken at Puerto Escondido (Velero III sta.
667-37) and Sullivan Bay, Galapagos Islands (Kim and Abele 1988). One hopes
that, in the future, a series of specimens with chelipeds can be collected and
compared with both descriptions to determine whether or not there are one or
two species agreeing substantially with the two descriptions.
IDENTITY OF SNAPPING SHRIMP 125
Discussion
Unless Lockington’s original specimens are found, identification of many of his
species is likely to remain uncertain. Even if some of his specimens are located,
identification of the species may remain questionable because, even today, the
range of morphological variation within a single species of alpheid shrimp is likely
to be unknown. Banner and Banner (1982), ina lengthy study of species of Alpheus,
contrasted the great variation in features such as length of the rostrum and details
of the movable finger of the major chela in various species, while other features
remained uniform over a range of the entire Indo-Pacific region. Features such
as length of the rostrum and shape of the dactyls of the pereopods in at least two
species were found to vary with age and maturity of the animal. Kim and Abele
(1988) used features such as rows of setae, spinules of the chelipeds, shape of the
ridges and grooves of the major chela and relative lengths of the stylocerite,
rostrum and segments of the first antenna to distinguish between species. To date,
no one has compared these features among large series of hundreds of animals
along the entire eastern Pacific coast. Color patterns of most species are unknown.
Some species are known from less than 10 specimens. The dispersal capabilities
of eastern Pacific alpheid larvae also are unknown.
In instances in which Lockington’s species can be recognized, the species occur
today in the localities from which he had specimens. The unidentifiable or ques-
tionable species at least are similar to species that still occur in the areas from
which Lockington’s material came. Lockington’s records indicate that at least a
few alpheids have occupied the same areas for over 115 years.
Literature Cited
Banner, D. M., and A. H. Banner. 1981. Annotated checklist of the alpheid shrimp of the Red Sea
and the Gulf of Aden. Zool. Verhand. Leiden, No. 190:1-99.
, and 1982. The alpheid shrimp of Australia. Part III: the remaining alpheids, prin-
cipally the genus Alpheus, and the family Ogyrididae. Rec. Aust. Mus., 34:1-362.
Bronson, W. 1959. The earth shook, the sky burned. Chronicle Books, San Francisco.
Chace, F. A., Jr. 1972. The shrimps of the Smithsonian-Bredin Caribbean expeditions with a sum-
mary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithson.
Contr. Zool., No. 98:1-79.
Christoffersen, M. L. 1984. The western Atlantic snapping shrimps related to Alpheus heterochaelis
Say (Crustacea, Caridea), with the description of a new species. Papeis Avulsos Zool., Sao Paulo,
35:189-208.
Coutiére, H. 1897. Note sur quelques alphéidés nouveaux ou peu connus rapportés de Djibouti
(Afrique Orientale). Bull. Mus. Hist. Nat., Paris, 3:233-236.
1899. Les ‘“‘Alpheidae” morphologie externe et interne, formes larvaires, bionomie. Théses
présentées a la Faculté des Sciences de Paris, sér. A, 321 (980):1-599.
1905. Marine Crustacea. XV. Les Alpheidae. Pp. 852-921 in Fauna and geography of the
Maldive and Laccadive Archipelagoes 2. (J. S. Gardiner, ed.), Cambridge Univ. Press, Cam-
bridge.
1909. The American species of snapping shrimps of the genus Synalpheus. Proc. U.S. Natl.
Mus., 36:1—-93.
Dana, J. D. 1852. Crustacea, Part I. In United States Exploring Expedition during the years 1838,
1839, 1840, 1841, 1842, under the Command of Charles Wilkes, U.S.N., 13:1-685.
Guise, W. V. 1854. Upona new species of “Alpheus” discovered upon the coast of ““Herm” (Channel
Islands). Ann. Mag. Nat. Hist., ser. 2, 14(82):275-280.
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. Natl. Mus., 115:431:466.
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Holmes, S. J. 1900. Synopsis of California stalk-eyed Crustacea. Occ. Pap. Calif. Acad. Sci., 7:1-
265.
Kim, W., and L. G. Abele. 1988. The snapping shrimp genus A/pheus from the eastern Pacific
(Decapoda: Caridea: Alpheidae). Smithson. Contr. Zool., No. 454:1-119.
Kingsley, J. S. 1878a. A synopsis of the North American species of the genus Alpheus. Bull. U.S.
Geol. Geogr. Survey Terr., 4:189-199.
1987b. Notes on the North American Caridea in the Museum of the Peabody Academy of
Science at Salem, Massachusetts. Proc. Acad. Nat. Sci. Phila., 1878:89-98.
Knowlton, N.,and D. K. Mills. 1992. The systematic importance of color and color pattern: evidence
for complexes of sibling species of snapping shrimp (Caridea: Alpheidae: Alpheus) from the
Caribbean and Pacific coasts of Panama. Proc. San Diego Soc. Nat. Hist., 18:1—5.
Lockington, W. N. 1877a. Remarks on the Crustacea of the Pacific coast with description of some
new species. Proc. Calif. Acad. Sci., 7:28-36.
1877b. Description of seventeen new species of Crustacea. Proc. Calif. Acad. Sci., 7:41-48.
—. 1878. Remarks on some new Alphei, with a synopsis of the North American species. Ann.
Mag. Nat. Hist., ser. 5, 1:465-480.
Miers, E. J. 1881. Ona collection of Crustacea made by Baron Hermann-Maltzan at Goree Island,
Senegambia: Macrura. Ann. Mag. Nat. Hist., ser. 5, 8:204—220, 259-281, 364-377.
Milne Edwards, A. 1878. Description de quelques espéces nouvelles de Crustacés provenant du
voyager aux iles du Cap Vert de MM. Bouvier et de Cessac. Bull. Soc. Philom. Paris, ser. 7,
2:225-232.
Rathbun, M. J. 1900. The decapod Crustacea of West Africa. Proc. U.S. Natl. Mus., 22:271-316.
Say, T. 1818. An account of the Crustacea of the United States. J. Acad. Nat. Sci. Phila., 1:57—80.
Wicksten, M. K. 1983. A monograph on the shallow water caridean shrimps of the Gulf of California,
Mexico. Allan Hancock Monogr. Mar. Biol., 13:1—59.
. 1984. New records of snapping shrimps (family Alpheidae) from California. Proc. Biol. Soc.
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Accepted for publication 12 February 1994.
Bull. Southern California Acad. Sci.
93(3), 1994, pp. 127-134
© Southern California Academy of Sciences, 1994
Amphisamytha fauchaldi: A New Species of Ampharetid
(Annelida: Polychaeta) from the
Hydrothermal Vents at Guaymas Basin, Mexico
Vivianne Solis-Weiss and Pablo Hernandez-Alcantara
Laboratorio de Ecologia Costera, ICMyL, UNAM,
Apdo. Postal 70-305, México, D.F. 04510
Abstract. —A new species of the polychaete family Ampharetidae, Amphisamytha
fauchaldi, is described from the hydrothermal vents at Guaymas Basin in the Gulf
of California, Mexico, at a depth of 2020 m. This is a very common species in
the sampling site. The environment is also described and comments are given
about the species inhabiting the area.
The hydrothermal vents have been studied intensely since they were discovered
in 1977 (Corliss et al. 1979). They constitute a most unusual habitat where tectonic
activity and toxic emissions (mostly H,S) at high temperatures (270—400°C) com-
bine to harbor an exotic and dense fauna (Tunnicliffe 1992).
At Guaymas Basin, the hydrothermal vents differ from others known in the
East Pacific Rise, in that there are sediments entering the Gulf of California from
the Colorado River. Sediments accumulating at a rate of more than 1 m/1000
years, have covered the rift floor to a depth up to 400 m. Hydrocarbons are formed
and percolate through the area (Simoneit 1985). Dense accumulations of organ-
isms, dominated by the giant tubeworm Riftia pachyptila Jones, occur in close
relationship with the hot springs.
Among the abundant biological material obtained from the Riftia washings
collected during the Guaymas Basin expedition of February 1988, an undescribed
ampharetid was found in large numbers.
The holotype is deposited in the collection of the National Museum of Natural
History, Smithsonian Institution (USNM). Paratypes and representatives of the
species described were deposited in the collections of the Australian Museum
(AM), the British Museum (Natural History) (BM), the Hamburg Zoologische
Museum (HMZ), The National Museum of Wales (NMW), the Zoological Mu-
seum at the University of Copenhagen (ZMUC), the Muséum National d’ Histoire
Naturelle de Paris (MNHN), the Los Angeles County Natural History Museum
(LANHM), and the Instituto de Ciencias del Mar y Limnologia, UNAM, collection
(ICML-UNAM), as well as the Dr. J. Frederick Grassle (FG) collection.
Study Area
Guaymas Basin is located in the Central region of the Gulf of California, Mexico,
approximately at 27°00’North latitude and 111°25’West longitude. The sampling
site is located in the Southern Basin in 2000-2020 m depths.
The site was discovered in 1980 (Lonsdale et al. 1982), and has been studied
since 1982 (Lonsdale 1984; Grassle 1986). Previous geophysical studies have
127
128 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
shown that in the Gulf of California two large spreading centers exist: the Northern
and the Southern Basins.
The Guaymas Basin differs from other hydrothermal active sites in the Pacific
Rise in the particular geographic and geological conditions of its spreading centers.
Because of land erosion and sediments from the Colorado River and planktonic
blooms, there is an organic rich sediment deposit over the area about 400 m deep
which prevents therefore lava eruption. One of the consequences of the transport
of hot fluids through the organically enriched sediments is the geologically rapid
formation of hydrocarbons which percolate through the area (Lonsdale et al. 1980;
Simoneit and Lonsdale 1982; Lonsdale 1984).
In the Southern Basin where the samples were taken, the vertical temperature
gradients around the bottom are greater than 4°C/m. The sulphur rich fluids that
discharge through the chimneys do so at temperatures ranging from 270-—314°C.
The site of collection was on the side of a seamount and from soft sediments
away from other visible megafauna.
Dense thickets of Riftia pachyptila dominate the biota at the seamount. They
are often lined with abundant mucus which provides an adequate habitat for a
large array of organisms, among them A. fauchaldi, a polynoid polychaete and
limpets. In addition, there are large mats of light orange and yellowish bacteria
known as Beggiatoa, several galatheid crabs and some clams. The clams found
around the area are small and do not dominate as in other hydrothermal vents.
Materials and Methods
The specimens were collected by one of the authors (VSW) in February 1988
using DSRV “ALVIN” Dive 1979, during the Guaymas Basin expedition of 1988
to the Gulf of California. Collections included Riftia washings. The specimens
were washed and sieved through a 0.3 mm mesh, then fixed in buffered formalin
and later preserved in 70% alcohol (Fauchald 1977).
Results
Amphisamytha fauchaldi new species
Figures 1 A-E
Material examined. —Guaymas Basin, Southern Trough, Riftia washings, Alvin
Dive 1979, 18 Feb 1988, 2014 m, holotype (USNM) and 124 paratypes (USNM
holotype 168087 + 20 paratypes 168088); AM 10 paratypes (W21709), BM 15
paratypes (1993:5-14); HMZ 10 paratypes (P21986); NMW 10 paratypes
(NMW.Z.1993.027); ZMUC 5 paratypes (POL-00020); MNHN 5 paratypes (UD
240 (vial A 923)); LANHM 10 paratypes (LACM-AHF1655); ICML-UNAM 29
paratypes (PO-68-002); and FG 10 paratypes (no numbers assigned).
Description.—The holotype is a complete specimen measuring 20 mm long.
The paratypes vary from 3.5 mm to 20.5 mm, the most abundant size class being
13.5-14 and 15-16 mm. The smallest ovigerous females are 9.5 mm in length.
In Table 1, some morphometric measurements are given from 22 specimens
chosen among the total (which was too high to be convenient to include here) so
as to give an idea of the variability of the characters present in the different size
classes of the mature specimens. Color in life as well as in preserved specimens
is light brown. The deep-red heartbody could be seen dorsally over the first seven
segments through the translucent body wall in living material.
A NEW SPECIES OF AMPHARETID FROM GUAYMAS BASIN, MEXICO 129
0.5 cm
0.5 mm
Fig. 1. 1A. A. fauchaldi, holotype; entire worm, dorsolateral view. 1B. A. fauchaldi, holotype;
dorsal view of the prostomium with branchial disposition. 1C. A. fauchaldi, holotype; thoracic uncinus.
1D. A. fauchaldi, holotype; abdominal uncinus. 1E. A. galapagensis, holotype; dorsal view of prosto-
mium with branchial disposition.
The eggs found in the gravid females are small, subcircular with a white central
vessicle about one third of the length of an entire brownish egg.
There are 17 thoracic setigers (14 uncinigers) and 13 to 15 abdominal uncinigers
(14 in the holotype); (Fig. 1A).
Prostomium indistinctly trilobed and slightly notched with no glandular ridges.
Mouth bilobed, oral tentacles numerous, deciduous, ventrally grooved and re-
tractile.
Four pairs of finely annulated long branchiae (approximately 15 mm in holo-
type), narrowly grooved ventrally, individually inserted across the dorsal surface
SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
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A NEW SPECIES OF AMPHARETID FROM GUAYMAS BASIN, MEXICO 131
of segments 2 to 5 so that the first pair is associated to the last asetigerous segment.
The inner pair is associated with the last asetigerous segment. The inner pair is
associated with the 5th segment in the normal ampharetid fashion. There is no
gap between the branchial groups (Fig. 1B).
Segments | and 2 fused, ventrally forming the lower lip which shows two
ventrolateral grooves. Segment 3 asetigerous. Paleae absent.
First thoracic setiger reduced with a bundle of 10-12 capillary winged smooth
notosetae. Second and third setigers are dorsal to the others.
Notopodial lobes bear 15—18 winged smooth capillary setae in two rows with
about 7-8 long setae behind 8 short ones.
Neuropodial lobes (uncinigerous pinnules) each with a single transverse row of
uncini from setiger 4 to the end of abdomen. Number of thoracic uncini per setiger
varies with length of the organism. It can be as high as 68 in longest specimens
and around 38 in small specimens (Table 1). The shape of the thoracic uncini is
shown in Fig. 1C. They bear four denticles above a squared-off base with a distinct
prow; subrostral tip tiny, uppermost denticle smallest.
Abdomen consisting of 13 to 15 (directly related to size) gradually tapering
setigers bearing only neuropodial lobes, distinctly different from thoracic ones,
rounded, with a glandular pad covering a few uncini and prolonged dorsally.
Shape of thoracic and abdominal uncini shown in Fig. 1C and 1D. Four denticles
present. Number of abdominal uncini per setiger is also a function of length,
varying from 18 in the smallest to 38 in the largest organisms (Table 1).
Pygidium rounded, bearing two large lateral papillae, three small dorsal papillae
and a smoother ventral pad, but no true anal cirri. Anal aperture terminal.
Tubes. —Specimens embedded in clumps of many tubes parallel to each other
and held together by a mucous substance. There were as many as 20 tubes in a
clump. Outer tubes formed by aggregates of fine sediments and different debris,
arranged as transverse white and dark brown areas. Inner part of tubes formed
by whitish, translucent, vertically oriented linings. Only part of the anteriorly
oriented branchiae protruding from the tubes which are simple or branched. Some
specimens were found attached to the outer walls of the Riftia tubes to which
they adhered by similar mucus.
Remarks. — Along with Amphisamytha fauchaldi, which was by far the most
abundant organism, we collected in the clumps several black limpets (7, one of
them very small), 26 specimens of Ophryotrocha akessoni Blake, six specimens
of O. platykephale Blake (Solis-Weiss and Hilbig 1992) and one large alvinellid:
Paralvinella grasslei.
Etymology. —This species is dedicated to Dr. Kristian Fauchald, an outstanding
polychaetologist, as a small attempt to acknowledge all the help and friendship
provided for so many years.
Discussion
This species belongs to the group of ampharetids with four rather than three
pairs of branchiae. It is easily distinguished from the species A. japonica (Hessle)
and A. bioculata (Moore) as mentioned by Zottoli (1983) by the lack of anal cirri
and by the possession of glandular pads in the abdominal segments.
A. fauchaldi is closely related to A. galapagensis Zottoli, also a vent dweller
found in the Galapagos site, from which it differs mainly in that there is a clear
132 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Table 2. List of the Annelid Polychaetes reported from the Guaymas Basin.
Family Alvinellidae s 44
Paralvinella grasslei Desbruyeres & Laubier, 1982
Family Cossuridae
Cossura sp. 1 Grassle et al., 1985
Family Dorvilleidae
Exallopus jumarsi Blake, 1985***
Ophryotrocha akessoni Blake, 1985
Ophryotrocha platykephale (Blake, 1985)***
Family Euphrosinidae
Euphrosine rosacea Blake, 1985
Family Glyceridae
Glycera profundi Chamberlin, 1919
Family Hesionidae
Nereimyra alvinae Blake, 1985
Orseis grasslei Blake, 1985***
Family Nereididae
Ceratocephale pacifica Hartman, 1960***
Nereis sandersi Blake, 1985
Family Polynoidae
Bathykurila guaymasensis Pettibone, 1989***
Branchinotogluma grasslei Pettibone, 1985b
Branchinotogluma sandersi Pettibone, 1985b
Branchiplicatus cupreus Pettibone, 1985a
Lepidonotopodium williamsae Pettibone, 1984
Lepidonotopodium riftense Pettibone, 1984
Macellicephaloides alvini Pettibone, 1989***
Opisthotrochopodus alvinus Pettibone, 1985b
Family Sigalionidae
Neoleanira racemosa (Fauchald. 1972)
Family Spionidae
Lindaspio dibranchiata Blake & Maciolek, 1992***
Spiophanes sp. 1 Grassle et al., 1985
*** Species found so far only in the Guaymas Basin.
branchial gap between the two groups of branchiae in A. galapagensis (Fig. 1E);
such a gap is absent in A. fauchaldi. This is a much larger species than A. gala-
pagensis, which measures 3-10 mm in length compared to 14-20 mm in A.
fauchaldi.
General remarks about the polychaete fauna of Guaymas Basin. —So far there
have been 22 species of polychaetes found in the Guaymas Basin site (Table 2).
From these, 17 were newly described from the area (Desbruyeres and Laubier
1982: Blake 1985; Grassle et al. 1985: Pettibone 1984, 1985a,. b, 1989; Blake and
Maciolek 1992) and two presumably undescribed (Cossura sp. 1 and Spiophanes
sp. 1) (Grassle et al. 1985). Eight species of polynoids mainly in the genus Bran-
chinotogluma Pettibone, and dorvilleids with three species represent the highest
species richness for this group. The diversity was low as is normal for these
A NEW SPECIES OF AMPHARETID FROM GUAYMAS BASIN, MEXICO 133
environments but endemism was high. Seven species are known only from Guay-
mas. Closer examination of additional material may prove that species formerly
assigned to already described ones will turn out to be new as in the case of A.
fauchaldi. The Gulf of California is already known for its endemism, brought
about in great part by its configuration with its relatively small opening to the
Pacific Ocean.
Acknowledgements
We wish to thank Dr. J. F. Grassle for inviting us to participate in the project
and for his constant help and encouragement and Dr. K. Fauchald for his con-
tinuous help. The project on the Mexican part was supported by the Instituto de
Ciencias del Mar y Limnologia, UNAM.
Literature Cited
Blake, J. A. 1985. Polychaeta from the vicinity of deep-sea geothermal vents in the Eastern Pacific.
1. Euphrosinidae, Phyllodocidae, Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae
and Maldanidae. Biol. Soc. Wash. Bull., 6:67-101.
, and N. Maciolek. 1992. Polychaeta from deep-sea hydrothermal vents in the Eastern Pacific.
III. A new genus and two new species of Spionidae from the Guaymas Basin and Juan de Fuca
Ridge with comments on a related species from the Western North Atlantic. Proc. Biol. Soc.
Wash., 105(4):723-732.
Corliss, J. B., J. Dymond, L. I. Gordon, J. M. Edmond, R. P. Von Herzen, R. D. Ballard, K. Green,
D. Williams, A. Bainbridge, K. Crane, and T. H. Van Andel. 1979. Submarine thermal springs
on the Galapagos Rift. Science, 203:1073-1083.
Desbruyeres, D., and L. Laubier. 1982. Paralvinella grasslei, new genus, new species of Alvinellidae
(Polychaeta, Ampharetidae) from the Galapagos Rift geothermal vents. Proc. Biol. Soc. Wash.,
95(3):484—494.
Fauchald, K. 1977. The Polychaete worms. Definitions and keys to the Orders, Families and Genera.
Allan Hancock Scientif. Series, 28:1-388.
Grassle, J. F. 1986. The ecology of deep-sea hydrothermal vent communities. Advances in Marine
Biology, 23:301-—362.
, L. S. Brown-Leger, L. Morse-Porteous, R. Petrecca, and I. Williams. 1985. Deep-sea fauna
of sediments in the vicinity of hydrothermal vents. Biol. Soc. Wash. Bull., 6:443-452.
Lonsdale, P. 1984. Hot vents and hydrocarbon seeps in the Sea of Cortez. Oceanus, 27(3):21-25.
, J. L. Bishoff, V. M. Burns, M. Kastner, and R. E. Sweeney. 1980. A high temperature
hydrothermal deposit on the seabed at a Gulf of California spreading center. Earth and Planetary
Science Letters, 49:8-20.
, R. Batiza, and T. Simkin. 1982. Metallogenesis at seamounts on the East Pacific Rise. Marine
Technology Society Journal, 16(3):54—-60.
Pettibone, M. 1984. Two new species of Lepidonotopodium (Polychaeta: Polynoidae: Lepidonoto-
podinae) from the hydrothermal vents of the Galapagos and the East Pacific Rise at 21° N.
Proc. Biol. Soc. Wash., 97(4):847-863.
1985a. An additional new scale worm (Polychaeta: Polynoidae) from the hydrothermal rift
area off western Mexico at 21° N. Proc. Biol. Soc. Wash., 98(1):150-157.
1985b. Additional branchiate scale-worms (Polychaeta: Polynoidae) from Galapagos hy-
drothermal vent and rift area off Western Mexico at 21° N. Proc. Biol. Soc. Wash., 98(2):447—
469.
. 1989. Polynoidae and Sigalionidae (Polychaeta) from the Guaymas Basin, with descriptions
of two new species, and additional records from hydrothermal vents of the Galapagos Rift, 21°
N, and seep-sites in the Gulf of Mexico (Florida and Louisiana). Proc. Biol. Soc. Wash., 102(1):
154-168.
Simoneit, B. 1985. Hydrothermal petroleum composition and utility as a biogenic carbon source.
Biol. Soc. Wash. Bull., 6:49-56.
134 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
—. and P. Lonsdale. 1982. Hydrothermal petroleum in mineralized mounds at the seabed of
Guaymas Basin. Nature, 295:198-—202.
Solis-Weiss, V..and B. Hilbig. 1992. Redescription of Ophryotrocha platykephale Blake (Polychaeta,
Dorvilleidae) from the Guaymas Basin Hydrothermal Vents. Bull. South. Cal. Acad. Sci., 91(2):
92-96.
Tunnicliffe, V.
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1992. Hydrothermal-vent communities of the deep sea. American Scientist, 80(4):
1983. Amphisamytha galapagensis, a new species of ampharetid polychaete from the
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ecosystems. Proc. Biol. Soc. Wash., 96:379-391.
Accepted for publication 27 September 1993.
Bull. Southern California Acad. Sci.
93(3), 1994, pp. 135-136
© Southern California Academy of Sciences, 1994
INDEX TO VOLUME 93
Amphisamytha fauchaldi, n. sp., 128
Bloom, Peter H.: The Biology and Current Status of the Long-eared Owl in
Coastal Southern California, 1
Braden, Gerald T., see Cynthia H. Stubblefield
Bradley, Richard A.: Cultural Change and Geographic Variation in the Songs of
the Belding’s Savannah Sparrow (Passerculus sandwichensis beldingi), 91
Choanodera moseri n. sp., 111
Dailey, Murray M., see Patrick J. Frost
Demetrion, Robert A., and Richard L. Squires: Middle Miocene Pholadid Borings
at the Base of the Isidro Formation, Arroyo Mezquital, Baja California Sur,
Mexico, 83
Frost, Patrick J.,. and Murry M. Dailey: Helminth parasites of Some Southern
California Fishes with a Redescription of Proctoeces magnorus Manter, 1940
(Digenea: Fellodistomidae) and Description of Choanodera moseri sp. n.
(Digenea: Apocreadidae), 110
Gobalet, Kenneth W.: Additional Archaeological Evidence for Colorado River
Fishes in the Salton Basin of Southern California, 38
Grismer, L. Lee, Jimmy A. McGuire, and Bradford D. Hollingsworth: A Report
on the Herpetofauna of the Viscaino Peninsula, Baja California, Mexico, with
a Discussion of its Biogeographic and Taxonomic Implications, 45
Hernandez-Alcantara, Pablo, see Vivianne Solis-Weiss.
Hollingsworth, Bradford, D., see Lee L. Grismer
Lea, Robert N., and Florence McAlary: Occurrence of the Swallow Damselfish,
Azurina hirundo, from Islands off Southern California, 42
McAlary, Florence, see Robert N. Lea
McGuire, Jimmy A., see Lee L. Grismer
Shane, Susan H.: Occurrence and Habitat Use of Marine Mammals at Santa
Catalina Island, California from 1983-91, 13
Soiseth, Chad R.: Occurrence of the Anostracan Branchinecta lindahli (Packard)
on the California Channel Islands, 81
Solis-Weiss, Vivianne, and Pablo Hernandez-Alcantara: Amphisamytha fauchal-
di: A New Species of Ampharetid (Annelida: Polychaeta) from the Hydro-
thermal Vents at Guaymas Basin, Mexico, 127
135
136 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Squires, Richard L, see Robert A. Demetrion
Stubblefield, Cynthia H., and Gerald T. Braden: Denning Characteristics of Black
Bears in the San Gabriel Mountains of Southern California, 30 /
Wicksten, Mary K.: On the Identity of Snapping Shrimp Described and Identified
by W. N. Lockington, 1878, 118
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CONTENTS ,
Cultural Change and Geographic Variation in the Songs of the Belding’s
Savannah Sparrow (Passerculus sandwichensis beldingi). By Richard
A. Bradley ee
Helminth Parasites of Some Southern California Fishes with a Redescription
of Proctoeces magnorus Manter, 1940 (Digenea: Fellodistomidae) and
Description of Choanodera moseri sp. n. (Digenea: Apocreadidae). By
Patrick J. Frost and Murray M: Dailey 2 Eee
On the Identity of Snapping Shrimp Described and Identified by W. N.
Lockington, 1878. By Mary K. Wicksten 220 2 Se
Amphisamytha fauchaldi: A New Species of Ampharetid (Annelida: Poly-
chaeta) from the Hydrothermal Vents at Guaymas Basin, Mexico. By
Vivianne Solis-Weiss and Pablo Hernandez-Alcantara_ ss t—“‘CS™;C«*dr
Index to Volume 93 135
COVER: Belding’ Savannah Sparrow and an audiospectrogram of its song. Drawn by Richard A.
Bradley.