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ISSN 0038-3872 


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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 
California State University, Fullerton 


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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 


0.740 5 
e 
>> 
e 
wu 0.672 
fq) 
i e 
Oo & @ 
D ° i 
Be é e e @ S 
@ 
6D 0.605 5 e 
5 
= e 
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=) 
= 5 
SG O87 4| 
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@ 
0.4707 
T I | T 
(0) 150 300 450 600 


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 


A ae 8 K By L 
Ss =) % at 
2 2 | 
z,| “7 z.| & z.| © ze] * 
= = = === ape —— 7A a 
nt Date MU nde ss eaten ree ree | ee eel get ANN 
as as as as 
N 
c 2 D an M = 
C 2 2 | 24 
2, wis z,/ win =,) =.) &1 
= % = = al = — : a ~ Z x 
om joee 00 — 
as as as as 
Pp 
E F = ie) 
2 2 2 2 
= — =,| ™ = =) 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. 


Literature Cited 


Adret-Hausberger, M. 1986. Temporal dynamics of dialects in the whistled songs of starlings. Ethol- 
ogy, 71:140-152. 

Andrew, R. J. 1962. Evolution of intelligence and vocal mimicking. Science, 137:585-589. 

Baker, M. C. 1975. Song dialects and genetic differences in White-crowned Sparrows (Zonotrichia 

leucophrys). Evolution, 29:226-241. 

,and M. A. Cunningham. 1985. The biology of bird-song dialects. The Behavioral and Brain 

Sciences, 8:85—133. 

, and D. B. Thompson. 1985. Song dialects of White-crowned Sparrows: historical processes 

inferred from patterns of geographic variation. Condor, 87:127-141. 

Baptista, L. F. 1985. The functional significance of song sharing in the White-crowned Sparrow. 
Canadian Journal of Zoology, 63:1741-1752. 

, and J. R. King. 1980. Geographical variation in song and song dialects of Montane White- 

crowned Sparrows. Condor, 82:267-284. 

, and M. L. Morton. 1982. Song dialects and mate selection in Montane White-crowned 

Sparrows. Auk, 99:537-547. 

,and M. L. Morton. 1988. Song learning in Montane White-crowned Sparrows: from whom 

and when. Anim. Behav., 36:1753-1764. 

, and L. Petrinovich. 1984. Social interaction, sensitive phases and the song template hy- 

pothesis in the White-crowned Sparrow. Anim. Behav., 32:172-181. 

, and L. Petrinovich. 1986. Song development in the White-crowned Sparrow: social factors 

and sex differences. Anim. Behav., 34:1359-1371. 

Bowman, R. I. 1979. Adaptive morphology of song dialects in Darwin’s finches. J. fir Ornithol., 
120:353-389. 


108 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 


Bradley, R. A. 1973. A population census of the Belding’s Savannah Sparrow, Passerculus sand- 

wichensis beldingi. Western Bird Bander, 48:40—43. 

1977. Geographic variation in the song of Belding’s Savannah Sparrow (Passereulus sand- 
wichensis beldingi). Bulletin of the Florida State Museum. Biological Sciences, 22:57—100. 
Bradley, D. W.. and R. A. Bradley. 1983. Application of sequence comparison to the study of bird 

songs. P. 209 in Time warps. string edits, and macromolecules: the theory and practice of 
sequence comparison. (D. Sankoff and J. B. Kruskal, eds.). Addison-Wesley Publishing Co. 
Inc., Reading, Mass. 

Cade, T. J.. and G. A. Bartholomew. 1959. Sea-water and salt utilization by Savannah Sparrows. 
Physiol. Zodl., 32:230—238. 

Chew. L. 1981. Geographic and individual variation in the morphology and sequential organization 
of the song of the Savannah Sparrow (Passerculus sandwichensis). Can. J. Zool.. 59:702—713. 

Chilton, G., M. R. Lein, and L. F. Baptista. 1990. Mate choice by female White-crowned Sparrows 
in a mixed-dialect population. Behav. Ecol. Sociobiol.. 27:223-227. 

Cooper, B. A., and E. C. Murphy. 1985. Savannah Sparrow sings a White-crowned Sparrow song. 
Anim. Behav... 33:330-331. 

Dawkins, R. 1976. The selfish gene. Oxford Univ. Press, New York. viii + 224 pp. 

DeWolfe. B. B., L. F. Baptista, and L. Petrinovich. 1989. Song development and territory estab- 
lishment in Nuttall’s White-crowned Sparrows. Condor, 91:397—-407. 

Garrett, K.. and J. Dunn. 1981. Birds of Southern California: status and distribution. Los Angeles 
Audubon Society, Los Angeles. 

Gibbs, H. L. 1990. Cultural evolution of male song types in Darwin’s Medium Ground Finches, 
Geospiza fortis. Anim. Behav., 39:253—263. 

Gill, F. B. 1990. Ormmithology. W. H. Freeman and Co., New York. 660 pp. 

Gish, S. L.. and E. S. Morton. 1981. Structural adaptations to local habitat acoustics in Carolina 
Wren songs. Z. Tierpsychol., 56:74-84. 

Grinnell, J.. and A. H. Miller. 1944. The distribution of the birds of California. Pacific Coast 
Avifauna, 27:1-608. 

Hafner, D. J.. and K. E. Petersen. 1985. Song dialects and gene flow in the White-crowned sparrow, 
Zonotrichia leucophrys nuttalli. Evolution, 39:687-694. 

Handford, P. 1988. Tmill rate dialects in the Rufous-collared Sparrow. Zonotrichia capensis, in 
northwestern Argentina. Canad. J. Zool., 66:2658—2670. 

Immelmann, K. 1969. Song development in the Zebra Finch and other estrildid finches. Pp. 61-74 
in Bird vocalizations. (R. A. Hinde, ed.). Cambridge Univ. Press. xiii + 394 pp. 

Jenkins, P.F. 1977. Cultural transmission of song patterns and dialect development in a free-living 
bird population. Anim. Behav., 25:50—78. 

Kroodsma, D.E. 1974. Song learning. dialects, and dispersal in the Bewick’s Wren. Z. Tierpsychol.. 
35:352-380. 

Lougheed, S. C., and P. Handford. 1992. Vocal dialects and the structure of geographic variation in 
morphological and allozymic characters in the Rufous-collared Sparrow. Zonotrichia capensis. 
Evolution, 46:1443-1456. 

Marler, P..and M. Tamura. 1962. Song “dialects” in three populations of White-crowned Sparrows. 
Condor, 64:368—377. 

Massey. B. W. 1977. A census of the breeding population of the Belding’s Savannah Sparrow in 

California, 1977. State of California, Department of Fish and Game. Sacramento. 

1979. The Belding’s Savannah Sparrow. U.S. Army Corps of Engineers, Work Order No. 
6, Los Angeles. 

McGregor, P. K.,and D.B. A. Thompson. 1988. Constancy and change in local dialects of the Corn 

Bunting. Ornis Scandinavica, 19:153—159. 

. V. R. Walford, and D. G. C. Harper. 1988. Song inheritance and mating in a songbird with 

local dialects. Bioacoustics, 1:107—129. 

Morton, E. 1982. Grading, discreteness, redundancy, and motivation-structural rules. Pp. 183-212 
in Acoustic communication in birds. Vol. 1. (D. E. Kroodsma and E. H. Miller, eds.), Academic 
Press. New York. 

Mundinger. P_C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations 
of birds. Pp. 147-208 in Acoustic communication in birds. Vol. 2. (D. E. Kroodsma and E. H. 
Miller, eds.), Academic Press, New York. 


SONG EVOLUTION IN BELDING’S SPARROWS 109 


Nicolai, J. 1959. Familientradition in der Gesangsentwicklung des Gimpels (Pyrrhula pyrrhula L.). 
J. Ornith., 100:39—-46. 

Nottebohm, F. 1969. The song of the Chingolo, Zonotrichia capensis, in Argentina: description and 
evaluation of a system of dialects. Condor, 71:299-315. 

Payne, R. B. 198la. Song learning and social interaction in Indigo Buntings. Anim. Behav., 29:688- 

697. 

. 1981b. Population structure and social behavior: models for testing the ecological significance 

of song dialects in birds. Pp. 108—120 in Natural selection and social behavior. (R. D. Alexander 

and D. W. Tinkle, eds.), Chiron Press, New York. 532 pp. 

. 1983. Bird songs, sexual selection, and female mating strategies. Pp. 55—90 in Social behavior 

of female vertebrates. (S. K. Wasser, ed.), Academic Press, New York. 

1985. Behavioral continuity and change in local song populations of Village Indigobirds 

Vidua chalybeata. Z. Tierpsychol., 70:1-44. 

——.,, W. L. Thompson, K. L. Fiala, and L. L. Sweany. 1981. Local song traditions in Indigo 

Buntings: cultural transmission of behavior patterns across generations. Behaviour, 77:199— 

DONe 

, and L. L. Payne. 1993. Song copying and cultural transmission in Indigo Buntings. Anim. 

Behav., 46:1045-1065. 

Petrinovich, L. 1985. Factors influencing song development in the White-crowned Sparrow (Zo- 

notrichia leucophrys). J. of Comparative Psychol., 99:15-29. 

1988. Individual stability, local variability and the cultural transmission of song in White- 

crowned Sparrows (Zonotrichia leucophrys nuttalli). Behaviour, 107:208—240. 

, and L. F. Baptista. 1984. Song dialects, mate selection and breeding success in White- 

crowned Sparrows. Anim. Behav., 32:1078-1088. 

Poulson, T. L., and G. A. Bartholomew. 1962. Salt balance in the Savannah Sparrows. Physiol. 
Zo6l., 35:109-112. 

Rothstein, S. I., and R. C. Fleischer. 1987. Vocal dialects and their possible relation to honest status 
signalling in the Brown-headed Cowbird. Condor, 89:1-23. 

Slater, P. J. B., L. A. Eales, and N. S. Clayton. 1988. Song learning in Zebra Finches (Taeniopygia 
guttata): progress and prospects. Adv. Stud. Behav., 18:1-34. 

Sokal, R. R., and F. J. Rohlf. 1981. Biometry. 2nd ed. E. H. Freeman and Co., San Francisco. 

Trainer, J. M. 1983. Changes in song dialect distribution and microgeographic variation in the song 
of White-crowned Sparrows Zonotrichia leucophrys nuttalli. Auk, 100:568—-582. 

Wheelwright, N. T., and J. D. Rising. 1983. Savannah Sparrow (Passerculus sandwichensis). In The 
birds of North America, no. 45. (A. Poole and F. Gill, eds.), Philadelphia: The Academy of 
Natural Sciences; Washington, D. C.: The American Ornithologists’ Union. 

Wiens, J. A. 1982. Song pattern variation in the Sage Sparrow (Amphispiza belli): dialects or epi- 
phenomena? Auk, 99:208—229. 

Zann, R. 1990. Song and call learning in wild Zebra Finches in south-east Australia. Anim. Behav., 
40:811-828. 

Zink, R. M., and G. F. Barrowclough. 1984. Allozymes and song dialects: a reassessment. Evolution, 
38:444-448. 


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. 


126 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 


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. 

Wash., 97:186-190. 

1990. On the status of Alpheus barbara Lockington (Crustacea: Caridea: Alpheidae). Proc. 

Biol. Soc.. Wash., 103:100-102. 


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. 
336-349. 
Zottoli, R. 


1992. Hydrothermal-vent communities of the deep sea. American Scientist, 80(4): 


1983. Amphisamytha galapagensis, a new species of ampharetid polychaete from the 


vicinity of abyssal hydrothermal vents in the Galapagos Rift, and the role of this species in rift 
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.