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7) 







SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 




ULLETIN 



Volume 70 



Number 1 




w 



BCAS-A70(1) 1-56 (1971) 



APRIL 1971 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Board of Directors 



John J. Baird 

Russell E. Belous 

Donald B. Bright 

Jules Crane, Jr. 

John E. Fitch 

Takashi Hoshizaki 

John L. Mohr 



William J. Morris 
Donald J. Reish 
Charles E. Rozaire 
Elbert L. Sleeper 
Andrew Starrett 
David L. Walkington 
Stewart L. Warter 



Charles R. Weston 



Officers of the Academy 



Andrew Starrett 

Elbert L. Sleeper 

Jules Crane, Jr. 

Charles E. Rozaire 

Russell E. Belous 

Donald J. Reish 

Robert J. Lavenberg 

Roberta S. Greenwood 



President 

First Vice President 

Second Vice President 

Secretary 

Treasurer 

Editor 

Managing Editor 

Index Editor 



Membership in the Southern California Academy of Sciences 

Membership is open to all scientists of all branches and to any person interested 
in the advancement of science. 

Annual Members $ 8.00 

Student Members 5.00 

Life Members 150.00 

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



The Bulletin is published three times a year by the Academy. Address all communica- 
tions to the appropriate officer in care of the Natural History Museum of Los Angeles 
County, Exposition Park, Los Angeles, California 90007. 



BULLETIN OF THE SOUTHERN CALIFORNIA 
ACADEMY OF SCIENCES 

Volume 70 August 27, 1971 Number 1 



CONTENTS 

Changes in the intertidal algal flora of the Los Angeles area since the survey by 

E. Yale Dawson in 1956-1959. By Thomas B. Widdowson 2 

A new species of Simopelta from Costa Rica (Hymenoptera: Formicidae). By Roy 

R. Snelling 16 

A revision of the genus Loandalia Monro with description of a new genus and 
species of pilgargiid polychaete. By Raymond R. Emerson and Kristian 
Fauchald 18 

Taxonomy and distribution of the Arctic species of Lucicutia (Copepoda: Cal- 
anoida). By Julio Vidal 23 

A review of the genus Boccardia Carazzi (Polychaeta: Spionidae) with descriptions 
of two new species. By James A. Blake and Keith H. Woodwick 31 

Parasitic mites of Surinam. VIII. A new genus and species of chigger, Fauranius 
atecmartus, and additional records of species (Acarina: Trombiculidae). By 
James M. Brennan and F. Lukoschus 42 

Variation and geographic distribution in some Argentine and Chilean Osmylidae, 
with a new species of Kempynus (Neuroptera). By Phillip A. Adams 45 

RESEARCH NOTES 

Distribution of some southern California Kangaroo rats. By Blair A. Csuti 50 

An evaluation of two methods of measuring metacarpal length in Artibeus lituratus 
(Olfers) (Chiroptera: Phyllostomatidae). By Donald R. Patten 51 

A new synonymy in the genus Chilicola (Hymenoptera: Colletidae). By Roy R. 

Snelling 52 

Cuvier's beaked whale, Ziphias cavirostris, from Barbados. By David K. Caldwell, 

Warren F. Rathjen, and Melba C. Caldwell 52 

Additional specimens of the colubrid snake Amastridium veliferum Cope from 
Costa Rica, with comments on a pseudohermaphrodite. By Larry David 
Wilson and Douglas C. Robinson 53 

News and Notes 55 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(1): 2-16, 1971 

CHANGES IN THE INTERTIDAL ALGAL FLORA OF THE 

LOS ANGELES AREA SINCE THE SURVEY 

BY E. YALE DAWSON IN 1956-1959 

Thomas B. Widdowson 1 



Abstract: Each of the 15 stations set up by Dawson between Pt. Dume 
and Dana Point were reoccupied two or three times during the period 1968 to 
1970. The data obtained were analysed in a manner comparable to Dawson's. 
The most conspicuous reductions were found in the flora of stations exposed 
to direct human interference, and in the species which characteristically grow 
half buried in sand. Methods are suggested by which problems associated with 
sand movement, seasonal variation, and taxonomy may be minimized. 



From the years 1956 and 1959, the late E. Yale 
Dawson surveyed the intertidal algae in southern 
California for the State Water Pollution [Quality] 
Control Board. He examined the collections which 
W. A. Setchell and N. L. Gardner preserved from 
numerous visits to the White Point area between 
1895 and 1912, and from these deduced that the 
flora at that time contained at least 60 species of 
conspicuous benthonic algae (Dawson, 1965a: 
223). With a view to providing an accumulation of 
information for future workers, Dawson set up 44 
stations along the exposed rocky shores between 
Point Conception and San Diego (Fig. 1). These 
stations consisted of a fixed shore base point and a 
transect running down the beach at a certain bearing 
from this point. Dawson's published field notes 
(1965b), color transparencies and annotated aerial 
photographs (deposited in the Allan Hancock Foun- 
dation Herbarium) served to identify the base point 
and the bearing of each transect. His field notes 
provided qualitative information about the fre- 
quency of algae found during each visit in an unde- 
fined narrow strip along each transect. He visited 
each station between one and four times for a total of 
103 visits. During each visit he made collections 
which he filed in duplicate under station and species 
in the Herbarium of the Allan Hancock Foundation. 
In general, these collections probably provide a 
more complete and accurate list of the species at 
each station than do the notes, but the notes provide 
the only information about frequency. 

Dawson published two brief discussions of his 
results (1959a, 1965a). In these, he calculated the 
number of conspicuous species observed at each 
station, averaged over several visits! 1965a. Fig. 8.1). 
He then assigned a numerical weight of 1 for a spe- 
cies reported to be rare or scant at a station for a 
particular visit, 3 for occasional to frequent, and 10 
for common to abundant. These weights, totalled 



over all visits and stations, provided the basis for a 
list of the "46 most common marine plants of south- 
ern California in order of their relative frequency" 
(1965a, Table 8.1). He also presented these weights 
in two Jarge tables ( 1 959a, Tables I and II) with 1 89 
species on the Y-axis and a selection of station 
visits, grouped in four seasons, along the X-axis. 
He planned a more explicit analysis of seasonal 
variation (1959: 181) but apparently never com- 
pleted this (1965a: 224). 

Dawson based some of his conclusions on a 
comparison of his 1956-9 data with the data he 
derived from Setchell and Gardner's 1895-1912 
collections at White Point. In the same way, the 
conclusions in this study are based on a comparison 
of Dawson's 1956-9 data with those gathered in 
1968-70. These comparisons are necessarily limited 
by the extent and precision of the earlier study 
(which cannot now be improved) and by the avail- 
ability of valid statistical techniques for use in 
making the comparison. 

One long term comparison similar to the present 
study is that of Bellamy, Bellamy, John, and Whit- 
tick (1967). The effects of pollution along about 
50 miles of the Durham and Northumberland coast 
are measured by a reduction of 9-70% in species 
number since 1861. It is only recently that other 
quantitative measurements, such as percentage 
cover and numbers of individuals per unit area, 
have become common practice in the study of 
intertidal plants. This means that comparisons 
using such quantitative measures are short term 
studies, with the earlier data taken in the recent 
past (Bellamy, Clarke, Jones, Whittick and Darke, 
1967; Cowell, 1969). These short term studies also 



'Department of Biology, California State College at Long Beach, 
California 90801 



1971 



SOUTHERN ( A I, HORN I A ALGAE 



have a practical advantage over long term studies in 
that both the earlier and the later surveys can be 
made by the same observer. 

Conclusions derived from the observations oi' 
populations before and alter, with and without 
pollution, should be confirmed by experimental 
study of the effects of individual pollutants on 
individual species. The enormous number of prob- 
lems here can only be reduced to manageable size 
by prior observational work, so it is not surprising 
to find that little has been done thus far. Honey 
(1968) investigated the effects of various detergents 
on Ascophyllum, Cladophora, Bryopsis, Prasino- 
cladus, Porphyra, Acrochaetium, and Polysiphonia. 
Okamura, et al. (1926) correlated heavy mortality 
of intertidal Porphyra in Tokyo Bay with episodes 
of heavy fog. They demonstrated experimentally 
the lethal effect of soot particles from the fog con- 
taining 70% to <0.7% adsorbed sulfur dioxide and 
also of air containing more than 0.0 1 % free SO2. 

Any survey designed to be the first part of a long 
term comparison study should certainly use the 
best current techniques, and these now include 
quantitative measurements and experimental work. 
However, it is perhaps inevitable with the develop- 
ment of new techniques that the person who comes 
to make the second part of the study should be 
dissatisfied with the first. In this context, it is perhaps 
ironic that the paper by Okamura, et al. (1926) 
should seem so contemporary. 

METHODS 

Choice of stations. — The 15 Dawson stations 
between Point Dume and Dana Point were each 
visited two or three times during the years 1968 to 
1970. This particular set of stations is the maximum 
available for a study centered on Los Angeles, since 
a larger set would be affected by the infrequency 
of stations southeast of Dana Point, producing a 
bias in favor of the stations northwest of Los 
Angeles (Fig. 1). These 15 stations occur in three 
groups; along the Malibu coast, on the Palos Verdes 
Peninsula, and along the Orange County coast. 
They are separated by stretches of sandy beach 
along Santa Monica Bay and Long Beach which 
form a physically unsuitable substratum for large 
benthonic algae. The stations near Point Dume 
and Dana Point appear to enjoy natural conditions, 
while varying degrees of water and air pollution, 
and intensity of collecting activity, can be observed 
at the stations in between. 

Of the 32 station visits made in this study, 16 
were made by students, whose contributions are 
acknowledged below. Specimens were collected to 
substantiate each record and. in the case of student 



work, were used by the authoi to check the identi- 
fications made. These specimens were deposited in 

the herbarium of the ( alifornia State ( ollege at 
Long Beach. 

Taxonomic problems. — Many intertidal alj;ac 
cannot be identified to species or, sometimes, even 
to genus in the field. Sometimes this problem arises 
because such identification requires laboratory 
techniques, and sometimes because described 
species appear to intergrade or show the need for 
further taxonomic work in some other way. Since 
many of the areas studied are now marine preserves. 
and since indiscriminate collecting in the past may 
have been a factor in the decline of the algal flora, 
it is desirable to reduce the need for collections as 
much as possible in the future. Species which 
cannot readily be distinguished in the field have 
been grouped together in this study. The taxonomic 
categories used in analyzing both the 1956-9 and 
the 1968-70 data include single species (e.g. Gigar- 
tina leptorhynchos), pairs of species (e.g. Gelidium 
coulteri or G. crinale), genera (e.g. Viva spp). 
pairs of genera (e.g. Gracilaria or Gracilariopsis). 
or even larger groupings such as diatoms or blue- 
green algae. These groupings are listed in Table I. 
and are referred to as 'entities' below, wherever the 
word species would be inapropriate. The taxonomic 
system used is basically the same as that published 
by Dawson (1959a, Tables I and II). The marine 
grass Phyllospadix is, in effect, included with the 
algae in this study. 

Number of species per station. — The easiest 
analysis of the algal flora, and the one least subject 
to taxonomic problems, is an estimate of the number 
of entities per station. Dawson (1965a. Fig. 8.1) 
used the arithmetic mean of his observations for 
stations which he visited more than once. This does 
not seem to be the best statistic available, since 
observations of this type are subject to a biased 
error. (It should be easier to miss a species which 
is there, than it is to record one which is not there.) 
The statistic presented in Figure 2 is the total 
number of entities observed during all the visits 
made to a station. If entity A. B. and C were 
observed on one visit, while entity B. C. and D were 
observed on a second, the number shown in Figure 
2 would be 4. This is substantially the same method 
which Dawson himself used in analysing Setchell 
and Gardner's records from White Point (Dawson. 
1965a: 223). This statistic will tend to increase 
with each successive visit, so in Figure 2 the value 
for 1956-9 and for 1968-70 is calculated for the 
same number of visits at each station. However, the 
number of visits used varies from station to station, 
so one station should not be compared with another. 
Dawson derived the figure of 60 species for Setchell 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 



J* " 










Pt. ^**»jj&; 










Conception ^l| 


F 1 "" ' ' 


. •.. • 






-*W- 




.v.-.Mqljbu;. 

pt. \v. 

Dume •/.■■,.*}."} 


Los 
Angeles 

Long . 
"^^Beach 









Palos ^tbrange 

Verdes •V :: -r« 
not *ii.-S-.°\ ' 






%> 


Dana Pt. >&•.'• • 




<N> 




y & 


\. 


'• . . 


• Reoccupied 1968-70 




^v 




\iV • 


O Best 1956-9 




V 




J*'." 


X Others 




N^ 




JivSan Diego 


50 




- 




IK.* .' - 


MILES 








\V:. • • 



Figure I. Stations for sampling intertidal algae, established by E. Yale Dawson during the period 1956-9. 



and Gardner's collections from many visits, so it 
probably represents a flora not greatly larger than 
that found at Station 19 or 22 today. 

Scoring species for frequency. — Dawson's 
numerical scoring for qualitative observations of 
frequency were changed in two ways. First, the 
score for 'common to abundant' was changed from 
10 to 9, so that it could be accommodated in one 
column on the punched cards used for this analysis. 
Second, for each visit and species a score was 
assigned to each zone (upper, middle, and low inter- 
tidal, and tide pools) instead of to each station as a 
whole. In this way, the maximum score per entity 
per visit was increased from 10 to 36, and more 
information was derived from the original notes. 

Dawson (1959a, Table I and II) tried to present 
variations in frequency by species, station, and 
season simultaneously. These modes of variation 
are presented separately in this study (Figs. 3 and 
4, 5 and 6). 

In Figure 3, the scores totalled over all stations 
are compared for the 10 most frequent entities, 
using data from the 15 stations in the Los Angeles 
area in 1968-70, (right), the same stations in 1956-9 
(middle), and the 15 stations outside the Los Angeles 
area where Dawson found an average of more than 
30 species in 1956-9 (left). The scales are adjusted 
to allow for the fact that 32 visits are involved in 



the data on the right, 34 visits for those in the 
middle, and 39 visits for those on the left. The 
numbers in circles show the rank of a certain species 
when it is not in the top 10 for that particulai 
column. The data on the left are included as the 
best available indication of the original compo 
sition of the algal flora of southern California. Tota 
scores in the Los Angeles area for species below 
the top 10 are presented in Table I. The position ol 
the stations used are shown in Figures 1, 7 and 8 

Placing species in non-taxonomic groups. — | 
Dawson remarked (1965a), that articulated coral- 
lines increased and leafy red algae decreased ir 
areas exposed to sewage pollution. In Figure 4, the 
total scores for a number of such groupings are 
shown. Data from Dawson's best 15 stations outside 
the Los Angeles area (A), Dawson's 15 stations ir 
the Los Angeles area (B), and these same station; 
during the period 1968-70 (C) are compared in the 
same way as in Figure 3. The values for the Lo; 
Angeles area are weighted to compensate for the 
fact that the first set were obtained in 39 visits, the 
second in 34 visits, and the third in 32 visits. 

In grouping entities by zone, each was assignee 
to high interfidal, mid-intertidal, low intertidal, oi 
tide pools according to where it had the highesi 
total score. In respect to geographical distribution 
species were grouped into those which appear to be 



SOI II II I. RN CALIFORNIA Al.C.AL 




34 10 

T. 
? 18 

MALIBU 



-i — rr 

S. MONICA 7 33 16 LONG BEACH 

PALOS VERDES 

STATIONS 



9 21 

ORANGE CO. 



Figure 2. Overall number of species reported from stations in the Los Angeles area. Open bars: 
1956-9. Solid bars: 1968-70. Stippled bar: Dawson's analysis of Setchell and Gardner's collections 
(1895-1912). 



mainly northern forms near their southern limit in 
southern California, mainly southern forms near 
their northern limit, or cosmopolitan. A species 
was counted as endemic if its distribution seems 
to be centered on southern California and does not 
range outside California or Pacific Baja California. 
Information was mainly taken from Dawson ( 1 96 1 ). 
In the classification by habit, only the articulated 
corallines represent a taxonomic category. The 
massive species are mainly kelps. The turf category 
was given precedence over the leafy in the case of 
Ulva, because of the characteristic habit of the most 
common species in southern California, U. cali- 
fornica. The leafy category includes a few brown 
algae as well as red. The crustose forms include 
encrusting corallines as well as Ralfsia and encrust- 
ing non-coralline red algae. A few species are found 
characteristically coming up through sand while 
firmly attached to rocks buried in the sand. This 
habit seems to be distinct from that of the great 
number of species which are occasionally found 



half buried by rising sand levels. The determination 
of the month of maximum occurrence of entities is 
discussed below (p. 10). The assignment of each 
entity in these various categories is given in Table I. 

Total scores by station. — In Figure 5. the total 
scores for each station in the Los Angeles area in 
1968-70 are compared with the scores for the years 
1956-9. In the case of stations visited more than 
once, the arithmetic mean is used. The standard 
deviation between different total scores for the 
same station is 40 for the 1968-70 observations and 
35 for the 1956-9 ones. 

Before using the arithmetic mean of multiple 
observations for Figure 5. I needed to investigate 
the possibility that there are systematic seasonal 
variations in the score at any one station. This prob- 
lem was made difficult by the fact that Dawson's 
work shows a strong bias with respect to the 
seasons of January-March. April-June. July-Sep- 
tember, and October-December (1959a. Tables I 
and II). He made very few visits durine the summer 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 



and very few visits to the same station during the 
same season in a different year. 

The sign test for the paired case (Alder and 
Roessler, 1964: 150) was used to test for the pres- 
ence of systematic seasonal variation in the total 
scores at each station. In this statistic, all the re- 
peated visits made by Dawson to the same station at 
different seasons were classified by the season of the 
earlier and the later visit. Thus all the differences 
between the pairs of same station-different season 
visits were grouped over all stations as Spring-Sum- 
mer, Spring-Autumn, Spring-Winter, Summer- 



Autumn, Summer-Winter, and Autumn-Winter dif- 
ferences. The positive or negative sign of these 
differences was determined, and a greater than 
random preponderance of a particular sign would 
indicate the presence of a systematic variation, 
although it would not determine its size. 

No systematic variation could be detected by the 
sign test at the 5% level of significance. Moreover, 
the 1968-70 visits were found to have much the 
same bias against summer observations as did the 
1956-9 study. So it was thought safe to use the 
arithmetic mean for the values shown in Figure 5. 



oncouvsMentil 



400 



LU 
H 

O 

o 

to 



200 



BEST 56-9 




400 



200 



L. A. 68-70 



Figure 3. Total scores for the most common species (or groups of species) compared for the origi- 
nal southern California flora (as indicated by Dawson's best 15 stations outside the Los Angeles 
area), the 15 Los Angeles stations 1956-9, and the same stations 1968-70. 



1971 



SOUTHERN CALIFORNIA ALGAE 
iNciii oiouriD ■' 



DI5THIIUIION 



HAII T 



• AX I MUH 

MONTH 



1000 




Pool 




A » 

= BEST 56-9 



■ c 

= L.A.56-9 




C A 

= L. A. 68-70 



Figure 4. Total scores for various non-taxonomic groups of species compared for: A — Dawson's 
15 best stations outside the Los Angeles area; B — the 15 stations in the Los Angeles area in the 
period 1956-9: and C — the same stations as B in the period 1968-70. 



Seasonal variations — Nevertheless. I have the 
subjective impression that conspicuous seasonal 
variations, at least in quantity, do occur in the 
algal flora, with a maximum in late spring and a 
minimum in late summer. There are three possible 
ways in which such a variation could exist and 
not appear in the test. First, shifting sand appears 
to be the major factor in variation (Dawson. 1965a: 
225). If this shifting is random with regard to 
season at any one station, it could obscure season- 
al variation. Second, the period of any variation, or 
differences from year to year in the time at which 
peaks occur, could be such that Dawson's division 
of the year into four seasons would not be precise 
enough to detect seasonal variation. Third, a 
summer flora of small species could replace a 
winter flora of large species, so that the number of 
species remains much the same while the total bulk 
of algae changes from season to season. 

These possibilities were tested by an investigation 



of the total scores given each entity for each month 
(Fig. 6). Data were used from all Dawson's station 
visits throughout southern California, and from 
the first visit made to each station in the present 
study. (It was thought that the inclusion of the 
second and third visits made in the present study 
would bias the data too much in favor of stations 
in the Los Angeles area). The distribution of station 
visits by month in the data used was found to range 
from 20 in November and 18 in January and 
February, to 2 in August and 4 in July and Septem- 
ber. Data from the earlier visit in August were added 
to those in July to make a 'month' of 50 days with 
5 visits, and the later August visit was added to 
those in September to make a month of 40 days 
with 5 visits also. The score for each species in each 
month was then weighted by a ratio of 20 to the 
number of visits made in each particular month. 
Running means of three months were used to 
smooth the weighted data of each species. The 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 



Table I. Scores and classification given each species 
(or other taxonomic category). 



SPECIES OR ENTITY 


SCORE 


SCORE 




CLASSIFICATION ACCORDING TO: 






1956-9 


1968-70 


LEVEL 


DISTRIB. 


HABIT 


MAX. 
MONTH 


Coral Una vancouveriensisl 














gracilis 


568 


420 


MID 


North 


Art. coral. 


MAR. 


Gigartina canaliculata 


401 


285 


MID 


Endemic 


Other 


NONE 


Egregia laevigata 


332 


260 


LOW 


South 


Massive 


NONE 


Bossiella spp. 


280 


174 


LOW 


North 


Art. coral. 


NONE 


Phyllospadix spp. 


244 


238 


LOW 


North 


Other 


NONE 


Ralfsia spp. 


205 


241 


HIGH 


Cosmop. 


Crust. 


DEC. 


Gigartina leptorhynchos 


199 


173 


MID 


Endemic 


Other 


NOV. 


Rhodoglossum a/fine 


162 


45 


MID 


North 


Other 


JUNE 


Pelvetia fastigiata 


159 


91 


MID 


Endemic 


Massive 


JUNE 


Gigartina armatalspinosa 


154 


104 


LOW 


Endemic 


Massive 


FEB. 


Ulva spp. 


152 


257 


HIGH 


Cosmop. 


Turf 


MAY 


Pterocladia pyramidale 


137 


94 


LOW 


South 


Other 


MAR. 


Pachydictyon coriaceum 


114 


100 


LOW 


South 


Leafy 


JUNE 


Corallina officinalis var. 














ch ilensislpin natifolia 


114 


144 


LOW 


Cosmop. 


Art. Coral. 


APR. 


Gastroclonium coulteri 


111 


48 


LOW 


North 


Other 


NONE 


Lithothrix aspergillum 


97 


81 


LOW 


Endemic 


Art. Coral. 


MAR. 


Gelidium crinale/coulteri 


94 


150 


HIGH 


South 


Turf 


NONE 


Gelidium pulchrumjpurpurescens 


93 


44 


LOW 


Endemic 


Other 


JUNE 


Eisenia arborea 


91 


83 


LOW 


Endemic 


Massive 


MAY 


Prionitis lanceolata 


87 


114 


LOW 


Endemic 


Massive 


FEB. 


Enteromorpha spp. 


76 


83 


HIGH 


Cosmop. 


Turf 


MAY 


Ceramium spp. 


70 


23 


MID 


Cosmop. 


Turf 


MAY 


Gelidium robustum 


66 


109 


LOW 


Endemic 


Other 


FEB. 


Chaetomorpha aerea 


66 


10 


HIGH 


Cosmop. 


Sand 


NONE 


Laurencia pacifica 


66 


88 


LOW 


Endemic 


Other 


NONE 


GracilarialGracilariopsis spp. 


63 


13 


LOW 


South 


Sand 


JULY- 
OCT. 


Melobesia mediocris 


63 


102 


LOW 


North 


Crust. 


JUNE 


Plocamium coccineum var. 














pacificum 


60 


70 


LOW 


Endemic 


Other 


FEB. 


Porphyra spp. 


58 


30 


HIGH 


Cosmop. 


Leafy 


JULY- 
OCT. 


Cryptopleura spp. 


58 


18 


LOW 


North 


Leafy 


JAN. 


Halidrys dioica 


57 


47 


LOW 


Endemic 


Massive 


JUNE 


Petrospongion rugosum 


54 


20 


HIGH 


South 


Other 


MAY 


Cystoseira spp. 


52 


47 


LOW 


Endemic 


Massive 


MAY 


Rhodymenia pacifica/lobatal 














palmettiformis 


49 


69 


LOW 


North 


Leafy 


FEB. 


Chondria nidifica 


48 


3 


LOW 


South 


Sand 


DEC. 


Centroceras clavulatum 


45 


25 


MID 


South 


Turf 


MAY 


LithophyllumlLithothamnion spp. 


44 


58 


HIGH 


Cosmop. 


Crust. 


JAN. 


Zonaria farlowii 


42 


56 


POOL 


South 


Other 


MAY 


Colpomenia sp. 


37 


73 


MID 


South 


Other 


JUNE 


Gymnogongrus leptophyllus 


36 


16 


MID 


Endemic 


Other 


JUNE 


Pterosiphonia spp. 


36 


46 


MID 


Cosmop. 


Turf 


FEB. 


EctocarpuslGiffordia spp. 


34 


48 


POOL 


Cosmop. 


Turf 


APR. 


Acrosorium uncinatum 


33 


9 


LOW 


South 


Leafy 


MAR. 


Hildenbrantia sp. 


33 


22 


HIGH 


Cosmop. 


Crust. 


MAR. 


Smithora naiadum 


30 


72 


LOW 


North 


Leaf 


MAY 


Gigartina papillatalcristata 


25 


37 


HIGH 


North 


Other 


JULY- 
OCT. 



1971 



SOUTHERN CALIFORNIA ALGAE 

Table I (continued). Scores and classification given each species 
(or other Caxonomic category). 



SPECIES OR ENTITY 



SCORE 
1956-9 



SCORE 
1968-70 LEVEL 



CLASSIFICATION ACCORDING 10: 
MS1RIH. IIAF1II 



MAX. 
MONTH 



Prionilis corneal aust rails 
Dictyopterls zonarloldes 
Macrocystis sp. 
Cladophora trichotoma 
Spermothamnion snyderae 
Dictyota ftabellata 
Laurencia splendensldiegoensis 
Ghondria californica 
Rhodymenia rhizoldes 
Sargassum agardhlanum 
Gymnogongrus platyphyllus 
Lophosiphonia spp. 
Chondria decipienslpacifica 
Codium fragile 
Blue-green algae 
Taonia lennebackerae 
Scytosiphon lomentaria 
Nienburgia andersoniana 
Rhodymenia californica 
Hapterophycus canallculatus 
Hesperophycus harveanus 
Endarachne binghamiae 

Calliarthron spp. 
Anisocladella pacifica 
Botryglossum farlowianum 
Endocladia muricata 
Microcladia spp. 
Cruoriella sp. 
Nemalion helmlntholdes 
Bryopsis sp. 
Leathesla difformis 
Cladophoropsis fasciculata 
Jania natalensis 
Agardhiclla coulterl 
Plocamium vlolaceum 
Schlzymenla sp. 
Grateloupla spp. 
Herposiphonla spp. 
Laminaria sp. 
Cladophora graminea 
Callophyllis spp. 
Carpopeltis bushiae 
Filamentous diatoms 
Laurencia subopposita 
Polysiphonia spp. 
Ophidocladus californicus 
Pterochondria sp. 
Bangia sp. 
Ulothrix sp. 



27 


12 


LOW 


Endemic 


Other 


— 


24 


14 


LOW 


South 


Other 


APR. 


24 


3 


LOW 


North 


Massive 


MAR. 


24 


12 


HIGH 


North 


Other 


— 


24 


21 


LOW 


Endemic 


Turf 


FEB. 


22 


38 


POOL 


South 


Leafy 


JUNE 


19 


4 


LOW 


Endemic 


Other 


JAN. 


15 





LOW 


South 


Other 


DEC. 


16 





LOW 


Endemic 


Leafy 


— 


12 


19 


LOW 


South 


Massive 


APR. 


12 





LOW 


North 


Other 


— 


12 


27 


MID 


Cosmop. 


Turf 


NOV. 


12 





LOW 


Endemic 


Sand 


MAY 


1 1 


1 1 


LOW 


North 


Other 


DEC. 


10 


6 


— 


Cosmop. 


Turf 


— 


9 


22 


POOL 


South 


Leafy 


JUNE 


9 


10 


HIGH 


Cosmop. 


Other 


MAY 


9 


3 


LOW 


Endemic 


Leafy 


JAN. 


9 





LOW 


South 


Leafy 


— 


9 





— 


South 


Crust 


— 


6 





HIGH 


Endemic 


Massive 


— 


6 


23 


MID 


South 


Leafy 


JULY- 
OCT. 


6 


12 


LOW 


North 


Art. Coral. 


— 


6 





LOW 


Endemic 


Leafy 


NONE 


6 





LOW 


North 


Leafy 


— 


5 





HIGH 


North 


Other 


— 


4 


12 


LOW 


North 


Other 


— 


4 


6 


HIGH 


South 


Crust. 


— 


4 


5 


HIGH 


North 


Other 


— 


3 


1 


POOL 


Cosmop. 


Other 


— 


3 





MID 


North 


Other 


— 


3 





— 


South 


Turf 


— 


3 





POOL 


South 


Art. Coral. 


NONE 


3 





— 


Cosmop. 


Other 


MAY 


3 





— 


North 


Other 


— 


3 





LOW 


North 


Leafy 


— 


3 


11 


LOW 


Cosmop. 


Leafy 


NOV. 


3 


43 


MID 


Cosmop. 


Turf 


— 


1 





LOW 


North 


Massive 


— 


1 





— 


South 


Other 


— 


1 





LOW 


North 


Leafy 


— 


1 





— 


Endemic 


Other 


— 





25 


— 


Cosmop. 


Turf 


— 





12 


LOW 


South 


Other 


MAR. 





12 


MID 


Cosmop. 


Turf 


MAY 





12 


— 


Endemic 


Turf 


— 





3 


— 


North 


Turf 


— 





9 


HIGH 


Cosmop. 


Other 


APR. 





9 


— 


Cosmop. 


Other 


— 



10 



111 I II I IS SOI I III l<\ (II II OK \ I I \CADEMYOl SOU \( I S 



VOLUME 70 



species were then grouped according to the month 
in which the running means showed a maximum. 
The species in which the maximum running mean 
was not more than double the value of the minimum 
running mean were considered to have no maxi- 
mum. Then, the original weighted data, not the 
running means, of each species in each group were 
added together, and the result is shown in Figure 6. 

Ranking of stations. — The changes noted in 
the algal flora of the various stations between 1 956-9 
and 1968-70 are shown in Table II. These changes 
were ranked in order from greatest decrease to 
greatest increase. The ranks based on species 
number and average score showed the high rank 
correlation coefficient (Schefler, 1969: 133) of 
+ 0.7, so a combined ranking was determined using 
the changes in both the species number and the 
average score, and is shown in Table II. 

The stations were also ranked according to the 
degree they are exposed to human use and collecting 
activity, to air pollution, and to marine pollution 



from sewage outfalls. These are the three factors 
quoted by Dawson (1965a: 174. 181) as prob- 
able causes of deterioration in the algal flora. The 
ranking according to human use was made after a 
consideration of the length of walk from the nearest 
roadhead by public access to the station, the amount 
of parking available in the vicinity of the roadhead, 
and the degree to which the area is built up. The 
factors considered in the ranking according to 
exposure to air pollution are shown in Figure 7. 
The number of days per year when the measured 
concentration of oxidant exceeded the state 
ambient air quality standard was used as a measure 
of air pollution. All the monitoring stations, how- 
ever, are far from the beach, so the ranking was 
derived from a consideration of the prevailing 
winds in the area, the distribution of high ground, 
and the pattern shown by the incidence of higher 
than standard levels of oxidant at the inland 
monitor stations. 

The information used for ranking the intertidal 



300- 



8 200- 



< 



> 
< 



100- 







/,- 





34 10 1 



l l 

19 18 2 

MALIBU 



25 17 



42 
ii 



22 

-l— 



9 21 
ORANGE CO. 



S. MONICA 7 33 16 LONG BEACH 

PALOS VERDES 

STATIONS 



Figure 5. The average scores reported from the stations in the Los Angeles area compared for 
1956-9 (open bars) and 1968-70 (solid bars). 



1971 



SOUTHERN CALIFORNIA ALGAE 



II 



1000 



3000 




Figure 6. Seasonal variations in the intertidal flora of southern California. The top line repre- 
sents the total weighted score for each month, summed for all' species and all stations visited. 
The intermediate bands represent the contributions to this total made by all the species found to 
have a maximum in the month indicated (according to smoothed data, see p. 10). 



stations according to the effects of marine pollution 
from sewage outfalls is shown in Table III and 
Figure 8. It may be seen from Table III that, in the 
case of the three largest outfalls, the increase in 
discharge from 1956-9 to the present is roughly 
proportional to the present figures, so that a ranking 
based on either criterion is the same. Station 21 
(Laguna Beach) is ranked 15th because the outfall 
at this station was closed in 1959 and the new 
Laguna Beach outfall opened more than a mile to 
the southwest. Coliform bacteria counts taken 
recently near Station 2 1 are therefore much less 
than those found during 1956-9 (L. Burtman. per- 
sonal communication). 

The distribution of coliform bacteria around the 
three largest outfalls is shown by stippling in Figure 
8. For the area around the Hyperion outfall (Calif. 
Reg. Water Pollution Cont. Bd. No. 4, 1965, Tables 
D and H) the data are tabulated as the number of 
analyses which showed surface water to have 10 
or more MPN/ml of coliform bacteria. The dense 
stippling represents the area monitored by those of 
the 16 offshore stations which had the maximum 
of 2 out of 104 analyses showing 10 or more MPN/ 
t ml coliform bacteria. Among the 17 shore stations, 
dense stippling represents 9 to 12 cases out of 617 
analyses at each station. Off White Point (Ritten- 
berg, Mittwer. and Ivler. 1958. Fig. 3) the dense 
stippling represents counts of 1000 to > 10000 
MPN/cm 2 coliform bacteria in bottom sediments 



obtained at certain of 106 sampling stations. Off the 
Orange County outfall (Rittenberg el al., 1958, Fig. 
I) dense stippling represents counts of 3000 to 
> 12000 MPN/cm 2 coliform bacteria in bottom 
sediments obtained at certain of 100 sampling 
stations. In all these cases, the boundary of the area 
sampled is shown by a broken line, and the sam- 
pling stations are fairly evenly distributed within 
that area. 

Thus, the available measurements from the 
Hyperion outfall are not strictly comparable to 



Table III. Average dry weather discharge 

of sewage outfalls in area studied 

(millions of gallons per day). 



OUTFALL 


1957-9a 


1969 


INCREMENT 


Dana Point 


0.1 


1.0b 


+ 0.9 


South Laguna 


0.6 


0.5" 


-0.1 


Laguna Beach 


1.0 


2.0" 


+ 1 


Orange County 


55.0 


119c 


4-64 


White's Point (3) 


277.0 


360d 


4-83 


Hyperion (effluent) 


275.0 


340<J 


4-65 


Hyperion (sludge) 


3.6 


4.0d 





aAllan Hancock Foundation. 1965. Table 1.1. 
bBurtman. pers. comm. 

cBueermann. pers. comm. (data for period 1967-9). 
d Hertel. pers. comm. 



12 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Table II. Description of intertidal sampling stations. 



VOLUME 70 



STAT. 


GEOGRAPHICAL NAME 


CHANGE 


% CHANGE 


RANK ACCORDING TO MAXIMUM: 


NO. 


VISITS 


NO. 




IN NO. 


IN AV. 


















OF SPP. 


SCORE 


DECREASE 


USE 


SEWAGE 


OXIDANT 


1956-9 


1968-70 


19 


Latigo Shore Rd. 


-4 


-4% 


7 


12 


13 


8 


3 


2 


34 


Malibu Shore Rd. 


+ 7 


+ 18% 


14 


8 


12 


7 


2 


2 


18 


Malibu Beach 


+ 2 


-12% 


9 


1 1 


1 1 


4 


3 


2 


10 


Big Rock Beach 


+ 3 


-15% 


10 


9 


10 


3 


2 


2 


2 


Topanga Canyon 


-4 


-44% 


5 


10 


9 


2 


3 


2 


1 


Sunset Blvd. 


-7 


-48% 


2 


1 


8 


1 


4 


2 


7 


Flat Rock Pt. 


- 15 


-40% 


1 


6 


5 


5 


1 


2 


25 


Lunada Bay 


-5 


-32% 


6 


7 


4 


6 


3 


3 


33 


Portuguese Pt. 


+ 8 


+ 17% 


15 


15 


2 


1 1 


1 


2 


17 


White Pt. 





+ 16% 


11 


3 


1 


10 


3 


2 


16 


Pt. Fermin 




















(Cabrillo Beach) 


- 14 


-23% 


4 


4 


3 


9 


2 


2 


9 


(Little) Corona 




















del Mar 


-12 


-43% 


3 


5 


6 


12 


2 


2 


42 


Crystal Cove 


+ 8 


-3% 


13 


13 


7 


13 


1 


2 


21 


Laguna Beach 


+ 6 


- 16% 


8 


2 


15 


14 


2 


2 


22 


Dana Point North 




















(Salt Creek) 


-2 


+ 31% 


12 


14 


14 


15 


2 


3 



those published about the White Point and Orange 
County outfalls, while the latter two are seriously 
out of date. Moreover, differences in sewage treat- 
ment (continuous chlorination is performed at 
Hyperion but not at White Point) affects the inci- 
dence of coliform bacteria but may not affect other 
deleterious components of sewage. Indeed, it is 
possible that chlorine itself is harmful to algae. 
However, the data presented seem sufficient to 
establish a ranking system for the stations occupied. 
The ranks assigned to each intertidal station 
according to each factor is shown in Table II. 

RESULTS AND DISCUSSION 

The reduction in the marine intertidal flora near 
Los Angeles since 1 956-9 is general and widespread. 
Figures 2 and 5 show a very similar situation from 
station to station even though they illustrate two 
different aspects; the diversity and the quantity of 
the flora. Almost all of the more frequent species 
shown in Figure 3 and the more general groupings 
shown in Figure 4 show a downward trend. 

In Figures 3 and 4, data from the best stations 
outside the Los Angeles area are used as an indi- 
cation of what the southern Californian flora may 
have been before the effects of human activity 
became apparent. While there is an obvious down- 
ward trend from the best stations through the Los 
Angeles stations in 1956-9 to the latter in 1968-70, 
in many individual species and groups of species 



the pattern of change is not consistent. Although 
the 'best' stations (different localities from the 
Los Angeles stations) might make the data for rare 
species or small groups incomparable, the total 
scores for the common species and the larger groups 
should be more consistent than shown. Thus, it is 
probable that the factor responsible for the decline 
before 1956-9 is different from that responsible for 
the decline after this period. There seems no reason 
to doubt Dawson's conclusion that the major factor 
in the deterioration prior to 1956 was marine 
pollution from sewage outfalls. 

Those entities which show a consistent decrease 
include Gigartina leptorhynchos, Gigartina armatal 
spinosa, Gastroc Ionium, and GracilarialGracilari- 
opsis. Similar species groups include the four 
largest groups; the low intertidal, the northern, 
those of miscellaneous habit, and those without a 
maximum month which tends to confirm the sug- 
gestion that the decrease is very general. The group 
showing the most dramatic decline is that which 
includes those entities which characteristically 
are found growing up through sand. These species 
seem to be relatively well protected from collectors. 
Perhaps the sand provides a mechanism by which 
pollutants are held in more effective contact with 
the plant than is the case with other habitats. On 
the other hand, these plants could be declining 
simply because their habitat is being reduced by 
the erosion of sandy beaches. The southern entities 
show much the same consistent decline as the 



1971 



SOU I III -RN ( Al.ll OKNIA ALGAE 



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*i ■ . • . * 












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\iv,: , l : ■•.%.• .•.;■;•/<. 




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*"•*' * :i • 








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95 


****».* 




"V .** *. ** *• . .' 




"'**■.""« • "•***.■*.**** ' • " 




•'•■:->.n.^~r-.- Jv:-:".\:«. ■•.-.: 


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100 
■•••.■'■—_ >'-••(. 




'* 123 


*•" '•. 


103 "\ : ':'.'.:'•'/;''•' 


.'« '.• ■ •'. 




0: 














•*"•" 


51 


67 






«*^« 87 




V< • 34 IB 10 2 










h 






19 










44 ^,;- i=?/.7-v-... 
K .;v.- ••.?•. •'••' 




N 




\ 9 










^> iiiE-»^/^ 












'**.... 'i 




i A 












■"VI 

52 <$MS 




yVC~ 


r L. A. Airport 




7 #/\.. 




7 




"**"■■*. • 

11 x N-?;:v 




N 




33 fNA 

' 7 .6 












v ./ 














/[ N.' 


y^-r^ 
















XII 












9 ^^. 




/ 









10 








San Pedro 




| 




1 




42 ^^ 

21 \. 






MILES 
















22 ^""N 





Figure 7. The information used to determine the ranking of intertidal stations (solid circles, small 
numerals) according to the amount of air pollution. (Stippled area: land over 1000 feet elevation; 
large numerals: number of days oxidant exceeded California State standards in the year 1968-9 
[Calif. Air Res. Bd. 1969a, b, c, d.]; wind roses: the prevailing winds at Los Angeles Airport 
[U. S. Weather Bur.. 1962. p. 15] and San Pedro [Stevenson, 1959, Fig. 9B]). 



northern; they might perhaps be expected to in- 
crease if thermal pollution were an important 
factor. The decline of leafy forms was mentioned 
by Dawson. In the groupings by maximum month. 
those with maxima in the summer, fall, and early 
winter (July to January) show a consistent decline. 

Those entities which show an inconsistent pattern 
of decline before 1956 and an increase after include 
Ulva, Gelidium crinalefcoulteri, and Pterosiplionia 
spp. The main feature these plants have in common 
is a turfy habit, which is particularly protected 
from collectors (as is also the crustose habit). The 
presence of Ulva and Gelidium crinalefcoulteri 
in the high intertidal category, and Ulva in the 
cosmopolitan and May maximum groups is mainly 
responsible for the similar appearance of the trend 
in these groups. 

Those species which show an inconsistent pattern 
of increase before 1956-9 and a decrease after 
include Corallina vancouveriensisl gracilis. Bos- 
siella spp.. Rhodoglossum affine, and Pelvetia. The 



first two are articulated corallines which Dawson 
mentioned particularly as being favored by sewage 
pollution. Rhodoglossum affine and Pelvetia, as 
well as Corallina vancouveriensis, contribute to the 
similar pattern shown by mid-intertidal forms, 
which are especially vulnerable to both collection 
and air pollution. Corallina vancouveriensislgra- 
cilis has a similar effect on the March group, while 
both Pelvetia and Rhodoglossum affine have a 
maximum in June. 

Finally, a small consistent increase is shown by 
Ralfsia, Corallina officinalis var. chilensis. and 
species with a maximum month in February. In 
the latter case, no one species appears to be respon- 
sible for the effect. 

No quantitative measurements are available for 
most of the stations with respect to human use. air 
pollution, or water pollution. In addition, these 
factors are complex in themselves, and may operate 
on any particular species indirectly, through their 
effect on another species which has an effect on the 



14 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OE SCIENCES 



VOLUME 70 




LAGUNA 

2 \"S. LAGUNA 

• 5 , 

DANA PT. 



Figure 8. The information used to determine the ranking of intertidal stations (solid circles, small 
numerals) according to pollution from sewage outfalls (named). (Large numerals: MGD. average 
dry weather discharge [see Table III]; stippled areas: distribution of coliform bacteria [see text, 
p. 11]; arrows: general off-shore ocean currents [Emery, I960. Fig. 87]). 



species in question, rather than directly. At the 
present stage of knowledge, non-parametric statis- 
tics are the best that can be used. Rank correlation 
coefficients were calculated between the rank for 
the change in the flora at each station between 
1956-9 and 1968-70, and the rank assigned each 
station with respect to human use, air pollution, 
and sewage pollution. These coefficients are +0.6, 
+ 0.4, and +0.2 respectively. 

Thus, of the main factors suggested for the 
decline of the algal flora, the most important since 
1956-9 are: first, human use; second, air pollution; 
and third, pollution from sewage outfalls. This is in 
contrast with the decline up to 1956 where, as has 
been said above, there is no reason to doubt Daw- 
son's conclusion that sewage pollution was the 
major factor. 

Even without a statistical analysis, it is obvious 
that there are some stations which are particularly 
high in one factor and low in others. Station 9 
(Little Corona) seems to have little air and water 
pollution, but it is a favorite spot for collectors. It 
has shown a drastic decrease in algae. It remains to 
be seen whether it will recover now that it has been 
made a marine preserve. Station 42 (Crystal Cove) 
is difficult of access but is close to Station 9 and 



presumably shares with it much the same level of 
air and water pollution; the flora there has improved. 
Station 33 (Portuguese Point) is totally shut off 
from the public, while it suffers a high degree of ! 
sewage pollution; the flora there has improved, 
although it is still at a low level. Station 2 (Topanga 
Canyon Rd.) suffers heavy air pollution, is difficult 
of access, and has at most moderate sewage pollu- 
tion; the flora there has obviously decreased. 

The data shown in Figure 6 indicate that there is 
indeed much seasonal variation in the algal flora 
of southern California, with a maximum in March- 
June and a minimum in August-September. Most 
of this variation is the effect of variation in species 
which are present throughout the year. Year to 
year variations probably make the grouping of 
observations by season unreliable but, if this is to 
be done, it would be better to divide the year into 
three four-month seasons: November- February; 
March-June; and July-October. There seems to be 
no justification for Dawson's statement first (1965a: 
224) that "In gross aspect the southern California 
benthic algal flora does not show conspicuous 
seasonal differences in quantity or quality," and 
second (1965a: 225) that "At most stations the 
flora as a whole shows a decline in growth and 



1971 



SOU I III, KN (ALII ORNIA Aid A I. 



/.5 



luxuriance during the winter and spring and an 
increase during ihe summer and fall." 

CONCLUSIONS 

The main purpose of an exploratory study such as 
the present one is to suggest the direction of future 
work. It is obviously desirable that the present 
non-parametric methods should be replaced by 
more precise quantitative ones. Practical considera- 
tions demand that a more precise study should be a 
more restricted one. 

Future work could be concentrated on certain 
species over wider areas than the stations sampled, 
or on the entire flora of certain areas more detailed 
than the present stations. Small areas studied in 
detail are extremely vulnerable to the effects of 
sand movement and man-made changes of sub- 
stratum (such as the building of a jetty across the 
transect line of Station I ). These factors are part of 
the scene, and no attempt was made to exclude them 
in the present general study, but they are difficult 
to evaluate in terms of 'pollution'. Studies of certain 
particular species over a wide area would be less 
vulnerable to such factors and also make them 
easier to evaluate. In conjunction with experimental 
work, the distribution of these species might be 
related to certain specific pollutants, and so make 
them valuable as indicator organisms. 

Grouping species into taxonomically hetero- 
geneous categories (Fig. 4) did not produce much 
information, except that plants coming up through 
sand appear to be particularly vulnerable. So, the 
entities chosen for detailed study should be the 
smallest taxonomic unit readily identifiable in the 
field. The use of taxonomic units which require 
identification in the laboratory would require 
extensive collections which would be undesirable 
in principle and would add greatly to the work 
needed for the study. (An exception would be 
needed in the case of many turf forming genera). 

The choice of species to be studied must also 
take into account seasonal variations (Fig. 6). If 
the study could not be carried on throughout the 
year, the species could be chosen which have only 
a slight seasonal variation (Table I). However, 
since most probable pollutants also have seasonal 
fluctuations in their occurrence, it would be much 
more desirable to monitor both species and pollu- 
tant throughout the year. Frequent observations 
would also be needed to suggest when dying and 
I damaged plants could be showing the effects of 
I pollution and when they are simply the effects of 
: drying winds. 

; It would also be advisable to use species which 
i appear to have decreased in response to pollution. 



rather than those which have increased, since there 
are so lew of the latter. 

Gastroclotiium loulteri and (,'haeiomorpha 
aerea are two species which are easily identified, 
have little seasonal variation, and have shown a 
consistent decline both before 1956 and alter. Their 
decline could perhaps be used as a general measure 
of 'pollution'. 

Rhodoglossum affine, Bossiella spp., Pelvetia 
fastigiata, Gigartina armata/spinosa, and (ielidium 
pulchrumlpurpurescens all have shown a relatively 
great decrease since 1956-9 and presumably are 
sensitive to the factors which are causing a decline 
in the intertidal flora at the present time. However, 
all except Bossiella show a marked seasonal varia- 
tion, and all except Rhodoglossum and Pelvetia 
contain more than one recognized taxonomic 
species. 

ACKNOWLEDGMENTS 

I wish to thank the 16 students in my algae course who 
made half the station visits used in this study: Ray 
Allen, Chris Beattie. Monica Beck, David Gray. Terry 
W. Howey, R. Hulbrock. Wayne D. Johnson. John 
Martz. Timothy Meenan, Louis Mikelson. Robert 
Mosley, Wayne Nelson. Tony Sciarrotta, Jim Slawson. 
Michael Sowby, and Gerald T. Wall. 

I am also grateful to Dr. L. Leamy, Biology Depart- 
ment. California State College at Long Beach, who 
discussed with me the statistical problems encountered 
in this study: to the staff of the State Water Quality 
Control Board who gave me access to unpublished 
data; and to the Department of Fish and Game, who 
gave us permission to collect in protected areas. 

The mechanical data processing used in this analysis 
is part of a system developed with the help of intra- 
mural funds from the University of California. Los 
Angeles; the University of California. Santa Barbara 
(in co-operation with Dr. M. Neushul): and from the 
Long Beach California State College Foundation. 



Literature Cited 

Alder, H. L.. and E. B. Roessler. 1964. Introduction to 
Probability and Statistics. 3rd ed. W. H. Freeman 
& Co.. San Francisco, xiv 3 13 pp. 

Allan Hancock Foundation. 1965. Chapter 1. Introduc- 
tion In An Oceanographic and Biological Survey 
of the Southern California Mainland Shelf pp. 1-28. 
State Water Quality Control Board, Pub. No. 27. 
Sacramento. California. 

Bellamy. D. J.. R. Bellamy. D. M. John, and A. Whit- 
tick. 1967. Some effects of pollution on rooted 
marine macrophytes on the northeast coast of 
England. British Phycological Bull. 3: 409. (Abstr.i 



16 



BULLETIN SOUTHERN C A LI LORN I A ACADEMY OF SCIENCES 



VOLUME 70 



Bellamy, D. J., P. H. Clarke, D. M. John, D. Jones, A. 
Wittick. and T. Darke. 1967. Effect of pollution 
from Torrey Canyon on littoral and sublittoral 
ecosystems. Nature 216: 1 170-1 173. 

Boney, A. D. 1968. Experiments with some detergents 
and certain intertidal algae. In J. D. Carthy and 
Don R. Arthur. Ed.. The Biological Effects of Oil 
Pollution on Littoral Communities. Field Slud. 
Council 2 (Suppl.) pp. 55-72. 

California Air Resources Board. 1969a. California Air 
Quality Data for September, October, November 
1968. Vol. 1 No. 1. Multilith 28 pp. 



. 1969b. California Air Quality Data for Decem- 
ber 1968, January and February, 1969. Vol. 1. 

No. 2. Multilith 27 pp. 



. 1969c. California Air Quality Data for March, 

April, and May 1969. Vol. 1. No. 3. Multilith 

28 pp. 

. 1969d. California Air Quality Data for June, 

July and August, 1969. Vol. 1. No. 4. Multilith 

29 pp. 

Cowell, E. B. 1969. The effects of oil pollution on salt- 
marsh communities in Pembrokeshire and Corn- 
wall. /. Appl. Ecology 6: 133-142. 

Dawson, E. Yale. 1959a. A primary report on the 
benthic marine flora of southern California //; 
Oceanographic Survey of the Continental Shelf 
Area of Southern California pp. 169-218. State 
Water Pollution Control Board, Pub. No. 20. 
Sacramento. California. 



. 1959b. Benthic marine vegetation. 

219-264. 



Ibid. pp. 



. 1 96 1 . A guide to the 1 iterature and distributions 

of Pacific benthic algae from Alaska to the Gala- 
pagos Islands. Pacific Sci. 15: 370-461. 



. 1965a. [not signed) Chapter 8 Intertidal algae 

In An Oceanographic and Biological Survey of 
the Southern California Mainland Shelf pp. 220- 
231. State Water Quality Control Board, Pub. 
No. 27. Sacramento, California. 



— . 1965b. 
351-438. 



Table IX. Ibid. Appendix Data pp. 



Emery. K. O. I960. The Sea off Southern California. 
Wiley & Sons, New York. 366 pp. 

Okamura, K., S. Oeda. and Y. Miyake. 1926. On the 
harmful action of deep fog on Porphyra tenera 
Kjellm. J. Imp. Fish. Inst. 21(6): 67-68. [In Japa- 
nese. Authors' English abstract Bios. Abs. 1:43. 
1927] 

Regional Water Pollution Control Board No. 4. Los 
Angeles Region. State of California Resources 
Agency. 1965. Report on Monitoring Programs in 
Santa Monica Bay January 1 through December 
31, 1964. Mimeograph 27 pp., 2 Appendices. 

Rittenberg, S. C, T. Mittwer and D. Ivler. 1958. Coli- 
form bacteria around three marine sewage outfalls. 
Limnol. Oceanogr. 3: 101-108. 

Schefler, W. W. 1969. Statistics for the Biological 
Sciences. Addison-Wessley, Reading, Massachu- 
setts. 23 1 pp. 

Stevenson, R. E. 1959. The marine climate of southern 
California In Oceanographic Survey of the Con- 
tinental Shelf Area of Southern California pp. 7-58. 
State Water Quality Control Board, Pub. No. 20. 
Sacramento. California. 

U. S. Weather Bureau. 1962. Decennial Census of* 
United States Climate. Summary of Hourly 
Observations. Climatography of the United States 
82-4 Los Angeles, California, International Air- 
port 1951-60. 10 pp. 

Accepted for publication September 26, 1970. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(1): 16-17, 1971 

A NEW SPECIES OF SIMOPELTA FROM COSTA RICA 

(Hymenoptera: Formicidae) 

Roy R. Snelling 1 

Abstract: Simopelta paeminosa, new species, is described from Pun- 
tarenas Province. Costa Rica. Diagnostic features of this species are: tridentate 
mandibles with acute basal tooth; no median clypeal spine; rugose cephalic 
and alitrunk integument. This species appears to be most closely allied to 
5. williamsi of Ecuador. 



The ponerine genus Simopelta was reviewed by 
Gotwald and Brown (1966), who discussed the 
generic characters, listed the known species and 



described two new species. They also published 



•Natural History Museum of Los Angeles County, Los Angeles 
California 90007. 



1971 



NEW SPECIES <>l AN IS 



17 



observations on the behavior of S. oculata Gotwald 
and Brown made by Brown in Costa Rica. These 
observations reaffirmed the army ant like behavior 
of the foraging workers. 

The following new speeies from Costa Rica was 
included in a vial containing three workers, two 
males and one mangled larva of Gnamptogenys 
simplex (Emery). The two males are not fully 
colored and one has the wings torn off on one side. 
It seems likely that the Gnamptogenys represent 
the prey of the Simopelta; unfortunately, the collec- 
tor after three years, was unable to recall details of 
the collection. 

Simopelta paeminosa, new species 
Figure 1 

Diagnosis. Clypeus without median spine; eyes 
small; mandibles tridentate. basal tooth acute; 
head, alitrunk and petiole coarsely rugose; integu- 
ment blackish brown. 

Holotype worker: TL 4.1; HL (occipital margin 
to anterior border of frontal lobes) 0.9; HW (with- 
out eyes) 0.7; WL 1.5; greatest diameter of eye 
0.06; scape L (chord, without basal neck) 0.8 mm; 
CI 81. Abbreviations and measurements as in 
Brown (1958). 

Similar to worker of 5. williamsi but differing as 
follows; 

1. Mandibles more slender, the basal tooth acute. 

2. Antennal scapes longer; in full face view, 
extending beyond occipital margin by about twice 
the apical breadth. 

3. Petiolar node longer, about as high as long. 

4. Rugulae of head and alitrunk not transversely 
oriented. In 5. williamsi, those of the occiput and 
pronotum, especially, are decidedly transverse; in 
5. paeminosa the rugulae are very irregular, mostly 
oblique, though a few may be partially or wholly 
transverse. The gaster is shiny, with well separated 
distinct punctures. 

Paratype series. Variation in ten randomly selec- 
ted individuals, but including apparently largest 
and smallest specimens: TL 3.8-4.2; HL 0.90-0.95; 
HW 0.70-0.73; ML 0.43-0.50; WL 1.40-1.50 mm; 
CI7I-81. 

Color, basically blackish brown, mandibles, fla- 
gellum, legs brownish. Pronotum with brownish 
areas of irregular extent. Gastric apex light brownish. 

Holotype and 2 1 paratype workers, 4 mi south 
of San Vito de Java. Puntarenas Province, Costa 






1 mm 

Figure I. Simopelta paeminosa, new species. Above. 
head in frontal view and enlarged outline of mandible. 
Below, lateral aspect. Figures by Ruth Ann DeNicola. 



Rica, 15 August 1967 (R. W. McDiarmid). Holo- 
type and most paratypes in the Natural History 
Museum of Los Angeles County. Paratypes 
deposited in American Museum of Natural History 
and Museum of Comparative Zoology. 

Etymology. Paeminosus, L., rough or uneven, in 
allusion to the roughened integument. 

The rugose integument of this species seems to 
indicate a relationship to 5. williamsi. That Ecua- 
dorian species, however, has the rugulae transverse 
on the occiput and dorsum of the alitrunk. In S. 
williamsi, too. the basal tooth of the mandible is 
truncate rather than acute. 

In the key to Simopelta workers by Godwald 
and Brown. 5. paeminosa fails at couplet 5 since it 
does not accord with either alternative. The other 
Central American species are punctate rather than 
rugulose. 

Literature Cited 

Gotwald. W. H.. Jr. and W. L. Brown. Jr. 1966. The 
ant genus Simopelta. Psyche, 73: 261-277. 

Accepted for publication September 25. 1970. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(1): 18-22, 1971 

A REVISION OF THE GENUS LOANDAL1A MONRO 
WITH DESCRIPTION OF A NEW GENUS AND SPECIES OF PILARGIID POLYCHAETE 1 

Raymond R. Emerson 2 and Kristian Fauchald 2 



Abstract: A re-examination of the genotype of Loandalia, L. aberrans 
Monro (1936) demonstrated several unique morphological features not pre- 
viously noted. A new genus, Parandalia, is established to contain the other 
species usually considered in Loandalia. Type species is Parandalia ocularis, 
new species; others include P. americana, P.fauveli, P. gracilis and P. indica. 



During the examination of some polychaetous 
annelids from the Santa Barbara Channel, Cali- 
fornia taken in connection with a study of oil spill, 
January 1969, a new species of a pilargiid poly- 
chaete was recognized. It became apparent that 
the genus Loandalia Monro (1936) as presently 
accepted (Pettibone, 1966) contained two different 
groups of species. The holotype of Loandalia aber- 
rans Monro (1936), which is the genotype, was 
borrowed from the British Museum (Natural His- 
tory) and re-examined. As a consequence of these 
studies it was found that Loandalia can be accepted 
only as a monotypic genus; all the species presently 
included in that genus are here removed to a new 
genus, Parandalia. 

The possibility that Talehsapia Fauvel (1932) 
could be a valid generic name for these species was 
considered and discarded. Talehsapia differs from 
all known pilargiids in the presence of jaws. Further, 
prostomial features do not resemble the pilargiids 
as presently accepted. Talehsapia is here considered 
incertae sedis and is not accepted as a valid genus in 
the family Pilargiidae. 

Ancistargis Jones (1961) was considered a syn- 
onym of Ancistrosyllis Mcintosh (1879) by Petti- 
bone (1966). A median antenna is present in 
Ancistrosyllis and absent in Ancistargis. The median 
antenna is often difficult to detect in species of 
Ancistrosyllis as remarked by Pettibone, but the 
character is here considered to be of sufficient 
importance to warrant maintaining the generic 
status of Ancistargis. 

The family includes nine genera; Ancistargis 
Jones (1961) from Gulf of Mexico, Ancistrosyllis 
Mcintosh (1879) from Greenland, Cabira Webster 
(1879), sensu Pettibone (1966). Loandalia Monro 
(1936) off Angola, Africa, Otopsis Ditlevsen (1917) 
from near Iceland, Parandalia, new genus. Pilargis 
Saint-Joseph (1899) from France, Sigambra Miiller 
(1858) from Brazil and Synelmis Chamberlin (1919) 
from the south Pacific Ocean. 

The holotype and paratype specimens of the 



new species have been deposited in the collection 
of the Allan Hancock Foundation. 

Key to Genera of Pilargiidae 

1 . Notopodia with stout emergent hooks 

or spines 2. 

Notopodia without emergent hooks 

or sp"ines 7. 

2. Notopodia with recurved 

emergent hooks 3. 

Notopodia with stout, straight spines 6. 

3. Peristomium dorsally entire Ancistargis 

Peristomium dorsally incised 4. 

4. Dorsal and ventral cirri reduced or 
absent; parapodia reduced, body 

subcylindrical Cabira 

Dorsal and ventral cirri well developed, 
parapodia well developed, body 
dorso-ventrally flattened 5. 

5. Antennae shorter than palps; 

integument papillated Ancistrosyllis 

Antennae longer than palps; 

integument smooth Sigambra 

6. Prostomial antennae and peristomial 
cirri present, parapodia sharply set off 

from the body Synelmis 

Prostomial antennae and peristomial 

cirri absent, parapodia distinct, but not 

set off from the body Parandalia 

7. Prostomial antennae and peristomial 

cirri absent Loandalia 

Prostomial antennae and peristomial 

cirri present 8. 

8. Prostomium with two antennae, 

biarticulate palps present Pilargis 

Prostomium with three antennae. 

palps without palpostyles Otopsis 



'Contribution from the Allan Hancock Foundation. 

-Allan Hancock Foundation. University of Southern California, 
Los Angeles. California 90007 



18 



1971 



NEW SPECIES OF POLYCHAl II 



19 



Parandalia, new genus 

Type species: Parandalia ocularis, new species. 

The anterior part of the body is somewhat 
inflated; median and posterior parts of the body are 
cylindrical. A pair of eyes may be present on the 
anteriormost segments. 

The small prostomium is indistinct: a pair of 
well developed, biarticnlate palps with button- 
shaped palpostylcs is present. The peristomium is 
separated from the prostomium by a shallow, indis- 
tinct segmental groove. Peristomial cirri are absent. 
The proboscis is cylindrical when everscd and may 
terminate with several sensory papillae. The proxi- 
mal surface ol the proboscis is smooth. 

The first parapodia are uniramous, further back 
all parapodia are biramous. A thick, stout, straight, 
jrystal-clear spine emerges from the median and 
posterior notopodia. All notopodia are poorly 
Jeveloped and have at most one or two capillary 
setae in addition to the spine. Neuropodia where 
fully developed, are slightly tapering with blunt, 
evenly rounded tips. Dorsal cirri are absent; ventral 
:irri are present as small, papillar lobes. Branchiae 
ire absent. 

The neuropodia have several geniculate, pointed, 
simple setae. Each seta has a cylindrical stalk and 
n blade provided with numerous transverse rows 
of slender teeth. 

The pygidium is an anal plaque which may have 
one to several short, blunt anal cirri. 

Parandalia as defined above includes all species 
previously assigned to Loandalia with the excep- 
tion of the type species of the latter genus. A 
re-examination of the type specimen of Loandalia 
■iberrans Monro (1936) showed that notopodial 
spines are absent and that branchiae are well 
jeveloped in the posterior part of the body. In 
addition, the two first neuropodia have thick, black 
acicula; a feature not found in any other described 
pilargiid. The species presently assigned to Paran- 
dalia include: P. americana (Hartman, 1947); 
P. gracilis (Hartman-Schroder. 1959): P. fauveli 
(Berkeley and Berkeley. 1941); P. indica (Thomas. 
1963); and the genotype, P. ocularis, new species. 

The different species in the genus can be differ- 
entiated as indicated in the key below and in the 
discussion of P. ocularis, new species. 

Key to Species of Parandalia 

1. All parapodia biramous 2. 

Anteriormost parapodia uniramous. 

all others biramous 3. 

2. A single notopodial seta present; 

four-five neuropodial setae present . . . P. gracilis 
Two notopodial setae present; four-twelve 
neuropodial setae present P. indica 



3. Eyes present; six-seven neuropodial 

setae in each parapodium /'. ,„ ularii 

Eyes absent; twelvc-liltccn neuropodial 

setae in each parapodium .4. 

4. Emergent notopodial spine from 

setiger 2 /• amer'u una 

Emergent notopodial spine from 

setiger 7 /'. fauveli 

Parandalia ocularis, new species 
Material examined: Alll VI I I KO I2K56. April 

3, 1969. 34°23'I2"N. 1 19°38'18"W, 24 fms silt) 
clay, gear: Campbell grab, paratype (I). AH I 
VELERO 12860. April 3. 1969, 34°22'58"N. 
1I9°38'05"W, 23 fms, silty-clay. gear: Campbell 
grab, HOLOTYPE. AHF VELERO 12864, April 

4, 1969,34°2I'I5"N. 1 I9°38'00"W. 25 fms. coarse 
sand, gear: Campbell grab, paratype ( 1 ). 
Description: Three complete individuals were 
collected. Length of the holotype specimen is 41 
mm. Width in the widest region at the third or 
fourth setiger is 0.5 mm without parapodia which 
are poorly developed in the anterior segments. 
Posterior to the inflated region at about setiger 20 
the width is 0.4 mm without and 0.6 mm with 
parapodia. The number of segments is 93. 

The body is a pearly white with some evidence 
of reddish brown colored patches on a few segments 
primarily near the parapodial lobes. Sections of an 
annulated skelopodium (chitinized secretion) cover 
some of the posterior segments producing a con- 
tinuous reddish brown appearance. The body is 
cylindrical, the anterior region through the first 
6 setigers is areolated and slightly dorsolateral!)' 
flattened. A pair of conspicuous, subcutaneous eyes 
is present on the dorsal surface between the second 
and third setiger. Each eye is subcircular. indicating 
the possible presence of a lens (Fig. 1 ). 

The small prostomium is inconspicuous and has 
a pair of Particulate palps. Each palp has a pal- 
pophore tipped with a single minute knoblike 
palpostyle. The palps are directed dorsally. an effect 
which may be somewhat accentuated by the par- 
tially everted proboscis. 

The peristomial segment is slightly emarginated 
laterally but less distinct on the dorsal and ventral 
surfaces. It is similar in length to the first setiger: 
tentacular cirri are absent (Fig. 1). 

A relatively large, cylindrical, muscular proboscis 
is present and terminates anteriorly with four small 
papillae (Fig. 4). The opening of the proboscis is 
horizontal. 

The first 6 setigerous segments are inflated and 
the surface epithelium may be weakly areolated 
delimiting a distinct anterior region. The parapodia 
in this region are reduced but increase in size up 



20 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OE SCIENCES 



VOLUME 70 






Figures 1-3. Parandalia ocularis, new species. Figure 
I. Anterior end in dorsal view x50. Figure 2. Para- 
podium 14 from left side in anterior view x 100. Figure 
3. Posterior end in ventral view x50. 



to the sixth, which is similar in size to the succeeding 
parapodia. The first podium of the first setigerous 
segment lie in notopodial positions but a gradual 
shift to the neuropodial positions is completed by 
setiger 3 or 4. 

The first 2 setigerous segments have uniramous 
parapodia. all subsequent parapodia are unequally 
biramous. The first 3 notopodia on setigers 3-5 are 
reduced relative to those in subsequent segments, 
each notopodium bears 1 or 2 small, simple seta. 
The notopodia have become fully developed by 
setiger 9 or 10. A crystal-clear notopodial aciculum 
begins to protrude at setiger 9 becoming more 
fully exposed in succeeding segments. 

When fully developed, each neuropodium is 
slightly tapering with a blunt, evenly rounded tip 
in all segments. The anteriormost neuropodia are 
reduced and do not become fully developed until 
setiger 6 or 7 (Fig. 1). The neuro-aciculum is fully 
embedded in most segments but may protrude 
slightly from the neuropodial lobe. A small papillar 
ventral cirrus at the outer basal edge of the neuro- 
podium begins to appear about setiger 8. The 
cirrus is somewhat larger posterior to setiger 10 but 
retains the same shape in all setigers from the tenth 
setiger on. (Fig. 2). 



The first two neuropodial lobes each bear 4 
geniculate, pointed, simple setae. The subsequent 
neuropodial setae are larger and may increase in 
number up to about 7. All ncurosetae are born in 
a similar pattern (Fig. 2). Each neurosetae has a 
cylindrical stalk with an upper blade provided with 
many transverse rows of slender teeth (Fig. 5). The 
notopodial setae are short and nearly smooth. 

The posterior end of the body tapers evenly and 
terminates in a rounded anal plaque (Fig. 3). Its 
margin is entire except for 3 anal cirrilike processes. 
Two cirri processes are lateral and one process 
is located mid-ventrally. The anal aperture is near 
the mid-ventral process and may be distinguished 
by a slight depression. 

Discussion: Five species are known in the new 
genus Parandalia. Three specimens of P. ocularis, 
new species, were collected from the Santa Barbara 
Channel off southern California in silty-clay at 
depths of 23-25 fms. P. fauveli, Berkeley (1941) 
is known from the single type locality record in a 
mud fia.t near Newport Bay, California and gener- 
ally from similar localities in southern California 
(Hartman. 1968). P. americana Hartman (1947) 
has been collected from sand flats at low tide near 
Biloxi, Mississippi and from the Gulf of Mexico 




Figures 4-7. Anterior end in dorsal view x50 (4, 6) and 
distal end of neuroseta x950 (5. 7). Figure 4 and 5. 
Parandalia ocularis, new species, proboscis everted (4). 
Figure 6 and 7. Loandalia aberrans, Monro. 






NEW SPECIES OF POLYCHAETE 



21 




M 8 

Figures 8, 9. Loam/alia aberrans, Monro. Figure 8. 
Distal end of notoseta x950. Figure 9. Posterior end 
in dorsal view x50. 



near San Jose Light, Guatemala in 12-13 fms. 
/'. indica Thomas (1963) is known from the West 
Coast of India from muddy substratum in 7-10 fms. 
P. gracilis Hartman-Schroder (1959) has been 
collected off El Salvador. 

P. ocularis is the only species having eyes; it is 
similar to P. americana and P.fauveli in that each 
has uniramous parapodia in the first segment and 
biramous parapodia continuous from the segment 
2 or 3. P. americana and P.fauveli have emergent 
notopodial spines from the second and seventh 
setiger. respectively. These two species along with 
P. gracilis have been considered by Pettibone ( 1966) 
as synonymous with P. fauveli, although only the 
holotype of P. fauveli and two paratypes of P. 
americana were examined. P. indica is considered 
by the same author to be a doubtful species. 

P. indica and P. gracilis are closely related species 
having biramous parapodia from the first segment. 
P. gracilis has 1 notopodial seta and P. indica has 
2 notopodial setae on each segment. 

Genus Loandalia Monro, 1936 
Loandalia aberrans Monro, 1936 

Loandalia aberrans Monro, 1936. 
Material examined: DISCOVERY 274. off St. Paul 
de Loanda, Angola. From 8°40' 15" S, 13°13'45" E 
to 8°38'15" S, 13°i3'00" E, 64-65 m, grey mud. 
trawl, one specimen, holotype British Museum 
Natural History type number 1936-2-8-3376. 
Remarks: A re-examination of the type species of 
the genus Loandalia showed that there are some 
details previously not reported by Monro (1936). 

The anterior part of the body including the first 6 
setigers is slightly inflated and the surface has a 
slight areolation; the median and posterior parts of 
the body are cylindrical. The posterior end was in 



regeneration when the specimen was collected, the 
specimen is dull yellow with dark brown ban on 
each side of the notopodia in median and posl 
setigers. 

The prostomium is short and wide; prostomial 
palps are large compared to the size <>l the pro 
tomium. A single pcristomial segment is present; 
peristomial appendages are absent dig. 7|. I he 
proboscis was dissected out by Monro and could 
not be described from the remnants present. 

The first 2 pairs of neuropodia are stout and 
conical; the first is dorsolateral in position; the 
second is in the normal position, somewhat ventral 
to the midline and directed ventrolaterally. All 
neuropodia from setiger 3 are similar; each has a 
long base with nearly parallel sides and a truncate 
distal margin; a small, button-shaped ventral cirrus 
is present near the inferior margin. The first indica- 
tion of notopodial structures is on setiger 3; the 
notopodia are fully developed from setiger 9 and 
all notopodia are similar in median and posterior 
setigers. Each is conical and has a digitate dorsal 
cirrus. Each notopodial seta is slender and is evenly 
dense with fine hairs. 

Branchiae are present from setiger 33 in all 
posterior segments except for the pygidium (Fig. 9). 
The first 10-15 pairs are short; all others are long, 
cirriform and project from the inner posteroventral 
margin of the neuropodia. 

The pygidium is a small circular disk; 2 short 
lateral anal cirri and a single, mid-ventral anal 
cirrus present. The anus is on a ridge near the 
middle of the pygidium. As noted above, the 
posterior end of the specimen is in regeneration; 
the pygidium is thus probably considerably smaller 
than normal for this species (Fig. 9). 

Each of the two first pairs of neuropodia has a 
single, thick, black aciculum with a rounded, blunt 
tip; setae are absent in these neuropodia. All other 
neuropodia have 2 closely similar kind of simple 
setae; near the superior margin is found 1 or 2 setae 
with a very coarse denticulation. The middle and 
inferior portion of the neuropodial fascicles consist 
of setae with fine denticles in closely packed whirls 
(Fig. 6). The neuropodial acicula are less than half 
as thick as the acicula in the two first neuropodia; 
each is yellow and bluntly cone-shaped. Acicula 
are absent in the notopodia; each has 2 to 3 short, 
slender setae. Each of the notopodial setae is 
cylindrical and finely pilose. Furcate setae were 
not observed. 

The re-description of the type-specimen of L. 
aberrans differs from the original description in 
the following features. The exact distribution of 
branchiae were not mentioned by Monro and may 
be of some importance in separating this from 



22 



mi lei i\ sot iiii.RX cm norma academy oi sciences 



VOLUME 70 



related species. Dorsal cirri or at least remnants of 
dorsal cirri, were found to be present on all notopo- 
dia. Such cirri were said to be absent in the original 
description. Monro reported the presence of thick, 
broken spines in the notopodia; no trace of any 
notopodial spines or notopodial acicula could be 
found in the re-examination; it is here suggested 
that Monro mistook the dorsal cirri for the noto- 
podial spines usually present in the pilargiids. 

Two thick, black acicula are present in each of 
the first neuropodia; these were mentioned by 
Monro, but he also assigned setae to the first 
neuropodia (Monro. 1936); no setae other than 
the acicula and no remnants of setae in the para- 
podial base were observed in the re-examination 
of the type. 

The prostomium, and the peristomial segment 
are clearly separated from each other and from the 
first setiger. 

The structure of prostomium and relationship 
between the different parts of the parapodia in 
L. aberrans are similar to those in other pilargiids. 
The genus Loandalia differs clearly from all other 
pilargiids in the presence of large, thick acicula in 
the anterior neuropodia and in the absence of 
notopodial spines and acicula. The presence of 
dorsal cirri in all notopodia separates it clearly 
from other species usually assigned to Loandalia; 
also branchiae have not been reported from any 
other member of this genus. 

Distribution: The holotype and only known speci- 
men of L. aberrans comes from Angola, West Africa 
in shelf depths. 



ACKNOWLEDGMENTS 

The authors wish to thank the Allan Hancock Founda- 
tion, Western Oil & Gas Association, and Dr. Dale 
Straughan for supporting this study; also Drs. Olga 
Hartman and J. David George for their assistance in 
making the holotype of Loandalia aberrans available 
for study. 



Literature Cited 

Berkeley. E. and C. Berkeley. 1941. On a collection of 
Polychaeta from southern California. Bull. So. 
Calif. Acad. Sci.. 40: 16-60. 

Chamberlin. R. V. 1919. The Annelida Polychaeta. 
Mem. Mus. Comp, Zool. Harvard, 48: 1-514. 



Day, J. H. 1957. The Polychaet fauna of South Africa. 
4: New species and records from Natal and 
Mozambique. Ann. Natal Mus., 14: 59-129. 

— . 1963. The polychaet fauna of South Africa, 8: 



New species and records from grab samples and 
dredgings. Bull. British Mus. (Nat. Hist.) Zool., 
|(): 183-445. 

Ditlevsen. H. 1917. Annelids, I. The Danish Ingolf 

Expedition, 4: 1-71. 

Fauvel. P. 1923. Polychetes errantes. Faune de France, 
5: 1-488. 

. 1932. Annelida Polychaeta of the Indian Muse- 



um, Calcutta. Mem. Indian Mus., 12: 1-262. 

Hartman, O. 1947. Polychaetous annelids. Pt. 8. Pilar- 
giidae. Allan Hancock Pacific Exped., 10: 483-523. 

. 1959. Catalogue of the polychaetous annelids 



of the world, parts 1, 2. Allan Hancock Found. 
Publ. Occ. Pap., 23: 1-628. 

Hartman-Schroder, G. 1959. Zur Okologie der Poly- 
chaeteten des Mangrove-Estero-Gebietes von 
El Salvador. Beitr. Neotrop. Fauna, 1: 69-183. 

Jones. M. L. 1961. Two new polychaetes of the families 
Pilargiidae Capitellidae from the Gulf of Mexico. 
Amer. Mus. Nov., 2049: 1-18. 

Mcintosh, W. C. 1879. On the Annelida obtained during 
the cruise of H.M.S. Valorous to Davis Strait in 
1875. Trans. Linn. Soc. London, 1: 499-51 1. 

Monro, C. C. A. 1936. Polychaete worms. Discovery 
Reports, 12: 59-198. 

Miiller, F. 1858. Einiges iiber die Annelidenfauna der 
Insel Santa Catharina an der brasilianischen kiiste. 
Arch. Naturg., 24: 21 1-220. 

Pettibone, M. H. 1966. Revision of the Pilargidae 
Annelida: Polychaeta including descriptions of 
new species and redescription of the Pelagic 
Podarmus ploa Chamberlin (Polynoidae). Proc. 
U.S. Nat. Mus., 118: 115-208. 

Saint Joseph, A. Baron. 1899. Annelides polychetes 
de la rade de Brest et de Paimpol. Ann. Sci. 
Nat. Paris., ser 8, vol. 10: 161-194. 

Thomas, P. J. 1963. Polychaetous worms from the 
Arabian Sea, 1: A new species of the genus Loan- 
dalia Monro. Bull. Dept. Mar. Biol. Oceanogr. 
Univ. Kerala, 1: 29-34. 

Webster. H. E. 1879. Annelida Chaetopoda of the Vir- 
ginian coast. Trans. Albany Inst., N.Y. 9: 202-272. 



Accepted for publication June 30, 1970. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(l):23-30, 1071 

TAXONOMY AND DISTRIBUTION OF II IF AIM l( SIM •( IIS Ol 
LUCICUTIA (COPEPODA: CAI.ANOIDA) 

JlJI. I(J VllMI. 1 



ABSTRACT: The (uxonomy ami distribution Of the Arctic species ol 
Lucicutia were studied. Analysis of approximately 400 plankton hauls from 
the Arctic Ocean revealed the presence of Lucicutia polaris Brodsky, /.. 
pseudopolaris Heptncr, and /.. anomala Brodsky in samples collected in deep 
waters. The two former species are closely related hut differ in morphological 
and biological characteristics of the adults and juvenile stages. The male of 
Lucicutia anomala is described for the first time. These species have a wide 
geographical distribution in the Arctic Ocean. They inhabit the Arctic bottom 
water and show different but overlapping ranges of vertical distribution. L. 
pseudopolaris is most abundant at a depth of about 1500 m; L. polaris prefer- 
entially lives at a depth of about 2000 m; and L. anomala is found in waters 
deeper than 2000 m. This species is most abundant at depths exceeding 3000 
m. Physical, chemical and biological factors which might restrict the distribu- 
tion of these species at different depths were discussed. 



The plankton analyzed in this study is part of the 
collections of the Artie project of the University of 
Southern California. The samples were collected 
from the drifting stations Arlis II and T-3 in. the 
western Arctic Ocean, during the years 1952-1955 
and 1964-1968, respectively. Horizontal tows at 
different depths of water and vertical tows in 

j increments from bottom to surface were made with 
0.5 m diameter closing nets of 62, 73, and 215 m 

' mesh apertures and open nets of 73 and 215 m 
mesh apertures. 

The male of L. polaris, and the adults and cope- 
podite stages IV and V of L. pseudopolaris are 

! described. Since L. polaris and L. pseudopolaris are 
closely related species, and since they are well 
represented in the Arctic Ocean, a detailed compari- 

; son of the external morphology of the adults and 
juvenile stages of these species was made. 

The study on the vertical distribution of the 
species was based on the analysis of closing net 
samples. For this purpose horizontal tows at depths 
of 100, 500. 1000, 1500, 2000 and 3000m, and 
vertical tows at depths between 100-500m, 500- 
1000m. 1000- 1500m, 1500-2000m. 2O0O-3OO0m. 
and below 3000m were considered. Since the verti- 
cal and horizontal tows were not quantitatively 
comparable, and the number of samples taken at 
different depths varied, the numerical values obtain- 
ed in the analysis of vertical distribution of the 
species have been expressed in terms of percentages 
of relative abundance. 



A list of stations at which plankton samples 
containing specimens of Lucicutia were collected 
has been deposited at the National Oceanographic 
Data Center. Reference specimens of the three 
species have been deposited in the collections of the 
Allan Hancock Foundation. 



DESCRIPTION OF THE SPECIES 

Lucicutia pseudopolaris Heptner. 1969 
Figures 2a-m. 4e-m. 

Lucicutia pseudopolaris Heptner. 1969. 

Lucicutia polaris Brodsky. 1950 ( 6? only): Hiilse- 
mann. 1966 ($ only). 

Measurements. Table 1 . 

Diagnosis. Total length 3.8 to 4.7 mm in the 
female and 3.6 to 4.4 mm in the male. Outline of 
the prosome ovate: anterior margin of the prosome 
rounded in dorsal and lateral view. Furcal rami as 
long as the abdomen. Antennules 24-segmented: 
in the female they exceed body length by the last 
2-3 segments; in the male they reach the end of the 
furca. First pair of legs with 2-segmented endo- 
podites; outer spine on the 1st exopodal segment 
reaches the distal third of the outer spine on 



department of Biological Sciences. University of Southern 
California, Los Angeles. California 90007 

Present address: Department of Oceanography. University of 
Washington. Seattle. Washington 9S105 



23 



24 



mill ll\ SOI III1RN CALIFORNIA ACADEMY OF SCIENCES 






VOLUME 70 



180° 




9 0° 

Figure 1. Occurrence of arctic species of Lucicutia in samples analyzed in this study. 



exopodal segment 2. Female inner seta on exopodal 
segment 2 reaches beyond the base of inner seta 2 
on exopodal segment 2. 

Description. Adult female. The general shape 
of the body is robust. The outline of the prosome is 
ovate. The anterior margin of the cephalosome is 
rounded in dorsal and lateral view (Figs. 2a, b). 

The urosome is 1.8-1.9 times as long as the 
prosome. The abdomen is nearly as long as the 
furcal rami. The genital segment is symmetrical in 
dorsal view (Fig. 2e); the genital protuberance is 
well developed and located in the proximal half of 
the segment (Fig. 2d). The furcal rami are long and 
thin, about 1 1 to 15 times longer than wide. 

The antennules, 24-segmented, extend beyond 
the furca by the last 2-3 segments. Segments 1 and 

2 fused. 

The mandibular palp has 4 setae on the basal 
segment. The exopodite is 5-segmented, the endo- 
podite 2-segmented; it is furnished with 4 setae on 
the proximal segment and 8 setae on the distal 
segment. The gnathobase of the mandible bears 9 
or 10 teeth and a seta; the 4 proximal teeth are 
larger than the others. 

The m&xillule has 14 spines on the 1st inner lobe; 

3 setae each on inner lobe 2 and 3; 3 setae on the 
basipodal segment 2; 9 setae on the endopodite; 1 1 



setae on the exopodite and 5 setae on outer lobe 1 . 

The first pair of legs (Fig. 2f) has 3-segmented 
exopodites and 2-segmented endopodites. The outer 
spine on exopodal segment I reaches the distal 
third of the outer spine on exopodal segment 2. 

Exopodites and endopodites have 3 segments on 
legs 2 to 4 (Fig. 2g). The outer distal margin of 
exopodal segments 1 and 2 of legs 2 and 3 are 
strongly projected. 

The fifth pair of legs (Fig. 2h) has 3-segmented 
exopodites and endopodites. The inner seta on 
exopodal segment 2 reaches beyond the base of 
inner seta 2 on exopodal segment 3. Exopodal 
segment 3 is as long as the combined length of 
exopodal segments 1 and 2. The terminal spine on 
the exopodite is slightly shorter than the length of 
exopodal segment 3. 

Adult male. Body shape (Figs. 2j, k) is as the 
female. The furcal rami are about 10 to 14 times as 
long as wide. The left antennule modified to serve 
as a weak clasping organ; the segments 1-2, 19-21, 
and 22-23 are fused. The right antennule reaches 
the end of the furca. 

The appendages of the cephalosome and legs 1 
to 4 of the metasome are as those of the female. 

The basipodal segment 1 of the right 5th leg 
(Fig. 21) is longer than the corresponding segment 



1971 



AKCTIC COI'I-.I'ODS 



25 



J.lt mm, o,b,j.k 
'■'■ mm c,d,o.g 
■ «■" mm , f.h.1,1 

.1.1* CDJIL m 




Figure 2. Lucie utia pseudopolaris Heptner. Female: a, 
dorsal view; b, lateral view; c, anterior part of cephalo- 
some, lateral view: d, posterior part of body, lateral 
view; e, posterior part of body, dorsal view; f, first leg; 
g, second leg; h. fifth leg; i. rostrum. Male: j, dorsal 
view; k, lateral view; 1, fifth leg; m, inner margin of 
second basal segment of right fifth leg in three different 
specimens. 



of the left leg. Basipodal segment 2 of the left leg 
has a protrusion on its inner distal corner. This 
protrusion carries a row of teeth, variable in number 
i| and size (3-13), and a small rounded protuberance 
(Fig. 2m). The right side of the fifth pair of legs has 
a 2-segmented endopodite and exopodite. Exopodal 
segment 2 has 2 apical spines and 1 outer subapical 
spine. The left side of the fifth pair of legs has a 
3-segmented exopodite and endopodite. Exopodal 
segment 3 has 1 outer, apical, and inner spine. 

Copepodite Stage V. Body shape (Figs. 4j, 1) is 
as the adult. The cephalosome and metasomal 
segment 1 are separated; metasomal segments 4 
and 5 are fused. The urosome is composed of 4 
abdominal segments and the furca. 

The antennules, 25-segmented. exceed the length 
of the body by the last 2-3 segments. 

The first pair of legs has 3-segmented exopodites 
and 2-segmented endopodites (Fig. 4j). Legs 2 to 4 
have 3-segmented exopodites and endopodites. 



The fifth pairol legs has 2-segmented endopi 
and exopodites in both sexes digs. 4k. mi ["hi 
distal segment corresponds to fused segments 2 
and 3. In the male Ihe right side of leg 5 is shorter 
than ihe left. 

Copepodite Stage IV. The specimens of this 
stage have a 3-segmented abdomen Oigs. 4c. I). 
Exopodites and endopodites of legs 1-4 have 3 
segments (Fig. 4g). The fifth pair of legs has 1- 
segmentcd endopodites and exopodites. 

Distribution. I., pseudopolaris has a wide geo- 
graphical distribution in the Arctic Ocean. It has 
been recorded by Heptner (1969) from an area 
between 76° 3 8' N and 8 1 ° 1 2' N and 1 73 ° 44' E and 
189°35' E, and at 88° 10' N 82' J 22' W. In our 
collections it was found between 72°42' N and 
83°27' N and 9° 15' W and 157° 12' W. 



<■» mm. obi jmnq 
, *■» mm — ,cdeg 
■ '■" mm I h kopr 
e , l.ll mm I 




Figure 3. Lucicutia polaris Brodsky. Female: a. dorsal 
view; b. lateral view; c. anterior part of cephalosome. 
lateral view; d, posterior part of body, lateral view: e. 
posterior part of body, dorsal view: f. first leg: g. second 
leg; h. fifth leg. Male: i. dorsal view: j. lateral view; k. 
fifth leg: 1. inner margin of second basal segment of 
right fifth leg in two different specimens. Stage V 
copepodite. female: m. dorsal view: n. lateral view: o. 
first leg; p. fifth leg. Stage V copepodite. male: q. dorsal 
view: r. fifth leg. 



26 



HI I / / 1 1\ SOI I III l<\ ( \l II OHM I l( I/)/ \/> Ol V II \t I \ 



VOLUME 70 



Lucicutia polaris Brodsky, 1950 
Figures 3a-r. 4a-d. 

Lucicutia polaris Brodsky. 1950 (9 only): John- 
son. !963;Heptner, 1969. 

Not Lucicutia polaris Brodsky, 1950: Hiilse- 
mann. 1966; Grice and Hiilsemann; 1967. 

Measurements. Table 1. 

The female of Lucicutia polaris was described 
by Brodsky (1950). Heptner (1969) transferred the 
male described by Brodsky under this name to a 
new species. L. pseudopolaris. In the same place 
Heptner described the male of L. polaris for the 
first time. 

L. polaris is similar to L. pseudopolaris. however, 
both are separate species with the following mor- 
phological differences: Adult female: Lucicutia 
polaris differs from L. pseudopolaris in the shape of 
the cephalosome. In dorsal view, the anterolateral 
margin of the cephalosome of L. polaris has slight 
lateral projections (Fig. 3a) which is not present in 
the cephalosome of L. pseudopolaris (Fig. 2a). In 
lateral view, the anterior margin of the cephalosome 
of L. polaris is depressed (Figs. 3b, c). whereas 
that of L. pseudopolaris is rounded (Figs. 2b, c). 

The length ratios of the prosome-urosome and 
abdomen-furca are smaller in L. polaris than in 
L. pseudopolaris (Table 1 ). 

The appendages of the cephalosome are not 
significantly different in both species. The external 
spines on the exopodal segments of the first 4 pairs 
of legs are slightly shorter in L. polaris (Figs. 3f-h) 
than in L. pseudopolaris (Figs. 2f-h). 

On the fifth pair of legs of L. polaris the inner 
seta on exopodal segment 2 does not reach the base 
of inner seta 2 of the exopodal segment 3 (Fig. 3p); 
whereas, in L. pseudopolaris it extends beyond the 
base of inner seta 2 of exopodal segment 3. Exopo- 
dal segment of L. polaris is longer than exopodal 
segments 1 and 2 together; whereas, in L. pseudo- 
polaris it is as long as the combined length of 
these segments. 

Adult male. In general, the morphological dif- 
ferences found in the female are also present in the 
male. The fifth pair of legs differs from that of L. 
pseudopolaris by the presence of 3 to 6 teeth on the 
inner margin of the left basipodal segment 2, and 
by the presence of an additional spine on the middle 
part of the distal segment of the right exopodite 
(Fig. 3k). 

Copepodite Stages V and IV. The overall appear- 
ance of the body is as that of the adults. The 
differences between the juvenile stages of L. polaris 
and of L. pseudopolaris are similar to those found 
in the adult of both species. They differ mainly by 
the shape of their cephalosome. as described for 



the adults. In /.. polaris. the total length of the body, 
and the length ratios of the prosome-urosome and 
abdomen-furca are smaller than those in the juvenile 
stages of /.. pseudopolaris (Table I ). 

Distribution. I., polaris has been recorded from 
the central part of the Arctic Ocean (Brodsky, 
1950: Johnson, 1963: Heptner. 1969) and from the 
north Pacific Ocean (Heptner, 1969). In our collec- 
tions it was found in the area between 75°03' N 
and 83° 15' N. and 90° 30' W and I 73° 36' W. L. 
polaris has also been reported from the Indian 
Ocean and southeastern Pacific Ocean (Hiilsemann, 
1966; Grice and Hiilsemann, 1967). These records 
probably correspond to species from the southern 
Pacific and Indian Ocean rather than to the species 
/_. polaris described by Brodsky from the Arctic 
Ocean. 

Lucicutia anomala Brodsky, 1950 
Figures 4n-s. 

Lucicutia anomala Brodsky. 1950; Grice and 
Hiilsemann, 1965; Hiilsemann. 1966. 

Measurements. Table 1 . 

Brodsky (1950) described the female of L. ano- 
mala. The male is here described for the first time. 

Adult male. The general shape of the body (Figs. 
4n. o) and of the appendages agree very closely with 
those described for the female (Brodsky, 1950). 

The left antennule is modified to serve as a weak 
clasping organ. The segments 1-2, 19-21, and 22-23 
are fused. The right antennule reaches the end of 
the furca. 

The 1st pair of legs has 3-segmented endopodites 
(Fig. 4q). 

The right side of the fifth pair of legs (Fig. 4r) 
has a 2-segmented exopodite and endopodite; basal 
segment 1 is slightly longer than the corresponding 
segment on the left leg; basipodal segment 2 of the 
right side has a smooth projection on the outer distal 
margin; exopodal segment 2 has 2 apical spines. 
The left leg has a 3-segmented exopodite; the inner 
projection of basipodal segment 2 has a row of 5 
to 1 1 small teeth (Fig. 4s); exopodal segment 3 
bears, 1 apical and 1 inner spine. 

Distribution. This species has been recorded 
from the Arctic Ocean (Brodsky, 1950) and north- 
eastern Atlantic Ocean (Grice and Hiilsemann. 
1965). In our collections it was found in the area 
between 75° 18' N and 77° 51' N. and 140° 40' W 
and 157°12'W. 

EPIZOIC SUCTORIANS 

Epizoic suctorians were observed on these three 
species of Lucicutia. The number of species of 
suctorians is uncertain. The rates of incidence on 



U97I ARCTIC COPEPODS 

I ABLE I. Measurements unci length ratios "I the Arctic species <>l Lucicutla 



27 



TOTAL LENGTH (MM]* 



I'ROSOMI -UROSOMI. 
LENGTH RA1 [0 



ABDOMI N I I V ft 

I I '.III HA IIO 



N Mean Range St. Dev. Mean Range St. Dev. N Mean Range 



Lucicutia pseudopolaris 




















Adult female 


37 


4.27 


3.8 1-4.75 


0.206 


1.87: 1 


1.70-2.15 


0.096 


20 


0.97: 1 


0.82-1.06 


Adult male 


20 


4.07 


3.68-4.43 


0.177 


1.73: I 


1 .56-2.00 


0.1 14 


18 


0.98: 1 


0.88-1.10 


Copepodite 
Stage V female 


y 


3.27 


3.10-3.56 


0.148 


2.11: 1 


1.90-2.27 


0.127 


9 


1.04: 1 


0.91-1.20 


Copepodite 
Stage V male 


7 


3.27 


3.13-3.41 


0.089 


2.07: 1 


1.86-2.31 


0.135 


7 


1.05: 1 


0.94-1.18 


Copepodite 
Stage IV 


5 


2.55 


2.46-2.64 


0.064 


2.32: 1 


2.26-2.41 


0.055 


5 


1.08: 1 


1.04-1.1 1 


Lucicutia polaris 


Adult female 


29 


3.76 


3.50-4.06 


0.132 


1.68: 1 


1.48-1.90 


0.097 


20 


0.82: 1 


0.67- 1 .00 


Adult male 


24 


3.68 


3.25-4.06 


0.197 


1.60: 1 


1.46-1.72 


0.079 


20 


0.83: 1 


0.72-0.96 


Copepodite 























Stage V female 12 2.84 2.76-2.98 0.060 1.87:1 1.75-2.00 0.085 12 0.89:1 0.77-0.94 



Copepodite 
Stage V male 



2.87 2.76-2.98 



1.91: 1 1.90-1.93 



2 0.89:1 0.87-0.91 



Copepodite 
Stage IV 



4 2.10 2.06-2.18 0.045 2.26:1 2.09-2.40 0.112 4 0.85:1 0.81-1.04 



Lucicutia anomala 



Adult female 9 2.50 2.43-2.58 0.043 1.75:1 1.62-1.85 0.058 9 0.74:1 0.66-1.06 



Adult male 



7 2.42 2.30-2.50 0.080 1.71:1 1.66-1.77 0.044 7 0.78:1 0.75-0.83 



Copepodite 

Stage V 2 2.01 2.00-2.03 — 1.97:1 1.86-2.09 



— 2 0.82: 1 0.79-0.86 



*N = number of specimens; St. Dev. = sample standard deviation 



their hosts varied on the three species of Lucicutia 
(Table 2). 

L. polaris carried at least 4 or 5 different species 
of suctorians and had the highest rate of incidence. 

L. pseudopolaris seemed to have 3 different 
species of epizoa and a lower rate of incidence. 

L. anomala carried only 1 species of suctorians 
and had the lowest rate of incidence. 

Suctorians were observed on adults and juvenile 
stages. In general, the rate of incidence was slishtlv 



higher in males than in females. The areas of the 
body with a higher number of epizoa were the furca 
and anal segment and. in the male, also the basipodal 
segments of the fifth pair of legs. Low rates of 
incidence were observed on other surfaces of the 
body and on metasomal appendages. 

VERTICAL DISTRIBUTION OF THE SPECIES 
The three species of Lucicutia discussed in this 
paper are widely distributed in the Arctic Ocean. 



28 



BULLETIN SOUTHERN CALIEORNIA ACADEMY OF SCIENCES 



VOLUME 70 



Table II. Rate [ r '< ) of incidence of suctorians 
on the species of Lucicutia 





L. 


polaris 


L. 


pseudopolaris 


L. 


anomala 


Female 




65 




1 1 




12 


Male 




77 




29 




12 


Stage V 




42 




32 




— 



They inhabit deep water, the Arctic Bottom Water, 
and show different but overlapping ranges of verti- 
cal distribution. 

L. pseudopolaris is abundant at a depth of about 
1500 m and diminishes gradually in abundance 
from there to disappear above 1000 m and below 
3000 m. L. polaris seems to inhabit depths between 
1500 m and less than 3000 m. It was found in 
greatest numbers at a depth of about 2000 m. L. 
anomala lives in deeper water from about 2000 m 
to more than 3000 m. It is more abundant at depths 
exceeding 3000 m (Fig. 5). 

The Arctic bottom water, as defined by Coach- 
man (1963), is a homogeneous water mass below 
about 900 m depth, with average temperatures 
from 0.0 to -1.0°C and salinities from 34.90 °/°° 
to 34.99 °/°°. Other studies and oceanographic 
data have demonstrated that chemical parameters 
such as oxygen, phosphate and silicate have a 
remarkable uniformity in this water mass (Kusunoki 
et al., 1966; Muguruma, 1961; English, 1961). 
Among the physical parameters, light is not likely 
to be a controlling factor for the vertical distribution 
of Arctic species at the depths in question. Conse- 
quently, it seems that among the measured physico- 
chemical parameters the only one which might 
have a major effect on the vertical distribution of 
these species could be pressure. 

Among the biological factors which might restrict 
these species to different ranges of depths we must 
consider food availability and competition for food. 
English ( 1 963) briefly discussed the possible sources 
of food for zooplankton in the Arctic Ocean, and 
suggested that the production of particulate food 
by secondary producers of the nannoplankton could 
play a more important role in the Arctic Ocean 
than in other areas. In regard to the Arctic bottom 
water, it is likely that food for filter-feeders can be 
provided by the local secondary producers of the 
nannoplankton. and by sinking of dead organisms, 
detritus, and aggregates of organic matter from the 
upper layers. Several studies on the formation, 
distribution, and role of detritus and aggregates of 



organic matter in the sea (Parsons and Strickland, 
1962; Riley, 1963; Menzel and Goering, 1966; 
J0rgensen, 1966; Johannes, 1967) concluded that 
they can be considered as a very important potential 
source of food for filter feeders. 

A close relationship between the feeding habits 
of calanoid copepods and the structure of their oral 
appendages was experimentally shown by Anraku 
and Omori (1963). The comparison between the 
oral appendages of the Arctic species of Lucicutia, 
and those studied by Anraku and Omori, indicated 
that Lucicutia has characteristics of herbivorous 
and omnivorous forms. The mesh formed by the 
setules of the maxillae, the filtering appendages, is 
of about 5 to 10,u in L. pseudopolaris and L. polaris 
and slightly smaller than 5/j. in L. anomala. This 
fine mesh and the structure of the oral appendages 




Figure 4 — Lucicutia polaris Brodsky. Stage IV cope 
podite: a, dorsal view; b, lateral view; c, first leg; d, fifth 
leg. Lucicutia pseudopolaris Heptner. Stage IV cope- 
podite: e, dorsal view; f, lateral view; g, first leg; h, fifth 
leg. Stage V copepodite, female: i, lateral view; j, first 
leg; k, fifth leg. Stage V copepodite, male: 1, lateral 
view; m, fifth leg. Lucicutia anomala Brodsky. Male: 
n, dorsal view; o, lateral view; p, mandibular gnatho- 
base; q, first leg; r, fifth leg; s, inner margin of second 
basal segment of right fifth leg in three different 
specimens. 



1971 



Al<( I K ( OI'I.I'ODS 



2't 



100 



Luclcutla pseudopolarit 
Luclcutla polarl» 
I I Luclcutla anomola 




1000 



1000- 
1500 



1500 



1500- 2000 

2000 



2000- 
3000 



3000 



below 
3000 



Depth 



i n 



meter 



Figure 5 — Percentages of relative abundance and number of specimens of the arctic species of 
Lucicutia at different depths. 



intermediate between herbivorous and omnivorous 
forms, would permit these species to feed on aggre- 
gates of organic matter and detritus, which range 
in size from about 1^ to several mm, and on small 
arganisms of the nannoplankton. The absence of 
morphological differences in the structure of the 
jral appendages (especially between L. pseitdo- 
fi oolaris and L. polaris) suggests that the above 
it species are feeding on the same or on a very similar 
f food supply. 

Related species can avoid or reduce competition 
or food by distributing themselves at different 
depths. Marshall (1954; 1963) discussed the vertical 
distribution of related species of fishes ( Vinciguerria 
;pp; Cyclothone spp: Stomias spp;) and gave exam- 
Dies of vertical exclusion. Other examples of sharp 



stratification of related species of pelagic orga- 
nisms were given by Banse (1964). In general, this 
behavior might permit a reduction of competition 
for food and a better utilization of the ecosystem. 

It is likely that the vertical distribution and 
stratification of the Arctic species of Lucicutia 
are the outcome of the interactions of all environ- 
mental factors through time — physical factors 
such as pressure, chemical, and more likely biologi- 
cal factors — tending to a better utilization of the 
food niches with consequent reduction of interspe- 
cific competition and better possibilities of survival. 

ACKNOWLEDGMENTS 

The author is indebted to Dr. John L. Mohr and Mr. 
Stephen Geiger (Allan Hancock Foundation). Drs. 



30 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCUM I S 



VOLUME 70 



Karl Banse and T. Saunders English (Department of 
Oceanography, University of Washington), and Dr. 
Michael M. Mullin (Scripps Institution of Oceanog- 
raphy) for their helpful criticisms, valuable suggestions, 
and careful reading of the manuscript. He is most grate- 
ful to Dr. John L. Mohr for providing the opportunity to 
do this study. The use of the arctic plankton collections 
and laboratory facilities provided by the Allan Hancock 
Foundation are also gratefully acknowledged. 

The present study was supported by a grant from 
the Office of Naval Research under contract NONR 
228(19), NR 307-270. 



Literature Cited 

Anraku, M. and M. Omori. 1963. Preliminary survey 
of the relationship between the feeding habit and 
the structure of the mouth-parts of marine cope- 
pods. Limnol. Oceanogr., 8: 1 16-126. 

Banse, K. 1964. On the vertical distribution of zoo- 
plankton in the sea. Progress in Oceanography, 

2:55-125. 

Brodsky. K. A. 1950. Veslonogie rachki Calanoida 
dal'nevestochnykh morei SSSR i polyarnogo bas- 
seina. Opred. Po Faune SSSR, Akad. Natik SSSR. 
35: 1-441. 

Coachman, L. K. 1963. Water masses of the Arctic. In, 
Proc. Arctic Basin Symp.. October 1962, Arctic 
Inst. N. Amer., p. 143-167. 

English. T. S. 1961. Some biological observations in 
the central North Polar Sea. Drift Station Alpha. 
1957-1958. Arctic Inst. N. Amer. Sci. Rep., 
15: 1-80. 

. 1963. Some remarks on the Arctic Ocean 



Plankton. In. Proc. Arctic Basin Symp.. October 
1962, Arctic Inst. N. Amer., p. 184-196. 

Grice, G. D. and K. Hiilsemann. 1965. Abundance, 
vertical distribution and taxonomy of calanoid 
copepods at selected stations in the northeast 
Atlantic. J. Zool., 146: 213-262. 

. 1967. Bathypelagic calanoid copepods of the 

western Indian Ocean. Proc. U. S. Nat. Mas.. 
122: 1-67. 



Heptner, M. V. 1969. Systematic status of Lucie tit ia 
polaris Brodsky. 1950. (Copepoda. Lucicutiidae) 
and description of L. pseudopolaris sp. n. from the 
Polar Basin. Zool. Zh.. 48: 197-206. 

Hiilsemann, K. 1966. A revision of the genus Lucicutia 
(Copepoda: Calanoida) with a key to its species. 
Bull. Mar. Sci.. 16: 702-747. 

Johannes, R. E. 1967. Ecology of organic aggregates in 
the vicinity of a coral reef. Limnol. Oceanogr., 
12: 189-195. 

Johnson, M. W. 1963. Zooplankton collections from 
the high Polar Basin with special reference to the 
Copepoda. Limnol. Oceanogr.. 8: 89-102. 

Jorgensen, C. B. 1966. Biology of suspension feeding. 
Oxford, New York Pergamon Press, 357 pp. 

Kusunoki, K., T. Minoda, K. Fujino, and A. Kawa- 
mura. 1967. Description of oceanographic observa- 
tions at Drift Station Arlis II in 1964-1965. Arctic 
Inst. N. Amer., Techn. Rep., 34 p. 

Marshall, N. B. 1954. Aspects of deep-sea biology. 
London. Hutchinsons. 380 pp. 

. 1963. Diversity, distribution and speciation of 

deep-sea fishes. In, Speciation in the Sea. The 
Systematic Association. Publ. No 5: 181-195. 

Menzel. D. W., and J. J. Goering. 1966. The distribu- 
tion of organic detritus in the ocean. Limnol. 
Oceanogr., 11: 333-337. 

Minoda. T. 1967. Seasonal distribution of Copepoda in 
the Arctic Ocean from June to December. 1964. 
Records of Oceanographic Works in Japan, 9: 160- 
168. 

Muguruma, J. 1961. Oceanographic observations on 
Fletcher's Ice Island T-3. Winter 1959-1960. Arctic 
Inst. N. Amer.. Res. Paper No 6: 1-18. 

Parsons, T. R. and J. D. H. Strickland. 1962. Oceano- 
graphic detritus. Science, 136: 313-314. 

Riley. G. A. 1963. Organic aggregates in seawater and 
the dynamics of their formation and utiliza'ion 
Limnol. Oceanogr., 8: 372-381. 



Accepted for publication September 4, 1970. 



BULLETIN SO. (ALU. ACADEMY OF SCIENCES 70(1): 31 42, l<;7l 

A REVIEW OF THE GENUS HOCC A RDIA CARAZZ1 (POLYCHAETA: 
SPIONIDAE) WITH DESCRIPTIONS OF TWO NEW SIM CIES 



James A. Blake 1 and Keith H. Woodwick- 



Abstract: Taxonomic characteristics "l 14 species of Boccardia are 
compiled in ihc form of a table. Each species is further distinguished in a key. 
Boccardia ligerica Ferronniere is redescribed with new synonymies. Two 
new species, B. berkeleyorum and /;. chilensis are described. 



Ihc present paper is fourth in a series on Boccardia 
taxonomy. The first three (Woodwick. 1963a; 
l§63b; Blake. 1966) dealt with B. tricuspa, B. 
•ohtmhiuna, B. proboscidea and B. hamata. Adults 
were described and observations on ecology record- 
ed. In the present study two species new to science 
are described. Boccardia berkeleyorum comes from 
jentral California while B. chilensis comes from 
the coast of Chile. In addition. B. ligerica, a 
European species, is redescribed and new synony- 
mies presented. Differential characteristics of 
<nown species of the genus are summarized in 
Table 1. Analysis of the critical characteristics 
eveals that several well-defined groups of species 
ire present. Discussion of these relationships follows 
he species descriptions. 

Specimens of Boccardia berkeleyorum were 

:ollected by us during the years 1961-63. The B. 

hilensis material was included among the collec- 

ions of the Lund University Chile Expedition 

1948-49, the sedentary polychaetes of which are 

low at the Allan Hancock Foundation. We are 

grateful to Dr. Kristian Fauchald for the loan of 

:his material. Specimens of B. ligerica were kindly 

isent by the Zoological Museum of Amsterdam 

ZMA) and Dr. Francois Rullier of the Universite 

Je Catholique. Angers. France. 

Information included in Table I comes mostly 
rom published sources. We are grateful to Dr. 
Bernard McAlice of the University of Maine for 
lis translation of the article by Khlebovitsch. 

This study was supported in part by N.S.F. Grants 
G- 17990) and (GB-1264). Additional studies on 
Boccardia larval development have been completed 
ind will be published in a subsequent paper. 

SYSTEMATICS 
Genus Boccardia Carazzi, 1893 

Type species: Boccardia polybranchia (Haswell) 
1885. 
Synonyms: ?Perialla Kinbers. 1866. 



Diagnosis: Prostomium rounded or bilid anteri- 
orly, extending posteriorly as a caruncle. Eyes 
present or absent. First setigerous segment with 
or without notosetae. Setiger 5 greatly enlarged, 
with heavy spines of either one or two types, 
arranged in a curved row. Posterior notosetae may 
include specialized spines, hooks and/or simple 
capillaries. Hooded hooks begin on setiger 7. 
Branchiae begin on segments anterior to setiger 
5 and are not fused to the notopodial lobes. Pygi- 
dium enlarged or reduced, saucerlike or divided 
into lobes. 

The following species of Boccardia are herein 
considered valid and are included in Table I. 
Synonymies are in parentheses; asterisk indicates 
a new synonymy. 
1. B. basilaria Hartman. 1961 

B. berkeleyorum new species 

B. chilensis new species 

B. columbiana Berkeley. 1927 

B. hamata (Webster). 1879 

(B. uncata Berkeley. 1927) 

(Polydora uncatiformis Monro. 

6. B. ligerica Ferronniere. 1 898 
(Polydora redeki Horst. 1920) 

7. B. natrix (Soderstrom). 1920 

8. B. perata (Khlebovitsch), 1959 

9. B. polybranchia (Haswell). 1885 
10. B. proboscidea Hartman. 1940 

(?Spio calif ornica Fewkes, 1889) 
(?Polydora californica Treadwell. 
1914 HOMONYM) 
11.6. pseudonatirx Day. 1961 

12. B. tricuspa (Hartman). 1939 

13. B. truncata Hartman. 1936 

14. B. sp. 



1938) 



'Pacific Marine Station. University of the Pacific. Dillon Beach. 

California 94929. 

'-Department of Biology. Fresno State College. Fresno. California 

93710. 



31 



32 



HI I I I II \ SOI I III RS CAI II ORNIA ACAOPM) <>/ S( II \< ES 



VOLUME 70 



(Polydora redeki scn.su Okuda, 1937. not Hoist. 
1920) 

Boccardia ligerica Ferronniere Emended 
Figure 1 

Boccardia ligerica Ferronniere. 1898; Fauvel. 
1927; ?Day. 1955. 1967. 

Polydora redeki Horst. 1920; Fauvel. 1927; 
Augener, 1939; Hempel. 1957a. 1957b; Rullier. 
1960. 

Material examined: Holland. Zuider Zee under 
rock, October 1 1. 1932, ZM A 1 156 (30 specimens); 
France. Benouville. April 25. 1953, coll. Francois 
Rullier. (6 specimens). 

Description: Complete specimens from Holland 
measure up to 12 mm and have about 70 seg- 
ments. The French material contains only anterior 
fragments. 

The prostomium is weakly bifid on its anterior 
margin (Fig. la). The caruncle extends posteriorly 
to setiger 2 or 3. There is no nuchal tentacle. There 
are 4 eyes; a widely spaced anterior cup-shaped 
pair, and a posterior pair closely spaced and oval 
in shape (Fig. la). 

Setiger 1 is reduced; it has small notopodial 
lobes, no notosetae, and only a small fascicle of 
capillary neurosetae (Fig. la). Setigers 2, 3, 4,-, 6 
and succeeding setigers contain spreading fascicles 
of winged capillary notosetae. The number of these 
setae diminishes in posterior setigers where they 
are replaced by stout capillary setae and a special- 
ized recurved hook (Fig. lg). The parapodia in 
posterior segments are modified in a manner similar 
to those of the closely related B. hamata (Blake, 
1966). They are laterally elevated leaving a dorso- 
median channel between them. The recurved noto- 
podial hooks project medially towards this channel 
(Fig. li). 

The neurosetae of setigers 2, 3, 4,-, and 6 contain 
fascicles of winged capillary setae. Bidentate hooded 
hooks which begin on setiger 7 largely replace the 
capillary setae. The hooks number 5 or 6 in a series 
with 1 or 2 ventral slender capillary setae. In 
posterior setigers the hooks number 2 or 3. The 
structure of the hooded hooks is similar to that of 
B. hamata (Blake. 1966). The 2 teeth have only a 
slight angle between them and there is no constric- 
tion on the shaft (Fig. lh). Angle between the shaft 
and the main tooth is obtuse. 

Setiger 5 is about twice as large as preceding 
and succeeding setigers. It is modified and contains 
a row of heavy spines (Fig. lb) alternating with 
lanceolate companion setae (Fig. lc-d). The heavy 
spines are simple, falcate and have no accessory 
structures. Dorsal to the row of heavy spines is a 
small group of stout geniculate setae (Fig. le) and 



ventral to it is a small tuft of pointed geniculate 
setae (Fig. 11). 

Branchiae occur on setigers 2. 3, -, -, -, 7 and 
succeeding setigers for about one-third the length 
of the worm. 

The pygidium consists of a flattened plate from 
which 2 anal cirri arise. The anus opens on the 
dorsal side (Fig. li). 

Remarks: In the past Polydora redeki and Boc- 
cardia ligerica have been maintained as separate 
species as a result of the lack of complete specimens 
and confusion among investigators on the reported 
distribution of the branchiae. Most descriptions 
of the posterior segments have been incomplete. 

Boccardia ligerica was described by Ferronniere 
(1898) from the Estuary of Loire, France. His 
general description and figures were adequate, but 
his observations on the branchiae were vague. His 
written description suggested that the anterior 
branchiae were distributed on setigers 2, 3, 4, -, -, 
7 . . . , although his figures show anterior branchiae 
only on setigers 2 and 3. He adequately described 
the pygidium and clearly illustrated the two anal 
cirri, one of which appears to have been broken. 
Fauvel (1927) repeated Ferronniere's earlier de- 
scription and Day (1955; 1967) in describing a 
South African specimen referred it with question 
to B. ligerica. 

Polydora redeki was described from Holland by 
Horst ( 1 920). His figures clearly show branchiae on 
setigers 2, 3, -, -. -, 7. Fauvel (1927) repeated Horst's 
description. Augener (1939) briefly described P! 
redeki from northern Germany. He figured the 
pygidium as a simple plate without processes 
Friedrich (1938) did not add to the description 
Hempel (1957b) did not describe the adult mor- 
phology in her paper on tube building. Rulliet 
( 1960) redescribed the species and for the first time 
elucidated the true structure of the pygidium. He 
reported the anterior branchiae as occurring or 
setigers 2, 3, 4, -, -, 7. The branchiae of setiger § 
were said to be smaller than on 2 and 3; however 
his figures do not show branchiae on setiger 4. Hi: ; 
descriptions and illustrations of the post-larva 
stage of 14 setigers show the branchiae as being or 
setigers 2, 3, -, -, -, 7. Rullier's description anc j 
figure of the pygidium are like Ferronniere's figun 
for B. ligerica. 

An examination of specimens from France col 
lected and sent by Rullier and additional material 
from the Zuider Zee of Holland collected in 193: i 
proved that specimens from the two localities wen j 
the same. Only the material from Holland contain 
complete specimens. The anterior ends of th'I 
specimens from the two localities were identical 
Branchiae were limited to setigers 2, 3, -, -, -, 7 1 



1971 



Nl-iW SI'l-X IIS Ol IUH ( A RDIA 



33 




Q05mm 



O 






0.02 mm 





Figure I. Boccardia ligerica Ferronniere: a, anterior end in dorsal view; b. heavy spine from 
setiger 5; c-d, companion setae from setiger 5: e. dorsal setae from setiger 5; f. neurosetae from 
setiger 5; g. posterior notopodial spine: h. hooded hook from setiger 10: i. posterior end in dorsal 
view. 



34 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 



With the absence of branchiae on setiger 4 of all 
specimens examined and the lack of any figures 
which show branchiae on that setiger from the 
published descriptions of Horsl ( 1 9201. Ferronniere 
(189S) and Rullier (I960) we conclude that Boc- 
cardia ligerica and B. redeki are synonymous. Since 
ligerica is the older name, it has priority over redeki. 

Okuda (1937) and Imajima and Hartman (1964) 
have described a species of Boccardia from Japan 
which was identified as B. redeki. It is closely 
related to ligerica, hamata and truncata but one 
important difference is the presence of a nuchal 
tentacle (Table I). 

Hartman (1941: 1961: 1969) referred some 
posteriorly incomplete California specimens to B. 
redeki. Boccardia ligerica is closely related to B. 
hamata. The two differ in the structure of the 
pygidium and the distribution of the branchiae 
(Table I). Because B. hamata is common in Cali- 
fornia waters, it is possible that the incomplete 
specimens described by Hartman are B. hamata. 
Ecology and distribution: Boccardia ligerica occurs 
in brackish waters of mud flats of western Europe 
including Holland, France and Germany: ?South 
Africa. Associated organisms include the amphi- 
pods Corophium volutator Pallas and C. lacustre 
Vanhoffen and the polychaetes Mercierella enig- 
matica Fauvel. Neanthes diversicolor (O. F. Miiller) 
and Neanthes succinea(Frey and Leuckart) (Rullier. 
1960). 

Boccardia berkeleyorum. new species 
Figure 2 

Material examined: California: Cayucos. August 
28. 1961. hermit crab shells, (8, TYPE); March 3 
(2). April 28 (5). July 3. 1962 (8) from coralline 
algae: Morro Bay. October 24. 1961, from shells of 
Pododesmus machroschisma (Deshayes) (1); May 
18. 1963. from Pododesmus (5): San Simeon Beach 
State Park. December 19. 1961, from hermit crab 
shells (1); Fort Bragg. June 20. 1962, from coralline 
algae (1): Eureka (Trinidad Head). June 21. 1962. 
from hermit crab shells ( 1 ). 

The holotype is deposited in the Allan Hancock 
Foundation. University of Southern California and 
was collected at Cayucos (San Luis Obispo County) 
from burrows in Tegula brunnea (Philippi) occupied 
by the hermit crab. Pagurus granosimanus (Stimp- 
son). The collections were made on August 28. 1961 
from rocks at mid-tide level. Additional specimens 
taken at this station have been designated as para- 
types and deposited in the Allan Hancock Founda- 
tion and the United States National Museum. 
Washington. D.C. Additional collections remain 
with the authors. 

It is a pleasure to name this species in honor of 



Edith and Cyril Berkeley who have made signilicant 
contributions to the systematics of polychaetes. 

Description: Specimens in the present collections 
measure 5-14 mm and have up to 100 setigerous 
segments. There is no noticeable body pigmentation. 

The prostomium is rounded on its anterior 
margin (Fig. 2a). The prostomial ridge is elevated 
and enlarged at the level of setiger I: it continues 
posteriorly as a stout caruncle to near the posterior 
border of setiger 3. Palps on preserved specimens 
extend posteriorly to about setiger 12. but are much 
longer on live forms. There are no eyes. 

Setiger 1 lacks both notosetae and a notopodial 
lobe (Fig. 2a). There is a small fascicle of slender 
capillary neurosetae and a bluntly rounded lobe on 
the first setiger. Setigers 2. 3. 4. -. 6 and succeeding 
setigers have posteriorly directed winged capillary 
notosetae arranged in 2 bundles. The second bundle 
has longer setae. In the posterior one-third of the 
body the winged capillary setae are replaced by 
longer capillary setae and heavy acicular spines 
(Fig. 2 j-k). The heavy spines are simple in structure 
and do not have the hooked appearance as in B. 
hamata and B. ligerica. The neuropodia of setigers 
2. 3. 4. -, and 6 have fascicles of winged capillary 
setae. Bidentate hooded hooks begin on setiger 7. 
There are 5 or 6 hooks at first, accompanied by a 
single capillary seta. The number of hooks does not 
exceed 7 per neuropodium and in posterior setigers 
is reduced to 3 or 4. The hooks have 2 teeth nearly 
equal in size separated by an angle of about 90° 
(Fig. 2 h-i). 

Setiger 5 is larger than preceding and succeeding 
setigers and contains modified setae (Fig. 2a). The 
dorsal setae are modified and include two types of 
heavy spines. The first is simple and falcate (Fig. 2 
f-g). The second is bushy-topped and has a small 
accessory tooth (Fig. 2 b-e). Ventral to the heavy 
spines is a small tuft of capillary setae. 

Branchiae are flattened and fingerlike in outline 
(Fig. 2a). They are small on setiger 2. 3. 4. -. 6 and 7. 
while those on 8 and succeeding setigers are much 
larger. Branchiae are reduced in size near the 
mid-body and are absent from the posterior one- 
fourth of the body. 

The pygidium is composed of 4 small individual 
lobes which surround the anal opening (Fig. 2k). 
Variability in size of the pygidial lobes in the present 
specimens may be an artifact of preservation. 

Remarks: The similarity between the modified 
setae of Boccardia berkeleyorum and Polydora 
anophthalma Rioja ( 1962) is apparent. Type mate 
rial for P. anophthalma is no longer in existence 
precluding verification of its branchial distribution 
Rioja did not describe the pygidium and ascribed 
the branchiae as beginning on setiger 8. He stated 



1971 



Nl-.W SPEC IES OF HO( ( A KhIA 



35 





d e 




0.1mm 



0.2mm 




Figure 2. Boccardia berkeleyorum new species: a. anterior end in dorsal view: b-e. bristle-topped 
heavy spines from setiger 5: f-g, falcate spines from setiger 5: h-i. hooded hooks from setiger 12 
seen at two angles; j. posterior setiger in posterior view: k. posterior end in dorsal view. 



36 



BULLETIN SOUTHERN CAL1EORN1A ACADEMY OF SCIENCES 



VOLUME 70 



that posterior spines were absent. The bushy-topped 
setae which he described for the fifth setiger are 
very similar to those of B. berkeleyorum; however, 
the second type of setae are somewhat different. 
Rioja's figure 91 suggests a bifid nature. In B. 
berkeleyorum the second type has a simple falcate 
structure, but if viewed from a certain angle the beak 
of the spine may give a pseudo-bifid appearance. 

Boccardia berkeleyorum is unique among species 
of the genus in the possession of special poste- 
rior spines which are acicular. In this respect it 
approaches certain species of the closely related 
genus Polydom. The species, P. quadrilobata, P. 
caulleryi, and P. armala each have awllike posterior 
spines but differ strikingly in the setation of the 
fifth setiger and lack of anterior branchiae, charac- 
teristic of species of the genus Boccardia. 

The occurrence of different types of posterior 
spines throughout species of the polydorid complex 
may be the result of convergent evolution of species 
of various genera. 

Ecology and distribution: Boccardia berkeley- 
orum was found in central California at Morro Bay, 
Cayucos, and San Simeon Beach State Park, and 
Fort Bragg and Trinidad Head in northern Cali- 
fornia. It was found in three different habitats with 
no apparent correlation with geographical location. 
This species was found in burrows in (1) the shells 
of Tegula brunnea Philippi inhabited by Pagurus 
granosimanus (Stimpson) (10 specimens at 3 sta- 
tions), (2) shells of Pododesmus machroschisma 
(Deshayes) (6 specimens at 2 stations at Morro Bay), 
and (3) in low encrusting coralline algae of the 
genus Lithothamnion (1 1 specimens at 3 stations). 
Other polydorids found associated with Boccardia 
berkeleyorum include B. columbiana, B. tricuspa, 
and Polydora near ciliata in Tegula; B. columbiana 
and B. tricuspa in Lithothamnion; and Polydora sp. 
and P. giardi in Pododesmus. Other Boccardia were 
not found with it in Pododesmus material. 

Boccardia chilensis, new species 
Figure 3 

Boccardia sp. Hartman, 1948. 

Polydora polybranchia Fauvel, 1916; Not Has- 
well, 1885. 

Material examined; Chile. Lund University Chile 
Expedition 1948-49; Sta. 22, July 16, 1948, Golfo 
de Ancud, Isla Quenu, Punta Pinto, western side. 
sheltered intertidal, Lat. 41°49'15" S, Long. 73° 10' 
15" W (2 fragments); Sta. M-121, June 9, 1949, 
Bahia San Vicente, Punta Liles just W. of San 
Vicente, semi-exposed intertidal, rocky, Lat. 36°43' 
36" S, Long. 73°08'10" W (3 specimens— HOLO- 
TYPEand2PARATYPES);Sta. 131, July 1, 1949, 
Iquique, southern part of town, exposed rocky 



intertidal. Lat. 20° 13' 10" S, Long. 70° 10' 19" W 
(1); Sta. M-133, July 7, 1949, Iquique, the harbor, 
sheltered rocky intertidal, Lat. 20° 12'30" S, Long. 
70°10'19" W (2); Bahia de Concepcion, central 
part, SE of Isla Quinquina soft bottom trawl, depth 
20 m, Lat.36°40'15" S, Long. 73°01'48" W ( 1 ). 

The holotype and 2 paratypes are deposited in 
the Swedish National Museum, Stockholm. 

Description: Of the 9 specimens available for 
study the 3 types are well-preserved and adequately 
show diagnostic characteristics. The remaining 
specimens are poorly preserved and show clearly 
only the setal characteristics. The holotype is 28 
mm long and has 125 segments. There is no body 
pigmentation. 

The prostomium is bifid on its anterior margin 
and gradually narrows posteriorly to a point near 
the palpal insertions (Fig. 3a). An enlarged and 
raised area at about the level of the palpal insertions 
bears a nuchal tentacle (Fig. 3a). The caruncle 
extends to the posterior of setiger 2. An enlarged 
dorsal ridge occurs on some specimens from the 
anterior border of setiger 5 to near the posterior 
border of 6. Palps were missing on all specimens 
and were presumably lost at the time of preserva- 
tion. There are no eyes. 

Setiger 1 has capillary setae in both the noto- and 
neuropodia. Setigers 2, 3, 4. -, 6 and succeeding 
setigers have spreading fascicles of winged capillary 
notosetae arranged in 2 rows. The posterior row has 
longer and thinner setae. The number of these setae 
and the distinction between rows rapidly decreases 
on more posterior setigers. In far posterior setigers 
only a few laterally directed long capillary setae 
remain. The neuropodia of setigers 2, 3, 4, -, 6 have 
fascicles of winged capillary setae. Bidentate hooded 
hooks begin on setiger 7. Initially, there are up to 
16 hooks per fascicle with the main fang of these 
hooks almost at a right angle to the shaft (Fig. 3g). 
In far posterior setigers the main fang is strongly 
bent and forms an acute angle with the shaft (Fig. 
3h). In those setigers the hooks number only 8 or 
9 per fascicle. 

Setiger 5 is large and well-developed (Fig. 3a). 
There are two types of heavy spines arranged in a 
curved double row. The dorsally placed setae are 
simple falcate hooks with a hump at the point of 
curvature on the convex side (Fig. 3b). The ventral 
spines have a distal concavity from which arises a 
small cone (Fig. 3 c-f). The end of the shaft contains 
randomly dispersed fine "hairs". In certain views 
one side of the concavity is seen to be elevated as a 
delicate, transparent rim (Fig. 3 e-f). A small tuft 
of capillary setae occurs ventral to the major spines 

Branchiae are fingerlike and occur on 2, 3, 4, -, 6 
and succeeding setigers and continue to the poste- 



1971 



ni:w snx us 01 nocc akdia 



37 




0.5mm 



1 



*m *&- -~- , 



0.05 mm 1 



t 


\ 




'i 

',: 

■; 
' ■; 




i 






"' 


b ' 




c 




i 0.1mm 



d' ! 





Figure S. Boccardia chilensis new species: a. anterior end in dorsal view; b. humped falcate spines 
from setiger 5: c-f. spines from setiger 5 with distal concavity: g. hooded hook from setiger 12: 
h, hooded hook from a far posterior setiger: i. posterior end in dorsal view. 



38 Hi ILLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

Table I. Some Taxonomic Characteristics of the Known Species of Boccardia 



VOLUME 70 













Spines of 










Prostomial 


Branchial 






Setiger 5 








Species of 


nuchal ridge 


distribution 


Notosetae 


Posterior 


a. number of 








Boccardia 


a. prostomium 


a. anterior 


on 


Notopodial 


types 


Pygidium 


Other Features 


References 




b. caruncle 


b. posterior 


Setiger 1 


Spines 


b. description of 










c. eyes 








types 











a. incised 


a. 2, 3, 4, -, 6 




b. to end of 


b. absent 


basilaria 


setiger 3 


posterior 




c. 4 


one-half 



absent 



a. rounded 


a. 2. 3, 4, -. 6 


b. to end of 


b. absent 


berkeleyorum setiger 4 


posterior 


c.O 


one-fourth 



chilensis 



Columbiana 



hamata 



a. incised 

b. into 
setiger 2 

c.O 



a. rounded 

b. to end of 
setiger 3 

c. 4 



a. incised 

b. to end of 
setiger 3 

c. 4 



absent 



a. 2. 3.4. -.6 

b. present present 



a. 2. 3, 4, -, 6 long 

b. absent last capillaries 
few setigers present in 

spreading 
fascicle 

a. 2. 3.-, -.6 

b. absent 

posterior absent 
one-fourth 



ligerica 



a. weakly a. 2, 3, -, -, 
incised -, 7 

b. to end of b. absent absent 
setiger 2 posterior 

c. 4 two-thirds 

a. incised a. 2. 3. 4, -. 6 

b. to end of b. present present 
setiger 2 

c. 4 



absent 



acicular 



absent 



absent 



long 
falcate 
recurved 
hooks 



long 
falcate 
recurved 
hooks 



absent 



a. two 

b. I . falcate 
2. bristle- 
topped with 
constricted 
neck 

a. two 

b. 1. falcate 
2. bristle- 
topped with 
accessary 
tooth 

a. two 

b. 1. falcate 
2. distal 

concavity 



a. two 

b. I. falcate 
2. bristle- 
topped 



a. one 

b. simple 
falcate 



a. one 

b. simple 
falcate 



a. two 

b. 1 . falcate 
2. bristle 

top with 
2 heavy 
smooth 
bosses 



semi- 
circular 
disc, with 
2 ventral 
lappets 



four small 
lobes 



a. pouched glands 
in neuropodia 
of setigers 
10-17 

b. hooded hooks 
unidentate in 
posterior 
setigers 

bores in 

calcareous 

materials 



Hartman 

(1961; 

1969) 



simple 


a. nuchal tentacle 




weakly 


present 




lobed 


b. up to 16 


This pape 


collar. 


hooded hooks 




ventrally 


per 




incised 


neuropodium 


Berkeley 










(1927) 


four equal 




Woodwic 




lobes 




(1963a) J 


small ring 


dorso-median 




with 2 


channel 




ventral blade- 


between 


Blake 


like lappets 


posterior 


(1966) 


each with a 


notopodia 




terminal process 






dorso-median 




flattened 


channel between 




plate with 2 


posterior 


This pap< i 


terminal cirri 


notopodia; 
inhabits 
brackish waters 


■ 



four 
lobes 



1971 



NE W SPEC I IS <>l Hi H < A HDIA 
Table I (Continued). Some I axonomic Characteristics ol the Known Species oi iim cardla 



19 









• ■• 


Spines of 










Prostomial Branchial 






Set liter 5 








ecics of 


nuchal ridge distribution 


Notosctae 


Posterior 


a. number of 








tfeartlia 


a. prostomium a. anterior 


on 


Nolopodial 


types 


Pygidium 


Other Features 


Reference* 




b. caruncle b. posterior 


Setiger l 


Spines 


b. description of 










c. eyes 






types 









a 


incised 


a. 2, 3, 4, -, 6 


b 


into 


b. absent 




setiger 5 


posterior 


c. 


3-6 


one-third 



a. two large forwardly- 

b. I. falcate simple directed "lateral 

present absent 2. bristle collar fans" on ventral Khlcbovitsch 

top and with several side of sctigcrs (1959) 

club-shaped weak lobes 1-4 



a. incised 

b. to end of 
ybranchia setiger 3 

c. 6-8 



a. 2. 3, 4, -, 6 

b. absent 
posterior 
one-half 



absent 



absent 





a. rounded 


a. 2, 3, 4. -. 6 






b. to end of 


b. absent 




boscidea 


setiger 3 


last few 


present 




c. 4 


setigers 


(short) 



absent 



a. two 

b. I. falcate 
2. inverted, 

bristle-topped 
cone with 
oblique 
base 

a. two 

b. I. simple 

falcate 
2. bristle- 
topped 



ventral oral 



disc-like 



4 lobes 

dorsal 

pair 

smaller 

than 

ventral 



groove to 


Carazzi 


anterior of 


(1893) 


setiger 3 


Mesnil 




(1896) 




Fauvel 




(1927) 


pigment along 




prostomium; 


Hartman 


diverse habitats 


(1940: 1941) 


of shell, sand 


Woodwick 


mud . . . 


(1963a) 



a. incised 

b. to end of 
■idonatrix setiger 2 



a. 2, 3,4, -.6 

b. absent 
posterior 
one-third 



a. rounded 




2,3,4,-, 6 


b. to end of 


b 


absent 


setiger 3 




posterior 


c. 4 




two-thirds 


a. incised 




2. 3, 4. -. 6 


b. to end of 


b 


absent 


setiger 2 




posterior 


c. 4 




one-fifth 


a. incised 




2. 3, -, -, 6 



s| redeki 

s(,u Okuda, b. to end of b. absent 
1'7) setiger 3 posterior 

c. 4 one-half 



present 



absent 



absent 



absent 



absent 



absent 



absent 



a. two 

b. 1. simple 

falcate 
2. central 
cone 
with 
raised 
margins 

a. two 

b. 1 . falcate 
2. tricuspid 



a. one 

b. simple 
falcate 



two 

flattened 

glandular 

cushions 



4 small 
lobes 



a. one 


disc-like 


b. simple 


with a 


falcate 


dorsal 




gap 



palps barred 



bores in 

calcareous 

materials 



nuchal 

tentacle 

present 



Day 

(1961: 1967) 



Woodwick 
(1963b) 



Hartman 
(1936: 1969) 



Okuda (1937) 



•to 



lillll ll\ SOT I III RX CAl II ORNIA ACADEMY Of SCIENCES 



VOLUME 70 



rior end. Anteriorly they arc longest on setigers 
2. 3, and 4; they are shorter posterior to setiger 5 
through the first one-third of the body and then are 
more elongate on succeeding segments with another 
gradual decrease in size near the pygidium. 

The pygidium consists of a simple collar with a 
ventral incision or notch and further subdivision 
into several weakly developed lobes (Fig. 3i). 

Remarks: The species is closely related to B. 
tricuspa (Hartman) and B. pseudonatrix Day. It is 
separated from those species and others of the genus 
by the characteristics listed in Table I. Fauvel 
(1916) reported Polydora poly bronchia from the 
Falkland Island. His descriptions and figures are, 
however, clearly those of B. chilensis. Two figures 
show the anterior end with a nuchal tentacle and 
another figure shows the characteristic spines of 
setiger 5. 

Ecology and distribution: The species was col- 
lected both intertidally and subtidally along the 
coast of Chile. In the intertidal it occurred both in 
protected and exposed rocky habitats. 



KEY TO THE KNOWN SPECIES OF 
BOCCARDIA CARAZZI 

la Heavy spines of setiger 5 of one 

type, simple, falcate 2 

lb Heavy spines of setiger 5 of two 

types, first type simple, falcate; 

second type highly modified 5 

2a Nuchal tentacle present B. sp. 

2b Nuchal tentacle absent 3 

3a Recurved posterior spines present; 

pygidium with lappets or cirri 4 

3b Recurved posterior spines absent; 

pygidium saucer-shaped B. truncata 

4a Pygidium with 2 broad ventral 

lappets, each with a short process . . B. hamata 
4b Pygidium with 2 long anal cirri ... 5. ligerica 
5a Second type of heavy spine of 

setiger 5 with dense bristles on the top .... 8 
5b Second type of heavy spine of 

setiger 5 without dense bristles 6 

6a Heavy spines of setiger 5 — falcate and 

tridentate; prostomium rounded . . B. tricuspa 
6b Heavy spines of setiger 5 — falcate 

and with a distal concavity from which a 

cone or tooth arises; prostomium incised . . 7 
7a Nuchal tentacle present; 

hooded hooks number up to 16 

per neuropodium B. chilensis 

7b Nuchal tentacle absent; 

hooded hooks number only 6 

per neuropodium B. pseudonatrix 



8a Prostomium rounded 9 

8b Prostomium incised II 

9a Posterior acicular spines present; 
bristle-topped spines of setiger 5 with 
an accessory tooth; notosetae absent from 
setiger I B. berkeleyorum 

9b Posterior acicular spines absent; 
no tooth on bristle-topped spines 
of setiger 5; notosetae present on 
setiger I 10 

10a Notosetae of setiger I long and in 

a fan-shaped fascicle B. columbiana 

10b Notosetae of setiger 1 short . . . B. proboscidea 
1 la Hooded hooks bidentate in anterior 

setigers, unidentate in posterior 

setigers; pygidium with 2 ventral 

lappets B. basilaria 

1 lb Hooded hooks bidentate throughout; 

pygidium otherwise 12 

12a Large forwardly directed lateral 

pouches on the ventral side of 

setigers 1-4 B. perata 

12b Lateral pouches absent 13 

13a Bristle-topped setae of setiger 5 

with 2 heavy smooth bosses; notosetae 

present on setiger 1 B. natrix 

13b Bristle-topped setae of setiger 5 

with a conical tip with the bristles 

continued basally below the thickest 

region; notosetae absent on setiger 1 

B. polybranchia 

DISCUSSION 

As discussed by Woodwick (1964: 156) "The 
generic breakdown of the polydorid forms has 
been unstable since the late 1800's and at the 
present time no one arrangement is accepted by 
all workers." According to Woodwick's scheme 
the polydorids may be divided into four genera: 
Polydora sensu stricto, Pseudopolydora, Tripoly- 
dora and Boccardia. 

Fourteen species of the genus Boccardia are 
here considered valid. Their characteristics are 
compared in Table I and the species are delineated 
in the key. Boccardia sp. is an unnamed Japanese 
species [ = B. redeki sensu Okuda, 1937]. The 
14 species may be conveniently arranged into two 
natural groups based primarily on the setae of 
setiger 5. The groups are as follows: 

I. Superior dorsal fascicle present on setiger 5; 
Spines of setiger 5 of one kind, simple and falcate. 

B. hamata B. truncata 

B. ligerica B. sp. [ = redeki sensu Okuda, 1937] 



1971 



Nl: W SPEC I IS OF BOt ( A Hill A 



■II 



II. Superior dorsal fascicle absent on setiger 5; 
Spines of setiger 5 of two kinds. 

a. One spine simple falcate and second with 
one or more projecting eusps. 

II. chilensis H. tricuspa 

II. pseudonatrix 

b. One spine simple falcate and second bristlo- 
topped. 

II. basilaria li. perata 

II. berkeleyorum II. polybranchia 
B. colunibiana li. proboscidea 
B. natrix 
Two species of Group I (ligerica and hamata) 
have recurved posterior spines, while one species 
in Group II b (berkeleyorum) has acicular posterior 
spines. 

Several species of the polydorid complex possess 
posterior notopodial spines. In some cases the 
species may be closely related but in others not, 
and their specialization in structure then suggests 
convergent evolution. Polydora hoplura Claparede 
and P. colonia Moore have recurved posterior 
spines similar to those of Boccardia ligerica and 
hamala. however the adult, and quite importantly, 
the larval morphologies are different in the four 
species. Posterior spines are such important diag- 
nostic characteristics it would be helpful to learn 
more about their relation to the ecology of the 
respective species. Other polydorids known to have 
posterior spines include; Polydora caulleryi Mesnil, 
P. quadrilobala Jacobi, P. armata Langerhans, P. 
cardalia Berkeley, P. caeca Oersted, Pseudopoly- 
dora corallicola Woodwick and Tripolydora spinosa 
Woodwick. 

Literature Cited 

Augener, H. 1939. Beitrag zur polychaetenfauna der 
Ostsee. Kieler Meeresforschungen, 3: 133-147. 

Berkeley. E. 1927. Polychaetous annelids of the Nanai- 
mo District. 3. Leodicidae to Spionidae. Contrib. 
Can. Biol, n.s., 3: 405-422. 

Blake. J. A. 1966. On Boccardia hamata (Webster), new 
combination (Polychaeta. Spionidae). Bull. So. 
Calif. Acad. Sci.. 65: 176-184. 

Carazzi, D. 1893. Revisione del genere Polydora Bosc, 
e cenni su due species che vivono sulle ostriche. 
Mitteilungen aus der Zoologischen Station zu 
Neapel, 11:4-45. 

Day, J. H. 1955. The polychaeta of South Africa. 
Part 3: Sedentary species from Cape shores and 
estuaries. J. Linnean Soc. London (Zool.), 42: 
407-452. 

. 1961. The polychaete fauna of South Africa. 

Part 6: Sedentary species dredged off Cape coasts 
with a few new records from the shore. J. Linnean 
Soc. London (Zool.). 44: 463-560. 



. 1967. a monograph or thi Po 

Southern Afrii a Pari 2 Sedentaria Brill h 
(Nat. Hist.) London, Pul -7x. 

Fauvel, P. 1916 \nn6lidca polycheb di Mi Falkland 
recucillies par M. Rupert Vallenlin I qn - 
1910). Archives de Zoologli Experimental! el 

t.rnrrale. 55: 417-482. 

. 1927. Polychetes sddentaires. Addenda aux 

Errantes, Archianne'lides, Myzoslomairei I num- 
de France, 16: 1-494. 

Ferronniere, G. 1898. Contribution a I'ctudc dc la faune 
de la Loire-inferieure (Polygordieru, Spionidieiu, 
Nemertien). Bulletin de la Sociiti dei Sciei 
Naturelles de I'Ouesl de In France, 8: If) I I l ' 

Fcwkes, J. N. 1889. New Invcrlcbrata from the 
of California, Bull. Essex Inst., 21: 99-146. 

Friedrich. H. 1938. Polyt haeta. Die Ticrwelt der Nord- 
und Ostsee. Gcgrundet von G. Grimpc und E. 
Wagler. Leipzig, 32 (Vl.b.): 1-201. 

Hartman. O. 1936. New species of Spionidae (Annelida 
Polychaeta) from the coast of California. Univ. 
Calif. I'ubl. Zool., 41: 45-52. 

. 1939. The polychaetous annelids collected hy 

the Presidential Cruise of 1938. Smiths. Misc. 
Coll.. 98: 1-22. 

. 1940. Boccardia proboscidea, a new species 

of spionid worm from California. J. Wash. Acad. 
Sci.. 30: 382-287. 

1941. Some contributions to the biology and 



life history of Spionidae from California. Allan 
Hancock Pac. Exped.. 7: 289-324. 

. 1948. The marine annelids erected by Kinberg 

with notes on some other types in the Swedish 
State Museum. Arkivfur Zoologie. 42A: 1-137. 

. 1961. Polychaetous annelids from California. 

Allan Hancock Pac. Exped.. 25: 1-226. 

. 1969. Atlas of the sedentariate polychaetous 



annelids from California. Allan Hancock Foun- 
dation. University of Southern California. Los 
Angeles. 812 pp. 

Haswell. W. A. 1885. On a destructive parasite of the 
rock-oyster (Polydora ciliata and P. polybranchia 
n. sp.). Proc. Linnean Soc. New South Hales. 
10: 272-275. 

Horst. R. 1920. Polychaete anneliiden uit net Alkmaar- 
der Meer door Dr. R. Horst. Zoologische Vlede- 
deelingen, Leiden. 5: 1 10-1 1 1. 

Hempel. C. 1957a. Zur Okologie einiger spioniden 
(Polychaeta Sedentaria) der deutschen Kusten. 
Kieler Meeresforschungen. 13: 275-288. 

. 1957b. Uber den Rohrenbau und die Nahrun- 

gaufnahme einiger Spioniden (Polychaeta Seden- 
taria) de deutschen kusten. Helgolander H'issen- 
schaftiiche Meeresuntersuchungcn. 6: 100- 1??. 



42 



HI III UN SOUTHERN CALIFORNIA ACADEMY OE SCIENCES 



VOLUME 70 



Imajima. M. and O. Harlman. 1464. The polychaetOUS 
annelids of Japan. Allan Hancock Foundation, 
Occasional Paper No. 26: 1-452. 

Khlebovitsch. V. V. 1954. [Species olpolychaete worms 
from the Kuril Islands which are new or recorded 
for the first time in the USSR fauna.) Zoologiche- 
skii Zhurnal, 38: 167-181. [In Russian]. 

Mesnil. F. 1896. Etudes de morphologie externe chez 
les annelides. Les Spionidiens des cotes de la 
Manche. Bulletin Scientifique de la France et de la 
Belgique, 29: 1 10-287. 

Monro. C. C. A. 1938. On a small collection of Poly- 
chaeta from Uruguay. Ann. Mag. Nat. Hist. ser. 
II. vol. 2: 3 11-3 14. 

Okuda. S. 1937. Spioniform polychaetes from Japan. 
J. Fac. 5c/. Hokkaido Imp. Univ. ser. 6, vol. 5: 

217-254. 

Rullier. F. 1960. Morphologie et developpement du 
Spionidae (Annelide Polychete) Polydora (Boc- 
carclia) redeki Horst. Cahiers de Biologic Marine. 
1:231-244. 

Rioja. E. 1962. Estudios Anelidologicos 26. Algunes 
Anelidos poliquetos de las costas del Pacifico de 



Mexico. Anales del Instituto </<■ Biologia Mexico, 
32: 131-229. 
Sodcrstrom, A. 1920. Studien iiber die Polychaeten- 
familie Spionidae. Dissertation. Uppsala. Almquist 
und Wicksells. 286 pp. 

Treadwell, A. L. 1914. Polychaetous annelids of the 
Pacific coast in the collection of the zoological 
museum of the University of California. Univ. 
Calif. Publ. Zool., 13: 175-234. 

Webster. H. E. 1879. Annelida Chaetopoda of the 
Virginian coast. Trans. Albany Inst., N. Y., 9: 202- 
269. 

Woodwick. K. H. 1963a. Comparison of Boccardla 
coliimhiana Berkeley and Boccardia proboscidea 
Hartman (Annelida. Polychaeta). Bull. So. Calif. 
Acad. Sci., 62: 132-139. 

. 1963b. Taxonomic revision of two polydorid 

species (Annelida, Polychaeta, Spionidae). Proc. 
Biol. Soc. Wash.. 76: 209-216. 

. 1964. Polydora and related genera (Annelida, 

Polychaetal from Eniwetok, Majuro, and Bikini 
Atolls. Marshall Islands. Pacific Sci. 18: 146-159. 

Accepted for publication September 25. 1970. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(1): 42-45, 1971 

PARASITIC MITES OF SURINAM. VIII. A NEW GENUS AND SPECIES OF CHIGGER, 

FAURANIUS ATECMARTUS, AND ADDITIONAL RECORDS OF SPECIES 

(ACARINA: TROMBICULIDAE) 1 

James M. Brennan 2 and F. Lukoschus 3 



Abstract: Twenty-six species of chiggers are now known from Surinam. 
Nineteen are recorded here. Fauranius new genus is described for F. atec- 
martus, new species off Philander opossum and F. myoproctae (Fauran, 1960), 
new combination. 



This paper augments a recent report (Brennan, 1970) 
in which 1 1 species of chiggers are recorded from 
Surinam. Fifteen additional species, one new, are 
recorded here. All material was collected by the 
junior author. 

Two specimens of each species, as available, will 
be deposited in Rijksmuseum van Natuurlijke 
Historic Leiden. The Netherlands, and National 
Collection of Surinam, Centraal Laboratorium. 
Paramaribo. 

Fauranius, new genus 

Type species: Fauranius atecmartus. new species. 
Neotropical trombiculine larvae. Leg segmen- 



tation 6-6-6 (unique in subfamily Trombiculinae); 
2 genualae I, sub- and parasubterminala, micro- 
tarsala I proximad of tarsala I, no genuala II and 
III, a tibiala III, no mastisetae. Scutum wider than 
long, the posterolateral setae extrascutal, sensillae 
probably expanded (as suggested by remaining 
stubs). Apparently, eyes may be present or absent. 



■Supported in part by Grant W83-1 from The Netherlands 

Foundation for the Advancement of Tropical Research. 

-U.S. Department of Health, Education, and Welfare, Public 

Health Service, National Institutes of Health, National Institute 

of Allergy and Infectious Diseases, Rocky Mountain Laboratory, 

Hamilton, Montana 59840. 

3 Zoologisch Laboratorium, Katholieke Universiteit. Nijmegen, 

The Netherlands. 



1V7I 



NEW SPECIES OF ( HIGGl l< 






Eheliceral blade with tricuspid cap. Galeal seta 
nude. Palpal tibial claw Irilurealc, palpal larsus 
jwith 5 branched setae and a tarsala. 

Distinguished from Pseudoschoengastia by leg 
segmentation 6-6-6 and absence of seiae between 
coxae II and III. Referred species: Pseudoschoen- 
gastia myoproctae Fauran, I960. 

Fauranius is named for Dr. Piene Fauran, 
Pasteur Institute of French Guiana, who has made 
significant contributions to the knowledge of neo- 
tropical chiggers. 

Fauranius atecmartus, new species 

Type data: Hololype and 2 paratypes, RML 
55406, from ears of Philander opossum, Coronie, 
Surinam, 28 November 1969, F. Lukoschus. 

Holotype to be deposited in Rijksmuseum van 
Natuurlijke Historic Leiden; paratypes in the 
Rocky Mountain Laboratory. 

Diagnosis: Leg segmentation 6-6-6. Extrascutal 
posterolateral setae; less than 30 dorsal setae; no 
genuala II and III, no mastisetae, microtarsala I 
proximad of tarsala I. Distinguished from F. myo- 
proctae by differently shaped scutum, PL>AM. 
smaller leg index because of shorter leg II, smaller 
tarsalae I and II. and microtarsala I much closer to 
tarsala I. 

Body: A small chigger, broad-ellipsoidal when 
engorged. Eyes not discernible. Anus at sixth row 
of ventral setae. Length and width of holotype. 
nearly engorged, 335 by 260,u . 

Gnathosoma: Sparsely punctate. Cheliceral blades 
with small tricuspid cap. Galeal seta nude. Palpal 
setae sparsely branched. B/B/BNB, tarsus 5B and 
a tarsala, tibial claw trifurcate. 

Scutum: As figured, lightly punctate; sensillae 
broken off in all specimens, but in one the remain- 
ing stubs are suggestive of expanded heads; extra- 
scutal PL>AM>AL. Scutal measurements of 
holotype; AW 47, SB 32. ASB IS. PSB 17, AM 27, 
AL 18, PL 34. 

Legs: Segmentation 6-6-6. Lightly punctate, 
specialized setae as figured: 2 genualae I. sub- and 
parasubterminala. no genuala II and III. a tibiala 
III. no mastisetae, microtarsala I proximad of 
tarsala I. Nonspecialized setae sparsely branched. 
Coxal III seta slightly anterior of center. 

Body setae: Dorsal setae; humerals 34^, dorsals 
25 to 3 l,u . arranged 2-6-6-6-4-4 (holotype). 2-6-6-6- 
4-2 (a paratype). Ventral setae 2-2 plus 36; postanals, 
paranals, and preanals of last 2 rows similar in form 
to dorsals. 

Remarks: Fauran (1960) described Pseudoscho- 
engastia myoproctae in detail from a single speci- 
men attached to teat of Myoprocta acouchy, May- 




Figure I . Fauranius atecmartus. Scutum and specialized 
setae of legs with measurements in microns. 



enne-Mana, French Guiana, April 1960. He noted 
the unique 6-6-6 leg segmentation. Geest and 
Loomis (1968) saw the holotype and retained the 
species in Pseudoschoengastia in their "Acci 
Group". 

Although we have not seen the holotype. it is 
clear from the description that F. myoproctae and 
F. atecmartus are closely related. Both are small, 
with corresponding small structures. Certain char- 
acteristics of F. myoproctae and F. atecmartus, 
respectively, are: leg index 570 and 519: dorsal 
formula 2-6-6-6-6-2-2 and 2-6-6-6-4-4: length of 
humeral setae 35,u and 34/* : length of other dorsals 
20 to 29,u and 25 to 3L*; eyes distinct 2/2 and 
eyes not discernible; lengths of tarsalae I and II 
18 tt . 17 u and 14u. 12u. 



O/i 



'/* 



/*• 



Crotiscus desdentatus tissoti Fauran. 1960 

Six specimens off Proechimys guyannensis, 
Lelydorp. 18 December 1969. 

Euschoengastia (s. 1.) desmodus (Brennan and 
Dal mat, 1960) 

Two larvae from ventral wing membrane ot 
Glossophaga soricina. Brownsweg. 9 February 
1970; 1 off Carollia perspicillala. Onverwacht. 5 
February 1970. 

This bat chigger and a new species from Venezu- 
ela are being referred to a new genus in a subsequent 
paper. 



44 



IU I I I IIS SOI I III RN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 



Eutrombicula alfreddugesi (Oudemans, 19 10) 

I i\l- larvae off Proechimys guyannensis, Lely- 
dorp, 18 December 1969; 3 off P. guyannensis, 
Uitkijk. 8 January 1970; 3 off 3 Didelphis marsupi- 
alis, Coronie, 29 November 1969 to 12 February 
1970; 1 off D. marsupialis, Brokobaka, 2 1 February 
1 970; 2 off Philander opossum, Coronie, 10 January 
1970; 6 off Rutins ran us frugivorus, Brokobaka, 
2 1 February 1 970; 4 off Nectomys squamipes 
melanius, Coronie, 12 February 1970. 

Eutrombicula batatas (Linnaeus, 1758) 

Six larvae from ears of 3 Rattus norvegicus, 
Paramaribo. 13 and 15 January 1970; 2 from ears 
of Herpestes auropunctatus, Meerzorg, 3 March 
1970; 1 from ears of Tamandua tetradactyla, Lely- 
dorp. 17 December 1969; 1 from scrotum of 
Eptesicus melanopterus, Meerzorg, 3 March 1970. 

This Linnaean species, the oldest recorded 
trombiculid mite, the "sweet potato chigger," is 
recorded again from the country of its discovery. 
For details concerning the history, biology and 
systematics of Eutrombicula batatas see Michener 
( 1 946). Jenkins ( 1 949). and Fuller ( 1 952). 

Eutrombicula goeldii (Oudemans, 1910) 

Three specimens off Proechimys guyannensis, 
Lelydorp. 18 December 1969; 2 off Didelphis 
marsupialis, Coronie, 12 February 1970; 2 off 
Epicrates cenchris (rainbow boa), Brokopondo, 2 
February 1970; 3 off Psarocolius decumanus, Lely- 
dorp, 23 January 1970. 

Eutrombicula tinami (Oudemans, 1910) 

Two larvae off Didelphis marsupialis, Brokobaka, 
21 February 1970. 

This form has not been recorded since it was 
described off Tinamou, Brazil. It is very close to, 
and may indeed prove to be Eutrombicula goeldii 
with branched palpogenual and palpoventrotibial 
setae. 

An additional record is 5 specimens off Proe- 
chimys semispinosus, Galeta Point, Panama, 23 
February 1950. 

Microtrombicula fragibarba 
(Brennan and Jones, 1960) 

Eight larvae from ears of Didelphis marsupialis, 
Brokobaka, 21 February 1970; 7 off 2 D. marsupia- 
lis, Coronie, 29 November and 4 December 1969 
7 off D. marsupialis, Lelydorp, 12 January 1970 
1 off D. marsupialis, Paramaribo. 3 1 January 1970 
1 off Philander opossum, Coronie, 10 January 1970 



Microtrombicula nr. tragulata 
(Brennan and Jones, 1961) 

Six larvae off Didelphis marsupialis, Brokobaka, 
21 February 1970. 

Nasicola annereauxi Brennan and Yunker, 1969 

Twenty-six larvae from nasal passages of Noctilio 
labialis, Lelydorp, 23 January 1970; 43 from 3 N. 
labialis, Meerzorg, 2 March 1970. 

This intranasal chigger of bats described from 
Phyllostomus hastatus, Venezuela is also known 
from P. hastatus and N. labialis, Colombia. 

Odontacarus tubercularis (Brennan, 1952) 

Seven larvae off Proechimys guyannensis, Uit- 
kijk, 8 January 1970; 1 off Philander opossum, 
Coronie, 28 November 1969; 2 off P. opossum, 
Coronie, 12 February 1970; 1 off Nectomys 
squamipes melanius, Coronie, 12 February 1970. 

Parasecia aitkeni (Brennan and Jones, 1960) 

Four specimens off Philander opossum, Coronie, 
10 January 1970. 

Parasecia manueli (Brennan and Jones, 1960) 

One larva off Proechimys guyannensis, Uitkijk, 
8 January 1970; 2 off P. guyannensis, Lelydorp, 18 
December 1969; 12 off 4 Philander opossum, 
Coronie, 28 November 1969 to 12 January 1970; 
4 off 3 Didelphis marsupialis, 29 November 1 969 
to 12 February 1970. 

Parasecia valida (Brennan. 1969) 

Two larvae off Proechimys guyannensis, Lely- 
dorp, 18 December 1969. 

Perissopalla barticonycteris Brennan, 1969 

Five larvae off 2 Carollia perspicillata, Brown- 
sweg, 10 February 1970. 

Perissopalla ipeani Brennan, 1969 

Six larvae off 3 Carollia perspicillata, Onver- 
wacht, 5 February 1970. 

Quadreseta flochi (Brennan and Jones, 1960) 

Two larvae off Proechimys guyannensis, Uitkijk, 
8 January 1970. 

Speleocola secunda Brennan and Jones, 1960 

Three larvae off Carollia perspicillata, Zandery, 
7 January 1970; 1 from ventral wing membrane of 
Saccopteryx bilineata, Lelydorp, 26 February 1970. 



1971 



NEW SPECIES OF < HIGOl R 






Tecomatlana (llooperella) vesperuginis 
(Brennan and Jones, I960) 

Thirteen larvae off 2 Car o Ilia perspicillata, 
Zandcry, 7 January and 1 February 1970; 2 off 
C. perspicillata, Brownsweg. 10 February 1970; 2 
from wing membrane of Saccopteryx bilineata, 
Lelydorp, 26 February 1970; 6 from ventral wing 
membrane of 3 Glossophagu soricina. Brokopondo. 
2 February 1970. 

The following ehiggers of Surinam, not listed 
here, were reeorded recently by Brennan ( 1970). 

Arisocerus amapensis Brennan, 1970 

Boshkerria punctata (Boshell and Kerr, 1942) 

Colicus icomi Brennan, 1 970 

Colic us oblonga (Fauran, 1959) 

Eutrombicula alfreddugesi (Oudemans, 1910), 
tropica form of Ewing, 1925 

Pseudoschoengastia tricosa (Brennan and Jones, 
1961) 

Tecomatlana (Hooperella) saccopteryx (Brennan 
and Jones, 1960) 

Trombicula (s. 1 .) palmigera Fauran, 1960 

Literature Cited 

Brennan, J. M. 1970. A small collection of ehiggers 



from Sin in. mi i a. ..i in. i i rombi utidae) tu,;. 
Calif. Acad. Set. 69 

Fauran, P. I960. Description de quatri 

especcs et d*unc notivcllc sous cspccc dc Irombi 
eulides ilc Guyanc francai* \rch in i r, 
Cuyane franq. et Intnl. Nm r 9, 20 p. 

Fuller, ll. s. 1952. I lie mite larvae ol the family 
Trombiculidae in the Oudemam collei tio I 
onomy and medical importance Zoologiichc 
Verhandellngen, No. ix. E.J. Brill, I eiden 261 p 

Gecst, J. C. and R. B. Loomis. 1968. < higgen ol the 
genus Pseudoschoengastia (Acarina; Trombicu- 
lidae) from Costa Rica. Los An Co Hut Sat 
Hist.. Contrib. Sci. No. 150: 1-49. 

Jenkins, D. W. 1949. Trombiculid mites affecting man. 
IV. Revision of Eulrombh ula in the American 
hemisphere. Ann. Em. Soc. Amer. 42: 2X9-3 IX. 

Michener, C. D. 1946. Observations on the habits and 
life history of a chigger mite. Eutrombk ulu batatas 
(Acarina: Trombiculinac). Ann. Ent. Soc. Amer. 
39: 101-118. 

Accepted for publication October 23. 1970. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(1): 45-49, 1971 

VARIATION AND GEOGRAPHIC DISTRIBUTION IN SOME ARGENTINE AND 
CHILEAN OSMYLIDAE, WITH A NEW SPECIES OF KEMPYNUS (NEUROPTERA). 

Phillip A. Adams 1 

Abstract: A similar color variation, which consists in longitudinally 
streaked wings, occurs in Kempynus falcatus Navas and Phymatosmylus 
caprorum Adams. These species are sympatric with the newly described 
Kempynus crenatus on the western slope of the Andes between 34° 30' S and 
40° S. but further south in Chiloe and Aysen, K. falcatus occurs on the coast. 



The material discussed below was not available for 
inclusion in my recent paper on South American 
Osmylidae (Adams, 1969). Kempynus is of par- 
ticular interest because it includes species from 
Australia-New Zealand and South American (Kim- 
mins, 1940). I am grateful to Ellis MacLeod of the 
University of Illinois, Urbana. for the loan of 
specimens and for helpful comments. 

Kempynus crenatus, new species 
Figures 1 ; 2A, C, E-H 



Description: Head pale with the following fuscous 
marks: dark border of antennal sockets confluent 
with large frontoclypeal triangle which has apex 
at anterior clypeal margin, spots anterior to anterior 
tentorial pits, borders of ocelli and vertex scars. 
Mouthparts fuscous, antennae pale. Pronotum pale, 
dark dotted, four black spots on transverse furrow. 
Mesonotum pale, dark-punctate, prescutum with 



'Department of Biology. California State College. FuIIenon. 
California 92631 



4b 



III I I 1 I IX SOUTHERN CALIFORNIA ACADEMY Ol SCIENCES 



VOLUME 70 







Figure 1 . Kempynus crenatus, wing venation. 



four dark marks, black triangle on prescutal- 
scutal suture, scutum and scutellum black laterally. 
Pleurae irregularly variegated, coxae fuscous, fore 
and mid-femora with pair of subapical central dark 
marks and dark dorsoapical spot, hind femora 
paler. Tibiae pale, dark bands at base, middle and 
apex. Tarsi brownish, apical tarsomere dark. Female 
fore coxa with an anterolateral field of numerous 
short pedestalled setae. Wing venation as in Figure 
1. MP2 of forewing dichotomously branched basad 
of union of CuA and CuP, posterior fork pectinate. 
Malculations brown. 

Abdomen fuscous at tip; setal bases infuscate. 
External male abdominal apex much as in K. incisus 
(Kimmins, 1940). Gonocoxites (Fig. 2A) resemble 
those of K.falcatus (Fig. 2B), but are more elongate. 
Hypandrium internum (Fig. 2C) more elongate 
than in K. falcatus. Female abdominal apex much 
as in K.falcatus. eighth sternite (Fig. 2F, G) setose 
anterolaterally, posterior lobes shorter and more 
rounded than in falcatus. Spermathecae slightly 
bent, constricted in middle, basal portion spherical: 
in one specimen, the spermathecae are ovoid (Fig. 
2E). Colleterial gland reservoir (Fig. 2H, cgr) and 



bulb slightly longer than in falcatus, and bursal 
gland duct (bgd) less tortuous. 

Measurements (mm). Forewing length, 9, 28.0, 
28.5, 29.0; 3 26.0. Antenna length. 9. 7.0, 8.0, 8.3; 
S 7.5. 

Holotype. Chile, Prov. Nuble, Cordilleras Chil- 
ian, Las Trancas, 12 km E. Recinto, 1-10 Dec. 
1964, 9, leg. L. E. Pena, Museum of Comparative 
Zoology, Harvard [ca. 800 mt., 71° 37' W, 36° 51' 
S]. 

Allotype. Argentina, Prov. Neuquen, Pucara, 
Parque Nacional Lanin, Jan. 1951, 8, leg. S. S. 
Schajovskoy, Museum of Comparative Zoology, 
Harvard (ex coll. E. G. MacLeod). 

Paratypes. I 9, same data as holotype, E. G. 
MacLeod collection. Argentina, Prov. Neuquen, 
Pucara, 30 Nov. 1959, 9, purch. ex F. H. Walz, P. 
Adams collection. 

Remarks. This species differs from falcatus in 
its oval wing shape, and shape of spermathecae. In 
K. longipennis, the wings are similarly shaped, but 
the wingtips are somewhat more acute, MP2 forks 
pectinately, and the marginal pale areas of K. 
crenatus are absent. In all the other species of 



NhW SPECIES <>l LACEWING 




Figure 2. Reproductive structures. A, C, E-H, Kempynus crenatus. B, D. K.falcalus. A. B. genital 
armatures of males, lateral view. C, D, hypandrium internum. E. spermathecae. F. eighth abdomi- 
nal sternite of female. lateral view. G, same, ventral view. H, female reproductive system viewed 
from the left side, abdominal apex downward. 



3«* V 





Figure 3. Striped wing pattern: A. Phymatosmylus caprorum: B. Kempynus falcatus. 



4S 



HI I II 1 1\ SOI I III R\ ( .1/ II ORN1A ACADEMY OE SCIENCES 



VOEUME70 



Kempynus the apical projections of the female 
eighth sternite are more slender and more separated 
basal I \ . 

Kempynus falcatus Navas 
Figures 2B. D;3B 

Kempynus falcatus Navas 1912. 1928. 1930, 
1936. Kimmins. 1949. Type: 9. Chile. Mulchen 
[Bio Bio, 37°44' S, 72° 15' W] , Jan. 1902. leg. H.J. 
Elwes, Brit. Mus. (Nat. Hist.), not seen. 

Kalosmylus falcatus, Kriiger, 1913, 1914. 

Remarks. The male genitalia have not previously 
been described. As Kimmins (1940) noted, these 
are similar in most species of Kempyninae, and 
reference to Fig. 2B demonstrates that K. falcatus 
constitutes no exception. Most of the apparent 
differences from K. crenatus are due to positioning. 
However, the hypandrium internum (Fig. 2D) is 
broader, with the winglike lobes fused on the mid- 
line, while they are separate to the base in K. 
crenatus. 

Distribution. The following specimens, unless 



34< 



46' 




• K. FALCATUS o K. CRENATUS 

+ P. CAPRORUM 

Figure 4. Known distribution of South American 
Kempyninae and Stenosmylinae. 



otherwise noted, are in my collection and were 
collected by L. E. Pefia. Chile: Curico, Buchen, 
Mar. 1956, 9, ("farm in preandean foothills"); 
Curico, La Jaula (Los Queries), Andes, 14-18 Feb. 
1965. <J ; Linares. Fundo Malcho ("in the high 
mountains") Dec. 1952, 9: Malleco Prov., Termas 
de Tolguaca, 30 km NE Curacautin, 20 Jan. 1959, 
$ : Cautin 30 km N Villarica, 1-30 Jan. 1965, 6 S 
4 9; Chiloe, Dalcahue, Feb. 1954, S ; Aysen, Pto. 
Cisnes. Feb. 1961, 9; Aysen, Rio Maniguales [vie. 
Puerto Aisen] 27 Jan. 1961. \S, 19. Argentina, 
Neuquen, Pucara, Parque Nacional Lani'n: 3 1 Nov. 
1959. 9. purch. ex. F. H. Walz; Feb. 1951, 9, Feb. 
1952, S , leg. S. S. Schajovskoy, E. G. MacLeod col- 
lection. 

Color variation. In my collection are single 
specimens of Phymatosmylus caprorum Adams 
(Fig. 3A 9, from Las Trancas, Cord. Chilian, Chile, 
1-10 Jan. 1964, L. E. Pefia leg.) and Kempynus 
falcatus Navas (Fig. 3B, 9, 30 km N. Villarica, 
Prov. Cautin, Chile, 1-30 Jan. 1965, L. E. Pefia leg.) 
which have a dark stripe on each wing instead of 
the usual mottling. While similar color patterns 
are common in other Neuroptera (e.g. the poly- 
stoechotids Platystoechotes and Fontecilla, and 
many Myrmeleontidae) they do not ordinarily occui 
in related Osmylidae ( Kimmins, 1 940). The capacity 
to develop similar aberrant color patterns may 
suggest a closer relationship between these osmylids 
than indicated by their present taxonomic position. 
They are now placed in different subfamilies (Phy- 
matosmylus in Stenosmylinae, Kempynus in Kem- 
pyninae) although Phymatosmylus is regarded as 
transitional toward Kempyninae (Adams, 1969). 

Geographic distribution (Fig. 4). Kempynus 
falcatus, K. crenatus, and Phymatosmylus caprorum 
are sympatric along the western slope of the Andes, 
from about 34° 30' S to 40° S, extending into 
Argentina in the lake region, where several passes 
exist at 1200m elevation. All three species have 
been taken are at Pucara, Lanin National Park, 
Argentina. Further south, in Chiloe and Aysen, 
the much commoner K. falcatus occurs on the 
coast, an elevational shift probably related to the 
cooler climate. Also plotted are localities for K. 
falcatus from Kimmins (1940) and from Navas 
(1928, 1930). 

None of these species appears to be sympatric 
with Isostenosmylus, which has several species in 
Ecuador, Peru. Bolivia, and Brazil, or with Pary- 
phosmylus, with one species in Ecuador. 

Seasonal abundance. Combined data from 50 
specimens of the three species indicate greatest 
frequency of capture near the time of the December 
solstice: November — 7. December — 21, January 
— 15, February — 6, March — 1. 



1971 



NEW SPECIES <>l I ACEWINC 



t; 



AKUKI-VIMIONS 



Bed — bursal gland duel, cgr - colleterial gland 
reservoir, cop b — copulatory bursa, CuA — 
cubitus anterior, CuP — cubitus posterior, fc — 
fertilization canal, gcx — gonocoxite, go — geni- 
tal opening, gs — gonarcus, hyi — hypanclrium 
internum, MP — media posterior, mu- ■mediun- 
cus, ovd — oviduct, sp — spermatheca. 



Literature Cited 

Adams, P. A. 1969. A new genus and species of 
Osmylidae (Neuroptera) from Chile and Argen- 
tina, with a discussion of planipcnnian genitalic 
homologies. Postilla, 141: 1-11, 

Kimmins, D. E. 1940. A revision of the osmylid 
subfamilies Stenosmylinae and KalosmylinaelNeu- 



roptera). Novltati r Zoologi pi 

1-8. 

KrUger, L, 1913, 1914. Beitr&gc zu • ini i Monographic 
der Neuropteren-Pamilii ttttnei 

entomologische Zeliun 

Navas, L. 1912. Insectus ncuroplcros ntl( 

conocidos. Memorial de la Krai AcadcmL 
Cienciasy Aries de Barcelona, io i 

. 1928. Insectoa neotropico 

Chilena Historia Natural, J2: 1')'. I2K. 

. 1930. Algunos insectos de C hile, 4a scric. 



Revlsta Chilena Historia Natural, }4 



— . 1936. [nsectos ncur6pteroa dc < hile, poco 
conocidos. Revista Chilian Historia Natural, 

40: 179-181. 



Accepted for publication September 4. 1970. 



50 BULLETIN SOI I HERN CALIFORN1 I ACADEMY OF SCIENCES 

RESEARCH NOTES 



VOLUME 70 






Bull. So. Calif. Acad. Sci. 70(1):50-51, 1971 

DISTRIBUTION OF SOME SOUTHERN 
< \l IFORNIA KANGAROO RATS 

The ranges illustrated by Grinncll (1922) and Hall 
and kelson (1959) for southern California kangaroo 
rats were drawn by encircling known records of 
occurrence in the latter ease without regard to 
physiography or habitat. While collecting speci- 
mens of Dipodomys agilis, D. panamintinus, and 
D. microps for a recent study (Csuti. 1969), it be- 
came apparent that the published ranges of these 
species in southern California were inaccurate. 
Conventional museum study skins were prepared 
from all specimens examined and are deposited in 
the collection of the Natural History Museum of 
Los Angeles County (LACM) unless otherwise 
noted. Plant communities referred to are those 
described by Munz and Keck (1959). 

Dipodomys panamintinus mohavensis (Grinnell). 
— Eleven specimens were taken from Soledad 
Canyon in the upper Santa Clara River valley (Los 
Angeles County). These specimens, captured in 
Pinyon-Juniper Woodland habitat, are from an 
area formerly indicated (Hall and Kelson, 1959) to 
be within the exclusive range of D. agilis agilis. A 
specimen of D. panamintinus (San Fernando Valley 
State College Vertebrate Collection [SFVSC] 746) 
was taken from Rye Canyon, near Castaic Junction, 
in similar habitat farther down the Santa Clara 
River valley. It appears that the presence of Pinyon- 
Juniper Woodland habitat along this river valley 
has allowed the penetration of D. panamintinus 
westward into the San Gabriel Mountains from the 
Antelope Valley for a distance of at least 28 miles. 

Dipodomys microps microps (Merriam). — Two 
specimens (LACM 28342, SFVSC 861) were taken 
6 miles north of Lancaster, Los Angeles County, in 
Alkali Sink habitat. This locality extends the known 
range of this species 50 miles west of the isolated 
population previously reported from Victorville 
by Grinnell (1922). The habitat at the Lancaster 
locality is typical for this race in its northern range 
in Owens Valley, since it is "dry sandy ground, 
sparsely grown to desert shrubbery such as salt- 
bush" (Grinnell, 1933: 164). Dipodomys microps 
probably has a discontinuous distribution in the 
Antelope Valley composed of relict populations in 
those low-lying areas where the Alkali Sink habitat 
persists. 

Dipodomys agilis perplexus (Merriam). — 
Vaughan (1954) has taken 17 specimens of this 
kangaroo rat in the eastern San Gabriel Mountains. 



Hall and Kelson (1959) indicated a connection 
between the populations represented by these speci- 
mens and those previously known from the 
Tchachapi Mountains. Since D. a. agilis occurs at 
Elizabeth Lake (Hall and Kelson, 1959) on the 
interface of the San Gabriel Mountains and the 
Antelope Valley, any connection between the two 
D. a. perplexus populations must lie to the north of 
this locality (as illustrated by Hall and Kelson, 1 959) 
and is necessarily confined to a path across the 
floor of the Antelope Valley. Dipodomys agilis 
perplexus typically occupies Sagebrush Scrub habi- 
tat at higher elevations (Vaughan, 1954), and is 
nowhere reported from desert areas. The absence 
of any suitable habitat in the Antelope Valley, 
together with the absence of specimens from inter- 
vening areas, makes it appear unlikely that a con- 
nection exists between the populations in the 
Tehachapi and the eastern San Gabriel Mountains. 
The populations of D. a. perplexus in the eastern 
San Gabriel Mountains seem to be ecologically 
distinct from D. a. agilis, according to Vaughan 
(1954). These populations may represent an iso- 
lated segment of the currently named D. a. perplexus 
which was formerly connected with the Tehachapi 
Mountains population, but due to a change in 
habitat is now restricted to the higher elevations 
of the eastern San Gabriel Mountains. Alternatively, 
the specimens in question may represent a popula- 
tion derived from D. a. agilis which adapted to a 
habitat similar to that of D. a. perplexus of the 
Tehachapi Mountains, and which responded mor- 
phologically to this habitat in a manner parallel to 
that of D. a. perplexus. 

These distributional changes emphasize the 
importance of considering habitat in determining 
ranges of small mammals. The complex pattern of 
plant communities in southern California has 
resulted in a similarly complex pattern of faunal 
distribution. This requires more than the usual 
caution when drawing ranges for local animals. 

ACKNOWLEDGMENTS 

I wish to thank Drs. George F. Fisler and Andrew 
Starrett for their assistance in this research. Specimens 
SFVSC 746 and 861 were collected by T. S. Kelly and 
M. Weigand respectively. This paper is part of a thesis 
submitted in partial fulfillment of the requirements for 
the degree of Master of Science at San Fernando Valley 
State College. 

Literature Cited 

Csuti, B. A. 1969. Interrelationships of five species of 
kangaroo rats (genus Dipodomys) in southern 



1971 



RESEARCH NOTES 






California. Unpublished M.S. Ihesis, . San lernando 
Valley State College, Northridge, < 'alifornia, 75 pp. 

Brinnell, J, I y 2 2 . A geographical study of the kangaroo 
rats of California. Univ. Calif. Publ. Zool., 24: I- 

124. 

1933. Review of the Recent mammal fauna <>i 



California. Univ. Calif. Publ. Zool., 40: 71-234. 

Hall, E, K., and K. R. Kelson. 1959. The mammals oj 
North America. Ronald Press Co., New York, 
1083 pp. 

Munz, P. A., and D. D. Keck. 1959. A California Mora. 

Univ. California Press, Berkeley, K>xi p. ■, 

Vaughan, T. A. 1954. Mammals of the San Gabriel 
Mountains of California. Univ. Kansas Publ., 

Mas. Nat. I list.. 1: 513-582. 

Blair A. Csuti, Department of Biology, San Fer- 
nando Valley State College. Northridge. California 
91324 (present address: Museum of Vertebrate 
Zoology, University of California. Berkeley 94720). 

Accepted for publication October 23, 1970. 



Bull. So. Calif. Acad. Sci. 70(1): 51-52. 1971 

AN EVALUATION OF TWO METHODS OF 
MEASURING METACARPAL LENGTH IN 
ART1BEUS L1TURA TUS (OLFERS) 
(CHIROPTERA: PHYLLOSTOMATIDAE). 

Two alternate methods are currently employed in 
measuring metacarpal length of bats. An extensive 
literature perusal of studies on bats utilizing the 
measurement of metacarpal length revealed that 
the method of measuring is seldom stated. Com- 
parative use of the data in such publications is 
therefore possible only when one has personal 
knowledge of the investigator and the type of 
method employed by him in the particular study. 
The distance measured is from the metacarpal-first 
phalanx articulation to either 1) the proximal end 
of the respective metacarpal, or 2) the apex of the 
wrist joint formed by the summit of thescapholunar. 
The carpals are therefore excluded in the first (e.g., 
Starrett and de la Torre, 1964) and included in the 
second (e.g., Davis. 1970) of the two methods. In 
either method, the measurement is usually taken 
between two parallel planes oriented perpendicular 
to the proximodistal axis of the metacarpal. 

Prior to extensive use of either method in a syste- 
matic study of the Neotropical bat. Artibeus litura- 
tus (Olfers). an attempt was made to determine the 
greater desirability, if any, of either method. One 
series of 50 adult specimens ( 10 males. 40 females) 
was subjected to both methods of measuring the 



length Hi the sc I third fourth and fifth n 

cat pals. 

I he series was obtained during ■> m •< i 
period, 10 to 17 Ma> 1963, from adj i 
the Rio ( iwi | Rio Segovia V/anl Ri 

which separates II luras and Nicaragua m a 

locality 7H mi. ENEE Danli, Honduras All i 
mens are deposited m the Texas ( oopcrativc Wild- 
life Collection, Texas A & M University. 

A statistical (comparison ol both mctho 
shown in Table I. Die variance ratio (I 
was used to lest the hypothesis that one method 
consistently yielded observations with smallci vari- 
ance and was therefore the more desirable None 
of the_F-values was significant at the 5 percent level. 
The method including the carpals consistently 
yielded smaller coefficient of variation values hut 
slightly higher variances for the third and fifth 
metacarpals. 

From a statistical point of view, it appears that 
either method can be employed without introducing 
significant bias. In practice, however, the method 
excluding the carpals is subject to greater error 
because of measuring technique. Determination of 
the exact proximal end of the respective metacarpals 
was difficult and time consuming. The topograph) 
of the wrist joint varied in study skins depending 
upon the quality of preparation which further 
hindered the development of a reasonably rapid, 
easy and consistent measuring technique. Unless 
prior usage or specific needs dictate otherwise. I 
suggest that the method including the carpals is 
the more satisfactory. 

Table I. Comparison of two methods of measuring 

the second, third, fourth and fifth metacarpals 

(MC) of one series of Artibeus lituratus. 



EXCLUDING CARPALS INCLUDING CARPALS 

MEAN MEAN 

MC RANGE S 2 * CV RANGE S 2 2 CV F 

2 53.4 5.53 4.40 56.5 5.29 4.07 1.05 
48.5-58.8 51.4-61.4 

3 64.7 5.18 3.52 67.9 5.28 3.38 0.98 
59.0-71.5 62.3-75.0 

4 64.0 5.97 3.82 66.2 5.15 3.43 1. 16 
57.6-69.9 60.0-72.0 

5 65.2 5.82 3.69 68.5 5.87 3.54 0.99 
59.6-71.6 62.6-75.6 

*s 2 and CV. variance and coefficient of variation, 
respectively. 
**The F-value is obtained from the variance-ration. 
s 2 i/s 2 2- Sample size is 50 throughout: measure- 
ments in mm. 



52 



HI I I I IIS \(>ri III AW CM HORN I A ACADEMY OE SCIENCES 



VOLUME 70 



ACKNOWLEDGMENTS 

Appreciation is extended to Gail Glick for clerical 
assistance, and to Dilford C. Carter, William B. Davis 
and Andrew Starrett tor suggestions concerning the 
manuscript. For permission to examine specimens 
under their curation, I thank William B. Davis and 
Dilford C. Carter. 

Literature Cited 

Davis. W. B. 1970. The large fruit bats (genus Artibeus) 
of Middle America, with a review of the Artibeus 
jamaicensis complex. Jour. Mamm., 5 1: 105-122. 

Starrett. A., and L. de la Torre. 1964. Notes on a 
collection of bats from Central America, with the 
third record for Cyltarops alecto Thomas. Zoolog- 
ica, 49: 53-63. 

Donald R. Patten, Section of Mammalogy, Natural 
History Museum of Los Angeles County, 900 
Exposition Blvd., Los Angeles 90007. 

Accepted for publication September 25, 1970. 



Bull. So. Calif. Acad. Sci. 70(1): 52. 1971 

A NEW SYNONYMY IN THE GENUS 
CHILICOLA (Hymenoptera: Colletidae) 

Cockerell (1918) described Prosopis howardiella 
from a male specimen taken at Oaxaca, Mexico on 
April 30 by L. O. Howard. The description is 
sufficiently detailed to indicate that the bee was not 
correctly assigned generically. 

In 1968 1 had the opportunity to examine the 
type of Prosopis howardiella, in the collection of 
the U. S. National Museum, and found it to belong 
to the genus Chilicola, in the subfamily Chilicolinae. 
This male is identical to those of C. ashmeadi 
(Crawford, 1906). The type of the latter species is 
a female, also in the USNM collections. Males 
have been associated with this name by Eickwort 
(1967) who reared both sexes in Costa Rica. 
Prosopis howardiella Cockerell, 1918, is clearly a 
junior synonym of Chilicola ashmeadi (Crawford, 
1906) (NEW SYNONYMY). 

Eickwort recorded a male from Chiapas, Mexico 
as probably belonging to C. ashmeadi. I have seen 
specimens of both sexes from several localities in 
Chiapas. I have also seen a male from Antigua, 
Guatemala and another from Mt. San Salvador, 
El Salvador. 

Literature Cited 
Cockerell, T. D. A. 1918. Descriptions and records of 
bees. LXXXI. Ann. Mag. Nat. Hist. (ser. 9) 2: 423- 
424. 



Crawford. J. C 1906. Some Costa Rican bees. Trans. 
Amer. lint. Soc, 32: 157-163. 

Eickwort, G. C. 1967. Aspects of the biology of Chili- 
cola ashmeadi in Costa Rica. Jour. Kans. Ent. 
Soc, 40: 42-73. 

Roy R. Snelling, Natural History Museum of Los 
Angeles County, Los Angeles, California 90007 . 

Accepted for publication September 25. 1970. 



Bull. So. Calif. Acad. Sci. 70(1): 52-53, 1971 

CUVIER'S BEAKED WHALE, ZIPHIUS 
CAV1ROSTRIS. FROM BARBADOS 

On 14 May 1968, Rathjen examined and photo- 
graphed a beached carcass of a female Cuvier's 
beaked whale, Ziphius cavirostris, at Cattlewash, 
Walker's Bay, on the northeastern (windward) shore 
of the Lesser Antillean island of Barbados. The 
whale was on the beach the day before as noted in 
the Barbados Advocate — News for 14 May, with 
one photograph. The condition of the carcass 
suggested that it had been dead for several days. 
Color photographs show the body to be moderately 
dark blue-gray except for a dirty white or light 
gray lower jaw and upper lip. 

Rathjen took several centimeter measurements 
on the right side as in Norris ( 1 96 1 ) and these were: 
tip of upper jaw to tip of right fluke (flukes were 
bent so that they were in a perpendicular rather 
than the normal horizontal plane), 454; tip of lower 
jaw to tip of left fluke, 470; tip of upper jaw to origin 
of dorsal fin, 294; origin of dorsal fin to tip of right 
fluke, 160; tip of dorsal fin to tip of right fluke, 144; 
origin of flipper to center of posterior edge of flukes, 
350; tip of upper jaw to anterior margin of eye, 
60; tip of upper jaw to angle of gape, 30; length of 
flipper, origin to tip, 53; width of base of flipper, 
23; width of flukes, tip to tip, 122; depth of flukes 
from anterior margin to center of posterior edge 
along axis of body, 37; height of dorsal fin, 18; 
length of dorsal fin base, 30; length of throat creases, 
20; length of eye, 4; height of eye, 1.5. 

The carcass was buried by surf action and 
unfortunately Rathjen was unable to relocate the 
exact spot for excavation and the record is now 
limited to several color and black and white photo- 
graphs and the measurements. 

A second Ziphius, in slightly fresher condition 
than the first, stranded on the beach at Bathsheba, 
St. Joseph, on the northeastern side of Barbados 
about 5-6 km from the first stranding, on 7 April 
1969. Circumstances prevented Rathjen's getting 



mi 



RESEARCH No 1 1 S 






to the site before the carcass was buried. I his 
carcass was unearthed briefly in early June and 
Rathjen was able to obtain the cranium shortly 
before the local health authorities insisted on 
reburying the remains. In early 1970, Joseph ('. 
Moore of the Field Museum of Natural History 
examined the cranium at the Caldwells' laboratory 
and noted that it was a female (we had not been 
able to tell from the carcass photographs, taken 
at a bad angle) and that it measured 893 mm in 
londylobasal length. 

The closest published western Atlantic records 
for Ziphius are from Argentina to the south (Hersh- 
kovitz, 1966) and Puerto Rico to the north (Erd- 
man. 1962). 

There are a few records of other cetaceans from 
Barbados. From specimens obtained in 1869, 
Turner (1912) listed the mandible of Globicephalus 
melas from Speightstown (probably Globicephala 
macrorhynchu, the only pilot whale now known 
in the Lesser Antilles) and the mandibles of Tursiops 
tursio (=T. truncatus) and of Prodelphinus sp. 
(= Stenella sp.) with no locality data other than 
Barbados. The latter, with a tooth count of 35, 
probably was one of the spotted dolphins of the 
frontalis group (Fraser, 1950). Brown ( 1942) listed 
sperm whales, Physeter catodon, and humpback 
whales, Megaptera novaeangliae from off the island. 

We wish to thank Dr. Albert Jones who arranged 
to have the cranium of the second specimen brought 
to Miami aboard the National Marine Fisheries 
Service's R/V UNDAUNTED and later tran- 
shipped it to the Caldwells. 

Literature Cited 

Brown, H. H. 1942. The sea fisheries of Barbados. 
Development and Welfare in the West Indies. 
Bull. 1. Advocate Printers, Barbados, 32 pp. 

Erdman, D. S. 1962. Stranding of a beaked whale, 
Ziphius cavirostris Cuvier. on the south coast of 
Puerto Rico. J. Mammal.. 43(2): 276-277. 

Fraser, F. C. 1950. Description of a dolphin Stenella 
frontalis (Cuvier) from the coast of French Equa- 
torial Africa. Atlantide Rept.. 1: 61-84. 

Hershkovitz, P. 1966. Catalog of living whales. Ball. 
U.S. Natl. Mus., 246: 1-259. 

Norris, K. S. (Ed.). 1961. Standardized methods for 
measuring and recording data on the smaller 
cetaceans. ./. Mammal.. 42(4): 471-476. 

Turner. W. 1912. The marine mammals in the Anatomi- 
cal Museum of the University of Edinburgh. 
Macmillan and Co.. London. xv+ 207 pp. 

David K. Caldwell. Communication Sciences Lab- 
oratory and The Florida State Museum. University 



of Florida, Gainesville, Florida ''.'//. Wai 
Rathjen Caribbean Flsherli i< lopmeni Project, 
Food and Agriculture Oi <n "I the t nlted 

Nations, Barbados {present addn Satlonal \lu 
nnr Fisheries Set i U e, Woods H 
and Melba ( ( aid well Communication s< «•„. , •. 
Laboratory, University "I Florida, Gainesville, 
Florida. 

Accepted for publication Septernbci 2'.. 1970. 



Bull. So. Calif. Acad. Sci. 70(1): 53-54, 1971 

ADDITIONAL SPI < 1M1 NSOl mi 
COLUBRID SNAKE /l/WAS/ft/OWA/ 
VFLIFFRUM COPE FROM ( OS I A k !< \ 
WITH COMMENTS ON A 
PSEUDOHERMAPHRODITE 

A review of the genus Amastridium (Wilson and 
Meyer, 1969) listed six specimens of this rare snake 
from Costa Rica, all from the Atlantic versanl. The 
locality, San Jose, given for the holotype of 
Fleischmannia obscura Boettgcr, the only specimen 
of Amastridium veliferum previously recorded 
from the Pacific versant of Costa Rica, has been 
questioned (Scott. 1969). By good fortune, two 
specimens of definite Pacific versant provenance 
have since been obtained. One of these (Museo 
Nacionial de Costa Rica, UCR 2819) was taken 
by Pedro Leon near the airport about 3 km south- 
west of Rincon, Peninsula de Osa. Puntarenas 
Province, on 6 August 1969. The other (MNCR 
3142) was obtained by Norman J. Scott. Jr. at 
Finca Las Cruces. 2 km south of San Vito. 
Puntarenas Province, on 16 March 1969. The first 
locality is not more than 100 m above sea le\el. the 
second approximately 1200 m. The only other 
known Pacific slope occurrence for the species is in 
Chiapas, Mexico. 

We first examined MNCR 2819 and noted that 
it is peculiar in a number of respects. Later, one of 
us (Robinson) located a second Pacific versant 
specimen which helps to clarify the status of 
Amastridium from this versant. The first specimen 
(MNCR 2819) is a male with 111 ventrals. the 
lowest count for the species. Wilson and Meyer 
(1969) recorded 119 as the lowest ventral count 
based on a male from southern Nicaragua. This 
Costa Rican specimen is also the only one known 
to show dorsal scale row reduction. All specimens 
examined by Wilson and Meyer have 17 dorsal 
scale rows throughout the length of the body. The 
scale reduction pattern of MNCR 2S19 is as 
follows: 17 ;^^ 15 (III). Furthermore. MNCR 
2819 has a "single anal plate (there is only a trace 



54 



Bl I I I ll\ SOI Till. R.X CAI. NORMA ,!( I/)/ A/) OI ■ SCIENCES 



VOLUME 70 



0< a groove on the anterior portion of the scale), 
all other specimens have a divided anal plate. The 
number ofsubcaudals in this specimen is 70, 2 less 
than the lowest count lor males from throughout 
the range (the subcaudal range is 72 to 86), given 
by Wilson and Meyer (I960), but the tail length- 
total length ratio (0.326) is greater than that 
previously reported lor male A. veliferum (0.277 to 
0.319). The anterior temporal is fused with the 
upper temporal in the second row on the left side 
(1 + 2, the normal condition, on the right), a 
condition seen in only one other specimen (from 
Panama). As is the case for specimens of A. 
veliferum from Nicaragua to Panama, there is no 
loreal. Infralabials number 9-9, and other cephalic 
scutellational features are typical for the species. 

One would certainly not recognize a subspecies 
in a known variable species on the basis of a single 
specimen. The second specimen, besides establish- 
ing the presence of a linking Atlantic-Pacific 
population, possesses enough "typical" character- 
istics to invalidate any suspicion that Pacific versant 
individuals might represent a taxonomically recog- 
nizable subgroup. In MNCR 3142 there are 130 
ventrals. the tail is incomplete, and other scutella- 
tional features are typical, except that there are a 
number of atypical additions and deletions of scale 
rows along the length of the animal. 

Coloration of both specimens is similar to that 
of others from Costa Rica, and shows no approach 
to the lighter Panamanian specimens (Wilson and 
Meyer, 1969). The right hemipenis is everted on 
MNCR 2819, and closely resembles that of a 
specimen from Honduras, described and pictured 
by Wilson and Meyer. 

That MNCR 28 1 9 should show so many external 
morphological peculiarities is, at first analysis, 
striking. The species Amasiridium veliferum, com- 
pared to most other colubrid snakes, is unusual, 
however, in the type of variation in several mor- 
phological features, such that the first four speci- 
mens collected were described as different genera 
and species. Since 1925, Amastridium has been 
considered to consist of two species, synonymized 
by Wilson and Meyer (1969). As Scott (1967) 
stated, "as intraspecific variation becomes better 
known, the fallacy of the constancy of 'key' charac- 
ters at the specific or generic level in colubrid 
snakes becomes increasingly apparent." 

While examining MNCR 3142 for stomach 
contents, the junior author noted certain peculiar- 



ities in the reproductive system of what, on the 
basis of external examination, appeared to be a 
male specimen. This specimen has one everted 
hemipenis. however, the internal reproductive 
system is that of a female. Although somewhat 
damaged during dissection, the specimen has a 
pair of swollen, convoluted oviducts and what 
appear to be elongate ovaries, which are in the 
correct relative positions for snakes (right ovary 
anterior to the left). Comparison of the hemipenis 
of this specimen with that of an unequivocal male 
(testes and vasa deferentia present) from Honduras 
(LACM 45150) demonstrates that the hemipenis 
of MNCR 3142 is aberrant. It is relatively small 
and thin, spines are absent the length of the organ, 
the sulcus spermaticus is shallow and poorly formed, 
and there are some atypical conical lobes on the 
absulcate surface. A m. retractor penis magnus 
is present. 

This pseudohermaphrodite specimen was prob- 
ably a functional female, but the presence of 
hemipenes and associated muscles would cause 
one to consider this animal a male when using the 
conventional means of sex determination and this 
might account for the preponderance of apparent 
males from the southern portion of the range, which 
was noted by Wilson and Meyer (1969). Only 
internal dissection of the specimens from this 
portion of the range can resolve this question. At 
any rate, MNCR 2819, a specimen from this area, 
is a typical male. 

Literature Cited 

Scott, N. J., Jr. 1967. The colubrid snake, Tropidodipsas 
annulifera, with reference to the status of Geatrac- 
tus. Exelencophis, Chersodromus annulatus, and 
Tropidodipsas malacodryas. Copeia, 1967: 280- 
287. 

. 1969. A zoogeographic analysis of the snakes 



of Costa Rica. Ph.D. Thesis, Univ. So. California, 
390 pp. 

Wilson. L. D., and J. R. Meyer. 1969. A review of the 
colubrid snake genus Amastridium. Bull. So. Calif. 
Acad. 5c/.. 68: 146-160. 

Larry David Wilson, Department of Biology, 
University of Southwestern Louisiana, Lafayette, 
70501, and Douglas C. Robinson, Museum of 
Zoology, University of Costa Rica, San Jose. 

Accepted for publication January 14,1971. 



News and Notks 



NEW 
ORMAT 



Willi I his issue. 
the Southern Cal- 



ifornia Academy of 
Sciences begins a new era of publications. The 
Bulletin has been redesigned and will appear three 
limes yearly. Hopefully, the Bulletin will now serve 
ils community better by providing more information 
to all members. During the period ol ^transition, 
from the old single column to the new format, more 
time than anticipated was required. The three 
numbers of the Bulletin will be issued on April 30, 
August 3 1 , and December 3 1 . 



COVER 
ILLUSTRATION 



A" 



n illustration will 
appear on the 
front of every issue of 
the Bulletin. The first number of each volume will 
always have a color frontispiece. The cover illus- 
tration will not necessarily be from or related to one 
of the contributions, although it is required to be 
related to our area. Authors and members of the 
Academy may submit to the Editor copies of art- 
work which they believe appropriate for the Bulletin 
cover. 



news a r : 

SECTION 



section has 
created to 
print relevant news 
relating to the southern California community. 
News of any nature or field of endeavor will be 
acceptable. The News and Notes section will also 
consider reviews, comments, and any type of 
announcement. This section will stress current news 
as well as that of particular interest to our area. 
Submit copy for publication to the office of the 
Managing Editor. 



INDUSTRY An April 19. 1971 

AND V^ Alec R. Stra- 

ENVIRONMENT chan, formerly with 

the California Department of Fish and Game, 



<sn Golden Shore, Long Beach accepted a 
industrial position with Southern < alifornia I 
Km i will be investigating the environmental 
of thermal discharges, antifouling equipment, and 
impregnation studies in prevent fouling. Ron is a 
fulltime employee ul Southern < alifornia I 'ii on 
as biologist -diver. 



OIL f~~* opies ol the 

POLLUTION y~s volume Univer- 

REPORT sity of Southern I 

fornia Oil Pollution Report arc now available. 

Copies may be purchased through Mrs. Dorothy 
Halmos, Librarian, Allan Hancock Foundation 
University of Southern California. Los Angeles. 
California 90007. 



ICHTHYOLOGICAL 1J dward Rar,L > and 

ASSOCIATES l ~ J Ed Bales - boIn 

of Ichthyological 
Associates, are consultant contractors to Southern 
California Edison to study the fishes in and about 
the generating plants in southern California. The\ 
will undertake experimental work and try to re- 
design intakes to keep fishes out during heat treat- 
ment operations. Marineland of the Pacific will also 
assist in the operations. 



RARE AND An May 21. 1971. 

ENDANGERED V^ the Fish and 
WILDLIFE Game Commission of 

the state of California declared that any species of 
wildlife considered to be rare or endangered would 
be prohibited to collect. The list of rare and en- 
dangered fish and wildlife adopted by the California 
Fish and Game Commission is available from B. E. 
Faist, Wildlife Protection Branch. Department of 
Fish and Game. 1416 Ninth Street. Sacramento. 
95814. 



55 



56 



BULI I ll\ SOI 11/ 1 RN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 



LLOYD VV. BARRKR 
APRIL 25. 1949 



In the early morning 
hours of Tuesday. 
June 29, 1971. a vessel 
under low bj the Coast Guard went down in rough 
seas between Santa Rosa and Santa Cruz Islands. 
Besides the numerous sea lions in cages, three 
individuals were aboard this ill-fated craft. This 
aeeident proved fatal to one young man and all the 
sea lions. 

The career of Lloyd W. Barker ended in the dark, 
cold waters of the Santa Barbara Channel. Lloyd 
was acting as an official observer on a commercial 
vessel catching California sea lions. He was repre- 
senting the California Department of Fish and 



Game, which had been under pressure by the Sierra 
Club to have an observer on each such commercial 
vessel. It seems that the issuance of a permit, and 
its implied regulations, were insufficient for the 
Sierra Club. 

Lloyd was a student at Humboldt State College 
and spent his summers working for the California 
Department of Fish and Game. He had an intense 
interest in the California fossil fish record as well as 
the recent fauna. Lloyd's cheerful outlook on life 
and warm personality will be missed for it had 
special significance to those of us fortunate to know 
him. 



Some Forthcoming Contributions 



New species of Polydora (Polychaeta: Spionidae) from the coast of California. By James A. 
Blake and Keith H. Woodwick 

The microhabitats of western wolf spiders of the genus Pardosa. By Donald C. Lowrie 

Two types of aggregation grouping in the large milkweed bug, Oncopeltus fasciatus (Dallas) 
(Hemiptera, Lygaeidae). By Steven R. Kutcher 

The larvae of Pagurus samuelis (Stimpson) (Decapoda, Anomura) reared in the laboratory. 
By Floy E. Mac M Ulan 

A new species of Iphitima (Polychaeta) from Cancer antennarius (Crustacea: Decapoda). 
By John Pilger 



INSTRUCTIONS H)K AUTHORS 

Contributions to the BULLETIN may be in any of the fields of seience, by any member of the 
Academy. Acceptance of papers will be determined by the amount and character of new infor 
mation and the form in which it is presented. Articles must not duplicate in any substantial way, 
material that is published elsewhere. Manuscripts that do not conform to HULLL'IIN style will 
bo returned to the author. All manuscripts will be handled through an editorial board working 
in cooperation with the Editor. 

Manuscript Form — (1) The 1964 AIBS Style Manual for Biological Journal-, i. recom- 
mended as a guide, but where there is a conflict, the BULLETIN style will prevail. (2) 'type- 
write material, using double spacing throughout, including Literature Cited, and leaving a mini 
mum of one inch margins, on only one side of 8'/2 x 1 1 inch standard weight paper. (3) Submit 
two copies. (4) Place tables on a separate sheet of paper. (5) Footnotes should be avoided if 
possible. (6) Legends for figures and unavoidable footnotes should be typed on separate paper. 
(7) Method of literature citation must conform to BULLETIN style — see Volume 70 and later 
issues. Spell out in full the titles of non-English serials and places of publication. (8) All manu- 
scripts, except research notes, must be provided with an abstract at the beginning of the article. 
(9) Acknowledgments should be placed just before literature cited. (10) Short reports may be 
submitted as research notes — see Volume 70 and later issues for style. 

Illustrations — All illustrations, including maps and photography, must be referred to as 
"Figures." Illustrations must not be larger than 8'/2 x 1 1 inches: they may be either the original 
or glossy print. All illustrations must be in proper proportions for reduction to BULLETIN 
page size. Submit one photoduplicated copy of each illustration with the manuscript. 

Cover photograph — A photograph will be printed on the cover of each issue. This may 
either be a photograph used in one of the articles or a photograph of something of scientific 
interest to southern California. Consult the Editor for selection of such material. 

Proof — Author will be sent proof which should be corrected and returned promptly to the 
Managing Editor. Changes, other than typesetting errors, will be billed to the author. Orders 
for reprints should be sent to the Managing Editor at the time corrected proof is returned: 
appropriate order forms will be included when proof is sent. The BULLETIN does not furnish 
free reprints. 



MEMBERS OF THE EDITORIAL BOARD 



1971 

Jack W. Anderson 
Robert M. Chew 
Kristian Fauchald 



1972 

George F. Fisler 
William J. Morris 
Nancy Nicholson 



1971-1974 
Phillip A. Adams Roberta S. Greenwood 

Joseph Arditti Daniel Gutherie 

James A. Blake I. M. Newell 



cover: Algal community at Eel Point, San Clemente Island, one of the offshore channel islands 
of southern California. The dominant algal forms include the surf grass. Phyllospadix, in the 
immediate foreground and background; sargassum weed, Sargassum agardhianum. hanging down 
from the top of the exposed rock; Halidrys dioica, along the face of the exposed rock with black 
abalones; and a patch of algal turf is exposed at the base of the exposed rock at the righthand 
margin. 

Photograph by Donald B. Bright 



LIBRARY 

MAR 1(3 1972 

NEW YORK 
30TAN1CAL GARDEN 



S U T HERN C A L I F O R N I A ACADEMY OF SCIENCES 




ULLETIN 



Number 2 




*K£3£.* 



D/^AC 1 A ^t\/*\ ff ^ -■/*-» /i A«^f\ 



A TTT'CT 1 mi 



SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



Board of Directors 



John J. Baird 

Donald R. Patten 

Donald B. Bright 

Jules Crane, Jr. 

John E. Fitch 

Takashi Hoshizaki 

John L. Mohr 



William J. Morris 
Donald J. Reish 
Charles E. Rozaire 
Elbert L. Sleeper 
Andrew Starrett 
David L. Walkington 
Stewart L. Warter 



Charles R. Weston 



Officers of the Academy 



Andrew Starrett 

Elbert L. Sleeper 

Jules Crane, Jr. 

Charles E. Rozaire 

Donald R. Patten 

Donald J. Reish 

James Dale Smith 

Roberta S. Greenwood 



President 

First Vice President 

Second Vice President 

Secretary 

Treasurer 

Editor 

Managing Editor 

Index Editor 



Membership in the Southern California Academy of Sciences 

Membership is open to all scientists of all branches and to any person interested 
in the advancement of science. 

Annual Members $ 8.00 

Student Members 5.00 

Life Members 150.00 

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



The Bulletin is published three times a year by the Academy. Address all communica- 
tions to the appropriate officer in care of the Natural History Museum of Los Angeles 
County, Exposition Park, Los Angeles, California 90007. 



BULLETIN OF THE SOUTHERN CALIFORNIA 
ACADEMY OF SCIENCES 

Volume 70 February 23, 1972 Number2 



CONTENTS 

The larvae of Pagurus samuelis (Decapoda: Anomura) reared in the laboratory. 

By Floy E. MacMillun 58 

Descriptions and notes concerning some oriental Aphelocheirus (Hemiptcra: 

Aphelocheiridae). By Ira La Rivers 69 

New species of Polydora (Polychaeta: Spionidae) from the coast of California. 
By James A. Blake and Keith H. Woodwick 72 

A new species of Hunterotrema (Digenea: Campulidae) from the Amazon River 

Dolphin (Inia geoffrensis). By Murray D. Dailey 79 

A new species of Funnel-eared bat (Natalus strarnineus) from western Venezuela. 

By Omar J. Linares 81 

A new species of Iphitime (Polychaeta) from Cancer antennarius (Crustacea: 
Decapoda). By John Pilger 84 

Two types of aggregation grouping in the Large Milkweed bug, Oncopeltus 
fasciatus (Hemiptera: Lygaeidae). By Steven R. Kutcher 87 

The invasion and distribution of the Asiatic clam (Corbicula manilensis) in a 
southern California reservoir. By Alio W. Fast 91 

Two bopyrids (Isopoda) from New Guinea. By Charles G. Danforth 99 

A new intranasal chigger of the subgenus Crypticula, genus Microtrombicula 
(Acarina: Trombiculidae) from Texas. By Richard B. Loomis and James P. 
Webb, Jr 102 



57 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(2): 58-68, 1971 

THE LARVAE OF PAGURUS SAMUELIS (DECAPODA: ANOMURA) 
REARED IN THE LABORATORY 

Floy E. MacMillan 1 



Abstract: Pagurus samuelis (Stimpson) larvae from northern California 
were studied. Larvae were raised at 15 and I7°C. Illustrations and descrip- 
tions of the four zoeal stages and glaucothoe are presented. The zoea are 
compared with descriptions of zoea of this species from Japan. 



Over 100 species of the genus Pagurus have been 
described. Few of these (Provenzano and Rice. 
1964) have had their larval stages described and 
many descriptions have been so general that it is 
impossible to distinguish planktonic larvae of 
different species. 

Four species of Pagurus are commonly found 
in the intertidal regions of the northern California 
coast: P. granosimanus (Stimpson), P. hemphilli 
(Benedict). P. hirsutiuculus (Dana), and P. 
samuelis (Stimpson). Larval stages of the first three 
species have been described by Mr. William Hall 
and will be published later. Coffin ( 1960) described 
the ovulation, embryology and developmental 
stages of P. samuelis; however, his description of 
the larval stages was of a general nature. Kurata 
(1968) also briefly described the larvae of this 
species from Japan, but his larvae differ significantly 
from California material. 

Provenzano and Rice ( 1 964), Greenwood ( 1 966), 
Scelzo and Boschi (1969) and Roberts (1970) have 
described larvae of Pagurus species comparing the 
setation and appendages of each larval stage as well 
as the overall size and appearance of the zoea and 
glaucothoe. My purpose is to redescribe the larval 
stages of Pagurus samuelis in a similar manner so 
that details will be available to help identify plank- 
tonic forms. Differences observed between Cali- 
fornia P. samuelis and the accounts of the species 
from Japan are compared. 

I wish to extend my thanks to the Pacific Marine 
Station for use of their facilities and to Mr. William 
Hall and Dr. James A. Blake for critically reviewing 
this manuscript. This summer study was supported 
by National Science Foundation Grant (GB- 1 9344). 

METHODS 

Seven ovigerous females were collected on July 28, 
1970 at Estero de San Antonio, two miles north of 
Dillon Beach, California. The specimens were 
found living in shells of Tegula funebralis (A. 



Adams) and Tegula brunnea (Philippi) in a pro- 
tected rocky intertidal area. In the laboratory each 
female was kept in a plastic aquarium containing 
sea water. On July 29, 1970, one of the females 
yielded over 250 larvae. 

The larvae were separated and placed in culture 
dishes. One hundred and eight larvae were placed 
in 5.5 mm bowls, 3 per bowl, with approximately 
25 ml of filtered sea water. One hundred sixty-eight 
larvae were placed in 10 mm bowls, 7 per bowl, 
with 200 ml of sea water. After the first molt, and a 
number of fatalities, all the larvae were maintained 
in the larger bowls with only 3 per 200 ml of water. 
Freshly hatched Anemia nauplii were added as 
food. The bowls were placed in refrigerators in 
which the temperatures were kept constant at 15 
and 17° C. Salinity varied from 30°/ 00 to 35°/ 00 . 
Water was changed every other day and fresh 
Anemia nauplii were added daily. 

Ten to 20 specimens of each instar and the 
exuviae were preserved in ethylene glycol. Draw- 
ings of the whole larvae were made from specimens 
narcotized with propylene phenoxytol (0.15%) and 
from preserved specimens. At least 5 specimens of 
each instar were dissected and studied in detail with 
respect to setation, segmentation, size and form. 
Drawings of dissected appendages were checked 
against the exuviae. 

Duration of each stage represents time spent in 
a given instar by larvae which successfully molted 
to the next instar. 

Measurements were made using an ocular 
micrometer. Total length (TL) represents a mea- 
surement from the tip of the rostrum to the posterior 
border of the telson exclusive of the telson processes. 
Carapace length (CL) represents a measurement 
taken from the tip of the rostrum to the tip of the 
posterolateral carapace spines. All measurements 
are accurate to within ±0.1 mm. 



Pacific Marine Station, University of the Pacific, Dillon Beach, 
California 94929. 



58 



1971 



THE LARVA I: OF I'ACIIRIIS SAMI/I I /', 



• 



r^ 



I 



I 





Figure 1. Pagurus sainuelis (Stimpson). Dorsal views of zoeal stages I-IV. Scale 0.5 mm. 



FIRST ZOEA (Figure 1 I, 2 I) 
Size:CL, 1.1 mm; TL 2.4 mm 
Duration: 6-7 days at 15 and 17 C C 

General Characteristics 

The eyes are sessile. The rostrum is long, extend- 
ing almost to the tip of the antennal endopodite. 
The carapace bears 2 short downward projecting 
hooks on the posterolateral margins. The abdomen 
is composed of 5 distinct somites and a sixth which 
is fused with the telson. Somite 1 lacks spines: 
somites 2-5 each bear 2 pairs of short postero- 
dorsal spines which increase in size from somite 
2-5. In addition, somites 2-4 have another pair of 
smaller dorsolateral spines. There is a pair of short 



lateral spines on somites 2-5. The telson (Fig. 3 li 
is longer than wide, with the longest setae as long 
as the telson itself. The posterior margin is straight 
with a distinct, almost v-shaped. notch. There are 
7 telson processes on each side: the outermost is a 
heavy spine, the second a delicate hair, the others 
are articulated plumose setae. 

Coloration 

The dorsal surface just posterior to the eye> 
contains a prominent yellow chromatophore. The 
eyes also contain a yellow pigment giving them an 
iridescent appearance. The base of each mandible 
contains a deep red chromatophore. There are red 
ehromatophores posterior to each antennule and S 



60 



lU'l.LETIN SOI I HERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 




Figure 2. Pugiirus samuelis (Stimpson). Lateral views 
of zoeal stages I-IV. Scale 0.5 mm. 




z 




small red-pigmented cells over the lateral surfaces 
of the carapace. The basipodite of maxillipeds 1-2 
bears a prominent red chromatophore. 

Appendages 

Antennule (Fig. 4 I) is uniramous and terminally 
bears 2 aesthetes and 2 fine hairs. Subterminally 
there is a long plumose seta. 

Antenna (Fig. 4 I) is biramous. The endopodite 
is fused to the basipodite. It lacks terminal setae 
and reaches almost to the tip of the antennal scale 
spine. There is a ventral untoothed spine on the 
endopodite near the base of the antennal scale. The 
antennal scale is 8 times as long as it is wide. It 
bears a long tooth terminally and 4 long plumose 
setae and I shorter plumose seta along the medial 
margin. 

Mandibles (Fig. 5 I) are asymmetrical. Each has 
an incisor, a molar process, and a number of 
corneous teeth. 

Coxal endite of the maxillule (Fig. 5 I) bears 4 
plumose setae and I nonplumose seta terminally. 
The basal endite has 2 very stout teeth and 2 smaller 
spines, with each tooth bearing a number of spin- 
ules. Endopodite bears 3 nonplumose setae on the 
distal segment and an additional nonplumose seta 
on the middle segment. 




n 




he 



Figure 3. Pagurus samuelis (Stimpson). Telson of zoeal stages I-IV. Scale 0.2 mm. 



E 



1971 



THE LARVAE OF EACH RIJS SAMU I I IS 



',1 



I 



H 





E 



/ 



I 







JZ 



Figure 4. Pagurus samuelis (Stimpson). Top, antennules of zoeal stages I-IV. Scale 0.1 mm. Bottom, antennae of 
zoeal stages I-IV. Scale 0.2 mm. 



62 



HI I I I ll\ sot ////.AW CALIEORNIA ACADEMY Of SCIENCES 



VOLUME 70 



Scaphognathite of the maxilla (Fig. 5 I) has 5 
plumose setae along the margin. The endopodite 
bc.us 3 plumose setae terminally and 2 setae sub- 
terminally on the medial margin. The distal and 
proximal lobes of the basal endite each have I 
subterminal and 3 terminal plumose setae. The 
distal lobe of the coxal endite has 2 terminal setae 
and I subterminal seta. The proximal lobe has 4 
terminal setae. 

Basipodite of the maxilliped I (Fig. 6 I) has a 
pair of plumose setae at the anterorhedial corner 
and 6-7 hairs on the medial margin. The exopodite 
is composed of 2 indistinct segments and terminates 
m 4 long plumose setae. Endopodite is five-seg- 
mented, the terminal segment bearing 4 terminal 
plumose setae and a fifth on the lateral margin. The 
penultimate segment has a heavy plumose seta and 
a finer plumose seta on the distal medial margin. 
The antipenultimate segment bears a single heavy 
seta on the distal medial margin and a row of fine 
setules on the lateral margin. The next proximal 
segment bears a heavy plumose seta and a non- 
plumose seta on the distal medial margin, and a 
fine row of setules on the lateral margin. The 
proximal segment has 3 nonplumose setae on the 
distal medial margin and, again, a row of setules 
on the lateral margin. 

Basipodite of the maxilliped 2 (Fig. 7 1) has I seta 
on the anteromedial corner and 2-3 hairs along the 
medial margin. The exopodite consists of 2 indis- 
tinct segments and bears 4 terminal plumose setae. 
The endopodite has 4 segments; the terminal seg- 
ment bearing 4 terminal plumose setae and 1 sub- 
terminally. The other segments each have 1 heavy 
plumose seta and 1 fine plumose seta on the distal 
margin. In addition, the antipenultimate segment 
has a row of fine setules on the lateral margin. 

Maxilliped 3 (Fig. 8 I) is a uniramous rudiment. 

SECOND ZOEA (Figure 1 II. 2 II) 
Size: CL 1 .4 mm; TL 2.6 mm 
Duration: 6-8 days at 1 5 and 17°C 

General Characteristics and Coloration 

The eyes are mobile and limb buds are visible 
under the carapace behind maxilliped 3. The telson 
(Fig. 3 II) remains fused with abdominal somite 6 
but the median notch has disappeared, and there is 
an additional pair of short medial telson spines. 

The coloration is essentially the same as in the 
previous stage except that the red chromatophores 
are more distinct. The pigmented cells on the 
carapace are now clearly red chromatophores. 

Appendages 

Antennules (Fig. 4 II) terminate in 1 long 
aesthete. 2 shorter aesthetes and 2 small hairs. 



Antennae (Fig. 4 II) have elongated but are 
identical to that of zoeal stage 1 in form and 
setation. There is an extremely small tooth at the 
base of each antennal scale. 

Mandibles (Fig. 5 II) are similar in shape to 
those of the previous stage, but there is an increase 
in the number of corneous teeth. 

Maxillules (Fig. 5 II) differ from those of the 
first zoeal stage with the addition of 2 heavy spicule- 
bcaring spines on the basal endite. There is also an 
additional small spine proximally on the basal 
endite and an additional small terminal spine on 
the coxal endite. 

Scaphognathite of the maxilla (Fig. 5 II) bears 
an additional seta on the lateral margin. The setation 
and form of the basal and coxal endites remain 
unchanged. 

Basipodite of the first maxilliped (Fig. 6 II) has 
2 fine setae and a short hair at the distal medial 
corner. There are 4 setae and 1-2 additional fine 
hairs on the medial margin. The exopodite termi- 
nates in 7 natatory setae. The setation of the termi- 
nal and penultimate segments remains unchanged 
but the antipenultimate and the 2 proximal segments 
each bear a long plumose seta on the lateral margin 
in place of the setule rows present in zoeal stage 1 . 

Basipodite of maxilliped 2 (Fig. 7 II) bears 2 
setae on the distal medial corner and 1-2 setae along 
the medial margin. The exopodite terminates in 7 
natatory setae. The setation of the endopodite 
remains unchanged except that a long plumose 
seta has replaced the row of setules on the second 
proximal segment. 

Maxilliped 3 (Fig. 8 II) consists of a basipodite 
and an exopodite with 6 natatory setae. 

THIRD ZOEA (Figure 1 III. 2 III) 
Size: CL 1 .5 mm; TL 2.6 mm 
Duration: 7 days at 17°C 

General Characteristics and Coloration 

Abdominal somite 6 is incompletely fused to the 
telson. There is no dorsal spine on this somite. The 
telson (Fig. 3 III) is twice as long as it is wide; the 
number and relative length of the telson spines is 
unchanged. Uropods consisting of a fused proto- 
podite and exopodite are present. The exopodite 
bears 3 small teeth and 3 plumose setae on the 
medial margin. Coloration is unchanged. 

Appendages 

Antennules (Fig. 4 III) are segmented and 
biramous. The distal segment terminally bears 1 
large and 2 small aesthetes. 1 plumose seta and 2 
hairs. Subterminally, there is an additional pair of 
aesthetes. The segment representing the ventral 



1971 



THE LARVAE <>l I'AdilKHS SAMlii.l is 




I 



- 



r - 






in 




n 




M 







1 .A\\\l 



Figure 5. Pagurus samitelis (Stimpson). Top, mandibles ofzoeal stages I-IV. Center, maxillules of zoea! stages [-IV. 
Bottom, maxillae ofzoeal stages I-IV. Scales 0.1 mm. 



fiagellum bears a single long plumose seta. The 
peduncle has a long plumose seta on the distal 
margin. 

Antennal scale (Fig. 4 III), excluding the termi- 
nal tooth, is 8 times as long as wide. On some 
specimens the terminal tooth was shortened, and in 
a few cases was forked at the tip. The ventral tooth 
on the endopodite is reduced and the margin 
beneath the antennal scale is untoothed. 

Mandibles and maxillules (Fig. 5 III) are un- 
changed. 

Scaphognathite of the maxilla (Fig. 5 III) bears 



7 plumose setae along the margin. The endopodite 
bears 4 terminal and 3 subterminal plumose setae. 
The setation of the basal and coxal endites is 
unchanged. 

Exopodite of maxilliped 1 (Fig. 6 III) bears 7 
natatory setae: the endopodite and basipodite are 
unchanged. 

Maxilliped 2 (Fig. 7 III) is unchanged except 
that the exopodite has S natatory setae. 

Maxilliped 3 (Fig. 8 III) has an endopodite 
terminating in 1 small plumose seta. The exopodite 
has 7 natatory setae. 



64 HCI I I ll\ SOI I III RN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 





Figure 6. Pagurus samuelis (Stimpson). First maxillipeds of zoeal stages I-IV. Scale 0.5 mm. 




Figure 7. Pagurus samuelis (Stimpson). Second maxillipeds of zoeal stages I-IV. Scale 0.5 mm. 



1971 



FOURTH ZOLA (Figure I IV, 
Size: CL 2.2 mm; TL 3.4 mm 
Duration: 7-8 days at I7"( 

General Characteristics and ( 'oloration 

Pleopod hutls are present on abdominal somites 
2-5. Telson (Fig. 3 IV) is almost 3 limes as long as 
wide. Protopodites of the uropods are articulated 
wilh Ihe exopodites but not with the endopodites. 
There is an additional yellow chromatophore on 
the rudimentary cheliped and a red chromatophore 
just ahove the basipodite of maxilliped 3. 

Appendages 

The process representing the dorsal llagellum of 
the antennule (Fig. 4 IV) bears 3 aesthetes, 1 
plumose seta, and 2 hairs. In addition, there is a 
longitudinal row of 4-5 subterminal aesthetes. The 
ventral llagellum has lost its terminal seta but has 
retained the plumose seta at its base. 

The endopodite of the antenna (Fig. 4 IV) is 
slightly wider than the antennal scale and extends 
beyond the tip of the antennal scale spine. 

Mandibles (Fig. 5 IV) are unchanged. No palp 
is present. 

Coxal endite of the maxillule (Fig. 5 IV) bears 2 
additional nonplumose setae. The basal endite 
bears 6 heavy spines, each with secondary spinules. 
The endopodite is unchanged. 

Scaphognathite of the maxilla (Fig. 5 IV) bears 
13 plumose setae. Setation of the endopodite and 
of the basal and coxal endites is unchanged. 

Exopodite of maxilliped I (Fig. 6 IV) bears 8 
natatory setae; the endopodite and basipodite are 
unchanged. 

Maxilliped 2 (Fig. 7 IV) is unchanged. 

Exopodite of maxilliped 3 (Fig. 8 IV) bears 8 
natatory setae. The endopodite bears 1 terminal 
plumose seta and 1 subterminally. 

GLAUCOTHOE( Figure 9, 10) 
Size: CL 1 .4 mm; TL 3.4 mm 

General Characteristics and Coloration 

Periopods are functional. The 4 pairs of pleopods 
are well developed and used in locomotion. The 
rostrum is blunt. The telson (Fig. 10 a) is only 
slightly longer than its width and has only 8 terminal 
plumose setae. There are 5 pairs of small setae on 
the dorsal surface and 1 pair on the lateral margin. 
The protopodites of the uropods are articulated 
with both the endopodites and exopodites. The 
endopodites bear a single terminal seta. The 
exopodites are subequal, the left being slightly 
larger than the right. Each has 10 long plumose 
setae along the margin. In addition, there are 5-7 
corneous granules and 4-5 smaller setae along the 
margin. 



THE I.AKVAI.oi PAGURUSSAMl I I is 



IV) 



' 






I 




\ 






Figure H. Pagurui samuelh (Stimpson) rhird rn ; > >. 1 1 

lipccls ofzoeal stages MV. Scale O.j mm. 



Red chromatophores are prominent on Ihe eye- 
stalks, antennulcs. antennal basipodites, and on 
the anterolateral region of the carapace. I here are 
a great number of pale yellow chromatophores 
scattered over the anterior dorsal region of the 
carapace, the eycstalks. and the fifth periopod. 

Appendages 

The ventral flagcllum of the antennule (I 
10 R) consists of 2 segments. The terminal segment 
bears 8 small setae; the proximal segment bears 2-3. 
The dorsal flagellum consists of 4 segments The 
most proximal is unarmed. The other segments 
each have 3-4 aesthetes. In addition, the terminal 
and the next most proximal segments bear several 
fine setae. 

Antenna (Fig. 10 Q) has a scale at its base which 
terminates in 2 nonplumose setae. There arc 10 
flagellary segments, each of which bears a few short 
setae. 

Mandibles (Fig. 10 P) are cuplike and have a 
two segmented palp which terminates with four 
small hooks. 

Endopodite of the maxillule (Fig. 10 O) has lost 
its segmentation and has 1 seta. The basal endite 
bears 12 spines distally and 2 proximally. The 
coxal endite bears 10-14 setae. 

Maxilla (Fig. 10 N) bears 27 setae on the margin 
of the scaphognathite which has increased in pro- 
portion to the rest of the maxilla. The scaphogna- 
thite and endopodite are no longer articulated with 
the endites. 

Maxilliped 1 (Fig. 10 M) is radically changed. 
The basipodite is broad and appears bilobed. It has 
15 setae on the distal lobe and 4 setae on the 
proximal lobe. The endopodite and exopodite are 
reduced and have only 3-4 short terminal setae. 

Exopodite of maxilliped 2 (Fig. 10 L) bears 5 
long plumose setae. The endopodite consists of - 
segments, each with a few short setae. 

Exopodite of maxilliped 3 (Fig. 10 K> has 6 



66 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OE SCIENCES VOLUME 70 




Figure 9. Pagurus samuelis (Stimpson). Glaucothoe. Lateral and dorsal views. Scale 1.0 i 



1971 



THE I.AIiVAl <)1 1'AdDKIIS SAMIII.I l\ 




Figure 10. Pagurus samuelis (Stimpson). Glaucothoe. A, telson; B-E. pleopods; F. fifth periopod; G. fourth perio- 
pod; H, third periopod: I. second periopod; J. cheliped: K. third maxilliped: L. second maxilliped: M. first maxilliped: 
N. maxilla; O. maxillule; P. mandible; Q. antenna; R. antennule. Scale 0.5 mm. 



plumose setae. The endopodite has 5 segments. 
The ultimate and penultimate segments are heavily 
armed with setae, many of which have setules. 

The chelipeds (Fig. 10 J) are subequal in size; 
the left being larger than the right. Periopods 2 and 
3 (Fig. 10 H-l) have dactyli as long as the propodi. 
Periopod 4 (Fig. 10 G) is nonchelate. The propodus 
has 4 corneous granules on the ventrolateral sur- 
face. Periopod 5 (Fig. 10 F) has 7 granules and 7 
long setae on the propodus. 



Pleopods (Fig. 10 B-C) are biramous. Endo- 
podite of each bears 2 small hooks. Exopodites of 
the pleopods on somites 2-4 bear 9 plumose setae, 
whereas the pleopod of somite 5 has onl> 8 

DISCUSSION 

The purpose of the present study is to extend an 
earlier work on Pagurus samuelis larval develop- 
men by Coffin (1960). Coffin did not make a 
detailed study of the larval appendages, making 



68 BULLETIN SOUTHERN CALIFORNIA ACADEMY Oh SCIENCES VOLUME 70 

Table I . Morphological differences between Japanese and California zoea larvae of Pagurus samuelis. 



Characteristic 



Japanese Species 



California Species 



Rostral length 



extends to base of 
antennal scale spine 



extends beyond tip of 
antennal scale spine 



Antenna 
endopodite 

scale 



tooth at base 



forked 

broad, concave outer 
margin with six plumose 
setae 

articulated 



not forked 

narrow, straight outer 
margin with five plumose 
setae 

not articulated 



Fifth abdominal 
segment 



prominent lateral spines 



short lateral spines 



Telson 



wide — convex outer 
margin 



narrow — straight outer 
margin 



Coloration 

chromatophores 
on carapace 

chromatophores 
at base of 
rostrum and on 
antennal 
protopods 



all on ventral margin 
present 



scattered on lateral 
surface 

absent 



differentiation of planktonic larvae difficult if 
species other than P. samuelis are encountered. 
He mentioned briefly the presence of setae on the 
antennule and antennal scale, the number of setae 
on the exopods of the maxillipeds, and the setae on 
the uropods and telson, but did not describe the 
mandibles, maxillules, and maxillae. Coffin did 
not state the number of setae on the antennal scale 
spine, but his drawings suggest only 4. In the 
present study 5 were found. The spine Coffin 
described on the end of the uropod in the third 
zoea is composed of 3 small spines. 

Pagurus samuelis larvae described by Kurata 
( 1968) differ from the California larvae in 5 aspects 
outlined in Table I. Differences in the telson shape, 
antennal structure and lateral abdominal spines 
are highly significant and it is probable that the 
California and Japanese P. samuelis are actually 
separate species. 

LITERATURE CITED 

Coffin, H. G. 1960. The ovulation, embryology and 
developmental stages of the hermit crab Pagurus 
samuelis (Stimpson). Walla Walla College Publ., 
25: 1-29. 



Greenwood. J. G. 1966. Some larval stages of Pagurus 
novae-zealandiae (Dana) 1852 (Decapoda. Ano- 
mura). New Zealand J. Sci. 9: 545-558. 

Kurata. H. 1968. Larvae of decapoda anomura of 
Arasaki. Sagami Bay Pagurus samuelis (Stimpson). 
Bull. Tokai Regional Fisheries Res. Lab., 55:265- 
269. 

Provenzano. A. J., Jr. and A. L. Rice. 1964. The larval 
stages of Pagurus marshi Benedict (Decapoda, 
Anomura) reared in the laboratory. Crustaceana, 

7:217-235. 

Roberts, M. H., Jr. 1970. Larval development of 
Pagurus longicarpus Say reared in the laboratory. 
I. Description of larval instars. Biol. Bull., 103: 
188-202. 

Scelzo, M. A. and E. E. Boschi. 1969. Desarollo larval 
del Cangrejo Ermitano Pagurus exilis (Benedict) 
en el laboritorio (Crustacea. Anomura. Paguridae). 
Physis, 29: 165-184. 



Accepted for publication February 18, 1971. 



BULLETIN SO. CALIF. ACADEM Y <>l S( IENCES 70(2): ',9-72. 1971 

DESCRIPTIONS AND NOTES CONCERNING SOME ORIEN I Al APHL I <" III IRl 
(HEMIPTERA: APHEI/X HI IKIDAI i 



Ika I. a Kivi-.ks 1 



Abstract: Two new species of Aphelocheirus are described from India 
and A. pallens Horvath is noted from additional localities in Nev i I 



During an investigation of the Naucoridae and 
Aphelocheiridae of New Guinea, specimens were 
examined from surrounding areas in an attempt to 
understand the relationships and origins of the 
very unique fauna which exists in New Guinea. 
This work turned up additional records of Aphe- 
locheirus pallens Horvath 1899 in New Guinea as 
well as two new species of Aphelocheirus from a 
related fauna in the southeastern Asiatic mainland. 



Aphelocheirus pallens Horvath 

In 1899, Horvath described this species from 
German New Guinea, in the northeast section of 
the island. In my perusal of the literature, I have 
not seen it recorded elsewhere, but in the Leiden 
and Bishop museums' collections, there are addi- 
tional specimens from the western part of New 
Guinea, as follows: Neth. Ind. -American New 
Guinea Exped., Araucaria Camp, 800 m, 1939 
Mar. 3, L. J. Toxopeus; Same, Mist Camp, 1 800 m. 
(Leiden Museum localities); New Guinea, Neth., 
Bodem, 10 m, 11 km SE of Oerber-faren, 1959 
July 7-17, T. C. Maa, and New Guinea, NW, 
Nabire, S. Geelvink Bay, 10-40 m, 1962 Oct. 13, 
N. Wilson (Bishop Museum localities). 



Aphelocheirus pygmaeus, new species 
Figure 1 

General: The smallest species that I have seen in 
the genus, measuring 4 by 2 mm, approaching the 
genus Potamocoris of the family Naucoridae in 
size and gross appearance. Darker in color anteriorly 
with some yellow bordering, hemelytra lighter. 
Venter blackish-brown with some yellow lightening 
anteriorly, legs yellowish. 

Head: Blackish brown, shiny with sparse pitting. 
Anteclypeus greatly expanded and protuberant 
before eyes, its anterior outline reasonably smoothly 
circular. Eyes blackish, essentially flush with 
surface; hyperocheal angle not discernible as such. 
Labrum large, prominent. Head ratios are: (1) Total 
length-to-width (including eyes) 25: 34 (74%), 
(excluding eyes and utilizing widest interocular 




Figure I. Aphelocheirus pygmaeus. new species, holo- 
typic female. 

space) 25: 21 (84%); and (2) anterior distance be- 
tween eyes to posterior distance between eyes 21: 
21 (inner margins are not parallel, but curved con- 
cavely inwardly). 

Pronotum: Angular in outline, disk brownish black, 
lateral and posterior borders widely yellow. Surface 
somewhat uneven. Lateral edges smooth, nearly 
straight, posterior angle rounded. Anterior edge 
between eyes essentially straight, posterior edge 
bow-shaped. Venter with weak median sternal 
keel, large coxal cavities and coxae, and propleural 
flaps widely separated by coxal cavities. Pronotal 
ratios are: (1) Width between anterior angles to 
width between posterior angles 35: 60 (58% 1: and 
(2) median length to greatest width 20: 60 (33% 
Scutellum: Large, blackish, shiny: ratio of three 
sides, anterior and two laterals 35: 25: 25. 



■Biological Society of Nevada, Verdi. Nevada 8*139. 



69 



-II 



Htlliri.XSOlllllKXl UIIOKM.t U I />/ W > ( '/ \( // A < / S 



VOLUME 70 



Hemelytra: Translucent brownish; claval sutures 
distinct; membranes very large, comprising almost 
halt' hemelytral length and milky transparent. 
Opaque region microroughened but shiny. 
Embolium indistinctly delineated, the inner margin 
discernible and apparently extending caudally to 
intersect outer hemelytral edge so that posterior 
emboliar border is not at right angles to the long 
axis of the body; outer edge weakly flared externally 
about at midline; ratio of length-to-width 31:4 
( 13% ). Hemelytra reaching caudally beyond tip of 
abdomen; connexival edge barely exposed in 
anterior half of abdomen. Wings fully functional. 
Venter: The prothoracic venter has been discussed 
above. Meso- and meta-thoracic ventra blackish, 
as is the abdomen. Connexiva not distinguishable, 
posterior abdominal angles non-spinose. Abdominal 
segment 1 with a prominent, whitish, narrowly 
elongate static sense organ (sta(exta) near external 
edge; spiracles on segments II, III, IV and V each 
in form of a short series of transversely-arranged 
dots. Female subgenital plate narrowing rather 
sharply to an almost pointed tip. 
Legs: Proleg — Coxa large, angularly-elongated. 
Trochanter prominent. Femur moderately incras- 
sate. flattened, ratio of length-to-width 30:15 (50% ), 
length I.I mm. Tibia long, narrow, tubular, bearing 
at its tip a two-segmented tarsus terminating in two 
small claws. 

Mesoleg — Coxa and trochanter similar to proleg. 
Femur intermediate between the predatory incras- 
sation of profemur and the elongated ambulation 
of metafemur, but not definable as incrassate; dense 
row of conspicuous yellow hairs on inner or 
posterior border; ratio of length-to-width 29: 10 
(34% ). length 0.9 mm. Tibia long, narrow, tubular, 
slightly widening distally, heavily beset with large, 
yellow spines, these clustered rather solidly at 
distal end; ratio of length-to-width 25:3 (12%), 
length 0.85 mm. Tarsus 3-segmented, first segment 
minute, remaining two long, tubular, terminating 
in two prominent claws. 

Metaleg — Much longer than other legs. Coxa 
and trochanter larger versions of mesoleg. Femur 
elongate, flattened, slightly bowed but not swollen 
as much as mesofemur; ratio of length-to-width 
38: 10 (26%). length 1.2 mm. Tibia long, narrow, 
rounded, slightly widening distally, sparsely spined, 
fringe of long swimming hairs on inner margin; 
ratio of length-to-width 33: 5 (17% ), length 1.1 mm. 
Tarsus very long and narrow, as long as tibia and 
consisting of 3-segments. the first minute; with a 
thick brush of swimming hairs on inside edge, and 
terminating in two large claws. 
Type Locality and Etymology: Holotypic female, 
allotype and one paratype from: East India. Assam, 
Kohara. Kaziranga, 1959 Oct. 7. E. S. Ross and 



D. Q. Cavagnaro. 1 10 m. Deposited in the Cali- 
fornia Academy of Sciences. The name pygmaeus 
is appropriate for this is the smallest of all known 
Aphelocheirus. 

Comparative notes: Aphelocheirus pygmaeus is 
virtually unique in the family by size alone. I know 
of nothing else which even approaches it in this 
respect. In fact, it bears more superficial resem- 
blance, on this basis, to the naucorid genus Potamo- 
coris, than it does to other aphelocheirids. 

Aphelocheirus nathani, new species 
Figure 2 

General: This description is based on the micropter- 
ous form as being the more common, with com- 
parisons, in appropriate places, with the winged 
form. Lighter in color anteriorly, the darker 
abdomen lighter along edges. Winged form darker 
mainly due to blackish hemelytra and more color 
development on pronotal disk. Venter yellowish, 
with no significantly darker areas. Winged form 
darker ventrally. Size 9.5 by 6.5 mm (winged form 
9.5 by 5.75 mm). 

Head: Shiny, smooth, yellow except for diffuse 
brownish posteriorly. Keystone-shaped, broadly 
and moderately protuberant between eyes in ante- 
clypeal region. Eyes darker, inner and outer margins 
subparallel, the hyperoche angle prominent 




Figure 2. Aphelocheirus nathani, new species, holo- 
typic female, micropterous form. 



1971 



ORIENTAL III.MII'II.RA 



71 



anteriorly, eyes essentially Hush with general 
surface. Labrum prominent, broad and weakly 
pointed, ratio of length-to-width 16:22 (73%), 
Head ratios are: (I) Total median length-to-widlh 
(including eyes) 65: 78 (83%) across hyperocheal 
angles, (excluding eyes) 65: 55 (85%) anteriorly; 
(2) anterior distance between eyes to posterior 
distance between eyes 55:40 (73%); and (3) antei km 
distance between eyes to inner eye length 55: 40 
(73%). 

Pronotum: Yellow, with two diffuse brown areas 
centrally, surface shiny but weakly roughened. 
Posterior border double-angulate, the outer angle- 
being the usual posterolateral one, the inner angle 
being a moderate, caudally-directed angulosity of 
the border; neither angle long or sharp. Lateral 
border smoothly curved; percent of curvature (as 
viewed perpendicularly to the frontal plane of 
section of the specimen as a whole) about 15 
(68: 10). Venter yellow, coxal cavities large, median 
area moderately keeled. Internal propleural flaps 
widely separated by coxal cavities. Ratios are: 

( 1 ) Width between anterior angles to width between 
posteroexternal angles 43:92 (47%); (2) width 
between anterior angles to width between postero- 
internal angles 43: 73 (59%); (3) median length to 
greatest width 25: 92 (27%); and (4) median length 
to parallel length from anterior angle to posterior 
border 25: 39 (64%). 

Winged form with straighter lateral borders, 
angles closer together, less pronounced, appearing 
more as the two edges of a longer, single angle 
which had been cut across diagonally, thus pro- 
ducing two reduced angles. Pronotum longer, 
broader, the proportions being: (1) Width between 
anterior angles to width between posteroexternal 
angles ( = widest part of pronotum) 43: 87 (49%); 

(2) width between anterior angles to width between 
posterointernal angles 43:78 (35%); (3) median 
length togreatest width 27: 87 (3 1 % ); and (4) median 
length to parallel length from anterior angle to 
posterior border 27: 39 (69%). 

Scutellum: Short, broad, yellow with some diffuse 
brownish areas anteriorly. Ratio of three sides, 
anterior and two laterals, 55: 36: 36. In winged 
form, scutellum proportionally larger, darker, 
more rugose, ratio 50: 38: 38. 
Hemelytra: Yellow, texture same as pronotum: 
rounded, abbreviated stumps not reaching midline 
and not reaching posterior border of first abdominal 
segment. Anterior region of embolium present, the 
outer angle being sharp and slightly recurved. 
Winged form with fully developed wings, blackish 
brown, texture different from pronotum. being 
smoother with widely scattered tiny blackish 
tubercles; claval and membranal sutures prominent, 
the membrane distinct; embolium well developed. 




Figure 3. Aphelocheirus nathani, new species, para- 
type, macropterous form. 

lighter laterally, long and narrow, the outer angle 
broad but distinct: embolium length-to-width 90: 
20 (22%), but narrower than this would indicate 
since the width of 20 includes the outer angle: 
wings narrow, rather markedly exposing the con- 
nexival edge. 

Venter: The prothoracic venter has been discussed 
above. Meso- and meta-thoracic ventra and abdo- 
men yellow, darker in the winged form. Static 
sense organ prominent, elongated, small tip on 
posterior end; spiracles typically baliopic. the dots 
spreading laterally in all segments except I. Cen- 
nexivum narrow, widening posteriorly in caudal 
segments, posterior angles sharp on all segments, 
becoming larger caudally. Female subgenital plate 
broadly wedge-shaped, narrowing sharply to a 
bluntly rounded tip. 

Legs: Proleg — Coxa large, angularly-globular. 
Trochanter prominent. Femur weakly incrassate. 
flattened, well-furred along tibial closure edge: 
ratio of length-to-width 65: 24(37%). length 2 mm. 
Tibia long, narrow, tubular, bearing a three- 
segmented tarsus whose first segment is minute, the 
remaining two elongate and terminating in two 
prominent claws. 



72 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OE SCIENCES 



VOLUME 70 



Mesoleg — Coxa and trochanter similar to proleg 

except larger. Femur weakly incrassale. similar to 
profemur but larger, prominently haired along 
inner or posterior margin: ratio of length-to-width 
63:20 (32%), length 2.1 mm. Tibia similar to 
protibia but proportionally broader, hairier and 
with a conspicuous armament o( reddish spines. 
particularly along outer edge and a loose, trans- 
verse, terminal row; ratio of length-to-width 48: 8 
(17 r f). length 2 mm. Tarsus a larger edition of 
protarsus. 

Metaleg — Much larger than pro- and meso-legs. 
with coxa more globular. Femur long, rather flat, 
not truly incrassate in any sense, thinly furred 
along posterior or inward edge; ratio of length-to- 
width 96: 28 (29<7). length 3 mm. Tibia relatively 
longer version of mesotibia, spined along edges 
and with a conspicuous fringe of long, yellow 
swimming hairs along internal or posterior border: 
ratio of length-to-width 90: 9(10%), length 3. 1 mm. 
Tarsus large, much elongated, at least twice the 
length of mesotarsus and heavily equipped with 
swimming hairs inwardly. 

Type Locality and Etymology: Holotypic female, 
allotype and 8 paratypes (2 being winged forms) 
from: South India. Madras, Anamalai Hills. 
Kadamparai. 1963 June. P. Susai Nathan. 1050 m; 
2 micropterous paratypes from Anamalai Hills, 
Cinchona. Nathan, 1959 April, 1050 m, under 
debris of stream; 1 winged paratype from Coimba- 
tore, Nathan. 1959 February. All specimens in the 



collection of the Biological Society of Nevada, 
Verdi, Nevada. I am pleased to name the species 
after its collector, who has diligently sampled the 
naucorid fauna of his region. 

Comparative notes: Aphelocheirus nathani appears 
to be related to the populations cast, rather than 
west, of India, although the continuous and con- 
stant land masses to the west would appear a more 
likely continuum from which it could be derived. 
Lacking the distinct and prominent connexival 
spines of the A. aestivalis (Fabricius) 1 803 group to 
the west, A. nathani is quite close to the Philippine 
species described by Usinger, A. philippinensis 
1938 and A. itichancoi 1938. particularly the latter, 
which is about the same size. From A. itichancoi, 
A. nathani differs in the lighter color, somewhat 
smaller size and much different head proportions, 
itichancoi having the anteclypeal region much 
more extensively developed anteriorly between 
the eyes. The emboliar flare is also more pro- 
nounced in nathani and is longer. Connexival 
spines are somewhat more prominent in itichancoi. 

LITERATURE CITED 

Horvath. G., 1899. Monographia generis Aphelochei- 
rus. Termeszetrajzi Fiizelek, 22: 256-267. 

Usinger, R. L. 1938. The Naucoridae of the Philippine 
Islands (Hemiptera). Philippine J. 5c/., 64: 299- 
309. 

Accepted for publication October 23. 1970. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(2): 72-79, 1971 

NEW SPECIES OF POLY DORA (POLYCHAETA: SPIONIDAE) FROM 
THE COAST OF CALIFORNIA 

James A. Blake 1 and Keith H. Woodwick 2 

Abstract: Four new species of Polydora are described. All species bore 
into calcareous substrata and occur in California, with one ranging as far 
north as British Columbia. 



During the years 1961-70 numerous collections of 
intertidal polychaetes were made along the Cali- 
fornia coast by us. Examination of these collections 
has resulted in the finding of four new species. These 
are added to the genus Polydora which is already 
well represented in California. Hartman (1969) 
recorded 1 3 species in her "Atlas of the Sedentariate 
Polychaetous Annelids from California." 

The four new species are described herein and 
information is presented on their distribution and 
general ecology. In addition, some information is 



provided on reproduction in P. elegantissima. 

The holotypes and one set of paratypes are 
deposited in the Allan Hancock Foundation, Uni- 
versity of Southern California. Additional para- 
types are deposited in the United States National 
Museum. Washington, D.C. 



■Pacific Marine Station, University of the Pacific, Dillon Beach, 
California 94929. 

2 Department of Biology, Fresno State College, Fresno, California 
93710. 



1971 



NEW SPECIES OF POL YDORA 






Genus Polydora Hose, I N02 
Polydora convexa, new speeies 
Figure l 
Material examined: California: Santa Barbara, 
August 27, 1961 (I), from shells of hermit crabs; 
Avila, March 3, 1962 (8, TYPE), from shell of 
Pododesmus macroschisma; Mono Hay, October 
24, 1961 (4), May IK, 196! (13), from hermit erah 
shells and Pododesmus; Cayucos, June 2K, 1961 
(5), August 28, 1961 (13), December 19, 1961 (4). 
March 3, 1962 (4), July 3, 1962 (9), from hermit 
crab shells, encrusting sponge, holdfast of Mac ro- 
cys/is, and Dodecaceria colonies; Bodega Harbor, 
March 3, 1970 (3), from Podoilesmus; Bodega Bay, 
September 30, 1970 (6), from Bryozoa, dredged in 
20 m; Eureka (Trinidad Head), June 21, 1962 (10), 
from Balanus and hermit crab shells. 
Description: Specimens of P. convexa from Bodega 
Harbor which were relaxed prior to preservation 
are largest in the collections and measure up to 30 
mm in length and have over 200 segments. Speci- 
mens collected from other localities are somewhat 
smaller, ranging in size from 4.2 mm (45 segments) 
up to 15 mm (160 segments), Live individuals are 
light tan with occasional dark pigment on anterior 
body segments. This pigment may be a vestige of 
larval pigmentation. 




Figure I. Polydora convexa, new species: a, anterior 
end in dorsal view; b-c, spines from setiger 5; d. pen- 
noned companion seta from setiger 5; e. hooded hook 
from an anterior setiger; f, hooded hook from a poste- 
rior setiger; g, specialized notopodial structure from a 
posterior setiger; h, posterior setiger in anterior view 
with specialized notopodial structure in place; i-j. 
posterior segments in dorsal view. 



I he prostomium it bifid on il ant* rioi n 
(Fig. la). I he i arum le i • tends to ihi 
border of setiger 4. ["hen il no nuchal i.'iii.i- I. 
Eyes may be absent, bui when presenl Ihi four cyc» 
are arranged in a trapezoidal mannci with the 
posterior pair closer together and imallci than the 
anterior pair. I he palps extend posteriori) onl) i" 
setiger 10 in preserved specimens; in life [hi 
longer and prehensile. 

Setiger I has capillary setae in both I 
neuropodia. Setigers 2, 3, 4, . 6 and succeeding 
setigers have well-developed notopodial lobes pre- 
ceded by posteriorly directed winged capillary setae 
arranged in three rows; an anterior row oi 
curved capillary setae, a second row ol longer, less 
curved capillary setae, and a third row ol Ion 
capillaries. The number of these setae per notopo 
diuni rapidly diminishes until in posterior segments 
only a few long slender capillary notosetae remain 
In the posterior one-half of the body these capillary 
setae are accompanied in each notopodium h\ an 
unusual setal structure which is difficult to interpret 
(Fig. lg-h). These structures do not protrude 
through the cuticle and resemble tight bundles of 
setal fibers. They are undoubtedly similar to the 
"organesen filieres" of Claparede (Mcintosh. 1915) 
found in Spiophanes bombyx Claparede. The 
neuropodia of setigers 2, 3, 4. _. and 6 have pos- 
teriorly directed capillary setae. Hooded hooks 
begin on setiger 7. They are bidentate (Fig. lei in 
anterior setigers with the secondary tooth gradually 
becoming smaller until it disappears entirely in 
posterior setigers. This results in a unidentate hook 
(Fig. If). The change-over from bidentate to uni- 
dentate is variable but usually takes place near the 
midbody region. In anterior setigers there are four 
or five hooks per neuropodium while in posterior 
setigers there are only 1-2 hooks. The hooks are 
accompanied by 1 or 2 very fine capillary setae 
throughout. 

Setiger 5 is larger than preceding and succeeding 
setigers (Fig. la). The setae include a small bundle 
of dorsal geniculate setae King anterior to a semi- 
circular row of alternating heavy spines (Fig. lb-c) 
and smaller pennoned companion setae (Fig. ldi. 
The heavy spines are falcate and have a broad 
collar on the convex side (Fig. lb-d. Veniral and 
posterior to the major spines is a small tuft of 
winged setae. 

The slightly tapered and strap-shaped branchiae 
begin on setiger 8 (Fig. la). They are small at first 
and reach full size on about setiger 15. Branchiae 
are absent from the posterior two-thirds of the body. 

The pygidium has four lobes (Fig. li. j). The 
dorsal pair are smaller than the ventral pair. The 
anus is located at the center of the lobes. 



74 



HI Ll.EIIN SOI I HERN CALIFORNIA ACADEMY Ol SCIENCES 



VOLUME 70 



A structure similar in appearance to the "gizzard- 
like" structure previously described for P. social is 
(Blake, 1969, 1971) was noted in P. convexa. It 
occurs at about setiger 18 on a 100 segment speci- 
men. The structure does not appear to be as 
muscular or as well developed as in /'. socialis. A 
similar type of structure was observed in P. flava 
by Carazzi (1893). 

Remarks: Polydora convexa is unusual in having 
the accessory structure, in this case a flange, on the 
convex side of the heavy falcate spines of setiger 
5. It is similar to Polydora langerhansi Mesnil 
(1896) declared inceriae sedis by Fauvel (1927). 
Rioja (1925) had reported P. langerhansi from 
Madeira and later from Acapulco (Rioja. 1939, 
1943). Mesnil reported no specialized notopodial 
setae in the posterior region, did not describe the 
pygidium, found no eyes, and made no mention of 
the unidentate hooded hooks in the posterior neuro- 
podia. The spines of setiger 5 also resemble those of 
P. armata (Mesnil noted" this for P. langerhansi) but 
members of the 2 species are not likely to be 
confused. 

Ecology: Polydora convexa was found at Santa 
Barbara, Avila. Morro Bay and Cayucos in central 
California, and at Bodega Harbor and Trinidad 
Head in northern California. This species was 
found in many different habitats, including gastro- 
pod shells inhabited by hermit crabs. At Trinidad 
Head and Cayucos, P. convexa occured in shells of 
Tegula brunnea with Pagurus granosimanus; T. 
brunnea with P. samuelis; T. funebralis with P. 
samuelis; Olivella biplicata with P. granosimanus; 
and at Santa Barbara in shells of O. biplicata with 
P. samuelis. Other habitats include Pododesmus 
macroschisma shells; Diadora aspera shells; scrap- 
ings of rocks from surf zone; Macrocystis pyrifera 
holdfast; Dodecaceria colonies; and encrusting 
red sponges. It was found in the hermit crab habitat 
with a new species of Boccardia and was associated 
with Polydora ciliata and B. columbiana in nearly 
all of the above habitats and in several of them with 
B. tricuspa. Bryozoa dredged from 18 m off 
Bodega Head contained P. convexa, a new species 
of Polydora, described herein, and Boccardia 
berkeleyorum. 

In Pododesmus shells at Bodega Harbor, P. 
convexa formed the branching type of burrow 
which is similar to that described by Evans (1969) 
for P. concharum which bores into shells of Placo- 
pecten magellanicus (Gmelin) in New England. 
Distribution: California; Santa Barbara to Eureka. 

Polydora elegantissima, new species 
Figure 2 

Material examined: California; Tomales Bay, 



March 30, 1970, intertidal from a hermit crab shell 
(I); Morro Bay, December 31, 1963, intertidal 
from shells of livela stultorum (Mawe) (8, TYPE); 
Malibu Beach. August 20. 1964. Subtidal in 2 m. 
from shells of hermit crabs (6). 
Description: A complete specimen from Tomales 
Bay measures 3 I mm in length, is 1 mm wide and 
has 170 setigerous segments. It was light tan (alive) 
with palps which were long and darkly pigmented 
along the margins of the ventral groove. There was 
no other body pigmentation. The specimen was 
first seen as it extended the long black palps from 
the shell of the hermit crab. The palps contracted 
greatly upon preservation. The holotype comes 
from Morro Bay, is incomplete, has 280 segments 
and is 44 mm long. 

The prostomium is divided into two widely 
divergent lobes which on the Tomales Bay specimen 
resemble anterior horns (Fig. 2a). The caruncle 
extends prominently back to setiger 3 or 4 and 
thereafter continues as a low nuchal ridge for 30 or 
40 segments; it may be indistinct on setiger 5. There 
are no eyes. 

Setiger 1 has capillary setae in noto- and neuro- 
podia. Setiger 2. 3, 4, _, 6 and succeeding setigers 
contain spreading fascicles of winged capillary 
notosetae arranged in 3 rows, each row with pro- 
gressively longer setae. In far posterior setigers 
these setae diminish in number and length. There 
are no specialized posterior spines. The notopodial 
lobes of setiger 1 are shorter and more finger-like 
than those of 2, 3, and 4, which are large and 
auriculate. The lobes of setiger 6 and following 
setigers are shorter and more finger-like, the length 
gradually diminishing in middle and posterior 
setigers. The neuropodial lobes of setiger 2, 3, and 
4 are also ear-like, although smaller than the 
notolobes, and they also diminish in size posterior 
to setiger 6. The neuropodia of setiger 2, 3, 4, _, 6, 
7, 8, and 9 contain spreading fascicles of winged 
capillary setae which are arranged in 2 groups; a 
large dorsal fascicle and a smaller ventral group. 
Replacement of the winged capillary setae with 
bidentate hooded hooks takes place over the next 
few setigers. Hooded hooksbegin on setigers 10, 11, 
or 12, but usually on 1 1. There are 5-7 hooks per 
neuropodium and only a few capillary setae. These 
remaining capillary setae are derived from the 
smaller, more ventrally situated fascicle which 
occurs on setigers 6, 7, 8, and 9. The capillary setae 
are absent about setiger 18, but are present again in 
the posteriormost setigers. The hooks have no 
constriction on the shaft and retain the same 
".tructure throughout the length of the body (Fig. 
2b). 

Setiger 5 is modified and larger than other 



1971 



NEW SPECIl s 01 POl ) DORA 



75 




Figure 2. Polydora elegantissima, new species: a. ante- 
rior end in dorsal view; b, hooded hook; c-h, heavy 
spines from setiger 5 showing different angles and 
degrees of wear; i, superior dorsal seta of setiger 5; j. 
companion seta of setiger 5; k-1, neurosetae from 
setiger 5; m, posterior end in dorsal view. 



setigers (Fig. 2a). The setae include a dorsal fascicle 
of petitioned setae (Fig. 2i) lying at the anterior end 
of a semicircular row of heavy modified spines 
(Fig. 2c-h) alternating with smaller pennoned com- 
panion setae (Fig. 2j). The deeper unworn heavy 
spines are sharply falcate and have a lateral sheath 
(Fig. 2c-d). Older worn spines are broader and 
more blade-like as a result of the erosion of the 
strongly falcate distal portion (Fig. 2g-h). Ventral 
to the row of heavy spines and companion setae is 
a neuropodial fascicle of stout geniculate setae 
(Fig. 2k- 1). Some of these setae appear to be worn 
and have blunt frayed ends. 

Branchiae begin on setiger 8 (occasionally 7) 
and continue to near the posterior end. They are 
relatively short and do not meet at the mid-line. The 
pygidium has 4 nearly equal lobes (Fig. 2m). 
Reproduction: Egg capsules were collected with 
the Morro Bay specimens. Each capsule contained 
about 80 eggs and each egg measured about 150/i 
in diameter. Cleavage and larval movement were 
observed with non-pigmented early larvae following 
cleavage by about 45 hours. The culture subse- 
quently became infected with bacteria. 
Remarks: Polydora elegantissima is closely related 
to P. commensalis. It differs from them in the 
prostomium, caruncle, modified spines of setiger 5. 
and in the arrangement of the branchiae. 
Distribution and Ecology: In Tomales Bay the 



specimen was found in a shell ol Ollvella blpllcata 
occupied by the hermit crab Paguru manut 

At Mono Bay it occurn d in hi Hi of 7 
stultorum .end al Malibu Beach in shells with 

hei init crabs. 



Polydora bioccipitalis, new species 
I igures I and 4 

Material examined: ( alifornia; Malibu Beach (22. 
TYPE), August 20. 1964 from hermil cral 
lected by James McHcnry; Santa Barbara (I), July 
30, 1965 from a shell ol Olivella hi/iluula occupied 
by a hermit crab 

Description of the adult: Since both adults and 
post-larval juveniles arc present in the collections 
and there are some features common only to the 
juveniles, the 2 groups will be treated separately. 
Adults measure up to 18 mm in length and have- 
about 1 20 segments. There is no body pigmental ion 
The prostomium is deeply notched on the ante- 
rior margin (Fig. 3a). The nuchal ridge extends 
past setiger 5 on most specimens and is visible on 
many anterior segments. On a few specimens a 
definite caruncle reaches only to the posterior 
margin of setiger 2. Two nuchal tentacles are 
present on large well-preserved specimens (Fig. 3a). 
The tentacles are well developed and one precedes 
the other. There are 4 eyes, rounded in shape: 
posterior pair is more closely spaced than the 
anterior pair. Palps are longer than those of Poly- 




Figure 3. Polydora bioccipitalis, new species: a. anterior 
end in dorsal view: b. hooded hook: c. fascicle of heavj 
spines and companion setae from setiger 5; d. posterior 
end in dorsal view. 



■v, 



HI I /./•. /7,Y .SO/ / ///-.K.V C.I I.I FORMA ACADEMY OF SCIENCES 



VOLUME 70 



iloru commensalis. but still relatively short when 
compared with most species of the genus. The 
Santa Barbara specimen was observed live and the 
first five setigers were narrower and showed less 
color (less blood) than succeeding setigers. When 
preserved, the anterior end contracted greatly and 
this combined with the shortening of the palps 
altered the specimen, causing it to appear fore- 
shortened. Setigers 1. 2. 3, 4 have well developed 
notopodial lobes. 

Setiger I lacks notosetae. Setigers 2. 3, 4, _, 6 
and succeeding setigers have fascicles of winged 
capillary setae. These setae are arranged in 2 rows 
with the longer setae in the posterior row. In 
posterior setigers the setae consist of a bundle of 
long laterally directed capillaries giving that part 
of the body a somewhat spinous appearance (Fig. 
3d). 

The neuropodia of setigers 2. 3, 4, _, 6, 7, 8 
have spreading fascicles of winged capillary setae 
arranged in the same manner as the notosetae. 
Bidentate hooded hooks usually begin on setiger 
9: however, on other individuals the hooks do not 
appear until setigers 10 to 14. Anteriorly there are 
2 or 3 hooks accompanied by capillary setae. 
Posteriorly the hooks increase to 16 per neu- 
ropodium and the capillary setae disappear. The 
angle between the main fang and the secondary 
tooth is not great and there is only a faint suggestion 
of a constriction in the shaft (Fig. 3b). 




Figure 4. Polydora bioccipilalis, new species, juvenile: 
a. anterior end in dorsal view; b, group of setae from 
setiger 5; c. posterior end in dorsal view. 



Setiger 5 is larger than other setigers. The setae 
include a curved row of heavy spines alternating 
with pennoned companion setae (Fig. 3c). There- 
is no anterior dorsal fascicle of setae and the ventral 
fascicle is lacking in the largest specimens, but is 
present in the smaller adults. The heavy spines arc 
falcate and have 3 accessory structures. A thin fin 
or sheath lies between the distal end of the spine 
and a large accessory tooth. Both the tooth and I'm 
may be so eroded as to be unrecognizable in older 
setae. A third accessory structure is a small trian- 
gular shaped tooth on the side of the spine. This 
small tooth can be seen only when the spine is 
viewed at certain angles. 

The broad and flattened branchiae begin on 
setiger 7 and continue to near the posterior end 
(Fig. 3a, d). 

The pygidium is a single thickened disk with a 
dorsal gap (Fig. 3d). The Santa Barbara specimen 
has in addition a slight ventral notch. 
Description of the Juvenile: Post larval juveniles 
present in the collections range in size from 2.5 mm 
to 4 mm and from 28 to 36 segments. The striking 
difference between these juveniles and the large 
adults is the retention of larval pigmentation (Fig. 
4a, c). The basic pattern of this pigment is the 
presence of lateral rows of black spots which begin 
on setiger 1 and continue to near the posterior end. 
Setiger 5 is unpigmented. A central row of chroma- 
tophores is found on the middle segments of the 
body and continues to the posterior end. Scattered 
pigment on the dorsal surface of setiger 7 and 
succeeding setigers suggests that in larvae the 
central row of chromatophores appeared on more 
anterior segments. Some individuals have additional 
pigment spots on the peristomium and pygidium. 
The pygidial pigment may form 2 dark spots on 
either side of the dorsal gap. 

The anterior margin of the prostomium is broad 
on the small juveniles but shows signs of a bifurca- 
tion on the larger individuals. The nuchal ridge 
extends into setiger 2. Only one nuchal tentacle 
was found in juveniles (Fig. 4a). This suggests that 
the development of the two nuchal tentacles may 
be related to the age of the specimen and the 
posterior growth of the nuchal ridge or caruncle. 
The anterior pair of eyes tend to be cup-shaped 
in the juveniles. 

Setation patterns are similar in juveniles and 
adults. The hooded hooks do not exceed 7 per 
neuropodium. Capillary setae are retained with the 
hooks on a greater number of setigers. 

In juveniles there are two additional setal types 
found among the modified spines of setiger 5 (Fig. 
4b). The first is a large curved spine with no 
accessory structures. The second is a more slender. 



1971 



NEW SPECIES OF POL I ix>KA 



77 



sharply hooked seta. The remaining heavy spines 
arc as in the adults. A similar situation has been 
found in other polydorids (Blake, 1969) and it is 
suggested that the two additional setal types are 
hold overs from the larvae. The heavy larval setae 
are the first to be formed and arc among the first to 
be worn down and replaced. Replacement is with 
the type of spine described for the adult. The 
branchiae and pygidium are as described for the 
adult. 

Remarks: Polydora bioccipitalis is unusual in the 
possession of two nuchal tentacles. Polydora macu- 
lata (Day, 1963) is the only other species of the 
genus which is known to have this feature, although 
several species have one nuchal tentacle. Polydora 
maculata is similar to P. bioccipitalis in other 
characteristics as well, including the lack of noto- 
sctae on setiger 1, the appearance of hooded hooks 
on setiger 9, and the lack of dorsal and ventral 
fascicles on setiger 5. The two species differ in the 
structure of the heavy spines of setiger 5 and the 
length of the nuchal ridge. The heavy spines of 
maculata have only a lateral flange whereas, bioc- 
cipitalis has three separate structures including two 
teeth and a flange. 

The morphology of P. bioccipitalis. especially 
the juveniles, is similar to the description of P. 
punctata from El Salvador (Hartmann-Schroder, 
1959). That species was described from juveniles 
which resemble those of P. bioccipitalis but the 
nuchal tentacles are not described nor figured. 
Further, the small lateral tooth of the specialized 
spines was not described. 

Distribution and Ecology: At present, Polydora 
bioccipitalis is known only from southern Cali- 
fornia at Malibu Beach and Santa Barbara. The 
species was found in gastropod shells occupied by 
the hermit crab, Pagurus hirsutiusculus. The shells 
included Ocenebra poulsoni, Olivella biplicata, 
Murex gemma and Polinices reclusianus at Malibu 
Beach where the species was found in association 
with other polydorids including P. elegantissima, 
P. commensalis, and Boccardia tricuspa. At Santa 
Barbara P. bioccipitalis was found in a shell of 
Olivella biplicata occupied by Pagurus hirsutiuscu- 
lus where it formed a shallow burrow which was 
covered dorsally with silt and mucous. This type of 
burrow is also formed by P. commensalis (Blake, 
1969). Polydora commensalis is known to be a 
commensal of hermit crabs and since P. bioccipi- 
talis has a similar habitat and burrow, it seems 
possible that P. bioccipitalis may also have a com- 
mensal relationship with hermit crabs. Other poly- 
dorids found in this habitat at Santa Barbara 
included B. proboscidea, P. commensalis. and P. 
ciliata. 



Polydora pygidialis. ... ■. 
I igun 

Polydora t illaia Berkeley and Berl 
1952: 19-20 (not Johnston, 1838) 
Material examined. California Santa Barbara, 
August 27, 1961 (4), from innei harboi piling 
material; Avila, March J, 1962 (3), from hermit 
crab shells; Mono Bay, Octobei 24, 1961 
May IX. 1963 (6); ' ayucos, June 28, 1961 
August 28, 1961 (23.TYPI I, Decembct 19 
(19), July 3, 1962 il). from hermit crab ihelb 
keyhole limpet shell, and piling material B 
Bay. September 2X. 1970 (10), from bl 
dredged from IS m. British ( olumbia; Departure- 
Bay, circa 1936 (4). from hermit crab shells (B 
ley and Berkeley, 1936). 

Description: The individuals of this species .ire 
very slender. A specimen 9 mm long and having \<K> 
segments is less than 0.5 mm in width. The anterior 
and posterior ends are slightly dusky in appearance 
but there is no strong body pigmentation. 

The rounded prostomium (Fig. 5a) continues 
posteriorly as a narrow caruncle to the posterior 
margin of setiger 2. Palps on preserved specimens 
extend posteriorly to about setiger 10. Eyes may 
be absent but when present, the number is variable 
to a maximum of four. The anterior pair are w idely- 
spaced and cup-shaped: the posterior pair are close 




Figure 5. Polydora pygidialis. new species: a. anterior 
end in dorsal view: b. hooded hook; c-d. fascicles of 
heavy spines and companion setae from setiger 5; e. 
individual heavy spine from setiger 5; f. superior dorsal 
setae from setiger 5: g. neurosetae from setiger 5; h. 
posterior end in dorsal view; i. posterior end in lateral 
view. 



78 



HI III 1 1\ SOI I III l<\ C.I I HORN I I K \DEM\ <>l S( II NCES 



VOLUME 70 



together and irregularly shaped. In some specimens 
only the anterior pair are present. 

Setiger I laeks notosetae but has a small conical 
notopodial lobe (Fig. 5a). The neuropodium of 
that setiger has a small fascicle of slender capillary 
setae and a bluntly rounded lobe. Setigers 2, 3, 4. _, 
6 and succeeding setigers have well developed 
posteriorly directed spreading fascicles of winged 
capillary notosetae in two rows; setae of anterior 
row are shorter and slightly bent. This same general 
setal arrangement persists throughout the length 
of the body. There are no specialized posterior 
notosetae. The neuropodia of setigers 2. 3, 4, _, and 
6 include fascicles of winged capillary setae. Biden- 
tate hooded hooks begin on setiger 7 (Fig. 5b) and 
are not accompanied by capillary setae. There are 
four or five hooded hooks anteriorly with an 
increase to 10 in median setigers and a reduction to 
the original number posteriorly. The main fang of 
the hook is almost at a right angle to the shaft which 
has a constriction about one-half the way down 
its length (Fig. 5b). 

The modified fifth setiger contains three groups 
of setae; a superior dorsal bundle of heavy, pointed 
setae (Fig. 50. a curved row of large modified 
spines and alternating pennoned companion setae 
(Fig. 5c-d), and a reduced neuropodial fascicle of 
small, pointed setae (Fig. 5g). The heavy spines are 
falcate and have a large lateral accessory tooth 
whose apex curves back toward the main axis of 
the spine (Fig. 5c-d). The branchiae which begin 
on setiger 7 are flattened but are fingerlike in 
outline (Fig. 5a). They are short on anterior setigers 
but increase in length and nearly meet at the 
mid-line on about setiger 15. They are absent from 
the posterior one-third of the body. 

In living specimens the pygidium is strongly 
scoop-shaped (Fig. 5h-i). Preservation may alter in 
part the shape of the scoop but the broad terminal 
end characteristic of the species remains easily 
identifiable. 

Remarks: Polydora pygidialis is most closely related 
to P. rickettsi Woodwick (1961) from Lower Cali- 
fornia. It is distinguished from that species by the 
length of the caruncle, structure of the pygidium 
and the modified spines of setiger 5. In California, 
P. pygidialis is easily confused with P. websteri 
with which it may occur. The new species is 
distinguished from P. websteri in that the latter has 
a bifid prostomium, and the modified spines of 
setiger 5 have only an accessory flange. Polydora 
pygidialis is also similar to Polydora limicola 
Annenkova but in the latter the prostomium is 
vaguely incised and more importantly the palps are 
crossed by four or five bars of black pigment 
(Hartman, 1961). 



Ecology: Polydora pygidialis was taken at Santa 
Barbara, Avila, Morro Bay and Cayucos in south- 
ern and central California. Most specimens were 
found in hermit crab shells, especially from the 
Tegula funebralisl Pagurus granosimanus relation- 
ship. It was also found in piling material in the 
warmer waters of the inner harbor at Santa Barbara 
and the more open pilings in the colder waters at 
Cayucos. It was commonly associated with Boc- 
cardia Columbiana. In the piling material from 
Cayucos it was also associated with Polydora 
websteri, a morphologically similar species. Bryo- 
zoa dredged from 18 m off Bodega Head contained 
Polydora pygidialis, P. convexa, and Boccardia 
berkeleyorum. Polydora pygidialis is a boring and/ 
or nestling form. Egg capsules were found with the 
Bodega Bay material in September. The larvae are 
currently being studied by the first author. 
Distribution: California (Santa Barbara) to British 
Columbia (Departure Bay). 

LITERATURE CITED 

Berkeley, E., and C. Berkeley. 1936. Notes on Poly- 
chaeta from the coast of western Canada. I. 
Spionidae. Ann. Mag. Nat. Hist. Ser. 10, 18:468- 

476. 

Blake, J. A. 1969. Reproduction and larval develop- 
ment of Polydora from northern New England. 
Ophelia, 7: 1-63. 

. 1971. Revision of the genus Polydora from the 



east coast of North America. Smiths. Contrib. 
Zoo/. 75: 1-32. 

Carazzi, D. 1893. Revisione del genero Polydora Bosc, 
e cenni su due specie che vivono sulle ostriche. 
Mitteilnngen aus der Zoologischen Station zti 
Neap el. 11: 4-45. 

Day, J. H. 1963. The polychaete fauna of South Africa. 
Part 8. New species and records from grab samples 
and dredgings. Bull. British Mus. Nat. Hist. 
(Zool.), 10: 383-445. 

Evans, J. W. 1969. Borers in the shell of the sea scal- 
lop, Placopecten magellanicus. Amer. Zool., 9: 

775-782. 

Fauvel. P. 1927. Polychetes sedentaires. Addenda aux 
errantes, archiannelides, myzostomaires. Faune 
de France, 16: 1-494. 

Hartman, O. 1961. Polychaetous annelids from Cali- 
fornia. Allan Hancock Pacific Exped., 25: 1-226. 

. 1969. Atlas of the sedentariate polychaetous 

annelids from California. Allan Hancock Found., 
Univ. So. California. Los Angeles, 812 pp. 

Hartmann-Schroder, G. 1959. Zur Okologie der Poly- 
chaeten des Mangrove-Estero-Gebietes von El 
Salvador. Beitrage zur Neotropischen Fauna. 1: 
69-183. 



1971 



NEW SPECIES <>l roi.YDoKA 



7V 



Johnston, (i. I K.IK. Miscellanea Zoologica. Ariciadac. 
Mag. Zool. Hoi.. 2: 63-73. 

Mcintosh, W. C. 1915. A Monograph of the British 
murine annelids. Polychaetu, Opheliidue to Am- 
phictenidae. Kay Soc, London. 3: I !68. 

Mcsnil, F, 1896. Etude ilc morphologic cxlcrnc the/ les 
Annclidcs. I cs Spionidicns des cotes dc la Marchc. 
Bulletin Scientifique tic In France et tic In Belgique, 
29: I IO-2K7. 

Rioja, E. 1925. Anelidos poliquetos dc San Vicente de 
la Barquera (Cantabrico). Trabajas del Mttseo 
Nacional dc Ciencias Naturales. Serie Zoologica, 
53: 1-62. 



19 19 i tudio im lidologico i 
accn a 'i' at ias lot ma li 
pciagii ' di Spionidai pro i di nt< 
con description de mm;, . • p. . j< n n 
Polydora Annulet del I" llmio di Blologla, Hi 
297-3 1 I 

,1943 Estudios anclidologico \ill 

acerca de las especies del genera Pol dot B 

de las coslas Mcxicanas del Paciflco .\t,n,,l 
tnstitutode Blologla, 14: 229 2-1 1 

Woodwick. K. H. 1961. Polydora ml, it, , t new 
species of spionid polychaete from Lowei Cali 
fornia. /'<« ifii Sci., 15: 7K-KI. 



Accepted for publication February IK. 1971. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(2): 79-80, 1971 

A NEW SPECIES OF HUNTEROTREMA (DIGENEA: CAMPULIDAE) 
FROM THE AMAZON RIVER DOLPHIN (INI A GEOFFRENSIS) 

Murray D. Dailey 1 



Abstract: A new species of Hunterotrema (Digenea: Campulidae) is 
described from the lungs of the Amazon river dolphin Unto geojfrensis). It 
differs from the single species in this genus, H. caballeroi. in body size, the 
lack of cuticular spines, placement of genital pore, and size of cirrus sac. 



During investigations on marine mammal hel- 
minths, numerous lung trematodes taken from the 
Amazon river dolphin (Inia geoffrensis) were given 
to me for identification by Dr. Sam Ridgway, Naval 
Undersea Research and Development Center, Point 
Mugu. California. The specimens were found to be 
similar to, but much larger than, Hunterotrema 
caballeroi Mcintosh 1960. Mcintosh (1960) de- 
scribed H. caballeroi from 3 entire worms and 2 
fragments of 2 additional specimens. The type 
material and additional specimens were obtained 
on loan from the USNM Helminth Collection. 
Beltsville, Maryland. A comparison of those 6 
specimens not designated as part of the type-series 
(USNM Helm. Coll. Nos. 56921, 56922) with this 
material indicated a similarity between these forms. 
Both groups differed from H. caballeroi sufficiently 
to warrant a new species description. 

The worms from Dr. Ridgway had been fixed in 
10% formalin, whereas, those received from Belts- 



ville were in 70% ethanol. Whole mounts uere 
stained in Semichon"s carmine or celes^ine blue B. 
dehydrated in ethanol. cleared in xylene and 
mounted in piccolyte. Drawings were made with 
the aid of a drawing tube. All measurements are 
given in millimeters unless otherwise stated. A\er- 
age measurements are presented with ranges m 
parentheses. 

Hunterotrema macrosoma, new species 

Figures 1-5 

Description based on measurements from 15 
specimens. 

Diagnosis: Body slender, elongate. 31 1 1 ■ 
long, with distinct forebody 3.7 (3.1-4.7). mid-body 
19.6 (12.2-25.4) and hindbody 7.S (6.0-9. S). Maxi- 
mum body width 1.1 (0.74-1.46) at acetabular level, 
hindbody 1.0 (0.53-1.5) in region anterior testis. 



'Department of Biology. California Stale College. Long Bea^i. 
California 90801. 



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VOLUME 70 




Figures 1-5. Hunterotrema macrosoma, new species. 
Fig. I. Ventral view of entire worm. Fig. 2. Forebody. 
Fig. 3. Mid-body. Fig. 4. Egg. Fig. 5. Hindbody. Abbre- 
viations: c — cirrus; cs — cirrus sac: gp — genital pore; 
i — intestinal ceca; lp — lateral pockets of esophagus; 
o — ovary; rs — receptaculum seminis; sv — seminal 
vesicle; t — testis; u — ulerus; v — vitellaria. 

Spines lacking. Oral sucker, well developed, cir- 
cular and subterminal 0.34 (0.30-0.40) in diameter. 
Prepharynx short, pharynx 0.32 (0.30-0.36) long by 
0.19 (0.17-0.24) wide. Esophagus expanded ante- 
riorly into two lateral pockets, 0.35 long (from 
midesophagus) by approximately 0.20 wide at 
ends (variable). Acetabulum, well developed, cir- 
cular, 0.58 (0.50-0.67) in diameter. Cirrus long, 
unarmed. Cirrus sac 2.0 (1.8-2.1) long by 0.52 
(0.43-0.60) wide, extending less than half its length 
posterior to acetabulum. Ovary oval, just cephalad 
to anterior testis, 0.33 (0.28-0.46) long by 0.35 
(0.30-0.50) wide. Receptaculum seminis large, lat- 
eral to ovary. Seminal vesicle large, genital pore 
anterior to acetabulum just posterior to cecal 
bifurcation. Testes confined to median and posterior 
half of hindbody. Anterior testis 0.92 (0.65-1.15) 
long by 0.69 (0.48-0.98) wide. Posterior testis 1.03 
(0.75-1.29) long by 0.71 (0.52-0.88) wide. Vitel- 
laria, uniformly distributed, confined to hindbody. 



extending from posterior end of intestinal cccae to 
anterior constriction of hindbody. Uterus coiled 
immediately anterior to ovary, then straightens to 
run as a sinuous tube to genital pore. Eggs oval 
108^ (105-1 I2)by62/i (58-66). 

Host: Inia geoffrensis 

Location: Lungs 

Locality: Amazon river basin, Leticia, Columbia 

and Iquitoz, Peru. 

Holotype and Paratypes: USNM Helm. Coll. Nos. 

71583 (holotype), 71584, 56921, 56922. 

DISCUSSION 

Currently the genus Hunterotrema is represented 
by a single species, H. caballeroi Mcintosh, 1960. 
Hunterotrema macrosoma differs from H. cabal- 
leroi in the following characters: (1) length and 
width of body {H. caballeroi 13.07 long by 1.64 
maximum width); (2) lack of cuticular spines (H. 
caballeroi heavily spined from anterior to region 
of reproductive organs); (3) placement of genital 
pore (immediately anterior to acetabulum in H. 
caballeroi); (4) cirrus sac not extending more than 
half its length posterior to acetabulum as in H. 
caballeroi. 

Since both familiar (Campulidae Odhner, 1926) 
and generic diagnosis included the presence of 
spines, an emendation to both is proposed to 
include H. macrosoma and should read "Cuticular 
spines present, or lacking." 

Woodard et al., (1969) described the pathology 
of pulmonary trematodiasis in an Amazon river 
dolphin from Iquitoz, Peru. The photograph and 
measurement (printed, in error, as "approximately 
250 mm in length" but corrected by Dr. Stephen 
Zam, University of Florida [pers. comm.] to be 
25 mm) indicates the trematodes involved in the 
report were H. macrosoma, not H. caballeroi. 

ACKNOWLEDGMENTS 

I would like to express my appreciation to Sam Ridg- 
way of NUC for collecting the specimens, and to the 
late W. W. Becklund for the loan of specimens from 
the USNM Helminthological Collection. Beltsville, 
Maryland. My sincere thanks to Lorraine Peterson 
and Barry Hill for their technical help. 

LITERATURE CITED 

Mcintosh, A. 1960. A new campulid trematode, Hunt- 
erotrema caballeroi n.g., n. sp. from an Amazon 
Dolphin, Inia geoffrensis. Libro Humanje al Dr. 
Eduardo Caballero y Caballero Jubileo 1930-1 960. 
Instituto Politecnico National Mexico, 207-208. 

Woodard, J. C, S. G. Zam, D. K. Caldwell, and M. C. 
Caldwell. 1969. Some parasitic diseases of dol- 
phins. Path. Vet.. 6: 257-272. 

Accepted for publication April 22. 1971. 



BULLETIN SO. (ALU. ACADEMY OF SCIENCE 70(2):81-84, W7I 



A NEWSUIiSIM'CIHSOl- FUNNI-I -HARI-DBA1 (NATALUS STRAMINl [ 
FROM WESTHRN VI Nl /I II, I A 

Omar J. Linares' 



Abstract: a new subspecies of the funnel-eared bal Natalus stramineut 
is described from a cave in the Guasare river, Zulia, Venezuela On the I asisof 
palatal Structure, the 3 recognized subspecies of N. major are assigned In V 
stramineus. The species status of N. tumidiroslris and its possible synonymy 
with N. stramineus are also discussed. Additional records of N. t. contint nli 
from northern Venezuela are included. 



In October 1967, an expedition of the Venezuelan 
Society of Speleology collected four specimens of a 
unique funnel-eared bat in a cave in the state of 
Zulia, Venezuela. These proved to be the first 
specimens of Natalus stramineus taken in Venezuela 
and northern South America. Comparison with 
material from Central America indicates that the 
Venezuelan bats represent a distinct subspecies, 
which is named and described below. 

Natalus stramineus tronchonii, new subspecies 

Holotype: An adult male, ale. with skull. Biology 
Museum, Central UniversityofVenezuela, MBUCV 
1-1578, collected by J. A. Tronchoni (original 
number L-281) from Gavilanes cave (Zu-1, Soc. 
Venezolana Espel.. 1968: 1 13-1 18), Guasare river. 
Zulia, Venezuela, 183 m, 20 October 1967. 

Paratypes: Three adult females, ale. with skulls, 
MBUCV 1-1577, 1-1579 and 1-1580 of the same 
locality, date and collector. 

Distribution: Known only from the type locality. 

Diagnosis: A small subspecies with the posterior 
border of the bony palate deeply emarginate to the 
level of the last molar. Very similar to N. s. mexica- 
nus Miller [ = N. s. saturatus Dalquest and Hall, 
1949], but smaller, and morphological characters 
intermediate between tumidirostris and stramineus. 

Description: Body size small (forearm 39.7 mm; 
greatest length of skull 16.1 mm). Dorsal coloration 
after one year in alcohol (capitalized color terms 
after Ridgway, 1912) Cream Buff, tips of hairs 
gradually darkened to a Pale Drab, underparts 
more yellowish than back. Vibrissae abundant and 
conspicuous on snout. Ears and lips grayish. Mem- 
branes, feet and legs, dark brown. Other somatic 
characters as in N. stramineus. Skull strictly similar 
to that of stramineus type, but with a palate inter- 
mediate in size between stramineus and tumidi- 





Figure I. Palatal emargination and dentition of upper 
right jaw of: a. Natalus stramineus tronchonii ( MBUCV 
1-1578. <? — holotype). and b. Natalus stramineus 
mexicanus (TCWC 8507. 9). 



rostris. Upper molar w ith the hypocone w idest 
lingually. The differences in the palate and dental 
characters between N. s. tronchonii and N. s. mexi- 
canus are shown in Figure 1 . 

Measurement: The holotype and the paratypes 
are similar (Fig. 2: Table 1). 



'Laboratorio de Vertebrados. Instituto de Zoologia Tropica). 

Universidad Central de Venezuela. Caracas. Api ' 

zuela (Present address: 3. Rue des Ecoffes. Paris-IV c -France). 



81 



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HI 7 / / 7//V SOUUIEKN CALIEORNIA ACADEMY OF SCIENCES 



VOLUME 70 



Table I. Measurements (mm) of Natalus stramineus tronchonii from Venezuela compared with selected 
measurements o( Natalus stramineus mexicanus from Mexico. Nicaragua and Guatemala. 







N. s. tronchonii 




N. 


v. mexicanus 






1577 


1578 1 


1579 


1580 


n 


Range 


X ± 1SD 




9 


S 


9 


9 








Total length (body + tail) 


97.0 


97.0 


92.0 


100.0 


— 








Length of tail 


52.0 


5 1 .0 


46.0 


53.0 


— 


— 


— 


Hind foot 


8.0 


8.5 


8.0 


8.5 


— 


— . 


— 


Ear from meatus 


14.0 


14.5 


13.0 


14.0 


— 


— 


— 


Forearm 


38.3 


39.7 


37.1 


38.3 


1(1 


(36.9-39.8) 


38.25±0.85 


Third ringer, metacarpal 


35.0 


36.6 


34.8 


36.1 


10 


(34.4-37.5) 


35.75 ± 1.00 


first phalanx 


16.4 


15.9 


15.3 


15.7 


10 


(14.0-17.0) 


15.35±0.70 


second phalanx 


21.8 


21.6 


20.7 


21.7 


10 


(19.3-22.3) 


21.14± 1.04 


Fourth finger, metacarpal 


35.2 


35.2 


33.8 


35.2 


10 


(33.3-37.3) 


35.20± 1.12 


first phalanx 


10.0 


9.0 


9.6 


9.4 


10 


(8.4-10.4) 


9.16 + 0.61 


second phalanx 


9.9 


10.6 


9.7 


10.3 


10 


(9.5-10.6) 


9.90 ±0.34 


Fifth finger, metacarpal 


35.3 


34.9 


34.2 


34.6 


10 


(33.4-37.3) 


34.93 ± 1.33 


first phalanx 


9.6 


8.9 


9.1 


8.9 


10 


(8.3-10.1) 


9.06 ±0.55 


second phalanx 


10.4 


10.5 


10.5 


10.4 


10 


(9.5-11.4) 


10.26±0.67 


Skull. 
















greatest length 


15.5 


16.1 


— 


15.9 


10 


(15.4-16.4) 


16.02 ±0.39 


condylo-basal length 


14.4 


14.9 


— 


14.8 


10 


(14.2-15.2) 


14.73 ±0.32 


condylo-canine length 


14.0 


14.3 


— 


14.3 


9 


(14.2-14.9) 


14.48±0.24 


basal length 


13.1 


13.5 


13.1 


13.4 


10 


(13.3-13.9) 


13.54±0.20 


palatal length 


7.7 


7.0 


7.8 


7.7 


10 


(8.7-9.6) 


9.02 ±0.29 


zygomatic width 


7.7 


8.1 


7.7 


— 


10 


(7.9-8.3) 


8.11±0.I9 


width of braincase 


7.4 


7.7 


7.7 


7.4 


10 


(7.5-8.0) 


7.73±0.14 


height of braincase 


6.1 


6.1 


— 


5.9 


10 


(6.1-6.5) 


6.22±0.12 


length of foramen magnum 


3.0 


3.4 


— 


2.9 


10 


(2.5-3.2) 


2.67 ±0.20 


width of foramen magnum 


3.3 


3.6 


— 


3.6 


9 


(3.2-3.9) 


3.43 ±0.20 


mastoidal width 


7.0 


7.3 


— 


7.1 


9 


(7.3-7.5) 


7.41 ±0.09 


interorbital width 


3.3 


3.2 


3.2 


3.2 


10 


(2.8-3.1) 


3.00±0.10 


width across molars 


5.0 


5.2 


5.1 


5.2 


9 


(5.1-5.3) 


5.26 ±0.06 


width across canine 


— 


3.5 


3.4 


3.5 


9 


(3.5-3.8) 


3.61-0.10 


upper toothrow. c-m 3 


6.5 


6.8 


6.5 


6.7 


9 


(6.8-7.0) 


6.86 ±0.08 


lower toothrow, c-m 3 


7.0 


7.1 


6.8 


7.1 


9 


(7.2-7.5) 


7.33 ±0.08 


length of mandible 


11.7 


12.0 


11.8 


11.9 


8 


(11.4-12.5) 


11.79±0.32 



1 holotype 



Comparison and discussion: There are no external 
characters by which N. stramineus can be distin- 
guished from N. tumidirostris. In cranial char- 
acters N. s. tronchonii can always be recognized 
by the small, swollen maxillary bones and semi- 
emarginated palate. 

The characters of this population give rise to new 
doubts about the 3 recognized species of the sub- 
genus Natalus (Dalquest, 1950). These 3 species 
are very similar (Goodwin, 1959); major is the 
largest (Jamaica, Cuba, Dominican Republic and 
Haiti), tumidirostris is intermediate in size (north- 
ern Venezuela, Curacao, Trinidad and Colombia) 
and stramineus is the smallest (Mexico, Central 



America, western Venezuela and northeast Brazil). 
The size differences in some cases are not significant 
and, in general, are no greater than 2 to 3 mm in all 
known forms. Recent comparisons of more than 
40 specimens from Venezuela and the 18 from 
Trinidad (some of which were reported by Goodwin, 
1959) have led me to concur with Goodwin (1959) 
and Handley (1966) in considering N. stramineus 
saturatus Dalquest and Hall a synonym of N. s. 
mexicanus Miller, and to place N. tumidirostris 
haymani Goodwin in synonymy with N. t. con- 
tinents Thomas. 

On the basis of an analysis of the form of the 
palate, only 2 groups can be admitted: stramineus- 



1971 



NEW SUBSPECIES OF HA I 



Hi 



Mil 




Figure 2. Comparisons of selected measurements of 
Natal us stramineus mexicanus, A (means connected by 
dotted line), and N. s. tronchonii, B (means connected 
by a dashed line). The means of measurements of the 
former have been arranged to form a base line for 
comparison with those of N. s. tronchonii. Horizontal 
lines indicate range, small boxes ± 1 standard error and 
large boxes ± 1 standard deviation. 



major with a very short palate, and tumidirostris 
with a very long palate. The intermediate palate of 
the Venezuelan population poses the interesting 
question of a possible cline, and makes a careful 
revision of the subgenus necessary. Tentatively, I 
suggest that the 3 subspecies of major (Goodwin, 
1959) be assigned to stramineus, thus bringing to 6 
the known forms of this latter species. 

Koopman (1968) assumes the presence of N. 
stramineus in the Lesser Antilles by way of South 
America. However, on the basis of this new cri- 
terion, it would be wiser to think of a mainland 
Neotropical center of origin, following a route from 
Yucatan (Mexico), Cuba. Jamaica, Hispaniola, to 
the Lesser Antilles; as the forms of stramineus have 
not been localized in northern South America. 
Goodwin (1959:4) mentions specimens from Brazil, 
Venezuela, and Trinidad, but he does not note any 
specific locality in either Venezuela or Trinidad. 



Based <>n the criterion "i palatal itructurc ii ii 

more likely thai tumldiri i U 

stramineus, thus rcdui ing the subgenus Natalui 

to only one species. In this paper. I will retain 
tumidirostris as ;i lull species which rcpn 

the maxim in the differentiation within the 

stramineus-major groups; more examples from 

South America should con fin negate this thesis. 

A revision ol the oilier species grouped in the 
various subgenera, and held investigations in die 
Greater Antilles, particularly Puerto Rico, would 
also provide new information concerning these 
delicate bats. 

Additional specimens examined < ompa 
material was kindly made available by the I 
Co-operative Wildlife Collection (TCWC), the 
American Museum of Natural Hislorv lAMMIi. 
the United States National Museum (USNM), (he 
La Salle Museum of Natural History (MHNI S. 
and the Museum of Natural Sciences. Caracas 
(MCNC). Specimens in the mammal collection. 
Biology Museum of the Tropical Zoological Insti- 
tute (Central University of Venezuela), are desig- 
nated by the abbreviation (MBUCV). 

Natalus stramineus stramineus: ANGI II I \ 
Northside Estate (AMNH. 2). 

Natalus stramineus major: DOM INK AN Kl - 
PUBLIC. Barahona (AMNH. 2). Maniel Viejo 
(AMNH, I). 

Natalus stramineus jamaicensis: JAMAICA. St 
Clair (AMNH. 3). 

Natalus stramineus mexicanus: MEXICO. Chia- 
pas: Cintalapa (TCWC, 2). Nayarit: Amatlan 
(AMNH, 8). Tamaulipas: El Pachon (AMNH 5 
GUATEMALA. Puerto Barrios (TCW< . 7); El 
Progreso (AMNH. 6). NICARAGUA. Rama 
(TCWC. 1). 

Natalus stramineus tronchonii: VENEZL ELA. 
Zulia: Guasare river. (MBUCV, 4). 

Natalus tumidirostris continentis: TRINIDAD. 
Oropuche caves (MCNC. 1); Mt. Tamana (AMNH, 
10). VENEZUELA. Carahobo: San Esteban 
(AMNH. 10). Araqua: Rancho Grande (MBUCN . 
1); El Limon (MNHLS. 1). Miranda: Baruia 
(MHNLS. 1); El Hatillo (MBUCV, 3); Petare 
(MHNLS. 1); El Encantado (MBUCV, 44. 
MHNLS. 24. USNM. 7); Araira (MHNLS, 3); 
Virongo (MBUCV. 2): Capaya (MHNLS. 2). Fal- 
con: Peninsula de Paraguana (USNM. 20). Bolivar: 
Caicara (USNM. 15). COLOMBIA. San Gil. Cueva 
del Nitro(AMNH. 2). 



ACKNOWLEDGMENTS 

I am grateful to the following persons who very kindly 



S-l 



111 I I I II, X SOI I III KX ( Al HORN I A ACADEMY Oh SCIENCES 



VOLUME 70 



permitted me to stud) comparative material from 
different localities: W. B. Davis (TCWC), K. F. Koop- 
man (AMNH), A. Muss,. (MHNLS), R. A. Lancini 
{Ml NO. and C. O. Handley. Jr.. (USNM). In addition. 
I wish to thank Osvaldo Reig. Andrew Starrett, Luis de 
la Torre. Edgardo Garcia, and Donald R. Patten, who 
read the manuscript and contributed useful suggestions. 
Juan A. Tronchoni, president ol'thc Venezuelan Society 
of Speleology, who has heen a companion on many 
expeditions in the country, permitted me to incorporate 
the material from that institution into the Biology 
Museum. Central University of Venezuela, and I take 
pleasure in naming this new bat in his honor. 

LITERATURE CITED 

Dalquest, W. W. 1950. The genera of the chiropteran 
family Natalidae. J. Mammal., 3 1: 436-443. 

Dalquest, W. W.. and E. R. Hall. 1949. A new subspecies 
of the funnel-eared bat (Nataltis mexicanus) from 



eastern Mexico. Proi . Biol. Soc. Washington, 

62: 153-154. 

Goodwin, G. G. 1959. Bats of the subgenus Natalus. 
Amer. Mus. Novit., no. 1977: 1-22. 

Handley. C. O., Jr. 1966. Checklist of the mammals of 
Panama. Pp. 753-759. in Ectoparasites of Panama 
(Wenzel. R. L. and V. J. Tipton, eds.). Field Mus. 
Nat. Hist.. Chicago. 861 pp. 

Koopman, K. F. 1968. Taxonomic and distributional 
notes on Lesser Antillean bats. Amer. Mus. Novit., 
no. 2333: 1-13. 

Ridgway. R. 1912. Color standards and color nomen- 
clature. Washington, DC, privately printed, 44 pp. 

Sociedad Venezolana de Espeleologia. 1968. Catastro 
espeleologicode Venezuela, CuevadelosGavilanes 
(Zu-1). Bolctin de la Sociedad Venezolana de 
Espeleologia, 1: I 13-1 18. 

Accepted for publication August 30, 1971. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(2): 84-87, 1971 

A NEW SPECIES OF 1PHITIME (POLYCHAETA) FROM CANCER 
ANTENNARIUS (CRUSTACEA: DECAPODA) 

John Pilger 1 

Abstract: A new species of Iphitime is described from Cancer anten- 
nariits. Notes are given on the morphological criteria used in separation of 
the five species of the genus. 



During fixation of a specimen of Cancer anten- 
nariits (Stimpson, 1856) at the Santa Catalina 
Marine Biological Laboratory, 2 specimens of a 
new species of Iphitime crawled out between the 
third maxillipeds of the host. These 2 specimens 
belong to a new species which is described below 
with notes on frequency of infection and some 
morphological features of the family. 

Iphitimidae Fauchald, 1970 

The genus Iphitime was originally referred to the 
family Lysaretidae (Marenzeller, 1902) but was 
recently raised to familial status by Fauchald (1970). 
This separation was made because members of 
this genus possess branchiae, composite setae, two 
antennae, three paired maxillae, maximally, and 
they lack an unpaired maxillary carrier. The 



lysaretids. on the other hand, lack branchiae, have 
simple setae, three antennae, five maxillae, and 
three maxillary carriers. 

Fauchald (1970:1 18) states that iphitimids have 
only simple falcate unhooded setae. However, this 
should be expanded to simple and composite 
unhooded setae. In addition, the two peristomial 
segments noted by Fauchald (1970:118) as diag- 
nostic of the family Iphitimidae should be broad- 
ened to one or two peristomial segments. This 
change in the familial definition is made to encom- 
pass the new species, which has only one peris- 
tomial segment. All species of Iphitime are found 
in the branchial cavities of decapod crustaceans 
(Fauchald. 1970). Hartman (1952) suggested that 



■Department of Biological Sciences, University of Southern 
California. Los Angeles. California 90007. 



1971 



NEW SPECIES OF I'OIX 11 \i 1 1 










Figures 1-6. Fig. 1, Dorsal view of anterior end, X48; Fig. 2. Mandibles, removed from maxillary apparatus, ventral 
view. XI 10; Fig. 3, Maxillary apparatus, mandibles removed, ventral view. XI 10; Fig. 4. Median parapodium. 
anterior view, X21; Fig. 5, Composite falcate setae. X245: Fig. 6. Simple falcate setae. X245. 



they may feed on small particles of food swept into have well-developed jaws for biting and chewing 
the branchial cavity by respiratory currents. How- which suggests that they may parasitize the bran- 
ever, it should be noted that all species of Iphitime chial tissue of the host. 



tS6 



BULLETIN SOUTHERN CALIFORNIA ACADEMY Oh SCIENCES 



VOLUME 70 



Iphitime holobranchiata, new species 
Figures 1-fi 

Material: The specimens were taken from a 
large male Cancer antennarius (Stimpson, 1856) 
caught near Little Harbor, Santa Catalina Island. 
California. The holotype. measuring 48 mm, and 
one paratype (hroken in two), 25 mm. are both 
deposited in the Allan Hancock Foundation. A 
number of individuals from an unknown locality 
measured from a few millimeters to 120 mm; all 
were found in the branchial cavity of the host crab. 

Description: The 2 specimens measured 25 mm 
and 48 mm. Each had 175-200 setigerous segments. 
The color in life is pink-orange. 

The prostomium (Fig. 1) is a rounded lobe with 
2 small antennae. These extend beyond the outline 
of the anterior margin of the prostomium when 
viewed from above. A shallow depression curves 
around the anterior and anterolateral portions of 
the prostomium. 

There is 1 peristomial segment. Between the 
prostomium and the peristomium is a narrow slit- 
like nuchal organ (Fig. 1). 

The mandibles (Fig. 2) are fused medially in 
their anterior portion for about one-sixth of their 
entire length. A conspicuous hole is located in the 
anterior portion of this suture. The maxillary 
carriers (Fig. 3) are fused medially with their free 
edges extending dorsally to form a V-shaped struc- 
ture. At their anterior margins they are fused with 
the large falcate maxillae I. Dorsal to this are the 
wavy-edged maxillae II. Distinct teeth are absent. 
Each maxillae III has 1 large and 2 smaller teeth. 



The long branchiae are dorsolateral. Branchiae 
are first present from the second setigerous seg- 
ment. Each is long and digitiform (Figs. I. 4). 

Parapodia (Fig. 4) consist of a small ventro- 
lateral presetal lobe and a large dorsolateral ly 
directed postsetal lobe. There are 5-7 composite 
falcate setae (Fig. 5) and about 29 simple falcate 
setae (Fig. 6) in each parapodium. All lack hoods. 
Fight acicula are present. 

Discussion: In contrast to other species of 
Iphitime, I. holobranchiata has only I peristomial 
segment. All other species have 2. Iphitime dihler- 
leinii, I. cuenoti, and /. paguri all have more com- 
posite falcate setae than simple ones. The opposite 
situation exists in /. holobranchiata which has 
many more simple than composite falcigers. 

Iphitime paguri and /. holobranchiata are the 
only 2 species in the genus with simple unbranched 
branchiae. In /. holobranchiata they are inserted 
dorsolaterally while in /. paguri they are dorsal. 
The specific name holobranchiata was chosen 
because it connotes simple, whole, unbranched 
branchiae. 

All species have 3 maxillae except for /. loxor- 
hynchi which has only 2. Maxillary formulas for 
all species are given in Table 1. which is expanded 
from Hartman (1952) by adding /. holobranchiata 
and diagnostic characters not contained in her table. 

Distribution: Cancer antennarius ranges from 
British Columbia to Magdalena Bay, Baja Cali- 
fornia. Examination of the branchial cavities of 
live C. antennarius from southern California 
showed the infection frequency to be quite high. 
From the 30 individuals inspected 142 /. holo- 



Table I . Diagnostic characters of species in the genus Iphitime. 



Name of Species 



Tola! Number of Number of First Presence of 

Length Host Species Setigerous Peristomial Branchiae and Parapodiai 

in mm and Locality Segments Segments Character Components 



Number 

of Maxillary 
Maxillae Formulae 



1. cuenoti 
Fauvel 



/. dbderleinii 
Marenzeller 



/. holobranchiata. 
new species 



/. loxorhynchi 
Hanman 



/. paguri 
Fage <i Legendre 



7-12 Maia squinado 

Porlunus spp. 
Gonoplax angulata 
Macropodia longirostris 
All from Europe 

61 Macrocheira kuempferi 

Japan 



Cancer antennarius 
Southern California 



Loxorhynchus grandis 200 or more 
Southern California 



Gonoplax angulata 
Macropodia longirostris 
Porlunus depuralor 
Eupagurus bernhardus 
All from Europe 



First; simple to 4-5 simple falcigers 

divided, lateral 20 composite falcigers 



First: simple and 14-15 simple falcigers 

palmate, midlateral Many composite falcigers 
About 4 acicula 



Second; simple 



Second; simple to 
divided, lateral 



Fourth; simple 
dorsal 



29 simple falcigers 
5-7 composite falcigers 
8 acicula 

26 simple falcigers 
20 composite falcigers 
6 acicula 

i simple falciger 

7-8 composite falcigers 



3 1+1- 1+1-1+ I 



3 1+ 1-1+ 1-1+1 



3 1+1-0+0-3+3 



2 1+1-0+4(5) 



3 l+l-l+l-l+l 



■Measurements of holotype and paratype only. 



1971 



NI-.W SPEC II SOI POL V< IIAI.II. 



H7 



branchiata were taken. I he maximum numbei 
found in any one crab was J9, 

Ecology: Small specimens commonly occupied 
the interlamellar areas of the phyllobranchs while 
larger ones were found on the walls of the cavity 
or on the surface of the branchiae. No visible 
damage to the gills was observed. 

ACKNOWLEDGMENTS 

I wish to thank the crew of the U.S. Naval Undersea 
Research and Development Center vessel Sea-See who 
collected (he Cancer crab in which the polychaetes 
were found. Research facilities were provided by the 
Santa Catalina Marine Biological Laboratory, Uni- 
versity of Southern California, and I would like to 
thank the staff for their encouragement. 
Reference materials were furnished by Olga Hartman 
and the Allan Hancock Foundation Library. A very 
special thanks goes to Kristian Fauchald who pro- 
vided me with equipment, information, criticism, and 
inspiration. 



Ill ERA I URE fill D 

Fauchald, K. 1970. Polyi hat I idt •>< the 

Familie i unii idai i umbi Inci idae, Iphitimii 

Arabcllidac, Lysarclidac and D lac from 

Western Mexico Mian //</;/<.»/ Mono \l<n 
Biol., 5: I 

llaii man. O, 1952. Iphltlme and ( i ""■» cphala 
chaetous Annelids; Ik. in ( alifornia Bull 
< alifornia Acad. Si I., 51: ')-20. 

Marenzeller, I von. 1902. Stidjapanischc Anncliden; 
Aphroditea, Eunicea. Denkschriften Dei > 
lichen Akademie l><> Wlssen chafien, Wcin. " 
563-582. 

Stimpson, W. ixsr,. On some California < mstacca. 
Proi . California Anal. Nat. & L, I: 9 



Accepted for publication March 12. 1971. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(2): 87-90. 1971 

TWO TYPES OF AGGREGATION GROUPING IN THE LARGE MILKWEED BUG, 
ONCOPELTUS FASCIA TUS (HEMIPTERA: LYGAEIDAE) 

Steven R. Kutcher 1 

Abstract: Two types of grouping behavior, parallel and linear groups, in 
the large milkweed bug Oncopeltus fasciatus (Dallas) seem to have survival 
value. Grouped bugs receive mutual benefit from the ability to perceive stimuli 
from more directions and are usually touching each other. Color pattern and 
uniformity of direction causes a striking pattern in aposomatic insects. A 
combination of the shape of the bugs, limited areas for grouping on the plant, 
and microclimates have influences on the type and shape of aggregations. 
Aggregation may also be influenced by the substrate, position of the heat or 
light source, and changing behavioral and physiological stages of the bug. 



The milkweed bug. Oncopeltus fasciatus (Dallas), 
has a gregarious behavior that has been documented 
by Weiss and Dickerson (1921). Andre (1934). 
Beck, Edwards and Medler (1958). and Barrett 
and Chiang (1967a). Gregarious behavior in milk- 
weed bugs is herein defined as the condition which 
exists when one bug is within one body length of 
another bug. When two bugs are of different sizes, 
the length of the larger bug is used as the measure 
between bugs. 

Aggregation or grouping is a normal occurrence 
in the bugs and they spend the majority of their 
lives in close proximity to one another. A "normal" 
aggregation consists of grouped bugs seemingly 
oriented in almost every direction. The bugs may 
be in compact groups where there is much contact 



between bugs or in loose groups where there is 
little contact. 

OBSERVATIONS 

Time-lapse pictures were taken of milkweed bugs 
grouped inside plastic containers. In the process 
of taking these pictures two grouping behavior 
patterns were observed (on the bottom of the 
container) which were different from normal aggre- 
gation. These two groupings have been observed 
on other substrates with varying conditions and 
are herein labeled "linear groups" and "parallel 
groups." 

In a linear group the bugs will line up in com- 



'Department of Biology. California Slate College, Long Beach. 
California 90801. 



88 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OE SCIENCES 



VOLUME 70 



binations of head to head, tail to tail, and head to 
tail so that their longitudinal axes line up in more 
or less one straight line. As many as 6 fifth instars, 
out of a population of 10, have heen observed in 
this type of grouping. The line of bugs may be 
straight or it may curve slightly (Fig. 2). 

In a parallel group the bugs will line up side by 
side, as if in a row. The longitudinal axes of their 
bodies will be parallel to one another. Their heads 
will usually be pointing in the same direction. As 
many as 5 bugs, out of a population of 10, have 
been observed in this type of grouping (Fig. I). 

Both linear, parallel, and combinations of these 
two types of groupings are found in the 5 instars 
and the adult stage. Linear and parallel groupings 
are usually a part of normal grouping activity, 
although they may appear attached appendicularly 
to a larger group. Both parallel and linear groups 
generally did not last longer then 20 minutes in the 
time-lapse experiments. 





Figures 1-2. Fig. 1. Fifth instars in a parallel group. 
Notice contact between legs, antenna, and body. Fig. 2. 
Fourth instars in a linear group. Notice contact be- 
tween forelegs, antenna, and head. Figures drawn from 
projected 16 mm color film. Line equal to 1 cm. 



DISCUSSION 

The explicit reasons for these types of grouping are 
not known, but there seems to be evidence for each 
of the following factors: copulation, protective 
coloration, physical properties of the milkweed 
plant, shape of the bugs, effect of light, temperature, 
physiological state of the bug, visual and tactile 
responses, and other external factors. Some of 
these factors will be briefly discussed in relation 
to survival value of the special kinds of groups 
previously discussed. 

Copulation: Andre (1934) has described the 
method of copulation in the milkweed bug. When 
a male attempts to copulate with a female, the male 
usually forms a very short-lasting parallel group. 
After the male and female attach genitalia, they 
move into a position in which both longitudinal 
axes lie on the same line and their heads are point- 
ing in opposite directions. This is a type of linear 
grouping. Other bugs were observed lining up with 
a copulating pair to form a linear group of 4 or 



more bugs. Temporary parallel and linear groups 
may also be formed when a secondary bug attempts 
to mate with a copulating pair. A copulating pair 
in a linear group will gain the protective advantage 
of potentially perceiving stimuli from more direc- 
tions than an individual bug. 

Both types of groupings are related to mating 
positions. Parallel and linear groups occur in all 5 
instars. Sexual behavior is probably not the only 
causative mechanism for this type of grouping 
since the instars are incapable of mating. The 
groupings may have influenced typical hemipteran 
mating behavior. 

Protective coloration: A specific color or color 
arrangement, adapted for recognition of undesir- 
able species, has been suggested as one means of 
protection for some insects by Ehrlich and Raven 
(1967) and Cott (1957). The bug is black with 
yellow to orange-red coloration. The percentage of 
black area increases with each succeeding instar. 
The pattern or color may be important in learning 
to avoid unpalatable bugs (Rettenmeyer, 1970). 
Gelperin (1968) stated, "The large milkweed bug, 
Oncopeltus fasciatus, served as the distasteful prey 
[for mantids] ." The milkweed bug will exude a 
pungent fluid when disturbed. 

Protection may be gained by easier recognition 
of large aggregations of aposematic animals (Cott, 
1957). A group of aposematic bugs may gain some 
advantage by being grouped in the most compact 
possible type of grouping or by producing a 
more striking pattern incorporating uniformity of 
direction. 

Grouping may be accomplished by normal com- 
pact grouping with some bugs on top of others. 
These layers will tend to cover up the empty spaces 
between the bugs and appear as a solid mass. 
Multiple layers of bugs have been observed in the 
field. Layers of bugs have also been observed in the 
laboratory. Barrett and Chiang (1967a) reported 
eight layers of bugs on a heat gradient. The layers 
of bugs may or may not be due to a lack of space. 
A pheromone may be involved in aggregation and 
group formation in the bugs. 

An efficient covering of the plant may be pro- 
duced by bugs lining up in parallel and linear 
combinations. Most bugs are found at angles to one 
another indicating other factors influencing group 
formation. 

Physical properties of the plant: The relationship 
of bugs to the substrate, surface texture, horizontal 
and vertical surfaces, and available room for group- 
ing are important in development of aggregations. 
Asclepias fascicularis is a glabrous milkweed, about 
1 m tall, and found in dry places in cismontane 
California and other parts of the western United 



1971 



IN SI. (I III.IIAVIOK 






Stales. Aggregations arc common on the upper 
parts of the plant, but the bugs have been observed 
at the base of the plant at night, before flowering, 
and after the plant has stalled i<> die back. The 
highest population densities occurred alter Dower- 
ing or while the follicles were developing. The bu^s 
were especially numerous along the suture line ol 
the follicle. 

The milkweed has both horizontal and vertical 
surfaces of varying degree, upon which bugs can 
aggregate. The bugs will aggregate more readily on 
certain parts of the plant. "Localization of feeding 
sites and the ability to mate while feeding may 
serve to bring pairs together, especially if a peak of 
activity occurs during one part of the day." (Cald- 
well and Dingle, 1967). Linear groups may be 
formed when bugs meet while moving up and down 
the stem. 

Aphis fabae will aggregate on plants because 
this aphid travels along major leaf veins and 
responds by probing when encountering a station- 
ary individual. Aggregation is produced by aphids 
probing longer in certain parts of the plant and 
mutual visual stimulus which plays an important 
part in gregarious behavior (Ibbotson and Kennedy. 
1951). 

The milkweed bug will walk along the midrib of 
a leaf. Bugs will line up on the midrib, a primary 
feeding site, or parallel with the midrib and on the 
stem (i.e., parallel and linear groups). 

Since there are many orientation lines on the 
plant (i.e., midribs, edges of leaves, stems, and 
petioles) it would seem that orientation lines of the 
plant, localized feeding sites, restriction of space, 
and the behavior of the bugs would be involved in 
the production of linear and parallel groups. 

Physiological condition: Beck el at., (1958) and 
Barrett and Chiang (1967a) report 3 intra-instar 
ages, pre-feeding, feeding period, and post-feeding. 
As the bugs grow they will go through periods of 
greater or lesser tendency to form groups. The 
tendency to group is due, in part, to the physiologi- 
cal changes taking place within the bugs. 

Shape of bugs: The shape of the bugs is impor- 
tant in the formation of aggregations. The instars 
are generally elongate-oval, tapering towards the 
head. The adults are generally elongate-oblong in 
shape. In most aggregations, especially post-feeding 
groups, bugs are usually in physical contact with 
one another. Contact involves the antenna, legs, 
beak, genitals, and the body itself. The legs are 
often used by overlapping or touching the leg or 
body of adjacent bugs. Overlapping legs are com- 
monly seen in parallel groups, whereas, the use of 
antenna and genitalia is commonly seen in linear 
groups. Body to body contact is usually found in 



large compai I a\ jn gationi in loo • gj .. 
association with one bug ll foi 

temporary grouping. In mo jationt 

bugs seem to require | ping with more than one 

bug 

Bugs will aggregate at feeding itcs If a mill 
seed is presented to some line's which have been 
deprived of seeds foi om da) thi will group 

around the seed ["he bug m a circle around the 
seed with their heads pointed towards the center ol 
the seed. II there are more bugs than feeding sites 
available, the extra bugs will climb on top ol 
ing bugs so that the seed may be reached by extend- 
ing their beaks between bugs. Since the he.nl is the 
smallest part of the body, compact groups are 
occasionally formed in this circular arrangement 

In compact aggregations the bugs were some- 
times in combinations of parallel and linear groups 
i.e., many grouped rows of bugs with the main bod) 
axis of each bug parallel to the bug lateral to it and 
on the same line with the axis of the bug anterior 
or posterior to it. 

Insects or other animals that aggregate and art- 
longer then they are wide, will form parallel groups 
if favorable conditions exist. Some lepidopterous 
larvae are examples (Olson, 1968) of this type ol 
grouping behavior. Parallel grouping does not seem 
to occur, to any great extent, in round insects, 
unless they are oriented by an external stimulus. 

Light and temperature: Changes in the behavior 
of the milkweed bug to humidity, light, and gravity 
have been reported by Barrett and Chiang 1 1967b). 
In addition, they (1967a) studied the effect of 
temperature on the bugs and found that, in labora- 
tory studies. "... where a thermal heterogeneity is 
present, the insects were attracted to a spot warmer 
than the ambient temperature." They also stated. 
"Under field conditions, the nymphs showed a 
certain behavior relative to the sun." The beha\ior 
of the nymphs depends on their period of develop- 
ment. 

The bugs are attracted to a temperature above 
the ambient and also to sunlight, if both sunlight 
and incandescent light are present. 

Bugs will aggregate more readily on certain parts 
of the plant due. in part, to the position of the sun 
or radiant energy source. Since a variety of dif- 
ferent temperatures occur on a single plant, the 
bugs can move to the most suitable microclimate. 
The change in the microclimate may be one reason 
for bugs not remaining in specific arrangements 
for prolonged periods of time. 

If a temperature range occupies a small area it 
will cause insects to cluster as in the case of a light 
bulb used to produce a concentric temperature 
gradient. This will cause insects to form a ring in an 



90 



Hi I I I 1 1 X S( >( llll RX ( I / II ORXIA AC A DEM y of sciences 



VOLUME 70 



optimum temperature range {e.g., blowflies, Fraen 
kel and Gunn, 1961). Barrett and Chiang (1967a) 
stated. "The nymphs around the gradients were 
usually lined up side by side in a broken circle 
around the heat at somewhat equal distance from 
the center and at about the same temperature on 
the gradient. A tew nymphs were situated behind 
these front ranks and in physical contact with 
them." 

Compact aggregations were found to occur under 
3 conditions: when a heat or light source is present, 
when the bugs are in a post-feeding stage (before 
moulting), and during the night hours. Compact 
groups may be involved with heat retention or 
control of temperature in the bugs. If the center of 
a temperature gradient is not too hot, the bugs will 
aggregate in the center of the gradient. If a temper- 
ature range occupies a large area or has homo- 
geneity, the insects may still cluster. In the case of 
desert locusts, they will turn their bodies so that 
they are perpendicular to the suns rays which will 
permit the locust to absorb more radiant energy. 
If conditions are too hot, the locust will climb a 
bush or rock and position its body parallel to the 
suns rays, in this way less energy will be absorbed 
from the sun. 

In the field, milkweed bug populations are usually 
of mixed ages. The changing physiological stages 
of the bug and the change in environment will cause 
the bugs to aggregate on certain parts of the plant, 
at certain times, and in certain ways. 



ACKNOWLEDGMENTS 

I wish to thank Elbert L. Sleeper and Richard J. John- 
son of California State College, Long Beach, Clay A. 
Singer, and Frank E. Strong of the University of 
California, Davis, for their help. Part of this work was 
undertaken at the University of California, Davis. 



LITERATURE CITED 

Andre, F. 1934. Notes on the biology of Oncopeltus 
fasciatui (Dallas). Iowa State Coll. ./. Sci., 
9: 73-87. 

Barrett. R. W., and H. C. Chiang. 1967a. The effects of 
thermal heterogeneity in the environment on the 
activity and development of Oncopeltus fasciatus 
(Dallas). Ecology, 48: 590-598. 

— . 1967b. Changes of behavior pattern within the 
fifth nymphai instar of the milkweed bug, 
Oncopeltus fasciatus (Dallas). Amer. Midi. Nutiir., 
78: 359-368. 

Beck, S. D., C. A. Edwards, and J. T. Medler. 1958. 
Feeding and nutrition of the milkweed bug, 
Oncopeltus fasciatus (Dallas). Ann. Ent. Soc. 
Amer.. 51: 283-288. 

Caldwell, R. L., and H. Dingle. 1967. The regulation 
of cyclic reproductive and feeding activity in the 
milkweed bug Oncopeltus by temperature and 
photoperiod. Biol. Bull.. 133: 510-525. 

Cott, H. B. 1957. Adaptive coloration in animals. 
Methuen and Co. Ltd.. London, 508 pp. 

Ehrlich, P. R., and P. H. Raven. 1967. Butterflies and 
plants. 5c/. Amer.. 216: 104-113. 

Fraenkel, G. S., and D. L. Gunn. 1961. The orientation 
of animals. Dover Pubis. Inc., New York, 376 pp. 

Gelperin, A. 1968. Feeding behavior of the preying 
mantis: A learned modification. Nature, 219: 399- 
400. 

Ibbotson, A., and J. S. Kennedy. 1951. Aggregation in 
Aphis fabae (Scop.). I. Aggregation on plants. 
Ann. Appl. Biol.. 38: 65-78. 

Olson, K. 1968. [Cover photo of Promethea moth 
larvae] . J. Econ. Ent., 61 (August): Cover. 

Rettenmeyer, C. W. 1970. Insect mimicry. Ann. Rev. 

Ent., 15: 43-74. 

Weiss, H. B., and E. L. Dickerson. 1921. Notes on 
milkweed insects in New Jersey. J. New York 
Ent. Soc, 29: 123-145. 



Accepted for publication June 5, 1970. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCE 70(2): 91 98, 1971 

THE INVASION AND DISTRIBUTION OF THE ASIATIC CLAM (CORBH I / i 
MANILENS1S) IN A SOU THERN CAI.II OKNIA RESERVOIR 



Akio W. Fas'i 



Abstract: Corbicula manilensis Philippi were probably established in I i 
Capitan Reservoir by 1962. Relatively few live specimens, and no empty shell' 
were observed during 1964. Empty shells were lirsl observed during the sum 
merof 1965 and outnumbered living specimens by 1967. Live clam populations 
increased from 234 x l() r ' individuals during the summer 1964 to a peal '.I 
10,816 x 10 B during January 1966. They declined to 6,744 x 10' during the 
summer 1967. These fluctuations were partually attributed to water level 
changes and reservoir stratification. Thermal and chemical stratification 
limited the depth distribution of Corbicula to the shallow, aerated depths 
during stratified periods. Their distribution extended to all depths following 
prolonged periods of artificial destratification. Corbicula densities appeared 
positively correlated with sediment mean partical size. 



Corbicula were reported established in this hemi- 
sphere by 1938 (Ingram, 1948) from an unknown 
source. Filice (1958) indicated they may have been 
introduced accidentally to the Columbia River 
estuary, Washington, with seed oyster (Ostreas 
gigas) importations. However, this possibility seems 
remote since C. manilensis is a freshwater species 
(R. O. Fox, pers. comm., 1970). Since their intro- 
duction, they have spread rapidly through the west 
coast states and the Ohio River basin (Sinclair and 
Isom, 1963). Fox (pers. comm.) now places their 
distribution "in 21 states, Baja California and 
Sonora, Mexico." 

While considered a delicacy in their native Japan, 
they are a nuisance organism in many local waters. 
They were a problem in the Coachella Valley canals. 
California, by 1953, (Ingram, 1959), and clogged 
pumps at the Tennessee Valley Authority steam 
plants in 1960 (Sinclair and Isom, 1963). Two to 
three feet of Corbicula bearing sediments were 
deposited in the Interim Canal of the South Bay 
Aqueduct, California, within a few years (California 
Dept. Water Resources. 1967). Clams greatly accel- 
erate sedimentation by their shell deposits and by 
removing suspended solids from the water and 
depositing them in a slime that is not easily sus- 
pended (Prokopovich, 1969). 

Corbicula constitute a biomass that is not readily 
available to higher trophic levels; only a few fish 
feed on Corbicula. No effective biological control 
of Corbicula is known, but chemical or mechanical 
control is possible in some cases (Sinclair, 1964; 
Isom, 197 1). Screens may prevent adult clam move- 



ments, and chlorination may control veliger larvae. 
These methods are expensive and more effective in 
closed water channels. No control is recommended 
for large open waters. 

El Capitan Reservoir impounds the intermittently 
flowing San Diego River, San Diego County. Cali- 
fornia. The reservoir was completed in 1935 and 
filled to capacity during 1938 (Fast. 1968). It is a 
euthropic and warm monomictic fluctuating reser- 
voir. Its water source is mostly runoff, although 
since 1958 it has periodically stored imported 
Colorado River aqueduct water. Corbicula were 
not reported in El Capitan Reservoir before 1964. 
although they were well established by this date in 
other San Diego County Reservoirs. This study 
describes the establishment of the clam, population 
increase, and some factors affecting its distribution 
within El Capitan Reservoir. 

METHODS 

Benthic samples were collected with an unscreened. 
15 cm square Ekman dredge along transects T-l. 
T-2, and T-3 (Fig. 1). Transects 1 and 2 extended 
perpendicular from the shore to maximum depth. 
Transect 3 followed the San Diego River channel 
from its shallowest point to the lakeward terminus 
of transect 2. Transect 1 is the deepest, and transects 
2 and 3 have the same maximum depth. Maximum 
depth varied between 20 and 2S meters. 



'Michigan State University. Department of Fisheries am 
life. East Lansing. Michigan 4SS2J i Present address 
Carbide Corp.. Tarrytown, New York 10591). 



91 



92 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OE SCIENCES 



VOLUME 70 



Three samples from each 2 m depth interval were 
collected at each transect during a collection period. 
Benthie collections were made on June I, 29, July 
26, and August 3 I. 1964; on January 30, May 27. 
June 30, July 28, and September I. 1965; on 
January 31 and July 15, 1966; and on August 2. 
1967. The summer samples represent pre-aeration, 
during-aeration, and post-aeration periods. The 
winter collections were made following a period of 
normal stratification, and following a period of 
artificial destratification. Sample size varied as a 
function of water depth. Increased water levels 
necessitated larger sample sizes. 

Benthie samples were partially washed in the field 
through a sieve with a 0.58 mm mesh. The sample 
was taken to the laboratory for preservation and 
examination. Samples were examined as described 
by Fast ( 1970) using Anderson's ( 1959) sugar flota- 
tion technique. Shell lengths were measured to the 
nearest 0.25 mm. Unbroken and disconnected empty 
shells were designated as dead clams. Most empty 
shells were disconnected. All living and dead clams 

San Diego R i 




Chocolate Creek 



were counted; measurements were made of both 
types from selected samples. 

Total population size was calculated for each 
collection period except July 1966. Average clam 
density/m 2 for each depth interval was multiplied 
by the estimated bottom area in m z for that interval. 
Bottom area per depth interval is calculated as the 
difference in lake surface area between the upper 
and lower limits of the depth interval. Sediment 
analyses, temperature measurements, oxygen deter- 
minations, and other techniques were described by 
Fast (1968). 

RESULTS 

California Corbicula are probably from the same 
genetic stock. Most previous identifications by 
Ingram and others call the clams C. fluminea. The 
Ohio River basin clams have characteristics of both 
C. fluminea and C. manilensis (Sinclair and Isom, 
1963). Dr. William J. Clench identified our El 
Capitan clams as "C. manilensis Philippi, which is 
now widely distributed throughout much of the 
Ohio River system [pers. comm.. 1968] ." 

Corbicula manilensis entered El Capitan Reser- 
voir before 1964. They were not numerous by this 
date, even though several large specimens were 
collected (Figs. 2, 3). Only 166 live specimens and 
no dead specimens were observed during 1964 
(Table 1 ). The largest 1964 specimen measured 13.7 
mm. Although the total estimated number of living 
clams increased greatly from 1964 through 1967, 

Total Number of Corbicula (xio 8 ) 




Figure 1. El Capitan Reservoir including sampling 
locations, compressor sites, and major tributaries. 



J J A S 'JAN. J J A S 'JAN. ' JULY 
1964 1965 1966 1967 

Figure 2. Total population estimates of El Capitan 
Reservoir Corbicula from June 1964 through July 1967. 
Total population estimates plus or minus one standard 
error are shown. 



1971 



ASIA IK CLAM INVASION IN ( ALII OUNIA 






250 
200 

150 

100' 
80 

60- 

40 

30- 

20- 



Longthi In Millimetori 






SL£S_^_^L_^ 



o I i ¥B dnod 



Augu»t 1 ee 7 

Coiblculo Mfomttd a ALIVI 



A = 400 
D = 73B 









^ F 1 ^ 



-D- 



yj ^ ^ 



January 196 6 



A = 21 1 
D = 6 



an 



" 100- 

,_ eo 

J! 60 

E 

3 40 




30- 



H 1 1- 



n 



Summer 1965 

A = 400 
D = 5 



771 



n 



n n 



H 1 P- 



January 1965 

A = 77 
D = O 



-l=l h-Q- 



H 1 I- 



Su m mer 196-1 

A = 38 
D = 



i~i n i—i 



.00 .02 .04 .06 .08 .10 .12 .14 .16 .18 .20 -30 .40 .50 .60 .70 .80 

Corbicula Shell Lengths In Inches 



Figure 3. Length distributions of live and dead Corbicula during summer 1964 through August 1 



<>4 



IHIIIII\S(>IIIIIR\( II II ( 'KM I U !/>/ \/> <>l S( // \( is 



VOLUME 70 



empty shell 1 - were not abundant until the summer of 
1967. No empt\ shells were observed until summer 
1965. rhe estimated live population ranged from an 
average minimum of 234 \ I0 5 (±384 x 10"') 
during summer 1 9b4 to an average of 10, 8 16 x I0 5 
(±1,661 x It)'') during January 1966. It subse- 
quently decreased to an estimated 6,744 \ I0 5 
(± 1,608 x I ()■"') during August 1 967. The July 1966 
estimate is not included in Table 1 because only 
transect I was sampled. 

Live Corbicula average shell lengths increased 
from 2.7 mm during summer 1964 to 3.1 mm 
during summer 1 965 (Table 1). A 9.5 mm maximum 
average shell length occurred during summer 1966. 
Average shell lengths then decreased to a low of 1 .7 
mm during summer 1967. Associated with this 
decreased average shell length was a smaller live 
population and an increase in empty shells. Mortal- 
ity rate apparently increased between 1965 and 
1967. Sinclair and Isom (1963) also reported mass 
mortality of unknown causes of Tennessee Cor- 
bicula on three occasions. 

Live Corbicula were restricted to the benthic area 
above 12 m during 1964 and mostly above 14 m 



during 1965 (Fig. 4). Their depth distribution was 
ubiquitous during 1966. and bimodal during 1967. 

El Capitan Reservoir stratified normally during 
1964 and 1967 (Fig. 5). A well-developed thermo- 
cline existed by late May 1964, and an oxygen 
deficit began at 8 to 10 m during that summer. 

The 1964 overturn occurred during November. 
The lake remained mixed until mid-March 1965 
when it again stratified. The lake remained stratified 
through mid-June 1965. Duringthisstratified period, 
hypolimnion oxygen concentrations approached 
zero. Artificial destratification by air injection began 
June 10, 1965. Compressed air was continuously 
injected until June 2 1 . The lake was well mixed and 
oxygen was plentiful at all depths. Air injection was 
discontinued from June 21 through July 1, 1965. 
The temperature and oxygen profiles began to 
resume their pre-aeration configurations, but the 
bottom water was much warmer and the oxygen 
concentrations were much greater than before mix- 
ing. Air injection resumed July 1, 1965 and con- 
tinued through October 4, 1965. The lake remained 
mixed without air injection from October 1965 
through mid-March 1966. 



Table 1. El Capitan Reservoir Corbicula manilensis total population estimates, maximum shell lengths, 
and average shell lengths of live clams and empty shells from summer 1964 through July 1967. El Capitan 
Reservoir average surface area and volume for each sample period is shown. 



Live Corbicula 



Empty Shells 



El Capitan 
Reservoir 



Sample Average Std. No. of Max. Mean No. No. Max. Mean No. Surface Volume 

period est. pop. error Samples shell shell meas. observed shell shell meas. area in in 

size in 10 5 10 5 length length in total length length 10 6 m 2 10 7 m 3 

individual (in mm) (in mm) samples (in mm) (in mm) 



Summer 

I964 1 234 384 288 



13.7 2.7 38 166 



1.4 



1.3 



January 

1965 2 5.546 1.653 72 



5.6 2.6 77 571 



1.4 



1.1 



Summer 

1965 1 5.249 6,220 387 



19.3 3.1 400 1,106 3.81 2. 



5 1. 



January 

1966 2 10,816 1.661 102 



17.0 2.8 211 1,714 3.81 3.2 6 1.9 



2.0 



July 



1966 3 



15.2 9.5 43 115 3.30 2.9 4 2.2 



2.5 



August 

1967 2 6,744 1.608 113 



3.8 1.7 462 645 19.30 4.4 735 2.3 



2.7 



'Estimates from four monthly sample collections from transects 1, 2, and 3. 
2 Estimates from one sample collection from transects 1, 2, and 3. 
3 Estimate from one sample collection from transect 1 only. 



1971 



ASIA I ' l( (I. AM INVASION IN ( A I II OHNIA 






Summer 64 
n - 166 
I - 20 m 




Figure 4. Depth distributions of live El Capitan Cor- 
bicula during the summers of 1964 through 1967. Only 
transect 1 was sampled during summer 1966. Maximum 
depth for each period is designated by (z), and (n) 
designates numbers. 

The average clam depth distribution during sum- 
mer 1965 was deeper than during 1964. but maxi- 
mum depth during both periods was about the 
same. Destratification beginning during June 1965 
did not significantly affect their maximum depth 
distribution during summer 1965. 

The lake began to stratify during mid-March 
1966. Air injection commenced March 19, 1966 
and continued through October 17, 1966. The lake 
was well mixed during 1966 and oxygen was abun- 
dant at all depths. The lake remained mixed from 
October 1966 through February 1967. Clams were 
well distributed to all depths during the summer 
of 1966. 

Stratification developed normally during March 
1967 (Fast, 1968). Hypolimnion oxygen concentra- 
tions declined, but did not fall below 0.1 mg/1 
by September 1967. Corbicula were again more 
restricted to the shallower depths. Over 70% of all 
clams collected during August 1967 were above 
6 m. The remainder were distributed from 12 m to 
maximum depth of 26 m. Few live clams were 



found between 6 and 12 m ["he reason i"i Ihcii 
absence in this zone is unknown Mosl "i ii 
empty shells were collected below <• m 

Water volume and maximum depth flu I 
markedly during the study period Watci volume 

fluctuated between a II \ ID 7 m' 1 minimum |., ., 

2.7 x I0 7 maximum during the study period 1 1 able 

I). Not shown are rapid drawdowns durinj 
summers 1966 and 1967. rhese drawdowns Colli 

large volume increases caused by wintei and spring 
rains. 

Corbicula concentrations were always greater .ii 
transect 2. Transect 1 had the next greatest concen- 
trations, and transect 3 was low (I ig. 7). 

DISC I SSION 

The size distribution of live clams, and the absence 
of empty shells indicate the oldest hi Capitan clams 
found during 1964 were 2 years old. Although the 
1964 sample size is small, 2 age groups appear 
present with one group ranging from 4.1 mm to 
13.7 mm, and the other from 0.1 mm to 2.5 mm 
(Fig. 3). The smaller group may be the progeny of 
the larger. Sinclair and Isom (1963) found that I 
year old Tennessee Corbicula (probably C. manilen- 
sis) reached sexual maturity at about 6.5 mm, and 
they aged a 29 mm clam at 4 years. Ingram (1959) 
estimated 50 mm Oregon Corbicula were about 
7 years old. 

There are indications that the population fluc- 
tuates yearly, reaching a maximum during the 
winter and a minimum during the summer (Fig. 2; 
Table 1). This pattern may be confounded h\ the 
initial population growth. This pattern is related 
to their reproductive cycle. Late fall and early 
winter reproduction would produce a maximum 
population density during the winter. The popula- 
tion would consequently decrease through natural 
mortality and reach a yearly minimum by the next 
fall. Average shell lengths demonstrate this relation- 
ship with January average lengths less than the 
preceding or following summer average lengths. 
These decreases indicate the January populations 
consist of younger clams and are evidence for a 
fall reproduction period. 

El Capitan Corbicula were probably established 
by veligers from imported waters. Fish and their 
native water from many sources were periodically 
stocked in El Capitan (Fast, 1966). Colorado River 
water via aqueducts was periodically stored in El 
Capitan since 195S. Corbicula were reported from 
the Colorado River aqueduct intake of the Metro- 
politan Water District since 1958 (Ingram. 1 959) 
and are well established in all San Diego County 
reservoirs routinely receiving this aqueduct water. 



96 



till II IIS SOll III- UN ( \l.llORNIA l( I DEMY OF SCIENCES 



VOLUME 70 




Surface 



1964 



1965 



1966 



1967 



Figure 5. El Capitan oxygen concentrations at the surface, 10 m depth, and the bottom from May 1964 through 
September 1967. 



Adult Corbicula are sometimes used as bait in 
California, but this practice is uncommon in San 
Diego County. Their most common natural means 
of dispersal are hydraulic movement of the adult 
clam, and passive movement of the veliger larval 
stage. Diverted or imported water within or between 
drainage systems greatly accelerates dispersal. 

Colorado River water was diverted into El Capi- 
tan Reservoir during October through January in 
1958, 1962, and 1963. Any of these importations 
could have introduced veligers. The California 
Department of Water Resources (1967) observed 
veliger larvae in the Interim Canal of the South 
Bay Aqueduct from October 29 through January 
20. They sampled from October 22, 1964 through 
May 6, 1965. Their adult clams had characteristics 
of both C. manilensis and C. fluniinea. 

Physical-chemical stratification and sediment 
composition are probably the two most important 
factors affecting Corbicula distribution within a 
reservoir. Hypolimnetic stagnation during 1964 
undoubtedly restricted clams to shallow depths. 
Still, some live clams were collected below the 



oxygen deficit. Although Corbicula have a high 
tolerance to low oxygen tension (Sinclair and Isom, 
1963) prolonged exposure to 0.0 mg/1 oxygen is 
fatal. Clams collected between 8 and 12 m may 
have been transients and move up and down the 
slope to avoid prolonged exposure to the stagnant 
water. Another possibility is that they reside in 
this depth zone. Internal seiches of the thermocline 
could have alternately bathed them with stagnant 
and aerated water, and although the oxygen deficit 
was relatively constant in the middle of the lake, 
fluctuations in its depth may have occurred at the 
lake bottom-thermocline juncture. 

Corbicula maximum depth distributions were 
about the same during the summers 1964 and 1965 
even though the reservoir was destratified after 
mid-June 1965 (Figs. 4, 5). This indicated their 
depth distribution was largely determined before 
mid-June and the adult clams did not actively 
migrate after this period. 

Clams were abundant at all depths during 1966. 
I attribute this dispersal to early air injection, since 
strong stratification was not allowed to develop 



1971 



ASIATIC CLAM INVASION IN CALII ohnia 



</7 




£ -5 



Depth intervals in meters 

Figure 6. Mean particle diameter of El Capitan sedi- 
ments collected from transects I, 2, and 3. 



anytime during 1966. The clams were still most 
abundant between 4 and 14 m. This was probably 
a result of a time lag in their dispersal, rather than 
a preference for the sedimental conditions, since 
they were scarce in this interval during 1967. How- 
ever, sediment conditions were quite varied at T-l. 
The shallow depths were predominately sand and 
gravel (Fig. 6), whereas, the deeper areas were pre- 
dominately fine colloidal matter with a high organic 
content (Table 2). 

Most Corbicula were again restricted to shallow 
depths by stratification during 1967, but about 30% 
were found within the hypolimnion. Hypolimnetic 
conditions during 1967 were not as severe as during 
1964. but oxygen concentrations approached zero 
by September 1967. While some Corbicula tolerated 
the low hypolimnion oxygen concentrations and 
associated stagnation, many may have perished 
after stratification developed. This could have con- 
tributed to the great population decrease between 
January 1966 and August 1967. 

The effect of reservoir volume changes on Cor- 
bicula are unknown. Increased volume increases 
the total bottom area available to the clams. This 
new area is predominately sand and gravel covered 
by sparse terrestrial vegetation. Decreased water 
volumes concentrate the clams and creates less 
desirable conditions. The rapid drawdowns during 
1966 and 1967 may also have contributed to the 
high mortalities, but Sinclair and Isom (1963) do 
not recommend water level change for controlling 
Corbicula. 

Several factors undoubtedly influenced the clam 
spatial distribution between transects. Foremost 
among these considerations are: (1) sediment par- 



ticle size and i ompi isition (2) n\ paid rn 
( u location ol initial introdui tion 

i ransects l and 2 have similai mean parti< li 
distributions with depth (I ig 6) However, tram 

2 has much coarsei sed :nts in the shallow depths 

There appears in be a direct relationship between 
the mean particle size of the shallow water sediments 
at each transect and the clam densitii I r .insect 3 
is mostly line colloidal m.iliiial at all depths and 

always had very low Corb'u ula densities. San I ran 
cisco Bay Corbicula prefer sandy substrates tn mud 
(Filice, 1958). Other authors observed similar pref- 
erences by Corbicula (Villadolid and Del Rosario 
1930; Cahn, 1951). 

Surface water currents usual I) flow from the dam 
towards the upper end of the reservoir (I ig. I). 
Veliger positive phototaxis would result in their 
transport from transect I towards transects 2 and 3. 
By this rationale, transect 3 receives the greatest 
concentration of young Corbicula, transect 2 the 
next greatest and transect 1 the least. Such a con- 
centration gradient among mature clams was not 
observed. The hypothetical larval distribution may 
be modified by active selection for sediment com- 
position. If the veligers select for larger sediment 
particle size, transect 2 may be the most desirable. 
Most of the larvae would reject transect 3 and delay 
settling until they were carried by the currents to 
transect 2 or 1. The fine sediments of transect 3 
could also cause high mortality during the matura- 
tion of those clams that settle there. 



400 
300 



I 3 3 
Winter 
1964-7 



JX4- 



: : ■ : -" 

Svmn.tr S»*i«ii 

1966 1967 



Figure 7. Numbers of El Capitan Corbicula collected 
at transects 1. 2. and ? from summer 1964 through sum- 
mer 1967. Transects 2 and 3 were not sampled during 
summer 1966. 



y,\ 



in I i i 1 1\ SOI / /// AW ( tUFORXl.t k mi \n <>l Mil \( i s 



VOLUME 70 



I Mil i 2. Total solids, chemical oxygen demand, and 
nitrogen content of F.I Capital) Reservoir bottom sedi- 
ments during August and November I «)64 at Transect 1. 



\l ,.i si I'M, ! 



NOVEMHKR I 964 



Depth % total C.O.D. 
(meters) solids (mg/gm) 



?N 2 % total COD. r .N, 
solids (mg/gm) 



0.5 


63.6 


43.27 


0.099 


79.0 


4.49 


0.014 


5.0 


67.6 


25.12 


0.082 


67.0 


. 49.78 


0.130 


10.0 


22.2 


105.48 


0.390 


46.0 


65.70 


0.186 


15.0 


25.8 


85.25 


0.302 


24.2 


101.70 


0.352 


20.0 


20.0 


125.05 


0.468 


23.2 


122.21 


0457 



ACKNOWLEDGMENTS 

Many people contributed to this study and their con- 
tributions are appreciated. William J. Clench, Museum 
of Comparative Zoology, Harvard University, identified 
the clams. Robert Doyle measured all Corbicula lengths, 
made many of the sample counts, and devoted much 
time to this study. R. O. Fox, California Academy of 
Science, reviewed the manuscript. 

LITERATURE CITED 

Anderson, R. D. 1959. A modified flotation tech- 
nique for sorting bottom fauna samples. Limnol. 
Oceanog., 4: 223-225. 

Cahn, A. H. 1951. Clam culture in Japan. Gen. Hdq. 
Sup. Comin. Allied Powers Nat. Res. Sec. Report 
No. 146: 1-103. Reprinted as Fisheries Leaflet No. 
399 USFWS. 

California Department of Water Resources. 1967. Water 
quality and biologic conditions. South Bay Aque- 
duct, 1962-1966. State of California, Dept. of 
Water Resources, mimeographed, 180 pp. 

Fast, A. W. 1966. Fisheries management of El Capitan 
Reservoir, San Diego County, California. 1960- 
1962. California Dept. Fish and Game. Inland 
Fish. Admin. Rept. (66-16), 16 pp. 



— . 1968. Artificial destratification of El Capitan 
Reservoir by air injection. Part I, effects on the 
chemical and physical parameters. California Dept. 
Fish and Game. Fish. Hull., 141, 97 pp. 



— . 1970. An evaluation of the efficiency of zoo- 
benthos separation by sugar flotation. Prof;. Fish- 
Cull.. 32: 212-216. 

Filice, F. P. 1958. Invertebrates from the estuarine 
portion of San Francisco Bay and some factors 
influencing their distribution. Wasmann J. Biol., 
16: 159-211. 

Ingram, W. M. 1948. The fresh-water clams of Cali- 
fornia, Oregon, and Washington. J. Ent. Zool., 
40: 72-92. 



. 1959. Asiatic clams as potential pests in Cali- 
fornia water supplies. J. Amer. Water Works 
Assoc.. 51: 363-370. 

Isom, B. G. 1971. Evaluation and control of macro- 
invertebrate nuisance organisms in freshwater 
industrial supply systems. Tennessee Valley Auth., 
mimeographed, 13 pp. 

Prokopovich, N. P. 1969. Deposition of clastic sedi- 
ments by clams. J. Sed. Pet., 39: 891-901. 

Sinclair, R. M. 1964. Clam pests in Tennessee water 
supplies. J. Amer. Water Works Assoc, 56: 592- 
599. 

Sinclair, R. M., and B. G. Isom. 1963. Further studies 
on the introduced asiatic clam (Corbicula) in 
Tennessee. Tennessee Stream Pollution Control 
Board, Tennessee Dept. Public Health, mimeo- 
graphed, 75 pp. 

Villadolid, D. V., and F. G. Del Rosario. 1930. Some 

studies on the biology of Tulla (Corbicula manil- 
ensis Phillippi), a common clam of Laguna de Bay 
and its tributaries. Philippine Agriculturist, 19: 
355-382. 

Accepted for publication June 16, 1971. 



BULLETIN SO. CALIF. A( A DEMY ()l SCIENCE 70(2) 99 102 1971 

TWO BOPYRIDS (ISOI'OI)A) I ROM NI-.W GUINEA 

( ' 1 1 a i< i i . s (i. Daniori ii ! 



Abstract; The female Parathelges weberi has been previously described 
but the male, host, and locale Were unknown. These data are hen pro ided 
and a new species of Pseudione is described from the same area. 



Four parasitized crabs were obtained from the 
collection of the "Alpha Helix" Expedition to New 
Guinea. From a study of these forms, it was deter- 
mined thai there were 2 epicarid isopod genera — 
one parasite represented a new species of Pseudione 
(a branchial bopyrid); while the other was the first 
record for the male, .the host, and the area for 
Parathelges weberi (an abdominal bopyrid). 



Parathelges weberi Nierstrasz 
and Brender a Brandis, 1923. 

Figure I A 

Host: Female of Calcinus laevimanus (Randall). 
Site: Lagoon, 22 miles north of Maiwara, Northeast 
New Guinea, at 0-2 feet, on broken coral bottom. 
Date: 21 September 1969. Collector: K. Kirk. I. 
Richards, and E. Ball. The host and site given here 
are the first record for the species. 

Description: Female. Essentially the same de- 
scription as that given by Nierstrasz and Brender a 
Brandis (1923). Located on the dorsal abdomen of 
the host, with the head of the parasite directed 
towards the posterior of the host, and the dorsum 
of the bopyrid against the host. The measurements 
of the holotype were 9 mm by 3.25 mm, whereas 
the new specimen measured 6 mm by 2.5 mm. No 
hyperparasites were found in the marsupium of 
the "Alpha Helix" form, whereas, a Duplorbis 
smithi was reported from the brood chamber of 
the "Siboga" specimen (Nierstrasz and Brender a 
Brandis, 1923). 

Male. 2.5 mm greatest length. 1.0 mm greatest 
width. Antennae I short, 2-3 articles. Antennae II 
long, 5 articles, with many flagellum subdivisions. 
Eyes obvious, though small. Cephalon definitely 
separated from thoracomere I. Pereopods in a group 
of 3 pairs, then a gap. followed by the posterior 4 
pairs. No mid-ventral tubercles. Scattered, weak. 
dorsal pigment spots. Lateral portions of tho- 
racomeres non-contiguous, with the largest gap 
between III and IV. All thoracomeres the same 
length. Pleon completely fused, with a clear "enve- 
lope" peripherally. Pleon length about equal to 



total width of 3 thoracomeres. No plcopods Slighl 

mid-ventral pigment spots 

Remarks: Whitelegge (1897) described a new 
bopyrid species from Funafuti (Ellice Islands) 
which he named Athelgue aniculi. Bonnier (1900) 
changed this to Parathelges nun uli (Whitelegge, 
1897), on the basis of the lack ol sudden size dimi- 
nution between the pcreon and the pleon. such as 
is found in Athelges (accepted spelling). Nierstrasz 
and Brender a Brandis 11923) recognized this 
generic difference, and erected the new species /' 
weberi for a specimen in the "Siboga" collection 
which showed neither host nor locale. These authors 
indicated that P. weberi was similar to P. aniculi. 
but gave adequate points of difference to justify the 
former species. 

There are 2 other species of Parathelges — P. 
racovitzai Codreanu ( 1 940) and P. whiieleggei 
Nierstrasz and Brender a Brandis (1931) — plus 
several related genera such as Anathelges Bonnier 
(1900), Athelges Hesse (1861), and Metathclges 
Nierstrasz and Brender a Brandis (1923). Pre\ious 
works in New Guinea by authors such as Nobili 
(1905) and Weber (1892), did not mention any 
form which could have been Parathelges weberi. 

Despite the wide European and Pacific distribu- 
tion of the "Athelges" types, there ace apparently 
none in North America (Danforth. 1970). 

The female and male described herein have been 
deposited in the Allan Hancock Foundation Collec- 
tion, University of Southern California. 

Pseudione novaeguineensis, new species 
Figure 1 B-G 

Pseudione Kossman 1881. 

Host: Clibanarius sp., aff. longitarsus (De HaanK 
Site: On mangrove roots, near Maiwara. New 
Guinea, lat. 5°4.4' S. long. 145°43.7' E. Date: 26 
October 1969. Collector: E. Ball. 

Description: Female. Length 3.5 mm. width 2.0 
mm. From right gill chamber of host: head of 
bopyrid directed downward and to rear. Marsupium 



'3612 Angelus Avenue. Glendale. California 91208. 



99 



100 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 




§ 



D 




E 




Figure 1. A. Parathelges weberi, dorsal aspect of male, 
scale equals 1 mm. B-G, Pseudione novaeguineensis, 
new species: B, dorsal aspect of female, scale equals 1 
mm; C, inner aspect of oostegite I; D, ventral aspect 
of uropoda of female; E. right pleopod I of female, 
exopodite is the larger; F, ventral portion of female 
pleon. right pleopods removed; G, dorsal aspect of male, 
scale equals 1 mm. 



towards host branchiostegite. Ccphalon slightly 
bilobed posteriorly. Eyes present, but tiny. Frontal 
margin of ccphalon upturned to form a "velum." 
Head sunken into thorax. 

Thorax of 7 thoracomeres, 1-1 V with anterolateral 
and post-lateral plates. Coxal plates very poorly 
indicated on first 4 right segments. Seven pairs of 
small, well-formed pereopods. No swelling on basal 
articles of the legs. Five pairs of oostegites, barely 
meeting over the marsupium. Oostegite I with 
pronounced teeth on inner aspect. Dorsum quite 
concave, without tubercles. Two epicaridea larvae 
in the marsupium. 

Abdomen of 5 pleomeres, tapering rapidly pos- 
teriad. Pleomeres strongly arched antcromedially, 
clearly demarcated, with pleural ends non-contig- 
uous. Segment V so arched as to appear bifurcated. 
Five pairs biramous pleopods. each ramus leaflike 
and subequal. One pair uniramous uropods on 
telson, which appears to be fused to mid-ventral 
surface of pleomere V. 

Male. Length 1.1 mm, width 0.3 mm. Body 
fusiform. Head sunken into, but not fused with, 
thoracomere I. Eyes obvious. Antennae I and II 
short (2-3 articles), barely evident in dorsal view. 

Thorax of 7 thoracomeres, having widely sep- 
arated terminations. Seven pairs of large pereopods. 
No mid-ventral tubercles. Scattered pigment spots 
dorsally. 

Abdomen of 6 pleomeres, sixth sharply arched. 
Lateral plates widely separated. No pleopods, al- 
though occasional swellings present and unpaired. 

The female holotype and the male allotype are 
deposited in the Allan Hancock Foundation Col- 
lection, no. 694 and 694a, respectively. 

Remarks: Two other crabs of the same descrip- 
tion, and parasitized by the same bopyrid species, 
were found at the collecting site. However, none 
was as clearly segmented as were the selected 
holotype and allotype. Both sexes of the host were 
parasitized. 

Descriptions and identifications of Pseudione in 
the literature leave considerable question as to 
uniformity of critical features. As summarized by 
Danforth (1970), Sars (1899) indicated that Pala- 
egyge hoylii was equivalent to Pseudione affinis; 
Barnard (1920) said that Palaegyge and Pseudione 
could be best separated by the warts on the female 
pleopods (which are variable); Caiman (1898) ap- 
parently felt that Palaegyge borrei was essentially the 
same as Pseudione giardi; Nierstrasz and Brender 
a Brandis (1929) felt that Palaegyge might be 
Probopyrus, and Van Name (1936) indicated that 
if this were so, Probopyrus had priority; Nierstrasz 
and Brender a Brandis ( 1 93 1 ) substituted Pseudione 
for the genus Cryptione; Stock (1960) felt that 



1971 



BOPYRIDS I ROM NEW (it UN I. A 



101 



Pleurocrypta porcellanae should be Pseudione con- 
vergens, but ihis was contested by Bourdon ( 1965); 
etc. 

Willi approximately (■>() species of Pseudione so 
far described, identification of a new form is quite 
difficult, and requires a survey of the original 
papers. The only original description not obtained 
for comparative study was that by Rodriguez y 
Pemenias (1886), in which Kossmann apparently 
described Palaegyge fraissei as a footnote on page 3. 
This form was then called Pleurocrypta balearica 
by Giard and Bonnier (1X87), and later Pseudione 
fraissei by Stebbing ( 1893). In view of the complex- 
ity and contusion surrounding the genus, Nierstrasz 
and Brender a Brandis (1923) attempted to sort 
Pseudione into 3 groups: (a) having no abdominal 
plates, (b) having distinct abdominal plates plus 
unseparated coxal plates, and (c) having distinct 
abdominal plates plus separated coxal plates. How- 
ever, there are many intermediate forms, which 
necessitates checking the original works. 

Pseudione novaeguineensis most nearly ap- 
proaches P. calcinii Shiino (1958), and P. diogeni 
Popov ( 1924). It is very close to the first, but differs 
from the female in the sharpness of the pleomerc 
plates, in the greater width and shorter length of the 
uropoda, in the greater breadth of the pleopoda. 
in the presence of teeth on the inner aspect of 
oostegite I, in the double flexion of the body axis, 
in the more narrow body width, and as to the host. 
There was no male reported for P. calcinii. 

Although the females of P. diogeni and P. nova- 
eguineensis are similar, differing only regarding 
oostegite I, the mid-dorsal pleon fusion, and the 
separation of pleopod rami, the males of the 2 
species are drastically different. [It is interesting 
to note that Caroli (1948) felt that P. diogeni might 
be the same as Gyge arcassonensis] . There are also 
points of similarity with the female of P. giardi 
Caiman (1898), and with the male of P. tattersalli 
Nierstrasz and Brender a Brandis (1923), but in 
each case the descriptions of the other sex do 
not coincide. 

As can be concluded from the foregoing the 
taxonomy of bopyrids as a whole, and of Pseudione 
specifically, should be critically reviewed. 

ACKNOWLEDGMENTS 

I am indebted to Eldon Ball of the University of Cali- 
fornia, Santa Barbara, and to Janet Haig of the Allan 
Hancock Foundation. University of Southern Cali- 
fornia, for the opportunity of studying these interesting 
parasites. 

LITERATURE CITED 

Barnard. K.. 1920. Contributions to the crustacean 



fauna ol South Afrii a No 6 i urthi i iddill 
tliu list "I iri.n n» Isopo 
Mus.. Cape Town, 17; 119 

Bonnier, J. 1900, ' onti ibution ■> I fcludi di i , . 
Les Bopyridae Publications <l< la Station / 
iogique de II Imi reux II I ravi Ic la St - 

8: 1-478. 

Bourdon, R, 1965. Remarqucs au sujcl de la nouvellc 
espece "Pseudione com ■ 
caride de la famille* des Bopyridae) Call lei 

Biologic Murine, (,. 17.1-179. 

Caiman, W. 1898. On a collection ol < rustacca from 
Pugel Sound. Ann. New Yorl Acad. Sir/., II 

259-292. 

Caroli, E. 1948. Identitadi Gyge arcassonen I 

con Pseudione euxini< a Popo\ I'uhhlic nrioni delta 
Stazione Zoolog'u a di Napoli, 2 I : US- 1 52. 

Codreanu, R. 1940. Sur un Epicaride nouveau 

thelges racovitzai n.sp., ectoparasite abdominal 
du Pagure Diogenes pugilator de la \lcr Noire. 
Comptes Rendus de I' Academic des v iem e, Paris, 

210: 679-681. 

Danforth, C. 1970. Epicaridea (Crustacea: Isopodai of 
North America. Univ. Microfilms. Ann Arhor. 
Michigan, 191 pp. 

Giard A., and J. Bonnier. 1887. Contributions a I'ctudc 
des Bopyriens. Travaux de /'Institute Zoologiqui 
de Lille el du Laboratoire de Zoologie Maritime 
de Wimereux. 5: 1-272. 

Hesse, M. 1861. Memoire sur deux nouveaux genres 
de 1'order des Crustaces Isopodcs. Annates des 
Sciences Naturelles, ser. 4, Zoologie, 15:91-116. 

Kossmann. R. 1881. Studien iiber Bopyriden. I. G 

tione moebii und Allgemeines iiber die Munducrk- 
zeuge der Bopyriden. Zeitschrifl fur Wissenschaft- 

licbe Zoologie, Leipzig. 35: 652-665. 

Nierstrasz. H.. and G. Brender a Brandis. 1923. Die 
Isopoden der Siboga-Expedition. II. Isopoda Gcn- 
uina. I. Epicaridea. Pp. 57-122. In Siboga-Expeditie 
1899-1900. 19 (monograph 32bl. E.J. Brill. Leiden. 

. 1929. Papers from Dr. Th. Mortensen*s Pacific 

Expedition 1914-16. Epicaridea I. Videnskabelige 
Meddelelser fra Dansk naturhistorish Forening i 
K0benhavn. 87(48): 1-44. 

. 1931. Papers from Dr. Th. Mortensen's Pacific 



Expedition 1914-16. Epicaridea II. Videnskabelige 
Meddelelser fra Dansk naturhistorisk Forening i 
K0benhavn.9\(51): 147-225 

Nobili. G. 1905. Decapodi e Isopodi della Nuova 
Guinea Tedesca. raccoltidal Sign. L. Biro. Annates 
historiconat urates Musei Nalionalis Hungarici. 

Budapest, 3:480-507. 

Popov. VV. 1929. Rhizocephala and Bop\ ridae of the 
Bay of Sevastopol. SevastopoKskaia biolociches- 



102 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 



kaia stantsiia-Trudy. Academiia Nauk SSSR Len- 
ingrad, I: 1-26. 

Rodriguez y Femenias, J. J. 1886. Historia natural de 
las Baleares. Zoologia. Adiciones a la Fauna 
Balear. Mahon, B. Fabregues, imp. 

Sars. G. 1899. An account of the Crustacea of Norway. 
Part II. Isopoda. Bergen Museum Publication. 

270 pp. 

Shiino. S. 1958. Note on the bopyrid fauna of Japan. 
Rept. Faculty of Fisheries, Prefect ural University 
ofMie, 3:29-73. 

Stebbing. T. 1893. A history of Crustacea. D. Appleton, 
New York, 466 pp. 

Stock J. 1960. Notes on Epicaridea, 1, 2. Crustaceana, 
1:28-33. 



Van Name, W. 1936. The American land and fresh- 
water isopod Crustacea. Bull. Amer. Mux. Nut. 
Hist., 7 1: 1-535. 

Weber, M. [ed], 1892. Die Siisswasser-Crustaceen des 
Indischen-Archipels, nebst Bemei kungen iiber die 
Siisswasser-Fauna in Allgemeinen. Pp. 528-571. In 
Zoologische Ergebnisse einer Reise in Nieder- 
landisch Ost-indien. Vol. 2. Brill, Leiden. 

Whitelegge, T. 1897. The Crustacea. Pp. 127-151. In 
The atoll of Funafuti, El lice group; its zoology, 
botany, ethnology and general structure. Part II, 
No. 6. Memoirs of the Australian Museum, Syd- 
ney, No. 3. 



Accepted for publication June 11, 1971. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCE 70(2): 102-103, 1971 

A NEW INTRANASAL CHIGGER OF THE SUBGENUS CRYPTICULA, 
GENUS MICROTROMBICULA (ACARINA: TROMBICULIDAE) FROM TEXAS 

Richard B. Loomis and James P. Webb, Jr. 1 

Abstract: A new species of Microtrombicula ( Crypt iculaj is based on 
intranasal larvae from Peromyscus pectoralis trapped near Del Rio, Val Verde 
Co., Texas. 



Webb and Loomis (1970) reported nine intranasal 
species which belong to the subgenus Crypticula 
Webb and Loomis of the genus Microtrombicula: 
five from North America and four from Korea. 
Microtrombicula diabola Webb and Loomis (1970) 
was taken from two Peromyscus difficilis nasutus 
(rock mouse) trapped near Del Rio, Texas and 
attempts to obtain additional larvae from near the 
type locality (now under water of the Amistad 
Reservoir) yielded a second new species which is 
described below. Studies upon which this paper 
is based were supported by the U.S. Public Health 
Service Research Grant AI-03407 from the National 
Institute of Allergy and Infectious Diseases. 

Microtrombicula welbourni, new species 
Figure 1 

Types. — Larvae: Holotype and 1 1 paratypes 
from the nasal passages of 2 Peromyscus pectoralis 
laceianus (white-ankled mouse) taken 10 mi N, 
8 mi W Del Rio, 1000 ft. Val Verde Co., Texas, 
28 July 1970, by W. C. Welbourn, Jr., R. B. Loomis, 
and R. C. Stephens, original numbers WCW700728- 
15 (holotype and 10 paratypes) and WCW700728- 



14 (1). The holotype and one paratype will be 
deposited in the collection of the Rocky Mountain 
Laboratory, Hamilton, Montana, and available 
paratypes will be sent to appropriate institutions. 

Diagnosis. — Larva differing from all other mem- 
bers of the subgenus Crypticula in having 3-5 setae 
on coxa III (M. ornata bisetose, other species uni- 
setose), and from M. diabola in possessing eyes, 
nude galeala, and dorsal knob on cheliceral blade. 

Description of holotype (all measurements in 
microns, with means and ranges of all 12 types in 
parentheses, unless otherwise indicated). — Body 
engorged, 580 by 410; color in life, pale yellow; 
eyes 2/2, anterior larger, ocular plate indistinct. 

Dorsal setae 2-6-6-6-6+14, total 40; measure- 
ments of humeral seta 43, anterior dorsal seta 29, 
posterior dorsal seta 27. 

Ventral setae 2-2 (sternals) + 48, total 52; meas- 
urements of anterior sternal seta 29, anterior ventral 
seta 28, posterior ventral seta 29. 

Scutum: Subpentagonal; with large puncta in 
reticulate pattern; sensilla flagelliform with 3-5 



■Department of Biology, California State College, Long Beach, 
California 90801. 



1971 



NEW SPECIESOF IN 1 KANASAl ( iik.ci i< 







Figure I . Microtrombicula (Crypticula) welbourni, new 
species. A. Scutum and eyes. B. Dorsal aspect of gna- 
thosoma. C. Ventral aspect of palpotibia and tarsus. D. 
Leg I, genu, tibia, and tarsus, showing nude setae (mea- 
surements in microns) and bases of branched setae. E. 
Leg II, as above. F. Leg III, as above. G. Body setae: 1 
ST, first sternal, H. humeral, PD. posterior dorsal. H. 
Coxa III. 



distal branches and several short proximal barbs. 

Scutal measurements: AW 36 (34. 32-36), PW 50 
(48.5, 47-50). SB 21 (20. 19-22). ASB21 (22.21-23). 
PSB 25 (26, 25-28), AP 28 (31, 28-33). AM 28 
(29, 25-33), AL 24 (21, 19-24; 1 1). PL 29 (30. 28- 
32), S 60 (57, 53-60; 8). 

Gnathosoma: Cheliceral blade with small tri- 



cuspid cap and dorsal 1 nob • hcli 
capitulai stci num lightly pum tati ( ialcala 
Palpal setae B/B/NbB palpotai \w vilh I nude 
and 5 blanched setae, and tarsals 9 palpotibii 
bifurcate, axial prong ilightl 
unisetose, 2 genualae and microgenuala ! tibialac 
and microtibiala, tarsala ! I I ! I 6) distal 

microtarsala, subterminala, parasubtcrminali 
pretarsala; leg II. coxa unisetose, 2 tibialac tarsala 
19(19, 17-20) and proximal nil- imi., i ,,|., |, g ||| 
coxa with 3 and 4 (3 to 5) In. niched setae, gcnuala 
tibiala, and I nude niaslitarsala I 19) 

Leg measurements: I 214 (209. 192-226), II 205 
(190, 174-205). Ill 236 (225. 211-239), total '. ; 
(625, 577-655). 

Remarks. — The cheliceral blade has a prominent 
knob characteristic of the lour Korean species 
described by Ah ( 1964), but not conspicuous among 
the American Crypticula. 

The geographic proximity of M. welbourni ' 
diabola (type localities less than 3 miles apart), may 
indicate an ecological relationship similar to the two 
sympatric species Microtrombicula nasal is and \l 
wrenni Loomis, recovered from the same individuals 
of several cricetid rodent species in Joshua I rcc 
National Monument. California (Loomis. I9( 

This new species is named alter Mr \\ I 
Welbourn, Jr. of California State College. Long 
Beach in recognition of his numerous contributions 
to the studies of trombiculid mites. 

Specimens examined (17). — -Type series (I2l 
plus 5 larval skins (WCW700728-I5). 

LITERATURE CITED 

Ah, H. S. 1964. New intranasal chiggers of the genus 
Microtrombicula from some bats in Korea (Aca- 
rina:Trombiculidael. J. Med. Ent.. 1: 186-191. 

Loomis. R. B. 1963. The discovery of chiggers (Acarina: 
Trombiculidae) in the nasal passages of cricetid 
rodents from California, with descriptions of two 
new species. J. Parasit., 49: 330-333. 

Webb. J. P. and R. B. Loomis. 1970. A new subgenus of 
intranasal chiggers of the genus Microtrombicula 
from North America and Korea. J. Med. Ent., 
7:655-663. 

Accepted for publication August 31, 197 1. 



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cover: A mass stranding of 28 Pacific pilot whales. Globicephala macrorhyncha Gray, at Pyra- 
mid Cove, San Clemente Island, California. The stranding occurred on January 8. 1971 and 
included 6 males, 21 females, and 1 of undetermined sex. Globicephala appears to be subject to 
mass stranding more frequently than any other cetaceans. Climatic and biological factors (high 
tide, no surf, slight wind, sloping beach with steep dropoff, and spawning squid) may have con- 
tributed to the stranding. 



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SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 

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BULLETIN () F THE SOUTHERN CALIFORNIA 

AC A DE M Y O I SC I I. N CIS 

Volumi; 70 March 24, i<?72 ;I) , ,, \ 



CONTENTS 

Taxonomic studies and notes on some Honduran amphibians and reptiles. 

By John R. Meyer and Larry David Wilson 106 

Observations on the distribution and eeology of the littoral ascidians ol the 
mainland eoast of southern California. By Rinimon C. Fay and Jeffrey V. 
Johnson 114 

Benthonic Foraminifora of the Ardath Shale and Stadium Conglomerate 

(Eocene), San Diego Basin, California. By James M. Gibson 125 

A second Recent species of Noetia, sensu stricto (Mollusca: Bivalvia) in the 

tropical eastern Pacific Ocean. By Barry Roth 131 



Errata/??: The date of publication for Vol. 70(2) should be corrected to March 1. 1972. 



105 



BULLETIN SO. CALIF. ACADEMY OF SCIENCE 70(3): 106-114, 1971 

TAXONOMIC STUDIES AND NOTES ON 
SOME HONDURAN AMPHIBIANS AND REPTILES 

John R. Meyer 1 and Larry David Wilson- 



Abstract: Fourteen species of amphibians and reptiles occurring in 
Honduras arc discussed. Anolis humilis, Anolis crassulus, Anolis laeviventris, 
Ninia atrata, Rhadinaea kinkelini, Sibon anthracops, and Thamnophis 
cyrtopsis are recorded from the country for the first time. Eleutherodactylus 
merendonensis, previously considered a synonym of Eleutherodactylus rugu- 
losus, is resurrected. Anolis humilis and Anolis uniformis are raised to specific 
rank from their former subspecific status. Anolis intermedins and Anolis 
nannodes are synonymized with Anolis laeviventris. and Anolis heteropholi- 
dotus is relegated to the synonymy of Anolis sminthus. New distributional 
and taxonomic data arc also given for Anolis anisolepis, Anolis houvierii. 
Anolis cobanensis, Sphaerodactylus dunni, Bothrops bicolor, Dryadophis 
dorsalis, Lampropettis trlangulum, and Sibon carri. 



For the past seven years we have been studying the 
herpetofauna of the Republic of Honduras, and in 
the course of this research have encountered speci- 
mens that have necessitated taxonomic revision or 
have contributed to the clarification of the status of 
little-known forms. Accounts of the distribution, 
relationships, and ecology of all of the Honduran 
amphibians and reptiles will be treated in a separate 
paper. 

Eleutherodactylus merendonensis Schmidt 

This species was described by Schmidt (1933) on 
the basis of a single specimen from an elevation of 
150 m in Santa Ana Canyon, Departmento de 
Cortes. Honduras. In August 1968 we collected 13 
specimens (LACM 48010-22) of Eleutherodactylus, 
which later proved to be referable to merendonensis. 
from the approximate type locality of the species. 
All 13 individuals were found at night in the open 
on boulders, logs, and rock ledges along the Rio 
Santa Ana at an elevation of 200 m; this locality is 
immediately below a small reservoir about 3 km 
west of San Pedro Sula. 

Lynch (1965). in a paper dealing with the rugulo- 
sus group of Eleutherodactylus in northern Central 
America, placed merendonensis in the synonymy of 
E. rugulosus. In synonymizing the two species. 
Lynch stated that "'the difference between merendo- 
nensis and rugulosus rugulosus are [sic] size or due 
to size. The type of merendonensis is a very large 
specimen (80 mm) thus paralleling E. r. natator in 
size. Direct comparison of the type and specimens 
of E. r. rugulosus from Mexico reveals that the two 
are identical." 



We have examined the holotype (FMNH 4672) of 
merendonensis, and find it and the 13 additional 
specimens to be distinguishable from rugulosus in 
the following features: 1) web between toes IV and 
V incised no deeper than distal tubercle of toe V in 
merendonensis (web virtually absent between toes 
IV and V in rugulosus), and 2) inner tarsal fold large 
and flap-like in merendonensis (inner tarsal fold 
ridge-like, not free at edge in rugulosus). In addition, 
merendonensis appears to be a larger and much 
stockier species than rugulosus, with the snout-vent 
length of adult merendonensis being 75-85 mm, 
whereas the maximum snout-vent length of adult 
rugulosus from Honduras is about 60 mm. The two 
species are sympatric (although apparently not in 
the sense of utilizing the same microhabitat), as we 
found rugulosus to be common along the Rio Santa 
Ana below the merendonensis locality at an eleva- 
tion of 100 m, and also in small seeps on the hillside 
above the merendonensis locality. 

Although the relationships of merendonensis are 
apparently with the rugulosus group of Eleuthero- 
dactylus, the species resembles E. punctariolus from 
lower Central America more closely than it does 
rugulosus. Both merendonensis and punctariolus 
are large, stocky frogs, but punctariolus differs 
from merendonensis in having considerably more 
webbing on the feet (toes about '/i to % webbed in 
punctariolus, as opposed to about V3 webbed in 
merendonensis). 



■The Research Ranch, Elgin, Arizona 8561 1. 
2 Horizon School, 9025 Sunset Drive. Miami, Florida 
33143. 



106 



1971 



AMI'IIIIIIANS AND Ul I'l II I Sol HONDl HAS 



107 



Anolis humilis Peters 

Stuart 1 1948) proposed recognition of a subspeci- 
fic relationship among populations of three small 
anoles previously considered as distinct species, 
Anolis humilis, A. quaggulus, and A. uniformis. 
Anolis humilis was described by Peters ( 1 863.) from 
Veragua, Panama, A. quaggulus by Cope ( I K s 5 ) 
from Rio San Juan, Nicaragua, and A. uniformis 
by Cope (1885) from Yucatan and Guatemala. 
According to Stuart's (1948) arrangement, A. Ii. 
uniformis ranges from Veracruz, Mexico to Hondu- 
ras, where it presumably integrades with A. Ii. 
quaggulus, which in turn ranges from Honduras to 
Costa Rica, where it intergrades with A. It. humilis. 
Taylor ( 1954a) described a fourth subspecies, A. h. 
marsupialis, from southeastern Costa Rica. 

In relegating humilis, quaggulus, and uniformis to 
subspecilic status, Stuart (1948) stated that unifor- 
mis differs from the other two races in having larger 
ventrals (35-45 between levels of axilla and groin in 
uniformis, more than 50 in quaggulus and humilis), 
and in having a brilliant purple spot on the dewlap 
(no spot present in quaggulus or humilis). Stuart 
stated further that he felt the lack of specimens of 
humilis (sensu lalo) from between Guatemala and 
Nicaragua was due to insufficient collecting in deep 
forest, and that future collecting in Honduras would 
reveal its presence there. 

We have collected throughout most of Honduras 
and examined all major collections from the region. 
but we have encountered only one specimen of 
humilis from the country (TCWC 23628, from ca. 
40 km E of Catacamas. Departamento de Olancho, 
located about 30 km from the Nicaraguan border). 
The nearest locality record to the northwest is 25 
km SSW Puerto Barrios, Departamento de Izabal, 
Guatemala (TCWC 23744). about 350 km from 
the Honduran locality. Examination of specimens 
of the humilis complex from throughout its range 
has caused us to question the advisability of follow- 
ing Stuart (1948) in synonymizing uniformis with 
humilis and quaggulus. We propose that the two 
subspecies, Anolis humilis humilis and Anolis 
humilis uniformis be raised to specific status, with 
Anolis quaggulus retained as a junior synonym 
of humilis. 

In our opinion, additional collecting in Honduras 
will not appreciably extend the ranges of A. humilis 
and A. uniformis towards one another. In Honduras 
the ecological roles of these species, both ground 
dwelling or low vegetation inhabitants of the deep 
forest, appear to be filled by a close relative of the 
two, Anolis tropidonotus. Anolis tropidonotus. 
which is absent from most of the range of humilis 
and appears to be ecologically separated from uni- 



formis, is the most common anole in the deep forests 
ul l londuras Stuart 1 1955) stated that tropidonotus 
is an inhabitant «>i savanna and dry fores) in ' iuati 

mala, whereas uniformis is restricted i<> the taller, 
more mesic forests in that country In occupying 
the mesic forests in Honduras, it may be that tropi- 
donotus prevents the spread oi humilli and itm/or- 
mis towards one another. 

Separation of the ranges ol humilis and uniformis 
is not in itsell sufficient evidence l"i considering 
the two as distinct species, and accordingly We 
undertook an examination ol morphological fea- 
tures of the various populations. We examined 
51 specimens ul uniformis ami I 17 specimens ol 
humilis from throughout their ranges, and tailed to 
find any feature of scalation by which the two 
species can be consistently separated, including the 
supposed difference in ventrals reported by Stuart 
(1948). There is. however, a constant difference in 
the male dewlap between the populations occurring 
northwestward of Honduras (uniformis) and those 
found from Honduras southward (humilis). This 
difference is not merely the presence versus the 
absence of a spot on the dewlap (Stuart. 1948), but 
rather the dewlap in uniformis consists of a bright 
red ground color with a more or less anterior blue 
to purple spot, whereas in humilis the ground color 
is reddish-orange with a bright yellow border, with 
the scales on the dewlap darkly pigmented so as to 
suggest obliquely transverse black stripes. 

Taylor (1954a) recorded Anolis humilis unifor- 
mis from Costa Rica and noted that the dewlap color 
and pattern was as described above for uniformis. 
Four specimens were reported by Taylor, all from 
Tenorio, Las Canas. Providencia de Guanacaste. 
and a detailed description of one of these (K.U 
40893) was given. In examining this collection, we 
find that this number is allocated to the holotype of 
Anolis humilis marsupialis, and the 4 specimens of 
A. h. uniformis mentioned by Taylor actually bear 
the numbers KU 40922-25. Of this series, only one 
(KU 40925) is a male and examination of this speci- 
men shows that the dewlap is typical of that of 
humilis, described above. In our opinion these 4 
specimens (KU 40922-25) represent Anolis humilis, 
and Taylor was in error in recording A. uniformis 
from Costa Rica. 

Anolis laevivenlris Wiegmann 

The name Anolis laevivenlris has previously been 
applied to a population of medium-sized anoles 
found in southern Mexico and western Guatemala, 
which according to Stuart ( 1948) is pan of a series of 
closely related species also including A. nannodes, 
A. dunni, and A. intermedins. In Stuart's arrange- 



/<« 



HI LLETIh SOI III! R\ (A I IIORXIA l( API \IY Oi SCII.NCIS 



I OI.VME70 



ment, the range of dunni is from Michoacan to 
Guerrero, Mexico, laeviventris from central Vera- 
cruz, Mexico to western Guatemala, nannodes from 
western to central Guatemala, and intermedius from 

northwestern Costa Rica to eastern Panama. We 
have not examined any specimens of A. dunni, 
and this species is excluded from the following 
discussion. 

Anolis laeviventris was described by Wiegmann 
(1S34) from "Mexico." and the type locality was 
subsequently restricted to Jalapa, Veracruz (Smith 
and Taylor, 1 950). Anolis intermediuswas described 
by Peters (1863) from Veragua. Panama. Anolis 
nannodes was described by Cope (1864) from 
Coban. Alta Verapaz. Guatamala; Arriba, Costa 
Rica; and Jalapa. Mexico, and was subsequently 
restricted to Coban by Stuart (1948). 

Dunn (1930) stated that he had compared the 
types of nannodes and intermedins and found them 
to be identical. Stuart (1948) stated that "in describ- 
ing nannodes. Cope unquestionably confused 3 
species intermedins, laeviventris. nannodes," but 
Stuart presented no data to indicate why he con- 
sidered the three species to be distinct. In a subse- 
quent paper. Stuart (1955) indicated that nannodes 
and laeviventris could be separated on the basis of 
the number of dorsal scales between the axilla and 
groin levels ( > 60 in nannodes, < 60 in laeviventris): 
he also indicated that the 3 species differed in head 
scalation (slightly keeled or rugose in laeviventris. 
less so in nannodes, and smooth in intermedius). 

In 1968 we collected 2 anoles (LACM 47291-92) 
obviously belonging to the laeviventris group, at La 
Esperanza. Departamento de Intibuca. These speci- 
mens and an additional one (KU 103239) from 
Cerro Uyuca, Departamento de Francisco Morazan. 
represent the only records for this group between 
central Guatemala and northwestern Costa Rica. 

We have failed to find a difference in the exami- 
nation of over 70 specimens of the three nominal 
species (intermedius, laeviventris, nannodes) from 
throughout the range. The supposed difference in 
number of dorsal scales between laeviventris and 
nannodes does not show any consistent pattern, nor 
is there any consistent difference in dorsal head 
scalation among the three. Examination of several 
other features of scalation shows that there is consid- 
erable overlap in various populations and indicates 
a lack of any clear clinial variation. Furthermore. 
in all three nominal populations, the males have a 
small, white dewlap, a character that is probably of 
considerable evolutionary significance. In view of 
the apparent lack of difference in scalation and the 
uniformity of dewlap coloration in this group of 
anoles, we propose that Anolis intermedius and A. 
nannodes be considered subjective junior synonyms 



o\ Anolis laeviventris. Anolis laeviventris at present 
appears to be restricted to intermediate elevations of 
both versants from southern Mexico to western 
Panama. 

Anolis smintlius Dunn and Kmlen 

This anole was collected by us 12 km SW San 
Juancito, Departamento de Francisco Morazan 
(LACM 47676-78, LSUMZ 21415-16), 3 km NE 
La Esperanza, Departamento de Intibuca (LACM 
47682), and 20 km E Nueva Ocotepeque, Departa- 
mento de Ocotepeque (LACM 47679-81). In the 
original description, Dunn and Emlen (1932) stated 
that A. smintlius is related to A. altae and A. con- 
color. In our opinion this was an erroneous associa- 
tion, and A. smintlius should be included in the 
group also including A. crassulus, A. haguei, and A. 
anisolepis as suggested by Stuart (1955) and Smith, 
Burley, and Frits (1968). Anolis crassulus was 
described by Cope (1864) from central Guatemala, 
A. smintlius by Dunn and Emlen (1932) from San 
Juancito, 2100 m, Francisco Morazan, Honduras, 
A. haguei by Stuart (1942) from 2 km S Finca 
Chichen, Alta Verapaz, Guatemala, and A. ani- 
solepis by Smith, et al., (1968) from 17 km SE San 
Cristobal de Las Casas, Chiapas Mexico. 

In 1968 we secured a series of what we consider 
to be Anolis crassulus, in sympatry with A. smintlius 
3 km NE La Esperanza. Departamento de Intibuca, 
and there is no problem in separating this series of 
crassulus from all known specimens of smintlius. 
Anolis smintlius may be characterized as follows: 
males with brick red to scarlet dewlap with white to 
cream scales in life; dorsal pattern of a broad dark 
brown stripe (S and 9), a narrow white mid-dorsal 
stripe (9 only), or a series of faint paravertebral 
inverted V's (<J and 9); tail rounded to oval in cross- 
section, scales of the dorsal caudal row of flattened 
aspect; mid-ventral scales smooth to faintly keeled, 
34 to 42 (9) and 41 to 46 (3) between axilla and 
groin. Honduran crassulus may be characterized as 
follows: males with orange dewlap with yellow 
scales in life; dorsal pattern a series of distinct, 
inverted paravertebral W's, bordered below by a 
white postocular stripe (S and 9); tail laterally 
compressed in cross-section (strongly so iniJ), the 
scales of the dorsal caudal row strongly keeled and 
ridge-like; mid-ventral scales strongly keeled, 29 to 
30 (9) and 30 to 36 ($) between axilla and groin. 

The above characters consistently separate smin- 
tlius from crassulus in Honduras; all other charac- 
ters of scalation show overlap or are too subjective 
to be of use. Despite the obvious differences between 
crassulus and smintlius in Honduras, specimens of 
the crassulus group from Guatemala and Mexico 



1971 



AMPHIBIANS AND REPTILES <>l HONDURAS 



IU'/ 



have a bewildering array of admixtures of the dis- 
tinctive characters observed in Honduras, except 
thai nowhere outside of Honduras and El Salvador 
(sue below) arc smooth or weakly-keeled mid- 
ventrals encountered. The inter-relationships ol the 
populations thai have received the names sminthus, 
crassulus, haguei, and anisolepis are exceedingly 
complex, and alter examining specimens ol all I 
supposed forms from Mexico to Honduras, we are 
unable to suggest a satisfactory arrangement. In our 
opinion extensive studies of live animals in the field 
will be necessary to clarify the status of these anoles. 

We have examined 2 specimens (U MM/. I I 7647, 
MCZ 57280) from Miramundo, Santa Ana, El 
Salvador, identified by Robert Mertens (Rand, 1957) 
as Anolis heteropholidotus, which was described by 
Mertens (1952a) from this locality. Miramundo 
lies just across the valley of the Rio Lempa from 
the A. sminthus locality in the Departamento de 
Ocotepeque, Honduras, and the 2 specimens men- 
tioned above from Miramundo arc definitely refer- 
able to sminthus. We suggest that Anolis heteropho- 
lidotus be tentatively considered a junior synonym 
of Anolis sminthus, until such time as a complete 
study of the crassulus group is undertaken. 

Two other species need to be mentioned in con- 
nection with Anolis sminthus. Anolis bouvierii, 
described by Bocourt (1873) from Escuintla, Guate- 
mala, was suggested by Stuart (1955) as a possible 
member of a group also containing Anolis cobanen- 
sis and A. heteropholidotus. Since we consider 
heteropholidotus to be a synonym of sminthus, it 
was necessary to examine specimens of both coba- 
nensis and bouvierii. Anolis bouvierii is known only 
from the holotype (MNHN 2464), and we were 
fortunate in being able to examine this specimen 
through the kindness of Jean Guibe, Museum 
Natural d'Histoire Naturelle. Paris, and Jay M. 
Savage. University of Southern California. The 
holotype is a male in excellent preservation, and 
differs in several features from the crassulus group. 
Anolis bouvierii has conical dorsal scales, not 
distinctly differentiated from the laterals (about 10 
rows of keeled, distinctly enlarged dorsals in the 
crassulus group); no enlarged postanal scales in 
bouvierii (row of enlarged postanals in males of 
crassulus group); 68 ventrals between axilla and 
groin in bouvierii (no more than 50 ventrals between 
axilla and groin in males of crassulus group); a very 
large dewlap, extending posteriorly to about halfway 
between axilla and groin in bouvierii (dewlap mod- 
erate in size, not or barely extending posteriorly 
beyond axilla in malesof the crassulus group); color 
of bouvierii. in preservative, light green with brown 
splotching (due to uneven color change) dorsally. 
the throat and sides of neck black, with some black 



extend i hi to shoulders (coloj and pattern variable 

in crassulus group, but never approaching thai "I 
bouvierii). Anolk bouvierii is clearly not related i" 

the (in wul us en nip. in i H 1 1 opinion does ii h 

any close relationship to cobanensis oi any other 
anole from northern Middle Amerii .i 

Stuarl (1955) based the supposed relationship o| 
A. cobanensis and A. heteropholidotus ( iminthu I 

on the presence ol smooth to weakly keeled ventrals 
in the two species I xamination ol more specimens 
ol sminthus than were available lo Smart indicates 
that there is a tendency toward strong keeling in 
all but the mid-ventral scales, so that sminthus 
approaches having the strongly keeled ventrals 
characteristic of all other species ol the crassulus 
group. In addition to the difference in ventrals 
between cobanensis and the < rassulus group, there- 
are no enlarged postanal scales in < obanensis iprcs- 
ent in all crassulus group males), the dorsals are not 
abruptly enlarged from the laterals in cobanensis 
(abruptly enlarged in crassulus group), the ventrals 
range from 53 to 6 1 between the axilla and groin in 
both sexes of cobanensis (no more than 50 ventrals 
between axilla and groin in either sex in crassulus 
group), and the loreals range from 8 to 10 in coba- 
nensis (maximum of 6 in crassulus group i. 

Sphaerodacty lus dnnni Schmidt 

This species was previously known only from the 
type, described by Schmidt (19631 from Rfo Naco. 
near Cofradia, Departamento de Cortes. Honduras. 
In July 1968. we collected 3 specimens (LACM 
47302-04) of this species. 12 km S LaCeiba. Depar- 
tamento de Atlantida. where they were found in the 
daytime among the leaves on the ground along a 
small stream. The junior author secured another 
specimen (LSUMZ 22450) in August 1969. along 
the Rfo Cristales at an elevation of 150 to 200 m in 
the mountains above Trujillo. Departamento de 
Colon. This individual was found in the morning 
moving among the leaves on the ground along a trail 
near the river. 

Scale counts and measurements for the holotype 
(Schmidt. 1936) and for the additional specimens in 
our collections are in close agreement. As the type 
description was based on a preserved specimen, we 
offer the following color and pattern description, a 
composite of notes made on two males (LACM 
47303). LSUMZ 22450) in life. Dorsum brown with 
light transverse markings, which decrease in inten- 
sity posteriorly. Nape region black, grading into 
orange-brown laterally and anteriorly, crossed by 
three white bands, which are more or less confluent 
with a lateral white stripe on each side. Dorsal sur- 
face of the head orange-brown to light brown, with 



110 



HI I II 1 1\ son III l<\ ( A I II OKM I l( \ni MY Ol ' SCI I. NCI. s 



VOLUME 70 



a reddish-brown nasal blotch and a dark reddish u> 
orange-brown parietal blotch, outlined by a Y- 

shaped white lo pinkish white marking, the arms oi 
which are confluent with the light ground color of 
the head: lateral head color orange-brown, with 

pinkish-white tipper labials. Chin white with orange 
to brown reticulate or streak-like markings; throat 
with a mixture of white, orange, and brown scales. 
Venter light brown. Tail orange-brown to reddish- 
brown dorsally and orange ventrally. Iris red. 

Bothrops bicolor Bocourt 

This species of arboreal pit viper is a relatively 
rare snake, and in his description of Bothrops row- 
Icyi, Bogert ( 1968) was able to gather data on only 
7 specimens of B. bicolor. At 9:00 P.M. on 28 June 
1968, we collected a juvenile male of bicolor (LSU 
MZ 23821). 23 km E Nueva Ocotepeque, Departa- 
mento de Ocotepeque, at an elevation of 1730 m. 
At the spot where this snake was collected, there is a 
small stream running through second growth cloud 
forest, and patches of apparently virgin cloud forest 
(Lower Montane Moist Forest formation of Hol- 
dridge, 1962) are present in the area. The snake was 
lying on a low branch overhanging the stream. 

Our specimen agrees with the description of 
bicolor given by Bogert (1968). There are 164 
ventrals, 61 subcaudals, 21-21-15 dorsal scale rows, 
9-10 supralabials. and 10-10 infralabials. 

In life the color pattern was: dorsum chartreuse 
green, dorsal blotches grayish-green, lateral blotches 
sky blue; venter very light green, finely peppered 
with darker flecks, lateral edges of most ventrals 
chartreuse green; head chartreuse green with two 
dark grayish-green bands extending posteriorly 
from the snout to the angle of the jaw, a similar but 
lighter band extending from the posterior edge of 
the eye to the angle of the jaw; some additional dark 
grayish-green mottling between the two bands on 
top of the head; eye chartreuse green with heavy 
black reticulations. 

Although Bogert (1968) stated that Bothrops 
bicolor lacks any markings, color notes made in 
life on LSUMZ 11638 (the only other specimen 
known from Honduras) by the collector. Burt L. 
Monroe, state that the dorsum was grass-green with 
powder blue chevrons and the venter was yellowish- 
green. 

Dryadophis dorsalis (Bocourt) 

Lynch and Smith (1966) suggested that Dryado- 
phis dorsalis be considered a subspecies of the wider- 
ranging Dryadophis melanolomus. This suggested 
change was made on the basis of the pattern of a 



juvenile from the mountains above Zanatepec, 
Oaxaca, Mexico, resembling the pattern depicted 
by Stuart ( 1941 I for dorsalis, and on the fact that 
adults from the same area are stripeless and arc 
allocated to melanolomus. The distribution of the 
two species as pictured by Stuart ( 1941 ) is. suggestive 
of a subspecifie relationship, but recent collecting 
in Honduras has demonstrated overlap in the ranges 
of dorsalis and melanolomus without intergrada- 
tion. In 1968 we collected a juvenile and an adult 
dorsalis (LSUMZ 23822-23) at Finca Los Naranjos 
(=Finca Fe), 2 km W El Jaral. Departamento de 
Cortes, and the same year Ernest A. Liner collected 
a juvenile melanolomus (EAL 2200), 6 km N Agua 
Azul, Departamento de Cortes. Both localities are 
on or near the edge of Lago de Yojoa, and are about 
4 km apart. 

Dryadophis melanolomus in Honduras occurs at 
sea level to 750 m in lowland rainforest, subtropical 
rainforest, scrub and thorn forest, and lowland pine 
savanna, whereas dorsalis occurs from 750 to 1900 
m in subtropical rainforest, upland pine and pine- 
oak forest, and cloud forest. The area where the 
ranges of the two overlap is dominated by forest that 
is ecotonal between the lower, warmer rainforest 
and the higher, cooler cloud forest (this intermediate 
type is designated the Subtropical Wet Forest for- 
mation by Holdridge, 1962). 

Adult dorsalis are olive green dorsally and yellow 
grading to green ventrally. A black mid-dorsal 
stripe occupies the mid-dorsal scale row. and runs 
the length of the body. Juvenile dorsalis are olive 
green on the anterior quarter of the body, grading 
to rust brown on the middle half, and grading again 
to olive green on the posterior quarter of the body. 
The blotches on the anterior portion of the body are 
the most distinct, dark olive green in color outlined 
with black, and the dorsal blotches alternate with 
the lateral ones, contrary to Stuart's (1941) descrip- 
tion. The dorsal blotches are connected medially to 
a wavy, black mid-dorsal line, which continues to 
the end of the tail. 

Adult melanolomus are chocolate brown, light 
brown, or grayish-brown with vague indications of 
darker stripes on scale row 3, and occasionally on 
rows 6 to 9. Juvenile melanolomus have a similar 
ground color to that of the adults, with light bars 
about one-half scale in width extending completely 
across the dorsum. This is the type of pattern figured 
by Stuart (1941, pi. 1. fig. 6) for D. dorsalis and 
D. melanolomus laevis. 

At least some juvenile dorsalis have a pattern of 
alternating dorsal and lateral blotches, and we have 
shown that melanolomus and dorsalis are sympatric 
and distinct in the Lago de Yojoa region of Hon- 
duras. Therefore, we prefer to regard dorsalis as a 



1971 



AMI'IIIIIIANS AND REPTILES <>l HONDl R \S 



III 



species distinct from melanolomus, especially in 
view of the somewhat confused situation presently 
existing with material from the Tehuantepec region 
of Mexico. Lynch and Smith (1966) have discussed 
this situation and regard the suhspceilic slat us of 
the circumisthmal populations of melanolomus as 
in doubt, and their allocation of the Zanatepec 
juvenile to dorsalis is in nc^\ of confirmation. 

Lampropeltis trlangulum Lacepede 

A specimen (LSUMZ 238241 from 4.5 km SE 
Catacamas, Departamento de Olancho, is unusual 
in having a color pattern composed only ol red and 
black rings. In life the dorsum was dark red with 4 I 
black body bands and 10 black tail bands. The 
venter was reddish-orange. 

We are aware of only one other such specimen 
having been reported; Dunn and Bailey (1939) 
recorded a bicolor Lampropeltis trlangulum micro- 
pholis from near the Canal Zone in Panama. Our 
specimen was taken at night in a partially Hooded 
cornfield. 

Ninia at rata (Hal lowed) 

Prior to our discovery of this species in Honduras. 
Ninia atrata was not known north of Costa Rica 
(Burger and Werler, 1954; Scott, 1969). During the 
summer of 1968, we collected 3 specimens (LSUMZ 
23825-27) from under logs in a clearing in cloud 
forest (Lower Montane Moist Forest formation 
of Holdridge, 1962). 21 km E Nueva Ocotepeque. 
Departamento de Ocotepeque. In life the color 
pattern was: dorsum gray; venter cream with some 
gray flecking on the posterior portion, especially 
the underside of the tail; supralabials cream; iris 
black; no evidence of a nuchal band. 

The absence of a light nuchal band in the Hondu- 
ran specimens is a point of dissemblance with typical 
atrata (Burger and Werler, 1954; Lynn, 1959; Roze, 
1966; Taylor. 1951). Roze (1966). however, noted 
that the nuchal band "at times, may be inconspic- 
uous or absent" (translation ours) in Venezuelan 
individuals. We have compared the Honduran 
material with two specimens of atrata from the 
provinces of Bocas del Toro (KU 112621) and 
Darien (KU 1 12622) in Panama, and can note no 
difference in color except for the presence of the 
typical nuchal band in the Panamanian specimens. 
The nuchal band of KU 1 1 262 1 is orange and that 
of KU 112622 is cream. In addition, comparison 
of the hemipenes of KU 112622 with those of 
LSUMZ 23825 and LSUMZ 23827 reveals no 
difference in structure. Scale counts for the Pana- 
manian and Honduran specimens fall within, or are 



close to, tin- i. mi"", given i"i atrata bj Burgei and 

Werler (1954) ("he Honduran »pc ins all males. 

have 142 u< I 16 vcntrals and 59 to 60 subcaudals 

In view of the facl thai onlj il" lai i "i a nuchal 
collar distinguishes tin- Honduran spe sna from 

typical atrata, and since some Venezuelan speci- 
mens also lack collars, we regard the Hondu- 
i.iii specimens as representatives ol Ninia atrata 
Whether the Honduran population is character- 
istically collarless cannol in- answered until more 
material becomes available. 

We accept the use of the name atrata instead "I 
lansbergi lor this species ol Ninia. in accordance 
with the decision presented in Opinion f>44 ol the 
International Commission on Zoological Nomen- 
clature (Bull. Zool. Nomencl.. 20; 2d) to supresslhe 
combination Coluber at rat us Omelin. 1768 and to 
validate the combination ( bluberatratus Hallowed, 
1845. 

Rhadinaea kinkelini Bocttgcr 

This snake has previously been recorded only 
from Nicaragua by Boettger (IK9K) in the type 
description. On 27 June 1968. we collected a single 
specimen (LSUMZ 23828) of R. kinkelini from 
under a log in a clearing in cloud forest (Lower 
Montane Moist Forest formation of Holdridge. 
1962). at an elevation of 1900 m. 21 km E Nueva 
Ocotepeque. Departamento de Ocotepeque. This 
specimen was identified for us by Charles W. Myers, 
who is currently revising the genus Rhadinaea. 

In life the color pattern was: dorsal ground color 
brown; dorsal scale rows 1 to 3 yellowish-cream 
with light brown flecking mostly along the edges of 
the scales, creating the impression of a stripe at the 
edge of each scale row: scale row 4 dark brown, row 
5 tan. and row 9 brown with a very' dark brown 
stripe down the center: head dark brown above with 
a tan collar about two scales long, two scales behind 
the parietals: supralabials mostly cream with dark 
brown edging; chin cream: anterior venter creamy 
yellow grading to cream posteriorly 

Our specimen is a male with 148 ventrals. 81 
subcaudals, 17-17-17 dorsal scale rows, and 8 supra- 
and infralabials. 

Sibon anthracops Cope 

A male specimen (LSUMZ 23S29) from 0.5 km 
N Coyoles. Departamento de Yoro. represents the 
first record for Sibon anthracops from Honduras 
and from the Caribbean versant: previously, the 
distribution was regarded as the Pacific slope of 
Costa Rica and Nicaragua ( Peters. 1 960). The range 
of 5. anthracops is poorly known, and only 3 of the 



//. 



hi 1 1 1 n\ sni i in i<\ i ii ii orxia ACAni.M) or sciences 



i on / mi: 70 



specimens cited bj Peters (1960) and one cited bj 
Taylor (1954b) have precise locality data (all four 
are from Costa Rica). 

Data for the Honduran specimen are: ventrals 
166; subcaudals 82; dorsal scale rows 13-13-13; 
anal plate single: supralabials 7-8, with the 3rd, 
4th. and 5th entering the orbit on the left side, and 
the 4th. 5th, and hth entering on the right; no 
preoculars; loreal single, elongate, entering the 
orbit: postoculars 2-3: temporals 14-2/1 + 2: maxil- 
lary teeth 15; total length 664 mm. tail length 
I 53 nun. 

In life the color pattern was: dorsum white with 
19 black body bands, some extending onto venter 
to form rings, and others not meeting on venter; 15 
black bands on tail: scale rows 3 to I 1 orange 
between the black body bands; orange dorsal colora- 
tion most pronounced and extensive posteriorly. 
and continuing onto tail: venter white: head mostly 
black, with thiscoloration extending laterally almost 
to lip. posterolateral ly as far as the anterior half of 
the anterior temporal, and posteriorly to the anterior 
three-fourths of the parietals: first white interspace 
extending to second scale behind parietals; chin 
white except for black markings on mental, first 
three infralabials. and anterior portion of anterior 
chin shields. 

Our specimen was collected at 8:35 P.M. on 28 
July 1 967. about 3 m in a tree ( Acacia) in an area of 
second-growth thorn forest (Tropical Arid Forest 
formation of Holdridge. 1962). 

Sibon carri (Shreve) 

Sibon carri was previously known only from 4 
specimens from El Zamorano, Departamento de 
Francisco Morazan. Honduras (Peters, 1960), and 
one specimen from San Salvador, EI Salvador 
(Mertens, 1952b). 

On the night of 18 June 1968, we collected 2 
females of this species, one from 5 km WNW 
Choluteca. Departamento de Choluteca (LSUMZ 
23830), and the other from 3 km E Goascoran, 
Departamento de Valle (LSUMZ 23831); both 
of these localities are in an area of mixed scrub 
forest (Tropical Dry Forest formation of Holdridge, 
1962). Both specimens agree well with the descrip- 
tion of the species given by Peters (1960). Data for 
our specimens are (LSUMZ 23830 given first, 
LSUMZ 23831 second): ventrals 169. 170; sub- 
caudals 43, 46; dorsal scale rows 13-13-13 in both; 
anal plate single in both; supralabials 6-6, 3rd and 
4th entering the orbit in both; infralabials 7-6, 7-7; 
no preoculars; loreal single, elongate, entering the 
orbit in both; postoculars 1-1 in both; primary 
temporal absent. 5th supralabial in contact with 



parietal and one secondary temporal in both: total 
length 379 mm. 42 I mm; tail length 6 1 mm. 68 mm. 

In life the dorsum was rust red with a series of 
irregular dark brown blotches, approximately equal 
in length to the interspaces. The venter was cream 
with scattered brown blotching. The vermiculate 
markings on the internasals and anterior portion of 
the prefrontals were cream, and the remainder of 
the dorsal head markings were reddish-orange. 

Both of our specimens were found on a paved 
highway at night during rainstorms. Each individual 
contained two well-formed, elongate eggs, which 
ranged from 34.7 mm to 36.2 mm in length and 
from 6.8 mm to 7.3 mm in width. 



Thamnophis cyriopsis (Kennicott) 

This gartersnake has not previously been recorded 
south of Guatemala (Stuart, 1963). Field work 
carried on in Honduras by Texas A and M Univer- 
sity in 1967 and by us in 1968 has demonstrated the 
presence of Thamnophis cyriopsis in the south- 
western part of the country. The specimens were 
collected 21 km E Nueva Ocotepeque. Departa- 
mento de Ocotepeque (LSUMZ 23832-33), at La 
Esperanza, Departamento de Intibuca (TCWC 
23817-19, LSUMZ 23835), and 5 km NNE La 
Esperanza, Departamento de Intibuca (LSUMZ 
23834). All localities are within cloud forest or a 
mixed pine-oak-sweetgum forest (Lower Montane 
Moist Forest formation of Holdridge, 1962), at ele- 
vations of 1700 to 1900 m. Individuals were found 
in and around streams and ponds in cleared areas, 
and one specimen from the Ocotepeque locality 
contained the remains of a frog. Plectrohyla sp. 



ACKNOWLEDGMENTS 

We wish to express our thanks to the following persons 
for the loan of specimens: James E. Bohlke, Academy 
of Natural Sciences of Philadelphia (ANSP): James R. 
Dixon, Texas A and M University (TCWC); William E. 
Duellman. Museum of Natural History, University of 
Kansas (KU); Robert F. Inger, Field Museum of Nat- 
ural History (FMNH); Douglas A. Rossman, Museum 
of Zoology, Louisiana State University (LSUMZ); Jay 
M. Savage. University of Southern California (CRE); 
Dorothy M. Smith, Museum of Natural History, Uni- 
versity of Illinois (UIMNH); Charles F. Walker. Muse- 
um of Zoology. University of Michigan (UMMZ); 
Ernest E. Williams. Museum of Comparative Zoology, 
Harvard University (MCZ); John W. Wright, Los 
Angeles County Museum of Natural History (LACM); 
Richard G. Zweifel, American Museum of Natural 
History (AMNH). We are also grateful to Ernest A. 
Liner, Houma, Louisiana, for allowing us to examine 
his private collection (EAL). 



1971 



AMPHIBIANS AND Kl Pill IS 01 HONDURAS 



II I 



Field work in I 967 was made possible by funds from 
the Biomedical Sciences Support Grant, IK 07012-01. 
from the National Institutes of Health (senior author) 
and a Field Research Fellowship from Louisiana State 
University (junior author), and in 1968 by a Graduate 
School Travel Fellowship from ihe University of South 
ern California (senior author) and a Sigma Xi Grant- 
in-Aid of Research (senior author). 

LITERATURE CITED 

Bocourt. M. F. 1873(1872-82). Rechercheszoologiques. 
Etudes sur les reptiles. In Mission Scientifique au 
Mcxiciiie et dans I'Amerique Centrale. Livr 3: 

28-1000. 



Merti ns, K. 1952a. Neues fiber die Rcptilicnfaui 
I I Salvadoi Zoologist Itt I) i igt » I i ; - : - 9 



. 1952b. Die Amphibicn und Reptilien ■■ I 
Salvadoi aul Grund dct Reiser von U Mcrtens 
und A Zilch, Ahhandlungen dei Senckenbei 
i tun NaliirforsclienJen Gesellschafl. 487: 1-120. 

Peters.J. A. I960. The snakes of t he subfamil Dipsadi- 
nae. Misc. Publ. Mu.i. Zool., Univ. Michigan, 114: 

1-224. 

Peters. W. 1863. Eine mittheilung iiher neuc Arten dcr 
Saurier-Gattung Anolis. Mnnut\hrricht Akademic 
Wissenschaftlichen Berlin, 1862 (1863): 135-149. 



Boettger, O. 1898. (Catalog der Reptilien-Sammlung 

im Museum der Scnckenbergisehen Naturfor- 
schenden Gesellschaft. Tiel II (Schlangen). Frank- 
fort; Gebruder Knauer. 160 pp. 

Bogert, C. M. 1968. A new arboreal pit viper of the 
genus Bothrops from the Isthmus of Tehuantepec, 
Mexico. Amer. Mus. Novit., no. 2341: 1-14. 

Burger. W. L., and J, E. Werler. 1954. The subspecies 
of the ring-necked coffee snake, Ninia diademata, 
and a short biological and taxonomic account of 
the genus. Univ. Kansas Sci. Bull.. 36: 643-672. 

Cope. E. D. 1864. Contributions to the herpetology of 
tropical America. Proc. Acad. Nat. Sci. Philadel- 
phia, 1864: 166-181. 



— . 1885. A contribution to the herpetology of 
Mexico. Proc. Amer. Philos. Soc, 22: 379-397. 



Rand, A. S. 1957. Notes on amphibians and reptiles 
from El Salvador, Fieldiana: Zool., 34: 505-534. 

Roze, .1 A 1966. La taxonomia y zoogeographia de los 
oftdios de Venezuela. Scrvicio de Distribuciones 
de Pulicaciones, Biblioteca, Universidad Central 
de Venezuela, Caracas, 362 pp. 

Schmidt. K. P. 1933. New reptiles and amphibians from 
Honduras. Field Mu.s. Nat. Hist., Zool. Ser., 20: 
15-22. 

. 1936. New amphibians and reptiles from 



Dunn, E. R. 1930. Notes on Central American Anolis. 
Proc. New England Zool. Club.. 12: 15-24. 



Honduras in the Museum of Comparative Zool- 
ogy. Proc. Biol. Soc. Washington, 49: 43-50. 

Scott, N. J.. Jr. 1969. A zoogeographic analysis of the 
snakes of Costa Rica. Ph. D. Thesis. Univ. So. 
California. 390 pp. 

Smith. H. M.. F. W. Burley. and T. H. Fritts. 1968. A 
new anisolepid Anolis ( Rept ilia: Lacertilia) from 
Mexico./ Herp.,2: 147-151. 



Dunn. E. R., and J. R. Bailey. 1939. Snakes from the 
uplands of the Canal Zone and of Darien. Bull. 
Mus. Comp. Zool.. 86: 3-22. 

Dunn. E. R., and J. T. Emlen. Jr. 1932. Reptiles and 
amphibians from Honduras. Proc. Acad. Nat. Sci. 
Philadelphia. 84: 21-32. 

Holdridge. L. R. 1962. Mapa ecologico de Honduras. 
Organizacion de los Estados Americanos. San 
Jose. Costa Rica. 

Lynch. J. D. 1965. A review of the rugulosus group of 
Eleutherodactylus in northern Central America. 
Herpetologica. 21: 102-1 13. 

Lynch, J. D., and H. N. Smith. 1966. New or unusual 
amphibians and reptiles from Oaxaca. Mexico. II. 
Trans. Kansas Acad. Sci.. 69: 58-75. 

Lynn. W. G. 1959. Some reptiles and amphibians from 
Trinidad. Herpetologica. 15: 113-117. 



Smith. H. M.. and E. H. Taylor. 1950. An annotated 
checklist and key to the reptiles of Mexico, exclu- 
sive of the snakes. Bull. U. S. Nat. Mas.. 199: 1- 

253. 

Stuart. L. C. 1941. Studies of Neotropical Colubrinae. 
VIII. A revision of the genus Dryadophis Stuart. 
1939. Misc. Publ. Mus. Zool.. Univ. Michigan. 
49: 1-106. 



. 1942. Comments on several species of Anolis 

from Guatemala, with descriptions of three neu 
forms. Occas. Pap. Mus. Zool.. Univ. Michigan. 
464: 1-10. 



— . 1948. The amphibians and reptiles of Alta 
Verapaz, Guatemala. Misc. Publ. Mus. Zool.. 
Univ. Michigan. 69: 1-109. 



. 1955. A brief review of the Guatemalan lizards 

of the genus Anolis. Misc. Publ. Mus. Zool.. L'niv. 
Michigan. 91: 1-31. 



114 



III I I 1.1 IX SOI I II FRN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 70 



. 1963. A checklist of the herpetofauna of Guate- 
mala. Misc. I'ubl. Mus. Tool., Univ. Michigan, 
122: 1-150. 

Taylor. E. H. 1951. A brief review of the snakes of 
Costa Rica. Univ. Kansas Sci. Bull., 34: 3-187. 

— . 1954a. A review of the Lizards of Costa Rica. 



Univ. Kansas Sci. Bull., 38: 3-322. 



-. 1954b. Further studies on the serpents of 
Costa Rica. Univ. Kansas Sci. Bull., 36: 673-800. 

Wiegmann. A. F. 1834. Herpetologica Mexicana. Bero- 
lini, 1834: 1-54. 



Accepted for publication May I I, 1971. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCE 70(3): 1 14-124, 1971 

OBSERVATIONS ON THE DISTRIBUTION AND ECOLOGY OF THE 
LITTORAL ASCIDIANS OF THE MAINLAND COAST OF SOUTHERN CALIFORNIA 

Rimmon C. Fay and Jeffrey V. Johnson 1 



Abstract: A study of the littoral ascidians of the mainland of southern 
California was conducted during the summer of 1970. The collecting sites are 
recorded and natural history notes on the 41 species listed are presented. Trie 
number of species found in each of the four areas studied along the coast 
(Santa Barbara. Ft. Dume, Palos Verdes-Los Angeles Harbor, and San Diego) 
suggests that the contemporary distribution of tunicates may be controlled 
more by water quality than by the availability of a physically suitable habitat. 



The distribution, natural history, and specific com- 
position of the ascidian fauna of southern California 
was first reported by Ritter and Forsyth (1917) and 
later reviewed and expanded by Van Name (1945). 
Both accounts were based on observations and 
collections made in the littoral zone and by dredging 
in the sub-littoral zone. A preliminary survey of 
this group of stenohaline organisms in southern 
California during the summer of 1970 resulted in 
some additions to the notes on the natural history 
of these organisms and a contemporary definition 
of their distribution. 

Millar (1960) and others have advanced reasons 
for the abandonment of the generic name Amarou- 
cium Milne Edwards, 1842. as used by Ritter and 
Forsyth (1917) and Van Name (1945), in favor of 
the use of the generic name Aplydium Savigny, 
1816, and this synonomy is adopted in this report. 



METHODS 

All collections from the intertidal zone to a maxi- 
mum depth of 50 m were made south-east of Pt. 
Conception Oat. 34°, 27'N, long. 120°, 28' W) from 
areas of protected water and on the open coast. 
Collections were made by trawling with an 8 m 
otter trawl, by diving with the aid of SCUBA or 
during low tides. Specimens were brought into the 



laboratory and narcotized with menthol overnight. 
Seawater formalin (5%) was added gradually over a 
2-3 hour period. Specimens were stored in 5% sea- 
water formalin until examined. The identification is 
based on Ritter and Forsyth (1917), Van Name 
(1945), and Abbott (1957). 

RESULTS 

The coast was arbitrarily divided into four collecting 
areas (Fig. 1-5), ranging from Santa Barbara south- 
east to San Diego. The species collected, collecting 
sites, and major firm substrate within each area are 
as follows: 

Santa Barbara area 

Figure 2 

1. Naples Reef, 13-15 m, shale reef and rubble: 21 
July 1970. 

2. Ellwood pier. 1-1 1 m. concrete, wooden, and steel 
pier pilings: 21 July 1970. 

3. Coal Oil Pt., 3-7 m, shale reefs; 21 July 1970. 

4. Stern's wharf. 1-7 m. wooden pier piling; 30 July 
1970, 18 August 1970. 

5. Santa Barbara Harbor, 0-3 m, floats and wooden 
piling; 18 August 1970. 



'Pacific Bio-Marine Supply Company, P.O. Box 536, 
Venice, California 90291. 



1971 



ASCIDIANS <>l SOUTHERN <M II oKNIA 



in 



Order Enterogona 



A /il itl in in californicum 
A. solidum 
A. arenatum 

i.iilir rdmania claviformls 
Polyclinum planum 
Synoicum par-fustis 
Didemnum carnulentum 
Trididemnum opacum 
Unidenl ilicd didemnid 
Diplosoma macdonaldi 
Clavelina huntsman! 
( ystodytes lobalus 
Distaplia occidental is 
Eudistoma psammion 
E. ritteri 
E. diaphanes 

Sigillinaria aequali-siphonis 
S. pulchra 
Ascidia ceratodes 
Chelyosoma productum 
Ciona intestinalis 
Perophora annectens 



Order Pleurogona 



Botryllus sp. 
Botrylloides sp. 
Styela montereyensis 
S. truncate! 



1.3 
1,3 


2.3,4 

3 

3 

1,2,3,4 

1,3 


3 

4 

2 

1.4 

1,3 

3 

3 

1 


1 
1,2,3,5 

4 
5 
2,3 


3,5 
4,5 

1.2.3,4,5 
2,3.5 



.V. sp. [burnlturti I 
S. gibbsii 

S. plu ilia 

I'' inn haustor 
Boltenia villosa 
Molgula vi an ifera 

Numbei "i species Found: ( .' 



5 

3,4,5 

5 

• 
2,3 
2.4 



I'l I )l Ml M(F A 

I igurc 3 

1. Paradise Cove. h;iil receiver. 1-2 m. nylon mesh 
fish nel: Id June 1970. 

2. Little Dume I't., 3-5 m. shale reef; 16 June 1970. 

3. I.eo Carrillo, 5 m, granitic rocks: 19 June 1970. 

4. Paradise Cove. No. X reef. 10 m. shale reef: 26 June 
1970. 

5. Malibu Pt„ 3 m, basaltic rocks: 2X June 1970. 

6. Pt. Dume rocks. 3-10 m, basaltic rocks: 2 July 1970. 

7. Little Dume Cove. 3 m, shale reef: 2 July 1970. 

8. Latigo Cove. 10 m. basaltic rocks; 26 August 1970. 

9. West end of Zuma Beach. 6 m. granitic rocks: I 
September 1970. 



Order Enterogona 



A pi i din in californicum 
A. solidum 



2.3.4.5 
3.7 



SAW/) BAffBflGA AREA 



PT. DUME AREA 

I co Carritlo Beach ~~~^ \ Venice 

' Marina del Reij 




PALPS VERDES-LOS AKJGELES AREA 
Uzuipor-t San 



MAIkJLAUD COAST 
OF SOUTH ERU CALIFORNIA 



ssion Bay 

said DIEGO 
AREA 



Fl GUISE I 



1 lf> lit ill 1 1\ Si >( 


llll UNCAL 


A. areiuittmi 


2.6 


A. propinquum 


2 


I ulierdinania clavifonnis 


2.5 


Polyclinum sp. 


8 


Synoicum par-ftistis 


9 


Dideinnum carnulentiim 


5 


Trididemntim opacum 


3,5,7 


Unidentified didemnid 


4 


/ issoi linttm caidleryi 


6 


Clavelina huntsmani 


5 


( ) ttodytes lobatus 


7 


Disiiipliu occidental is 


4 


Eudistoma psammion 


2.4.5.7 


E. ritlcri 


2.3 


Pycnoclavella Stanley! 


5.7 


Sigillinaria aeqnali-siphonis 


2 


Ascidia ceratodes 


1.4.5 


Chelyosoma production 


1.5.7 


Perophora annectens 


5 


Diplosoma macdonaldi 


8 



ACADEMY OF SCIENCES 



VOLUME 70 



Order Pleurogona 



Metandrocarpa taylori 

M. dura 

Styela montereyensis 

S. truncata 

Pyura haustor 

Number of species found: 27 



4 

2,7 

1.2.5 

1.3.4.5.7 

3 



Rl. HONDO. I'M OS \ 1 Kill s. 

Los ANGELES Harbor ari -a 
Figure 4 

I Palos Verdes, Long Pi. Marineland reef. 12 m, San 
Onofre Breccia; IX .lime 1970. 

2. Marineland pier. 3-5 m. wooden pilings; 15 June 
1970. 

3. King Harbor (Redondo Beach) boat slips. NE of 
harbor office, floats; 29 June 1970. 10 August 1970. 

I Last end of King Harbor breakwater. 7 m. breccia 
rocks; 29 June 1970. 

5. Rocky Pt., Palos Verdes, 7 m. San Onofre Breccia 
rocks; 29 June 1970. 

6. Alamitos Bay. 1-3 m, floats and wooden pilings: 3 
July 1970. 12 July 1970. 30 August 1970. 

7. Los Angeles Harbor breakwater, 4-10 m. granitic 
and breccia rocks and debris; 3 July 1970. 12 July 
1970. 30 August 1970. 

S. Watchorn Basin. San Pedro. 0-2 m, floats and wood- 
en piling; 15 August 1970. 
9. Pt. Vicente. 7 m, breccia rocks; 15 August 1970. 

10. Redondo, 45-50 m, sandy mud; 1 1 September 1970. 

ll.Abalone Cove, 6 m. breccia rock; 21 September 
1970, 26 September 1970. 

12. Resort Point, 5-10 m. breccia rock; 28 September 
1970. 

13.Abalone Cove. 3 m, breccia rock; 28 September 
1970. 



Maples 



K.Jreef 




_ Elluood 
pier 



Coal Oil Pt. 



Go/era Pt. 




HEY MAP 

UO SCALE 



^Stearns 
Santa Barbara /_* ,~, Wharf 

l-iarbor ■ 



Pt. Castillo 



Santa Barbara Pt. 



Pacific 



SAKJTA BARBARA AREA 



Ocean 



FIGURE 2 



1971 



ASCIDIANS OF SOU I III. UN ( :il II oKNIA 



117 



Order Enterogona 



A pi i ilium culifomicuin 
A. stil id ii in 

A. pilljlilHjIIIIIII 

Eiiherdnumki claviforinis 
Synoictim par-fiistis 
Didemnum curnulentum 
Clavelina lutntsinani 
Distaplia occidental is 
Cysiodytcs lobatiis 
Eudistoma psammion 
Ascidia ceratodes 
( 'iona inlestinalis 
Diplosotna macdonaldi 



Order Pleurogona 



Metandrocarpa taylori 
Botrylloides sp. 
Bolryllus sp. 
Unidentified styelid 
Styela truncata 
S. sp. (ho in ha rii) 
S. gibbsii 
S. plicata 

Halocynthia igaboja 
Pyura haustor 
En gym arenosa 

Number of species found: 24 



1.4.6,7 

2 

2.13 

13 

12 

3 

9. 12 

X 

2,5,7,1 1,12 

13 

3.7.1 1,12 

3,6,10 

3 



I I 

3.6.7 

3,6 

6 

2.7,11,12 

3,6 

7, S.I 1 

3.6,8 

I 1 

1.5.7.12 

10 



Sa-. Dll 00 AIM \ 

I i U| 

I Dana mai ina, Mission Ba> I m, float! and 
en pilings; 5 June 1970. 

2. Pi. I. «ima. 27 m, shale rod transect opposite inlal c 
line ini naval station 'I ig ill June 1970 

3. 23 in. shale nick: I I June 1970. 

4. 20 m, shale rock; 1 1 June 1970. 

5. 14 m. shale rock; I I .Inn,- 1970. 

6. 7 in. shale rock; I I .lime 1970. 

7. Entrance channel cast jelly inside Mission Hay. 
2-3 m. breccia rock: 1 1 June IV o 

X. Ventura Bridge, i m, wooden pilings; 14 June 1970. 
9. Ventura Bridge. 0-5 m. rock rip rap: IX .lime 1970. 
12 September 1970. 

10. New Hope Rock I ranscct line Pt. I.oma. 25-30 m. 
shale rock; 21 October 1970. 

11. 15 m, shale rock; 2 1 Octobei 1970. 

12. 10 m. shale rock; 21 October 1970 
13.3 m, shale rock; 2 1 October 1970. 

14. Bird Rock. La .lolla. intertidal. cobble bed; 2X 
November 1970. 



Order Enterogona 



A pi i din in californicum 
A. soli dn in 

Euherdmania claviforinis 
Polyclinum planum 



1.3.7 

2 
7 
2 



Zu 



m a 



3 



a ° c h 



Pacific 
PT. DUME AQEA 




KEY MAP 

UO SCALE. 



Latiqo Pt. 



Malibu Pt. 



'Paradise 
Core. 

''little Dume Pt. 



Pt. Dume 



Ocean 



FIGURE 3 



118 



111 I I I I IX SOI I III R.X CAl.ll ' ORNIA ACADEMY O I- SCIENCES 



VOLUME 70 



Kino 
Harbor 




Marina/end 
Pt. 
l/izznt* -Cfi babni 

Cat/a 




KEV MAP 
ua scan 



Alamitos Bay 



San Pe dro 3 a y 



r al brcakuotar 



Pt. Farmin 



Port of Los Anaeles - Lona Beach 
Pacific <S> c e a /z 

PALOS VER.DE5- LOS AUGELES AREA 




FIGURE 4 



Xj 
& 



Bird Rock. 



<3> 



»• Mew Hope 
■a Rock — 

Pt. Loma 



SAU DIEGO AREA 





key map 

UO SCDLS 



FIGURE 5 



1971 



ASC1DIANS <)1 SOU I III KN ( Al // oKNIA 



119 



Synoicuin par-fustis 
Didemnum carnulentiitn 
Clavelina Ituntsmani 
Trididemnum opacum 
Diplosoina macdonaldi 
Lissoclinutn catdlevyi 
Distaplia occidental is 
Eudistoma psammion 
E, diaplianes 
E. rltteri 

( ystodytes lobatus 
Ascidia ceratodes 
Chelyosoma productuin 
Ciona intestinalis 
Perophora annectens 



Order Pleurogona 



2,10 
1.4,7 

I I 
2 



3.4.7 
4 

4.5 
I I 

2,4.7,9 
3,10,1 I 
I 
3 



13,14 
1,6 

1,7 

1,5,7,10.1 I 

8 

I 

7,9.10 

1 

2,7,8,9 

9.10,1 I 

7 



Metandrocarpa taylori 

Botryllus sp. 

Botrylloides sp. 

Sly da montereyensis 

S. truncata 

S. sp. (barnharti) 

S. gibbsii 

S. plicata 

Pyura haustor 

Boltenia villosa 

Molgula verrucifera 

Number of species found: 30 



ANNOTATED SPECIES LIST 

Aplidiwn californicum (Ritter and Forsyth). 
Specimens were collected from each of the four 
areas, often in abundance. Colonies were taken in 
the bathymetric range of 2-27 m and were found 
growing over rocks, worm tubes, algae, and other 
substrates. Zooids reached a size of 5-8 mm in 
length and were often organized into definite sys- 
tems. The colonies ranged in color from opalescent 
white to yellow-orange. The breeding season is at 
least throughout the summer months as larvae 
and/or embryos (4-6) were found in some individ- 
uals in each collection. 

Aplidiwn solidum (Ritter and Forsyth). Speci- 
mens were collected from the coastal areas surveyed 
and were found growing over worm tubes, kelp 
holdfasts, and rocks. The zooids measured to 10 
mm in length; they ranged in color from orange to 
nearly white. Some animals in each colony had 
larvae and/or embryos in their atrial cavities: the 
embryos in the colony collected 15 June appeared 
to be more advanced than any of the other speci- 
mens of this species collected during the summer 
of 1970. 

Aplidiiim propinquum (Van Name). Colonies, 
made up of several sand covered lobes, were taken 



al I ittlc Diiiik |'i .. al 5 m growing over worn lubes 

and al Palos Verdes, Mai inciand Pii i m 

and pier piling rhi zooids measured 17-27 mm in 

length (the posl abdoi anged from 10 !0 mm) 

and were a ange-red coloi l*hts represents an 

extension ol the range ol ibis species to southern 
( alifornia (Abbott, 1957). 

Aplidiiim arenatum (Van Name). < olonieswerc 
collected from Santa Barbara and Pi Dume down 
to a depth <>l 5 m: they were found growing over 
sponge, algae, and other surfaces, Zooids reached an 
overall length of 14 mm (the thorax and abdomen 
measured about 5 mm) and were white. Animals 
collected in June and July were found with larvae in 
their atrial cavities. 

Euherdmania claviformis Ritter. Specimens were 
collected at each of the areas surveyed, from the 
intertidal down to a depth of 12m; only one colony 
was found in the Palos Verdes region. These social 
ascidians, with zooids measuring 15-20 mm. were 
taken growing with hydroids and bryozoans. The 
animals from Fllwood pier were covered only very 
lightly with sand. Trason (1957) gives the breed- 
ing season as July to September, with the peak in 
August. Animals collected on 16 June had a charac- 
teristic arrangement of embryos in a longitudinal 
series in the oviduct. 

Polyclinum planum (Ritterand Forsyth (.Colonies 
were taken from Pt. Loma. Latigo Cove, and Coal 
Oil Pt. The bathymetric distribution ranged from 
5-30 m. The colonies are reddish in color and lightly 
covered with sand; they often become quite large, 
measuring up to 20 cm in width. The zooids have a 
length of 7 mm. The colonies narcotized well \\ ith 
menthol. 

Polyclinum sp. An unidentified species of this 
genus was collected at a depth range of 5-10 m at 
Latigo Cove in the summer of 1970 and 1971. The 
colony is of 4-15 cm in height and of a more glob- 
ular form than P. planum. It is grayish white in 
color. 

Synoicum par-fastis (Ritter and Forsyth). Speci- 
mens were collected at Pt. Loma (27 m), Coal Oil 
Pt. (5 m). and at Zuma Beach (6 m). Zooids aver- 
aged 10 mm in length: the colonies were 60-1 15 
mm in height. A colony collected at Zuma Beach 
was nearly twice as long as reported for the maxi- 
mum size of colonies of this species (Van Name. 
1945). Colonies taken from Coal Oil Pt. and Zuma 
Beach were orange-red: some from Pt. Loma were 
white. This species narcotized well with menthol. 
The breeding season is at least June and July as 
embryos were found in the atrial cavities of individ- 
uals at this time of year. Generally, zooids had 1 or 
2 well-developed tadpoles and several embryos in 
the atrial cavity. 



120 



HI III 1 1\ SOI I III l<\ (II NORMA ACADEMY OE SCIENCES 



I O1.UME70 



Didemnum carnulentum Ritter and Forsyth, 
specimens were collected in each of the areas sur- 
veyed, down to .1 depth of 20 in. Colonies, encrust- 
ing and less than 4 mm thick, were found growing 
on various algae, with a colony of Euherdmania, 
and on rocks. I he yellow-orange zooids were less 
than 2 mm in length. Colonies were also taken in 
abundance growing on the carapace of decorator 
crabs, Loxorhynchus crispatus, at Santa Barbara. 
No indication of breeding activity was seen although 
Ritter and Forsyth (1917) mentioned that the zooids 
contained tadpoles in June. 

Trididemnum opacum (Ritter). Specimens were 
taken from all areas but the Palos Vcrdes-L.A. 
Harbor area. The bathymetric range is from surface 
to 27 m. Colonies of this gray ascidian were found 
growing on kelp holdfasts, hydroids, and worm 
tubes; they ranged in thickness from 1-4 mm. 

Diplosoma macdonaldi Herdman. This species 
was found from Santa Barbara to San Diego, at all 
areas surveyed, on floats, pier pilings, mussel shells, 
and kelp down to 10 m. The zooids are brown and 
measure about 1 mm in length. A colony taken from 
King Harbor (10 August) had numerous zooids in 
various stages of budding. Animals collected on 21 
July had larvae free in the common test. This species 
is characterized by a very fragile, gelatinous test. 

Lissoclinum caulleryi (Ritter and Forsyth). Colo- 
nies were taken from San Diego and Pt. Dume 
growing on kelp, hydroids, and rocks. The zooids 
measured to 1.5 mm in length. A colony from San 
Diego on 1 1 June had many larvae and large eggs 
free in the test; those specimens collected in July 
appeared to be sexually mature. 

Clavelina huntsman! Van Name. This species was 
taken in each of the areas surveyed. It is character- 
ized by a clear test and a bright pink pharynx. It 
grows on rocks in areas of vigorous water move- 
ment. A colony from Santa Barbara was surrounded 
by a sponge. The zooids grow to over 30 mm in 
length. Van Name ( 1945) found this species to occur 
on an annual basis and to be in reproductive condi- 
tion at Pacific Grove from June to September. 

Cystodytes lobatus (Ritter). This species was 
collected in each of the areas surveyed. Colonies 
of up to 25 cm in width were found growing on 
worm tubes, kelp holdfasts, shells, and rocks; indi- 
vidual zooids were 2.5 mm in length. Individual 
colonies ranged in color from off-white through 
orange-tan to pink. Animals collected in June and 
July had larvae and eggs in their atrial cavities. 
This is the dominant species found in the collec- 
tions from the Los Angeles-Long Beach Federal 
breakwater. 

Distaplia occidentalis Bancroft. Colonies were 
found in all of the areas surveyed down to 15 m on 



mussel shells, rocks, piling, and other surfaces. 
Zooids were about 2 mm in length. Colonies were 
found with larvae in them throughout the summer. 
Ritter and Forsyth (1917) pointed out that the 
color of the colony becomes darker with age. The 
colors of observed colonies of this species ranged 
from cream-while to dark red-brown. 

Eudistoma diaphanes Ritter and Forsyth. Colo- 
nies of this opalescent white ascidian were taken 
from San Diego and Coal Oil Pt. down to a depth 
of 20 m. They were found growing on algae and 
over clam shells. Colonies reached a width of 25 
mm. zooids a length of 3 mm. Colonies collected on 
2 1 July had larvae and eggs present in them. 

Eudistoma psammion Ritter and Forsyth. This 
species was collected in all of the areas surveyed. 
The bathymetric range extended from the intertidal 
to a depth of 27 m. The sand-encrusted colonies 
are of a deep burgundy color and were found 
growing on rocks. Chaetopterus tubes, and kelp 
holdfasts. Colonies of Euherdmania were found 
growing with colonies of Eudistoma psammion 
taken from Malibu Pt. while these two joined with 
Aplidium propinquum at Abalone Cove. Zooids 
measured to 5 mm in length. Van Name (1945) 
gives the breeding season as June-July when the 
atrial cavities are often distended by large eggs and 
larvae. The colonies from Coal Oil Pt. had very 
little sand on their surfaces. 

Eudistoma ritteri Van Name. Colonies were 
collected from Santa Barbara and San Diego be- 
tween the depths of 3-20 m. Colonies were found 
with some sand over their surfaces, contrary to 
Van Name's description, and are dull gray in colora- 
tion. Zooids measured up to 3 mm in length. Van 
Name (1945) gave the breeding season as beginning 
in February-April, peaking in July, and declining 
in August. 

Pycnoclavella stanleyi Berrill and Abbott. Speci- 
mens were found commonly at Little Dume Pt. 
on rocks and on Cystoseira, and in abundance at 
Malibu Pt. where they formed golden-orange mats 
up to 50 cm in width. This species was taken in 
depths ranging from 2-7 m. 

Sigillinaria aequali-siphonis (Ritter and Forsyth). 
Colonies were found at Pt. Dume and Santa Barbara 
in 3-15 m of water. Light colored colonies were 
growing on algae and worm tubes. Zooids measured 
up to 6 mm in length. Eggs and larvae were present 
in the atrial cavities of animals in June and July. 
This species was reported from Pt. Loma by Turner, 
Ebert, and Given (1968). 

Sigillinaria pulchra (Ritter). Two light colored 
colonies were taken in 15 m of water at Naples Reef, 
west of Santa Barbara; they were found growing 
on worm tubes. The zooids measured up to 15 mm 



1 97 J 



ASCIIUANS Ol SOU I III, KN ( ,11 II < iKNIA 



121 



in length and there were lour larvae in the atrial 
eavities of some individuals. This collection site 
serves as a range extension lor the species as it is 
farther south than reported by Van Name (1945). 

Ascidia ceratodes (Huntsman). Specimens were 
taken from every area studied in 0-27 m of water. 
The translucent light green or yellow animajsjmeas- 
ured 20-30 mm in length and were found growing 
on their sides on kelp holdfasts, pebbles, boat floats, 
pilings, and rocks. The external appearance is highly 
variable as it is controlled by the confinement ol 
the surroundings. Only those specimens growing in 
the absence of confinement exhibit the form figured 
in Van Name (1945). Several of the specimens had 
a copepod crustacean living in their branchial 
basket; many of the copepods were carrying eggs. 
The gonad tissue of A. ceratodes appeared to be 
mature during the summer of 1970. 

Chelyosoma productum Stimpson. Specimens 
were taken from Santa Barbara to San Diego but 
they were not found in the Palos Verdes-L.A. 
Harbor area. The animals were collected in 2-27 m 
of water. The translucent individuals measured up 
to 15 mm across; specimens from Santa Barbara 
were noticeably larger than those from further 
south. The animals were found attached to abalone 
shells, kelp holdfasts, rocks, fish net, and corrugated 
aluminum sheeting. 

Ciona intestinalis (Linnaeus). This species was 
only taken at Santa Barbara, in the L.A. Harbor 
area (Alamitos Bay), and in Mission Bay; however, 
it may be expected to be present during the summer 
and fall in all of the harbors along the coast. This 
light green, translucent species is generally found 
growing on concrete pilings, wooden pilings, algae, 
floats, rocks, mussels, and other solitary ascidians. 
Many animals will fall from floats or pilings and 
live out their lives lying on the bottom. Others will 
attach to various objects and grow along the open 
coast at depths below the zone of active wave surge 
down to a depth of at least 50 m. Specimens col- 
lected from under floats in Newport Bay measured 
up to 25 cm in length. 

Perophora annectens (Ritter). This green or 
yellow-green species was collected in abundance 
from the Santa Barbara area; colonies were found 
growing on concrete and wooden pilings, algae, and 
rocks. A colony was also taken at Malibu Pt. Two 
specimens were taken in 27 m of water at Pt. Loma. 
The zooids measured up to 1.5 mm in length. 

Unidentified styelid species. The individuals of 
this ascidian are joined together by stolons forming 
mats up to 4-6 cm in width which resemble colonies 
of Perophora. The animals measure 4-5 mm in 
length and are yellow in color. The colonies may 
have a lisht sand covering if taken near the surf 



zone, a raphe is presenl along the intestine and the 
Stomach has I I 12 ridgCS. The ovaries are in .i row 
of 5 units along the right side ol the bod) I here 

were numerous eggs and lat ae in the peribranchial 
area, Specimens were collected al a depth ol i m 

from the exposed side ol the Venice brcakwatCI "ii 

22 August 1970. and from piei pilings in the pi 
tected waters ol Alamitos Has al depths to 4 m on 
30 August 1970. I his Styelid closely resembles the 
description ol Alloeocarpa bacca, which has been 
reported from the coast ol < hile (Van Name. 1945). 

Botryllus spp. I hese animals were collected in 
protected waters at Santa Barbara, Alamitos Bay. 
and Mission Bay from the surface to a depth "i I 
m. This genus is also common in the Manna Del 
Rey in Venice. California. Specimens were lound 
on any available firm substrate and narcotized well 
with menthol. Botryllus luberatus and possibly 
other species were present. I he colonies ranged in 
color from black to orange. 

Botrylloides spp. Representatives of the genus 
were collected where protected water was found in 
each of the areas surveyed; they were lound growing 
on mussels, hydroids. and other firm substrates from 
the surface to a depth of 4 m. Botrylloides diegense 
and possibly other species were present. 

Matandrocarpa dura (Ritter). Specimens were 
collected only from the Pt. Dume area and were 
found growing over bryozoans, Cystoseira, and 
Styela montereyensis, down to 7 m. The zooids 
reached 4 mm in length: the colonies were a vivid 
red-brown color. 

Metandrocarpa laylori Huntsman. This orange- 
red species was collected at Paradise Cove (Pt. 
Dume area), Palos Verdes from 0-1 1 m. Pt. Loma 
in 3 m of water, and from the intertidal at Bird 
Rock, La Jolla. This tunicate was found growing 
on wood, shells, rock, and Chaetopterus tubes. 
Zooids were 3 mm in length. Two tadpoles and 
some advanced eggs were present in June. Abbott 
(1953) gave the breeding season as peaking in July 
and August. This record represents a southern 
range extention for this species on the mainland. It 
has also been reported from Catalina Island (Given 
and Lees, 1967). 

Styela montereyensis (Dall). Specimens of this 
pinkish-red ascidian were not taken in the Palos 
Verdes-L.A. Harbor area but were found routinely 
elsewhere from the intertidal down to 17 m. The 
individuals were up to 25 cm in length and were 
growing on kelp. Pyura hauslor. pilings, rocks. 
floats, shells, or any available hard substrate. Styela 
montereyensis is found with 5. sp. (barnhani) in 
Santa Barbara Harbor and is the main channel at 
Mission Bay: thus 5. montereyensis may grow in 
protected water. Many animals appeared sexually 



122 



lit I I I I IX SOI I ill RX CM IIORXIA ACADEMY OI- SCIENCES 



VOLUME70 



mature during the summer mouths. A specimen was 
taken at Pt. loma with a gravid pea crab in the 
branchial basket. 

Styela plicata (Lesueur). This species was col- 
lected in protected waters from Santa Barbara to 
San Diego but isolated individuals may also be 
found in unprotected locations. The animals were 
found growing on pilings, boat tloats, rocks, and 
hard objects on the bottom to a maximum depth of 
S m. They often reached a length of 10 cm. The 
species is opalescent white with brown striped 
siphons. It narcotized well with menthol. Styela 
plicata was one of three species of tunicates col- 
lected at Watehorn Basin in L.A. Harbor. It is 
recognized as having been introduced to southern 
California about 1945 (Abbott, pers. comm.). and is 
now locally abundant in protected waters through- 
out southern California. 

Styela sp. (bamharti). Thanks to the insistence of 
J. Richard Whittaker (pers. comm.) that a problem 
exists in the accepted designation of this animal as 
5. sp. {bamharti), a close examination of the litera- 
ture revealed that this species was not accurately 
described by Ritter and Forsyth (1917) nor Van 
Name (1945) (Abbott, pers. comm.). A description 
of this distinct species is now in preparation. 

Individuals of this species were collected on 
floats, pilings, and rocks in Mission Bay and Santa 
Barbara Harbor from the surface to a depth of 7 
m. This species is predominantly brownish in color. 
It may be found in protected waters anywhere in 
southern California where suitable water quality 
exists. Individuals grow to a length of about 20 cm. 
Sexually mature animals were seen at most times 
throughout the summer but spawning could not be 
induced in the laboratory. 

Styela truncata Ritter. This species was taken 
from protected and exposed locations in all areas 
surveyed from the surface down to a depth of 12 m. 
Individuals were found growing on floats, worm 
tubes, shells, and sponges, as well as on other 
tunicates. The specimens were about 20 mm in 
length. 

Styela gibbsii (Stimpson). Specimens were taken 
from semi-protected to protected sites at all four 
areas surveyed, from the surface to a depth of 30 m. 
They may be found attached to any firm substrate. 
Individuals reached 40 mm in length, and ranged 
in color from old rose with brown siphons to orange- 
red with red siphons. An animal from Pt. Loma 
carried the same type of copepod in its branchial 
basket as those found in Ascidia ceratodes. 

Pyura haustor (Stimpson). This species was col- 
lected in all areas surveyed from the intertidal 
(Mission Bay) to 27 m (Pt. Loma). Animals ranged 
up to 12 cm in length and were generally lodged in 



crevices, under rock ledges, in rip rap. or on pier 
pilings. This animal is always covered with a com- 
plex biota of fouling organisms and the only indica- 
tion of the presence of this tunicate is its bright 
pink-red siphons. The animals from San Diego 
carried an amphipod in their branchial basket. 
This species probably breeds in the summer as 
indicated by the appearance of the gonads. 

Halocynthia igaboja Oka. One specimen was 
collected at a depth of 6 m from under a rock at 
Abalone Cove, Palos Verdes. Van Name (1945) 
comments that this ascidian is fairly common about 
Catalina Island at a depth of 20-80 m. This finding 
is the first reported from the mainland of southern 
California, and it represents a new shallow depth 
record for the species. 

Boltenia villosa (Stimpson), Specimens were taken 
in the Santa Barbara region, at Ellwood pier, and 
Coal Oil Pt. in 2-12 m of water, in Mission Bay at 
a depth of 4 m, and at Pt. Loma in 25 m of water. 
Individuals ranged up to 9 cm in length; Van Name 
( 1945) described the size range as 3-6 cm. The tunic 
has a yellow-orange stalk and is bright brick red 
over the main body. 

Molgula verrucifera Ritter and Forsyth. Individ- 
uals were collected from Santa Barbara, Venice, 
and San Diego. All specimens carried larvae in the 
peribranchial area. Specimens reached a maximum 
length of 7 mm. The cream color of the test is 
obscured by a gray-green layer of encrusting sand 
which makes the animals notably inconspicuous. 
Four of the specimens collected at Santa Barbara 
were taken from the carapace of a decorator crab, 
Loxorhynchus crispatus. 

Eugyraarenosa (Alder and Hancock). Specimens 
were taken in abundance with the aid of an 8 m wide 
otter trawl off Redondo Beach in the depth range 
of 44-50 m. Most individuals were 10 mm in length; 
the largest were up to 20 mm. This record extends 
the bathymetric range and maximum size recorded 
for this species on this coast. The gonad was ripe; 
eggs and sperm were present in the gonaducts in 
early September. The atrial cavity of one specimen 
harbored a white flatworm 6 mm in length. 

DISCUSSION 

The four delimited areas surveyed along the main- 
land coast of southern California yielded a total of 
40 recognized species of ascidians. This compares 
favorably with the 29 species noted as occurring in 
the littoral zone in this region by Ritter and Forsyth 
(1917). This listing does not include Pyura mirabilis 
Von Drasche, which has been collected by the 
senior author from the mainland of southern Cali- 
fornia, nor does it include one unidentified didem- 



1971 



ASCIDIANS Ol SOdllll. KN ( Al lioKNIA 



123 



nid. Inclusion of these iwo species would bring the 
total to 43 species of ascidians from the littoral zone 
of the mainland of southern California. Undoubt- 
edly further survey work in the field and additional 
descriptive work in the laboratory will add more 
ascidians to the list. 

II is interesting to note thai of the 39 species of 
ascidians listed by Abbott ( 1957) as occurring in the 
intertidal /.one or located so that they are available- 
near the surface (on floats) in central California, 16 
species were found in the same situation in southern 
California. These include: an undescribed styelid, 
Aplidium californicum, Ascidia ceratodes, Botryllus 
sp., Botrylloides sp., Metandrocarpa taylori, Ciona 
intestinalis, Distaplia occidentalis, Didemnum car- 
nulentum, Euherdmania claviformis, Eudistoma 
psammion, Pyura haustor, Styela sp., S. monte- 
reyensis, S. plicata, and S. truncata. Further, of the 
39 species listed by Abbott (1957) or described by 
Abbott and Trason ( 1968) only 5 species have not 
been noted in southern California (Cystodytes sp., 
Distaplia smithi, Eudistoma molle, Rittcrclla rubra, 
and Cnemidocarpa finmarkiensis); thus, the major- 
ity of the littoral ascidians reported from the inter- 
tidal zone of central California are found southeast 
of Point Conception in the subtidal zone. The 
primary controlling factor limiting the appearance 
of these animals in the intertidal zone is probably 
temperature, since both water and air temperatures 
southeast of Point Conception along the shore- 
line average higher than those northwest of this 
break point in the distribution of intertidal marine 
organisms. 

The present recorded difference between central 
and southern California with respect to the ascidian 
fauna amounts to about 10 species and one may 
question the quantitative significance of the state- 
ment of Van Name (1945) that Point Conception 
marks a "break point'" in the distribution of these 
organisms. At this time and subject to further study, 
it would appear that a continuum of change in 
distribution of ascidians exists along the coastline 
of southern California which continues into central 
California. 

Note has been made of the restriction of some 
species of tunicates to protected waters while others 
may be limited to the waters of the open coast. A 
comparison of the number of species found in either 
of these type habitats along the coast was made. 
A total of 10 species of tunicates was found in 
Santa Barbara Harbor and 16 species were found in 
Mission Bay, whereas 8 species were found in King 
Harbor, 8 species in Los Angeles Harbor, and 6 
species in Alamitos Bay. In the adjoining areas of 
open coast. 27 species of tunicates were found in 
Santa Barbara. 26 species at Pt. Dume-Malibu. 



and ' I species in San I )iego whereas only 10 species 
were found .u Palos Verdes. Since the conservative 
oceanographic parameters ol tempcraiun and alu 
ity, and the ranges i<>i these parameters are essen- 
tially the same lor all lour collecting areas surveyed 
(Barkely, 1968), and since the physical habitats. ip 

similar in .ill foui areas, these factors cannot account 
for the difference in numbers of species found in 
either the protected areas or on the open coast. 

Water quality conditions along the coast I 

the greater Los Angeles area have altered radically 
over the past 25 years. The reduction in abundance 
Of certain species of benthic marine algae has been 
observed and the extirpation ol Mat rot tstisp rifera 
from the Palos Verdes Peninsula has been mi 
significant (Young, 1964; Strachan and Koski. 
1969; Gregg and Kiwala. 1970). The decline ol 
Macrocystis and of other algae along the Pains 
Verdes Peninsula occurred during a period when 
the waste input into the waters bordering the penin- 
sula increased rapidly; it is now on the order of 1 .5 
billion liters perday. About 2 billion liters of heated 
sea water discharged into the King Harbor Marina 
per day from the Redondo Beach power plant of 
the Southern California Edison Company. Los 
Angeles Harbor receives 29.5 million liters of 
sewage per day (Crippen and Reish. 1969) in addi- 
tion to industrial wastes (largely brines) and some 
thermal wastes (Reish. 1959). 

Since qualitatively and quantitatively equivalent 
waste loadings are not introduced into the water 
masses about the region of Santa Barbara and San 
Diego, it may be that the comparatively reduced 
number of species of tunicates found in the Palos 
Verdes area and in the bays in the area of greater 
Los Angeles demonstrates the presence of biologi- 
cally unfavorable water quality in this area. It 
appears that the changes in water quality along the 
southern shoreline of Los Angeles County which 
have occurred and which currently exist are detri- 
mental and limiting to sensitive species of both algae 
and benthic invertebrates. 



ACKNOWLEDGMENTS 

The authors are grateful for the interest and criticism 
offered in the course of this work by Professor Carl 
Gugler. University of Nebraska, and for the invaluable 
aid of Professor Donald Abbott. Hopkins Marine 
Station, in the identification of specimens of tunicates. 
his interest in the study, and his review and criticism of 
the manuscript. Thanks are due to D. C. Barilotti. 
Richard Alexander. George Kelly, and Pat Broph\ for 
aid in the field and laboratory. Thanks are certainly due 
to Flare Drafting Service for the figures and to Mrs. 
.1. M. Fay for typing the manuscript. 



124 



HI I I I IIS sol llll I<\ ( II II OKXI.t ,l( ,!/>/ \/> Ol SCIENCES 



VOLUME 70 



LITERATURE CITED 

Abbott, D. P. 1953, Asexual reproduction in the colo 
nial ascidian Meiandrocarpa laylori, Huntsman. 
Univ. California Pnbl. Zool., 61: 1-78. 

1957. key to the littoral ascidians of the central 



California coast. Pp. 301-309. In Intertidal Inverte- 
brates of the central California coast: S. F. Light's 
laboratory and field text in Invertebrate Zoology. 
2nd edition. (R. I. Smith. F. A. Pitelka, D. P. 
Abbott, and F. M. Weesner. revisers), Univ. Cali- 
fornia Press, Berkeley, xiv + 446 pp. 

Abbott. D. P., and W. B. Trason. 1968. Two new colo- 
nial ascidians from the west coast of North Amer- 
ica. Bull. So. California Acad. Sci., 67: 143-154. 

Barkley. R. A. 1968. Oceanographic Atlas of the Pacific 
Ocean. Univ. Hawaii Press. Honolulu. 20 pp. 

Crippen. R. W.. and D. J. Reish. 1969. An ecological 
study of the polychaetous annelids associated with 
fouling material in Los Angeles Harbor with 
special reference to pollution. Bull. So. California 
Acad. Sci.,68: 169-186. 

Given. R. R., and D. C. Lees. 1967. Santa Catalina 
Island Biological Survey. Survey Report No. I. 
Allan Hancock Foundation, Univ. So. California. 
Los Angeles, mimeographed report. 126 pp. 

Gregg. R. W.. and R. S. Kiwala. 1970. Some ecological 
effects of discharged wastes on marine life. Cali- 
fornia Fish Game, 56: 145-155. 



Millar. R, II. I960. Ascidcacca. Discovery Reports 
XXX: 1-160. 

Reish, D. J. 1959. An Ecological Study of Pollution 

in Los Angeles-Long Beach Harbors, California. 
Allan Hancock Foundation Puhl. Occas. Paps., 
no. 22. 1 19 pp. 

Ritter. W.. and R. Forsyth. 1917. Ascidians of the 
littoral zone of southern California. Univ. Cali- 
fornia Puhl. Zool.. 16: 439-512. 

Strachan. A. R., and R. T. Koski. 1969. A survey of 
algae of Palos Verdes Point. California. Culifomia 
Fish Game. 55: 47-52. 

Trason. W. B. 1957. Larval structure and development 
of the oozooid in the ascidian Eiiherdmania clavi- 
formis. J. Morph., 100: 509-526. 

Turner, C. H., E. E. Ebert. and R. R. Given. 1968. The 
marine environment offshore from Pt. Loma. San 
Diego County. California Dept. Fish Game, Fish. 
Bull., no. 140, 79 pp. 

Van Name. W. 1945. North and South American Ascid- 
ians. Bull. Amer. Mas. Nat. Hist.. 84: 1-476. 

Young. P. H. 1964. Some effects of sewer effluent on 
marine life. California Fish Game. 50: 33-41. 



Accepted for publication August 30. 1971. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCE 70(3): 125-130, 1971 

BENTHONIC FORAMINIFERA OF THE ARDATH SHALE AND 
STADIUM CONGLOMERATE (EOCENE), SAN DIEGO BASIN, CALIFORNIA 

Jami'.s M. Gibson 1 



Abstract: Suites of samples collected from the Ardath shale ami stadium 
Conglomerate, San Diego, California were analysed for benthonic loiamini- 
fera. The Ardaih Shale contains a classic "Ulatisian" henthonic foraminiferal 
assemblage characterized by large percentages of Cyclammina, Reophax, 
Gyroidina, Bulimina, and Eponides. The henthonic assemblage suggests 
deposition of the Ardath Shale in upper middle bathyal water depths (600 
1500 m). Planktonic foraminifera indicates an early Middle Eocene age. 

A "Narizian" henthonic foraminiferal assemblage is characteristic ol 
the Stadium Conglomerate. Arenaceous genera arc few in number (mainly 
Gaudryind). Species of Anomalina, Hanzawaia, Nonion, Melonis, and Cibi- 
cides dominate the henthonic fauna. Such a faunal composition is suggestive of 
inner shelf deposition (0-60 m). A late Middle Eocene age is indicated by 
planktonic foraminifera. 



The pre-Tertiary geology of the San Diego Basin 
(Fig. 1) was discussed by Hertlein and Grant (1944, 
1954), Milowand Ennis(1961), Peterson and Nord- 
strom (1970), and Moore and Kennedy (1970). 
Hanna (1926) proposed the term "La Jolla Forma- 
tion" for the entire Paleogene sedimentary sequence 
of the San Diego Area. Milow and Ennis (1961) 
divided the "La Jolla Formation" into the Delmar, 
Torrey, Rose Canyon, unnamed unit, and Poway 
Formations. Peterson and Nordstrom (1970), how- 
ever, retained the usage of "La Jolla Formation" 
for the lower Tertiary strata. Kennedy and Moore 
(1971) divided the La Jolla group into the Mount 
Soledad Formation, Delmar Formation, Ardath 
Shale, Scripps Formation, Friars Formation and 
also divided the Poway Formation into the Stadium 
Conglomerate and Mission Valley Formation. 

This study is concerned with the benthonic fora- 
minifera of the Rose Canyon Formation and the 
superjacent Poway Formation. The planktonic fora- 
minifera, age relationships, and general correlation 
of the Rose Canyon and Poway Formations are 
discussed more fully elsewhere by Steineck and 
Gibson (1971, 1972), Steineck, Jervey, and Gibson 
(1971) and Steineck (1971). 

Hanna (1926) and Cushman and Hanna (1927) 
established an Eocene age for the Rose Canyon 
member of the "La Jolla Formation" and the Poway 
Conglomerate. A late "Tejon" age was assigned to 
the Poway Formation by Stock (1937) on the basis 
of vertebrate remains. Foraminiferal faunas studied 
by Cushman 1 and Dusenbury (1934) were inter- 
preted also as being late Eocene in age. Clarke and 
Vokes (1936), evaluating molluscan faunas, and 
Laiming (1943), evaluating foraminiferal assem- 



blages, ascertained a "Domengine" age (upper 
Middle Eocene) for the Rose Canyon Formation. 
Laiming (1943) retained a "Tejon" age for the 
Poway Formation. Mallory (1959) presented data 
which assigns the Rose Canyon Formation to the 
"Ulatisian" stage (Middle Eocene) and the Poway 
Conglomerate to the "Narizian" stage (late Eocene). 
Milow and Ennis (1961) suggested a synchroniza- 
tion of both the Rose Canyon and Poway Forma- 
tions with the middle Eocene of the Gulf Coastal 
Plain; however, the authors do not make explicit 
the reasons for such an age assignment. Bukry and 
Kennedy (1969) assigned an early Middle Eocene 
age to the Rose Canyon Formation on the basis of 
nannoplankton floras. Kennedy and Moore (19711 
divided the La Jolla group and Poway Formation 
into various formations and assigned the Mount 
Soledad Formation. Delmar Formation. Ardath 
Shale, and portions of the Scripps and Friars For- 
mations to the Middle Eocene and the remaining 
portions of the Scripps and Friars Formations as 
well as the Stadium Conglomerate and Mission 
Valley Formation to the Upper Eocene. 

METHODS 

Natland (1933) established benthonic foraminiferal 
assemblages which are characteristic of distinct 
water depth ranges. Phleger and Parker (1951), 
Bandy (1963a). and Bandy and Arnal (1957) con- 



'Department of Geological Sciences. University of 
Southern California. Los Angeles. California 90007 
(Present address: Texaco Inc.. 3350 Wilshire Boulevard. 
Los Angeles. California 90010). 



125 



126 IU I II 1 1\ SOI I III R.X ( '.-(/ // OKMA ACADEMY OE SCIENCES VOEUME70 




Figure 1. Map of San Diego Basin showing sample localities (see text for exact locations of sample sites). 



ducted similar studies; they also found certain 
assemblages to be characteristic of distinct water 
depths. Bandy and Arnal ( 1960) used modern depth 
distributions of benthonic foraminiferal assemblages 
to interpret the paleoecology and paleobathymetry 
of Neogene deposits in the San Joaquin Basin, 
California. 

The principles developed by Natland (1933), 
Bandy (1963a), and Bandy and Arnal (1957, 1960) 
are used herein to delineate the depositional site of 
the Eocene Ardath Shale and Stadium Conglom- 
erate. Although the same species present in modern 
seas were not present during the Eocene, the general 
assemblages of modern forms when compared with 
like Eocene associations can be used to indicate the 
environs of deposition of the Eocene formations. 
The forms with the lowest depth ranges are used to 
determine the depositional site, so shoaler forms 
do not give erroneous results. Approximately 500 
specimens were counted for each sample; percent- 
ages were computed on this basis. 

Samples of the Ardath Shale were collected from 
the type locality in Rose Canyon behind the Rose 
Canyon Gas and Electric Company Power Sub- 



station, which is located near the intersection of 
Ardath Road and interstate 5 (locality 1-1, Fig. 1). 
This suite of samples lies in the upper siltstone mem- 
ber of Milow and Ennis ( 1 96 1 ) or the Ardath Shale 
of Kennedy and Moore ( 1 97 1 ). 

Stadium Conglomerate samples were collected in 
the Fenton Material Company Quarry in Murray 
Canyon, near the intersection of Friars and Murray 
Canyon Roads (locality 2-1. Fig. 1). Samples were 
collected from a megafossiliferous mudstone lens 
in the upper Poway Conglomerate unit of Milow 
and Ennis (1961) or the Stadium Conglomerate of 
Kennedy and Moore ( 1 97 1 ). 

LITHOSTRATIGRAPHY AND 
BIOSTRATIGRAPHY 

The La Jolla group consists of tan-colored shales, 
sandstones, and mudstones with minor limestone 
and conglomerate. A southwestern source area for 
Ardath Shale sediments is suggested by Milow and 
Ennis (1961). Abundant molluscan and foramini- 
ferid remains indicate a marine depositional site 
for the sediments. 



1971 



EOCENE IOKAMINII IRA 



127 



Table I. Dominant groups and species of foraminifera in samples ol Ardath Shale and Stadium ( onglomcralc. 
STADIUM CONGI.OMI RAI I AKDA'I II SI I.M I 



OROUP/SPECIES 


% TOTAL 1 AUNA ' 


<,uo[]|'/si"i (us 


rOTAI I At NA 


Arenaceous 


1. 


Arenaceous 


13 


Gaudryina sp. 


1 


Cyclammina i lurk i 1 1 lanna) 


5 


Calcareous 


W 


( '. pacifica Heck 


1 


Anomalina affinis I Rcuss) 


22 


Reophax curtus ( ushman 


4 


A. coalingensis Cushman ami 1 lamia 


1 


Textularia lajollaensis Lalicker 


1 


lliiliiiiinii all sp. 


1 


( alcareous 


7K 


Cibicides sassesi Cole 


9 


Anomalina regina minor (Smith) 


5 


Dental ina all sp. 


1 


Bolivina all sp. 


4 


Elphidium schencki Cushman anil 








Dusenbury 


1 


Bulimina robcrtsi Howe and Ellis 


3 


Eponides dorfi Toulmin 


3 


B. schwageri Yokayama 


12 


E. guaybalensis yeguaensis 


4 


Cibicides coalingensis minor (Smith) 


5 


Weinzierl and Applin 








Hanzawaia involuta (Cushman 




Elphidium sp. 


1 


and Dusenbury) 


23 


Eponides dorfi Toulmin 


6 


Lagena all sp. 


1 


E. minima Cushman 


14 


Lenticulina all sp. 


1 


Gyroidina orbicularis planum Cushman 


20 


Melonis planatus Cushman and 








Thomas 


!(. 


Lenticulina all sp. 


1 


Nonion florinensis Cole 


1 


Marginulina hantkeni Bandy 


1 


Trifarina advena californica Mallory 


3 


Noclosuria all sp. 


2 


Planktonic 


1 


Planktonic 


9 


Foram i nife ra/Ost racoda ratio 


< 100 


Foraminifera/Ostracoda ratio 


> 100 



*Ardath Shale percentages are the average of five samples from the same stratigraphic level: Stadium Conglomerate 
percentages are the average of four samples from the same horizon. 



The genesis of the Poway group, including the 
Stadium Conglomerate, is discussed by Bellemin 
and Merriam (1958) and Woodford, Welday, and 
Merriam (1968). A provenance in the Coast Range 
Batholith is suggested by Woodford el al., (1968). 
The Poway group consists of coarse sands and 
conglomerates with minor amounts of mudstone 
and siltstone. Marginal deposits yield terrestrial 
vertebrate and plant remains (Stock. 1937), while 
marine fossils (mollusks and foraminifera) occur in 
pebbly claystone lenses intercalated throughout 
the formation. 

The Ardath Shale is characterized by a Subbotina 
patagonica (Todd and Knicker) planktonic fora- 
miniferal fauna. The presence of Truncorotaloides 
densits (Cushman), Pseudohastigerina micra (Cole), 
5. patagonica, and Truncorotaloides spinitloinfiatus 
(Bandy) brackets deposition between the base of 
the Middle Eocene and medial Middle Eocene 
according to the work of Bandy ( 1 949), Bolli ( 1 957, 
1966), Pessango (1961). Saito (1962). Jenkins 
(1965a, 1965b), Berggren (1968). and Samuel and 
Salaj (1968). The Ardath Shale planktonic fora- 
miniferal fauna is equated with the Globigcrapsis 



kugleri-Hantkenina aragonensis interval of the 
standard zonation (Bolli, 1957) based on the age 
ranges of individual species. 

A Globorotaloides suteri Bolli planktonic fauna 
characterizes the Stadium Conglomerate. The cooc- 
curance of G. suteri and Truncorotaloides collacteus 
(Finlay) suggests equivalence with the Orbulinoides 
beckmanni Zone of the standard zonation according 
to the work of Bolli (1957. 1966). Pessagno ( 1 96 1 ). 
Saito (1962). Jenkins (1965a, 1965b), and Samuel 
and Salaj (1968). On this basis the Stadium Con- 
glomerate is late Middle Eocene in age. not Upper 
Eocene as reported by Kennedy and Moore ( 197 I ). 

BENTHONIC FORAMINIFERA 

The Ardath Shale is characterized by a "Ulatisian" 
{Amphimorphina californica Zone) benthonic fora- 
miniferal fauna (Mallory. 1959: Milow and Ennis. 
1961), although several of Mallory "s "Narizian" 
indices occur in the fauna I Bolivina pisciformis and 
Anomalina regina minor). The relative abundances 
of major foraminiferid groups and dominant ben- 
thonic species are listed in Table 1 . and the complete 



128 HI III IINSOl III URN CALIFORNIA ACADEMY OF SCIENCES 

I ami i 2. launal reference list 



I ()l.UMI:70 



ARDATll SlIAt E BKNTHONIC SPECIES: 

Ammopalmula sp. 

Anomalina regina minor Smith. 1957 

Bathsiphon eocenica Cushman and Hanna, 1927 

Bolivina huneri Howe, 1939 

Bolivina pisciformis Galloway and Morrey, 1929 

Bolivina sp. 

Bulimina schwageri Yokoyama. 1890 

Cibicides coalingensis minor (Smith) = Cibicidoidei 

coalingensii minor Smith, 1957 
Cibicides praecursorius (Sch\vager) = Discorbina 

praecursoria Sch wager. 1883 
Cibicides sandiegensis Cushman and Hanna. 1927 
Cyclammina clarki ( Hanna) = Nonionia? clarki Hanna. 

1923 
Cyclammina pacifica Beck. 1943 
Dental ina communis (d" Orhigny) = Nodosaria 

communis d' Orbigny. 1826 
Elphidium sp. 

Eponides dorfi Toulmin, 1941 
Eponides minima Cushman. 1933 
Fissurina sp. 
Globnlina sp. 

Gyroidina orbicularis planata Cushman. 1935 
Hormosina sp. 

Karreriella elongata Mallory, 1959 
Lenliculina anlipoda (Stache)= Rob id ina cult rata 

antipodum Stache. 1865 
Lenliculina inornata (A' Orbigny) = Robulinu inornata 

d" Orbigny. 1846 
Marginulina hantkeni Bandy. 1949 
Nodosaria cocoaensis Cushman, 1925 
Nodosaria latejugata Gumbel. 1868 
Nodosaria parexilis Cushman and Stewart. 1930 
Reophax curtus Cushman, 1920 
Texlularia lajollaensis Lalicker, 1935 
Textularia sp. 
Trochammina sp. 
Vaginulinopsis mexicana nudicostata (Cushman and 

Hanna )= Cristellaria mexicana nudicostata 

Cushman and Hanna. 1927 

Stadium Conglomerate benthonic species: 
Anomalina affinis ( Reuss) = Nonionina affinis Reuss. 1 85 1 



Inomalina coalingensis Cushman and Hanna, 1927 
Bulimina capitata Yokoyama. 1890 
Bulimina schwageri Yokoyama. 1890 
Buliminetla sp. 
Cibicides sassesi Cole, 1927 
Dentalina capitata (Bouvigner)= Nodosaria capitata 

Bouvigner, 1852 
Dentalina colei Cushman and Dusenbury, 1934 

Denial ina consobrina d' Obrigny. 1846 

Dentalina obliauistriata Reuss. 1851 

Dentalina substrigata (Stache) = Nodosaria substrigata 

Stache, 1865 
Dentalina vagina Stache, 1865 
Ellipsonodosaria sp. 
Elphidium californicum Cook. 1959 
Elphidium schenki Cushman and Dusenbury. 1934 
Eponides dorfi Toulmin, 1941 
Eponides guaybalensis yeguaensis Weinzierl and 

Applin. 1929 
Glohulina landesi (Hanna and Hanna) = Polymorph ina 

landesi Hanna and Hanna, 1924 
Guttulina hantkeni Cushman and Ozawa, 1930 
Hanzawaia involuta (Cushman and Dusenbury)= Val- 

vulineria involuta Cushman and Dusenbury. 1934 
Lagena becki Sullivan, 1962 

Lagena conscripta Cushman and Barksdale, 1930 
Lagena costata (Williamson)= Entosolenia costata 

Williamson, 1858 
Lagena hexagona (Williamson) = Lagena squamosa 

hexagona Williamson. 1848 
Lagena vulgaris Williamson, 1858 
Lenliculina articulata lexana (Cushman and Applin) = 

Cristellaria articulata texana Cushman and Applin, 

1926 
Lenliculina gyroscalpra (Stache) = Cristellaria 

gyroscalprum Stache, 1865 
Lenliculina rolula (Stache) = Cristellaria rotula Stache. 

1865 
Melonis planatus (Cushman and Thomas) = Nonion 

plantum Cushman and Thomas, 1930 
Nodosaria ewaldi Reuss, 1851 
Nonion florinensis Cole, 1927 
Trifarina advena californica Mallory, 1959 
Uvigerina sp. 



list of all species in Ardath Shale and Stadium 
Conglomerate samples are listed in Table 2. The 
presence of a substantial diverse arenaceous fauna 
characterized by abundant large Cyclammina is 
suggestive of deposition in bathyal water depths 
according to Bandy and Arnal (1960) and Bandy 
and Kolpack (1963). Bandy (1963b) and Theyer 
( 1 97 1 ) determined the upper depth limit of Cyclam- 
mina at 500 m. Size parameters of 50 specimens of 
Cyclammina (1.8 mm in diameter) indicate water 
depths approximating 600 m (Theyer. 1971). The 



foraminifera-ostracode ratio of greater than 100 
suggests shelf or deeper water deposition (Bandy 
and Arnal, 1957). The lack of miliolids and the 
high percentage of planktonic foraminiferid tests 
both point to deep water deposition (Bandy and 
Arnal. 1960). Large percentages of species of 
Eponides in the samples represent shelf species 
displaced into deeper waters, possibly by turbidites. 
The association of species of A nomalina, Gyroidina. 
Bulimina, and Cibicides is strikingly similar to the 
abyssal biofacies ( 1 500 m and deeper) of Bandy and 



1971 



I <>< ENE I OHAMINII I.KA 



129 



Arnal ( I960). All factors seem in indicate an upper 
middle bathyal depositional site (600-1500 m) for 
the Ardath Shale. 

The Stadium Conglomerate contains a "Nari- 
zian" benthonic foraminiferal assemblage. Domi- 
nant groups ami species of Stadium Conglomerate 
foraminifers are listed in Table I . Species of milio- 
lids (reported from the Poway Conglomerate by 
Cushman and Dusenbury, 1934) associated with 
species of Eponides, Cibicides, Nonion, Melonis, 
Anomalina, and Hanzawaia point to a shelf deposi- 
tional site for the Stadium Conglomerate according 
to work on modern distributions by Phleger and 
Parker (1951), Bandy and Arnal (1957, I960). 
Olsson (I960, 1963), and Nogan (1964). No niilio- 
lids were present in the samples analyzed for this 
study. Low percentages of planktonic tests and the 
presence of non-ornamented buliminids is further 
evidence of shell deposition (Bandy and Arnal, 
1957). A site of deposition approximating 0-60 m 
is suggested for the Stadium Conglomerate. 



ACKNOWLEDGMENTS 

The writer thanks the Fenton Material Company for 
permission to enter the Murray Canyon Quarry. O. L. 
Bandy. H. G. Lindenberg, and P. L. Steineck read the 
manuscript. Typing and editorial services were provided 
by B. A. Robertson. This study was partially supported 
by NSF Grant GB 8628, University of Southern 
California. 



Bcllcmin, G. .1.. and \< it Mcrriam 1958 Pctroli 

and Origin "I the Poway Conglo al< Hull 

Geol. Soc. Amur., 69: 199 220. 

Berggren, w. a. \'h,h. Phylogcnctic and Taxonomic 
Problems ol Some Tertiary Planktonii i it 
Ttilane Studies in Geol . 6 I 22. 

Bolli, II 1957. Planktonic Foraminifera from the 
Eocene Navel and S:hi Fernando Formatioi 
Trinidad, B. W, I. Bull. U.S. Nat Win .215: 155- 
172. 

. 1966. Zonation <>i the < retaceous to the Plio- 



cene Marine Sediments Based on Planktonic 
Foraminifera, Boletin Informative) Assoc. Vene- 
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Bukry. D., and M. P. Kennedy. 1969. ( retaceous and 
Eocene Coccoliths at San Diego ' alifornia. Cali- 
fornia Div. Mines aiul Geol. Spec. Rept., 100: 
33-43. 

Clarke. B. L.. and H. H. Vokes. 1936. Summary of 
Marine Eocene Sequence of Western North Amer- 
ica. Bull. Geol. Soc. Amer., 47: 851-878. 

Cushman, .1. A., and A. N. Dusenbury. 1934. Eocene 
Foraminifera of the Poway Conglomerate of Cali- 
fornia. Contr. Cushman Lab. Foram. Res., 10: 
51-65. 

Cushman. J. A., and M. A. Hanna. 1927. Foraminifera 
from the Eocene near San Diego, California. Trans. 
San Diego Nat. Hist.. 5: 47-64. 



LITERATURE CITED 

Bandy, O. L. 1949. Eocene and Oligocene Foraminifera 
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. 1963a. Dominant Paralic Formaminifera of 

Southern California and the Gulf of California. 
Contr. Cushman Lab. Foram. Res., 14: 127-134. 



— . 1963b. Larger Living Foraminifera of the Con- 
tinental Borderland of Southern California. Contr. 
Cushman Lab. Foram. Res., 14: 121-126. 



Bandy, O. L, and R. E. Arnal. 1957. Distribution of 
Recent Foraminifera off West Coast of Central 
America. Bull. Amer. Assoc. Pet. Geol., 41: 2037- 
2053. 

— . 1960. Concepts of Foraminiferal Paleoecology. 



Bull. Amer. Assoc. Pet. Geol.. 44: 1921-1932. 

Bandy. O. L., and R. Kolpack. 1963. Foraminiferal and 
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Tecolote Tunnel. California. Micropaleo., 9: 1 17- 
170. 



Hanna. M.A. 1926. Geology of the La Jolla Quadrangle. 
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Hertlein. L. G.. and U.S. Grant. IV. 1944. The geology 
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2:72 pp. 



. 1954. Geology of the Oceanside-San Diego 

Coastal Area. Southern California. Bull.. Cali- 
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Jenkins, D. G. 1965a. Planktonic Foraminiferal Zones 
and New Taxa from the Danian to Lower Miocene 
of New Zealand. New Zealand J. Geol. Geo- 
physics. 8: 1088-1126. 



. 1965b. A Re-Examination of Globorotaliu Col- 

lactea Finlay. 1939. New Zealand J. Geol. Geo- 
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Laiming. B. 1943. Eocene Foraminiferal Correlations 
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Kennedy. M. P.. and G. W. Moore. 1971. Stratigraphic 
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130 



in i i.i ns soi i in r.x ( ,1/ // ohm, i k tniM) or SCIENCES 



VOl.il Mil 70 



linns. Nan IIki'.i I nasi. J Au-.i. < aliloinia. 

Amer. Assoc. Pet. Geol.. >>: 709-722. 



/(»//. 



minifera Northwest Gulf of Mexico. Mem. Amer. 
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Mallory, V. S. 1959. Lower rertiary Biostratigraphj ol 
the California Coast Ranges. Amer. Assoc. Pet. 
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Saito, T. 1962. Eocene Planktonic Foraminifera from 

Hillsborough Island. Trans. Paleo. Sue. Japan, 
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Milow, E. D.. anil D. B. Hnnis. 196 I . Guide to Geologic 
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24-44. 

Moore. G. W.. ami M. P. Kennedy. 1970. Coastal 
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Natland. M. L. 19.13. The Temperature and Depth Dis- 
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Olsson. R. K. I960. Foraminifera of Latest Cretaceous 
and Earliest Tertiary Age in the New Jersey 
Coastal Plain. J. Paleo.. 34: 1-58. 



. 1963. Latest Cretaceous and Earliest Tertiary 

Stratigraphy of New Jersey Coastal Plain. Bull. 
Amer. Assoc. Pel. Geol.. 47: 643-665. 

Pessagno, E. A. 1961. The Micropaleontology and Bio- 
stratigraphy of the Middle Eocene Jacaquas Group, 
Puerto Rico. Micropuleo., 6: 87-105. 

Peterson, G. L.. and C. F. Nordstrom. 1970. Sub-La 
Jolla Unconformity in Vicinity of San Diego, 
California. Bull. Amer. Assoc. Pet. Geol., 54: 265- 
274. 



Samuel. O.. and J. Salaj. 1968. Microbiostialigiaphy and 
Foraminifera of the Slovak Carpathian Paleocene. 
Geologicky Ustav Dionyza Stum, Bratislavia: I- 

261). 

Steineck, P. L. 1971. Middle Eocene Refrigeration: New 
Evidence for California Planktonic Foraminiferal 
Assemblages. Lethaia, 4: 125-129. 

Steineck. P. L.. and J. M. Gibson. 197 1 . Age and Corre- 
lation of the Eocene Ulatisian and Narizian Stages. 
California. Bull. Geol. Soe. Amer.. 82: 477-480. 

. 1972. Age and Correlation of the Eocene Ulati- 



sian and Narizian Stages. California. Reply: Bull. 
Geol. Soc. Amer., 82: 2. 

Steineck. P. L.. M. T. Jervey. and J. M.Gibson. 1971. 
Paleoecologic and Biostratigraphic Implications of 
Eocene Planktonic Foraminiferal Assemblages. 
California. Bull. Amer. Assoc. Pel. Geo!., 55: 
365-366. 

Stock, C. 1937. An Eocene Titanothere from San Diego 
County, California with Remarks on the Age of the 
Poway Conglomerate. Proc. Nat. Acad. Sci., 23: 
48-53. 

Theyer, F. 1971. Size-Depth Variation in the Fora- 
minifer Cyclammina cancellata Brady from the 
Peru-Chile Trench Area. Antarctic Res. Ser., Vol. 
15, Antarctic Oceanology I: 309-3 13. 

Woodford, A. O.. E. E. Welday, and R. H. Merriam. 
1968. Siliceous Tuff Clasts in the Upper Paleogene 
of Southern California. Bull. Geol. Soc. Amer., 
79: 1461-1486. 



Phleger. F. B, and F. L. Parker. 1951. Ecology of Fora- 



Accepted for publication September 24. 1971. 



BULLETIN SO. CALIF. ACADEMY OF SCIENCE 70(3): 131-135, 1971 

A SECOND RECENT SPECIES OF NOETIA, SENSU SI RICTO (MOLU ISi A: 111 VAI.VIA) 
IN THE TROPICAL EASTERN PAC UK 0< IAN 

Barry Roth ] 



Abstract: Recent specimen^ of an arcoid bivalve differing in some partic- 
ulars from Noetia reversa (Sowerby) arc reported from (cniial Amen 
localities, and tentatively referred to Noetia magna MacNeil, described from 
the C'll.ate Pliocene of Peru, 



While sorting a suite of ark shells of the genus 
Noetia in the collection of the Geology Department, 
California Academy of Sciences, the author noted 
the presence of two distinct forms, the first being 
typical Noetia reversa (Sowerby, 1833), the second 
resembling the Late Pliocene or Pleistocene Noetia 
magna MacNeil, 1938, which bears considerably 
more and finer ribs on the exterior of the valves. 
An examination of all material in the collections 
showed the 2 forms to be present in lots from 2 
other localities; and the distinction was also found to 
hold in material from the Natural History Museum 
of Los Angeles County and Stanford University. 
Because the existence in the Recent fauna of two 
such similar species cast doubt on the synonymy of 
Noetia reversa and Noetia triangularis Gray, 1857, 
(type species of the genus Noetia), a search was 
instituted for the type specimens of those 2 species. 

Family Noetiidae 
Genus Noetia Gray 

Noetia Gray, 1840: 151 [nomen nudum] ; Gray, 
1857; MacNeil, 1938. 

Type species, by original designation, Noetia tri- 
angularis, "n.s." Recent, no locality given [ = Arca 
reversa (Gray MS) Sowerby, 1833; Recent, Tumbez, 
Peru]. 

Frizzell (1946) gave a critical discussion of 
factors affecting the suprageneric classification of 
arcoid pelecypods. MacNeil (1938) discussed the 
species of 8 genera and 1 subgenus which he placed 
in the subfamily "Noetinae." He gave the range of 
Noetia as "upper Eocene of the Indo-Pacific and . . . 
lower Miocene to Recent of America." Newell 
(1969) recognized 7 genera and subgenera in the 
subfamily Noetiinae and placed 3 subgenera under 
Noetia: Eontia MacNeil, 1938; Incanopsis Olsson, 
1944 (= Palestinarca Vokes, 1946); and Noetia 
sensu stricto. He characterized the typical sub- 
genus as follows: "Umbonal ridge carinate; radial 
ribs coarse, simple, not bifurcate over umbonal 



slope; posterior margin subangular. commonly 
truncate." 

For a discussion of the molluscan names pub- 
lished in the various editions of the "Synopsis of the 
Contents of the British Museum," in which Noetia 
appears as a nomen nudum, see Iredale (1913). 

Noetia (Noetia) magna MacNeil 

Area (Noetia) reversa Gray, Grzybowski, 1899: 
634-635, pi. 17, figs. 1,1a [Not Area reversa (Gray 
MS) Sowerby, 1833: 20]. 

(?) Area (Noetia) reversa Sowerby, — subsp.. 
Olsson, 1932: Miocene, Tumbez Formation, Zor- 
ritos, Peru. 

Noetia magna MacNeil, 1938. 

Noetia (Noetia) reversa magma [sic] MacNeil. 
Pilsbry and Olsson, 1941; Pliocene. Jama Forma- 
tion. Ecuador; Pleistocene, Panama. 

Noetia reversa magma [sic] MacNeil. Olsson. 
1942a; Pliocene and Pleistocene. Burica Peninsula. 
Panama. 

Type material: Holotype in Geological Institute 
of the Jagiellonian University. Krakow. Poland. 
Rubber cast of holotype. California Academy of 
Sciences, Department of Geology Type Collection 
No. 13630. 

Type locality: Paita (Grzybowski). ?Late Plio- 
cene. Mancora Tablazo Beds, Paita. Peru (fide 
Olsson, 1932). 

Original description: According to MacNeil 
(1938: 38), "The form from Paita which Grzybow- 
ski referred to Noetia reversa is much like it in 
shape but differs in size, being about twice as large, 
and in having more ribs (40-42) as compared with 
35 for N. reversa. Photographs of Grzybowski's 
specimens show it to have more elevated ribs, with 
less tendency to widen on the anterior part of the 



'Department of Geology. California Academy of 
Sciences. San Francisco. California 94118. (Present 
address: 1217 Waller St.. San Francisco. California 
94117). 



131 



132 



HI I I I I IN SOI I III l<\ CM II ORNIA ACADEMY Ol SCIINi I S 



\(>I.(/Mi:7() 



disk, although this might be a matter of individual 
variation. Ii compares with N. re versa in having 
strongly opisthogyrate, low. and very posterior 
beaks; these being at or even behind the posterior 

end of the cardinal area. The (wo species are 
extreme in this respect. Length SO mm. height 72 
mm, convexitj 37 mm. The margins are eroded, so 
that only approximate measurements can he made." 

DISCUSSION 

The existence of Recent specimens of Noetia, s. s. 
differing from typical Noetia reversa (Sowerby) 
was first noted by the author in a lot of beach valves 
from San Jose, Guatemala (Loc. 36678. CAS). 
Approximately 1 out of every 10 valves had between 
42 and 47 radiating ribs; the remainder gave rib- 
counts of 3 1-37. Most authors have reported 35 or 
36 as the number of ribs for N. reversa. [An appar- 
ent misprint in Olsson (1961) calls for 25 ribs] . 

Tabulation of rib-counts for all Recent Noetia. 
s. s. valves in the California Academy of Sciences, 
Natural History Museum of Los Angeles County, 
and Stanford University collections showed most 
falling between 3 1 and 37 (with 36 and 33 ribs most 
frequent), with another, smaller, concentration 
between 42 and 47. Counts of 38, 39, 40, and 41 
ribs were obtained from only one valve each. The 
fine radiating sculpture of the extreme posterior 
dorsal portion near the beaks was excluded from the 
counting. This fine sculpture is difficult to detect on 
a shell with periostracum intact, and is the first 
feature to be eroded from beach specimens. Number 
of ribs is not directly correlated with shell size. 

The results of the above tabulation seem to 
indicate the presence of two sympatric species. In 




the California Academy of Sciences collection, line- 
ribbed Recent specimens, resembling the holotype 
o\ Noetia magna MacNeil (1938), range from 
Corinto, Nicaragua (Loc. 27226. CAS) to near 
Ma/.atlan, Sinaloa, Mexico (Loc. 27583, CAS). The 
latter lot of 3 living specimens was trawled in 10-17 
fathoms by the Tcmplcton (rocker Expedition, 
July 29, 1932. One of the specimens is illustrated 
here (Figs. 1-5). The collection of the Natural 
History Museum of Los Angeles County contains a 
line-ribbed example (LACM No. HH-756) from 
Venado Island, Panama Bay, Panama. It seems 
likely that future collecting and examination of 
other specimens will extend the known range even 
farther. 

MacNeil ( 1938)distinguished Noetiamagna from 
N. reversa on 3 characters: greater number of ribs 
in N. magna, tendency of the ribs to remain narrow 
anteriorly (which is really a function of their multi- 
plicity), and the very large size of the shell. [Mac- 
Neil may have taken his measurements from a 
photograph. Dimensions of the plastotype (CAS 
No. 13630) match GrzybowskTs measurements: 
length 75 mm, height 68 mm, convexity of one 
valve 34 mm] . Pilsbry and Olsson ( 1941 ) reaffirmed 
the size difference while placing N. magna as a 
subspecies of N. reversa. Recent specimens larger 
than about 60 mm in length parallel to the hinge 
line have not been seen by the author. The largest 
seen have even more ribs than the 40-42 described 
by MacNeil. The application of the name N. magna 
to the Recent species, therefore, may be regarded 




Figures 1-5. Noetia cf. N. magna MacNeil. Hypo- 
type (CAS No. 13675). Recent, off Mazatlan. Sinaloa. 
Mexico. Length parallel to hinge line 56 mm; height 
52 mm. 



Figures 6-10. Noetia reversa (Sowerby). Holotype. of 
Noetia triangularis Gray, Reg. No. 1968856. Recent. 
locality unknown. Length parallel to hinge line about 
34 mm; height about 30 mm. Photographs courtesy of 
and copyright by British Museum (Natural History). 



1971 



IOSSII. PELECYPODS 



133 




Figures 1 1-15. Noelia reserva (Sowerby). "Possible syntypes" of Area reserva Sowerby. BM (NH) (unnumbered). 
Recent. (?) Tumbez. Peru. Length parallel to hinge line about 40 mm: height about 33 mm. Photographs courtesy of 
and copyright by British Museum (Natural History). 



as tentative. Olsson (1942a) reported that "the form 
[of N. magna] tends to be longer or more regu- 
larly rectangular." a statement which characterizes 
neither the holotype of N. magna nor most of the 
Recent specimens observed. 

A list by Bosworth (1922) of mollusks from the 
tablazos of northern Peru, assigned by him to the 
Quaternary, consists entirely of species still living 
in the same general area. The Mancora Tablazo. 
presumed source of the type of N. magna, contains 
several extinct species as well, and was referred by 
Olsson (1942b) to the Late Pliocene. 

The discovery that two similar species of Noelia 
occur in the Panamic province calls into question 
the synonymy, first advanced by Reinhart (1935), of 
Noetia triangularis Gray and N. reversa (Sowerby). 
Reinhart's study showed that N. triangularis is not a 
synonym of Area ponderosa Say, 1822, as had been 



assumed by some earlier workers. Photographs of 
the holotype of N. triangularis. B M (NH) Reg. No. 
1968856. supplied by Dr. J. D. Taylor. British 
Museum (Natural History), and reproduced here 
(Figs. 6-10) show a shell with 36 major radial ribs 
and a few smaller, intercalary ribs on the posterior 
slope of the valve. No undoubted type specimen of 
Area reversa Sowerby was located at the same 
institution, but photographs of a "'possible sy nt\ pe" 
supplied by Dr. Taylor (Figs. 1 1-15) show a speci- 
men with approximately 33 major ribs. Both names. 
Noetia triangularis and N. reversa. would therefore 
seem to apply to the same species, the common 
large Noetia of the Panamic province, which is 
reported to range from Puertecitos in the northern 
Gulf of California to Mancora. Peru. Its apparent 
geologic range is Pleistocene to Recent. 

Area hemicardium Koch in Philippi (1843) was 



134 



HI I I I 1 1\ SOI I ill R\ ( AI.IEORNIA ACADEMY Oh SCIENCES 



VOLUME 70 



described as having 20-22 ribs on the "posterior" 
(actual I) anterior) portion of the shell and 11-12 
on the (posterior) slope. The accompanying figure 

shows a sIk-II with about 38 ribs. Philippi (IN44) 
himself sj nonj mized Area hemicardium and Noetia 
reversa, and many later authors have repeated the 
synonymy. Synonymies for N. reversa were given 
bj MacNeil (1938) and Olsson (1961). 

One interesting feature revealed by the present 
inquiry, as pointed out by Dr. Leo G. Hertlein. is 
that the relationship between Noetia reversa and 
N. cf. N. magna in the Recent fauna replicates 
a situation existing in the Miocene of Central 
America. Typical Noetia macdonaldi Dall, 1912. 
(type locality. Miocene, near Banana River, Costa 
Rica), common in the Gatun Formation and its 
equivalents in Costa Rica, Panama, and Colombia. 
has low beaks and wide anterior ribs. Occurring 
alongside it is a high-beaked form in which the 
anterior and posterior ribs are more uniform in 
size. Olsson (1922). believing Dall's type to be 
a high-beaked specimen, proposed the varietal 
name subreversa for the low-beaked form. MacNeil 
(1938) synonymized Olsson's name with that of 
Dall and proposed both Noetia macdonaldi alia 
for high-beaked shells from Costa Rica and Noetia 
macdonaldi truncal a for Colombian specimens 
with very regular and comparatively narrow ribs. 
Whether the number and quality of ribs was more 
subject to individual variation in former times 
than at present, or if separate and sympatric species 
were involved remains to be shown by a statistical 
study. MacNeil also mentioned that specimens of 
N. macdonaldi average fewer and fewer ribs the 
higher one moves in the stratigraphic section at 
the type locality. 



Dall. W. II. 1912. New species of fossil shells from 
Panama and Costa Rica collected by D. F. Mac- 
Donald. Smithson. Inst. Misc. Coll.. 59: 1-10. 

Frizzell, D. L. 194ft. A study of two arcid pclccypod 
species from western South America. ./. Paleo., 
20: 38-51. 

Gray..). E. 1840. Mollusca. Pp. 146-152 /;; Synopsis of 
the contents of the British Museum. 42nd Edition. 
British Museum. U>ndon. 

— . 1857. A revision of the genera of some of the 
families of conchifera or bivalve shells. Part III. 
Arcadae. Ann. Mag. Nat. Hist., London, ser. 2, 
19: 366-373. 

Grzybowski. J. 1899. Die Tertiarablagerungen des 
nordlichen Peru und ihre Molluskenfauna. Nenes 
Jahrbuchfiir Mineralogie, Geologic und Palaeonto- 
logie, Stuttgart. 12: 610-664. 

Iredale. T. 1913. A collation of the molluscan parts of 
the synopses of the contents of the British Museum, 
1838-1845. Proc. Malacol. Soc. London. 10:294- 
309. 

MacNeil. F. S. 1938. Species and genera of Tertiary 
Noetinae. U.S. Geol. Sun.. Prof. Paper 189-A: 

1-49. 

Newell, N. D. 1969. Family Noetiidae Stewart, 1930. 
Pp. N261-N264/n R. C. Moore, et at., eds. Treatise 
on invertebrate paleontology, part N. I (Mollusca 
6, Bivalvia). Geol. Soc. Amer., New York. 

Olsson, A. A. 1922. The Miocene of northern Costa 
Rica, with notes on its general stratigraphic rela- 
tions. Part 2. Class Pelecypoda. Bull. Amer. Paleo., 
9: 341-482(169-310). 



ACKNOWLEDGMENTS 

For assistance during this investigation, the writer 
wishes to thank Leo G. Hertlein, Department of Geol- 
ogy, California Academy of Sciences; Myra Keen. 
Stanford University; James H. McLean and Gale 
Sphone. Natural History Museum of Los Angeles 
County; J. D. Taylor. British Museum (Natural His- 
tory), who supplied excellent photographs of two speci- 
mens in question: W. Nowak, Geological Institute. 
Krakow; and M. Ksiazkiewicz. Department of Geology. 
Jagiellonian University, Krakow, who furnished a cast 
of the holotype of Noetia magna. 

LITERATURE CITED 

Bosworth, T. O. 1922. Geology of the Tertiary and 
Quaternary periods in the north-west part of Peru, 
with an account of the palaeontology by Henry 
Woods, T. Wayland Vaughan. J. A. Cushman and 
others. Macmillian and Co.. Ltd., London. 434 pp. 



— . 1932. Contributions to the Tertiary paleonto- 
logy of northern Peru: Part 5. The Peruvian Mio- 
cene. Bull. Amer. Paleo.. 19: 1-272. 



— . 1942. Tertiary and Quarternary fossils from the 
Burica Peninsula of Panama and Costa Rica. Bull. 
Amer. Paleo.. 27: 153-259 ( 1-107). 



. 1942b. Tertiary deposits of northwestern South 

America and Panama. Proc. Eighth Amer. Sci. 
Cong.. Geol. Sci.. 4: 23 1-287. 

— . 1961. Mollusks of the tropical eastern Pacific. 



particularly from the southern halfof the Panamic- 
Pacific faunal province ( Panama to Peru). Panamic- 
Pacific Pelecypoda. Paleo. Res. Inst.. Ithaca, 574pp. 

Philippi. R. A. ( I842M845. Abbildungen und Beschrei- 
bungen neuer oder wenig gekannter Conchylien. 
Theodor Fischer, Kassel, 1: [i-x], 1-204. (Pages 
21-46 issued 1843; title and index issued 1844). 



1971 FOSSIL PEL El ) PODS 

— 1843, 1844. see Philippi, [1842] 1845, Say, T. 1822. An account of some of the marine ihcll 

ol the United States ./ I Phlladel 

Pilsbry, H. A., and A. A.OIsson. 1941. A Pliocene fauna phia, 2:221 248. 257-276. 

from western Ecuador. Proc, Acad, Nat. Sci. 

Philadelphia, 93: 1-79. Sowerby, G B 1833, Charactci ol new specie ol 

Molluscaand Conchifera Proi ZooI.Sch London, 
Reinhart, P, W. 1935. Classification of the pelecypod ixw 16-22. 

family Arcidae. Bulletin tin Muxie royal d'Histeire 
natnrelle de Belgiaue, II: 1-68. Accepted for publication August 10, 1971. 



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1971-1974 
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Daniel Gutherie 
I. M. Newell 



cover: The arborescent prickly pear cactus, Opuntia galapageia, growing on Bartholomew 
Island, one of the Galapagos group. This plant is believed to have evolved from a more shrubby, 
prostrate form in response to predation by lizards and the giant tortoise. 

Photograph by C. F. Anderson, 
Whitman College, Walla Walla, 
Washington. 



D 



SOUTHERN CALIFORNIA ACADEMY OF S C I E N ( E S 

1BRARY 

-■•- BULLETIN 

NEW YORK 
rANlCAJ. GARDEN 



Volume 71 



Number 1 




BCAS-A71(1) 1-56 (1972) 



APRIL 1972 



< 



Southern California Academy of Sciences 

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1970-1972 

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1971-1973 

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U U LLIiTI N () I' III L SO 111 II I R N (A I 
ACADEMY OF SCIENCES 



R N I A 



Volume 71 



May 5, 1972 



'.i MB! R I 



I IM I OK I \l 



A liii of History.' 



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(II Ni-RIt' PARI I HONING OF 1111': SOUTH AMERICAN LEPTODACTYL1D 
FROG GENUS EUPSOPHUS FITZINGER, 1843 (SENSU LATO) 

John D. Lynch 1 



ABSTRACT i The frog genus Eupsophus has been loosely defined and is heterogeneous. Study 
of osteological material revealed several major differences among the included species; these 
data when collated with published results of other workers on life history data and external 
morphology, provide the bases for definition and recognition of three genera of frogs now 
included in Eupsophus (Eupsophus. Ischnocnema, and Thoropa) and for placing some species 
in other genera (Batrachyla and N kef omnia). Owing to paucity of data, two species are 
not assigned to currently recognized genera (Cystignathus sylvestris and Phrynopus peruanus). 



According to Gorham (1966) the leptodactylid 
frog genus Eupsophus contains 16 species (E. 
bolitoglossus, E. columbianus, E. coppingeri, E. 
illotus. E. lutzi. E. miliaris, E. nodosus, E. per- 
vanus, E. petropolitamis, E. quixensis, E. roseus, 
E. sylvestris. E. taeniatus, E. versus. E. vertebralis. 
and E. wettsteini) . No author has dealt with the 
genus as a whole but reviews of several species 
groups are available ( Bokermann, 1965; Cei. 1960, 
1962; and Grandison, 1961). The limits of the 
genus have been debated by Bokermann ( 1 965 ) 
and Gallardo ( 1965 ) who recommended removing 
three Brasilian species (lutzi, miliaris, and petro- 
politamis ) from Eupsophus and using the genus 
Thoropa for these species. Gallardo (1970) and 
Barrio ( 1970) have generically isolated the species 
of Cei's (1960) and Grandison's (1961) nodosus 
group as Alsodes (including coppingeri. illotus. 
nodosus. and certain other names not mentioned 
in Gorham's list). Bokermann (1966) removed 
bolitoglossus from Eupsophus and considered the 
species identical with Craspedoglossa sanctae- 
catharinae. 

Some other names have been allocated to Eup- 
sophus. Rivero (1964) named Eupsophus ginesi 
from Venezuela; Lynch (1968a) allocated this 
species to Eleutherodactylus and tentatively as- 
signed columbianus to Niceforonia. Lynch (1968a) 
placed Syrrhophus juninensis in the genus Eup- 
sophus. Lynch (1968b) argued that Alsodes mon- 
ticola Bell was an older name for the species Cei 
(1960, 1962) and Grandison (1961) called Eup- 
sophus coppingeri. 

The genus Eupsophus has been taxonomically 
difficult to define. Definitions employed by 
earlier authors have not been complete, in that 
the genus could not be separated from all others 
and because the extent of intrageneric variation 



in some characteristics was not considered or 
known. In effect, Eupsophus was used for those 
leptodactylids not characterized by some distinctive 
features; evidence for the poor definition of the 
genus is provided by the submerged heterogeneity 
in osteological and life history data. Eupsophus 
was defined on the basis of narrow digital tips, 
absence of or very brief toe webbing, and pre- 
vomerine teeth lying between the choanae. 

The purpose of this paper is to point out the 
heterogeneity of the species included in Eupsophus 
and to subdivide the genus into definable units. 
The 16 species included in the genus Eupsophus 
by Gorham (1966) belong to at least six genera; 
Eupsophus in the restricted sense would include 
only species from Argentina, Chile, and central 
Peru. 

In the course of my generic review of the family 
(Lynch, 1971), I prepared skeletons of several 
species referred by various authors to Eupsophus 
and allied genera; study of the skeletal material 
and comparisons of the non-osteological traits of 
these species resulted in the conclusion that the 
generic arrangement of the frogs placed in Eup- 
sophus and allied genera was in need of review. 
This impression is enhanced by Bogart's (1970) 
paper on the chromosomes of several species of 
the group. 

Generic definitions of leptodactylid frogs are 
based on a number of osteological traits (arrange- 
ment of skull bones, shape of the vertebrae, and 
shape of the bones in the pectoral girdle), some 
myological traits (arrangement of the muscles of 



1 Museum of Natural History, University of Kan- 
sas, Lawrence, Kansas 66044 (Present address: Dept. 
Zoology, University of Nebraska, Lincoln, Nebraska 
68508). 



1972 



SOUTH AMERICAN LEPTODACTYLID FROGS 



the hyoicl, thigh muscles, and jaw muscles), ex- 
ternal morphology, and larva] traits (including egg 
I si/,e, deposition sites, and larval biology). Data 
for some traits are not available for all species hul 
enough data are available to group the species 
and to comment on generic status. 

METHODS 

Specimens were made available from the collec- 
tions of the American Museum of Natural History 
(AMNH), Field Museum of Natural History 
(FMNH), Kansas University Museum of Natural 
History (KLJ), Naturhistoriscb.es Museum zu Wien, 
Vienna (NMW), University of Illinois Museum of 
Natural History (UIMNH), University of Michigan 
Museum of Zoology (UMMZ), United Slates Na- 
tional Museum (USNM), and Univcrsitetcs Zoo- 
logiskc Museum, Copenhagen (ZMK). Preserved 
examples of the following species were examined 
(institutional abbreviations in parentheses indicate 
location of specimens): Batrachyla leptopus (KU, 
UMMZ), Eupsophus juninensis (KU, MCZ), E. 
Inlzi (KU), E. miliaris (KU, UIMNH, UMMZ), 
/:'. monticola (FMNH), E. nodosus (FMNH, 
UMMZ), E. peruanus (UMMZ), E. petropolitanus 
(KU), E. quixensis (KU, UIMNH, UMMZ), E. 
roseus (FMNH, KU), E. taeniatus (KU, UMMZ), 
E. versus (ZMK), E. wettsteini (NMW), Nice- 
foronia festae (KU, USNM), N. flavomaculata 
(KU, USNM), N. montium (MCZ), and N. nana 
(USNM). 

Skeletal maerial in the form of dry, dermestid 
cleaned skeletons and cleared and stained prepara- 
tions was used. The specimens used are listed 
elsewhere (Lynch, 1971); skeletons of the follow- 
ing species were studied: Batrachyla leptopus, 
Eupsophus juninensis, E. httzi, E. miliaris, E. pe- 
tropolitanus, E. quixensis, E. roseus, E. taeniatus. 
E. wettsteini, Niceforonia festae, N. flavomaculata, 
and N. montium. Superficial dissection was used 
to check certain cranial features (e.g., presence of 
quadratojugal, separation of nasal bones, presence 
of a frontoparietal fontanelle, and condition of the 
ear) in rare species (Eupsophus monticola, E. 
nodosus, E. peruanus, and E. versus). 

Literature reports were used as data sources 
for external characteristics and breeding biology. 
My primary sources were Cei (1960, 1962), Gal- 
lardo (1962, 1970), Grandison (1961), and the 
original descripions for species of which I saw 
no preserved examples — Barbour, 1922 (illotus). 
Tschudi, 1845 (sylvestris), and Grandison, 1961 
(vertebralis) . 



( HARACTERIS1 U S ' ►] GROI PS 

Cei ( I960, 1962) and Grandison I 1961 I di 

three specie, groups "I ' hilcan Eupsophut on th..- 
basis <ii position ol the prcvomcrinc teeth relative 
to the choanae, size ol tl mum, notching 

of the sternum, and expo ihe tympanum, 

Cei (1960, 1962) also commented on the other 
frogs sometimes included in Eupsophus and termed 
these the peruanus-wettstcinl group (central Peru) 
and Thoropa (southeastern Brasil) Gallardo 
( 1965) pointed out osteological bases for separat- 
ing Thoropa from Eupsophus 'shape Ol the ter- 
minal phalanges and arrangement ol atl.inlar 
cotyles ). 

Before listing the characteristics <>l the species 
groups of frogs included in Eupsophus by various 
authors, it is germane to comment on the U 
certain characteristics of the pectoral girdle in de- 
fining species groups ol / upsophus. All ol the 
species included in Eupsophus have arciferal pec- 
toral girdles, cartilaginous sterna, and an omo- 
Sternum. Several authors have used the presence 
or absence of a posterior notch in the xiphistcrnum 
(= sternum here) to distinguish Argentine-Chilean 
species groups. The notch is present in the species 
of the nodosus group (all relatively large frogs) 
and absent in the species of the roseus and tae- 
niatus group (smaller frogs). In species of the 
miliaris group, the large species (miliaris) has a 
notched sternum, whereas the smaller species have 
no notch in the sternum (lutzi and petropolitanus). 
I have not used this characteristic because it ma\ 
simply be size-dependent and thus agree with a 
classification based on average adult size. 

Three species (lutzi. miliaris. and petropolitanu\) 
from the mountains of southeastern Brasil make 
up the miliaris group. This group is frequently 
recognized as the genus Thoropa (Bokermann. 
1965: Gallardo. 1965) and is defined as follows: 
larvae aquatic, flattened and attenuate, labial pa- 
pillae incomplete anteriorly: eggs large (5 mm in 
diameter in lutzi, Bokermann. 1965). pigmented: 
prevomerine dentigerous processes lying between 
choanae: tympana visible externally: fingers not 
bearing terminal pads, first finger equal in length 
to or shorter than second, males bearing spiny 
nuptial pads on thumb and sometimes second 
and third fingers: males lacking pectoral plates of 
spines: toes not bearing terminal pads, tips slightly 
expanded, lacking webbing, bearing slight lateral 
fringes: two metatarsal tubercles, inner much 
larger than outer: plantar surface lacking distinct 
supernumerary tubercles: quadratojugal bone pres- 



Ill I II ll\ SOI llll.li\ < .11 II OHM. t ACADEMY OF SCIENCES 



VOLUME 7 1 





Figure I. Plantar views of feet of Thoropa miliaris 
(left, KU 92852) and Ischnocnema quixensis (right, 
KU 99010). 



ent; nasal bones separated; frontoparietal fonta- 
nels present; occipital condyles widely separated: 
terminal phalanges T-shaped. 

Three species (Mollis, monticola, and nodosus) 
from western Argentina and central Chile make 
up the nodosus group (Cei, 1962; Gallardo, 1962). 
This group is defined as follows: larvae aquatic. 
not flattened or attenuate, labial papillae incom- 
plete anteriorly; eggs large (3.2-3.6 mm in diam- 
eter), not pigmented (Cei, 1962; egg data only for 
nodosus); prevomerine teeth on processes lying 
between choanae; tympana not evident externally 
(lost in monticola and illotus); fingers not bearing 
terminal pads, first finger longer than second, 
males bearing nuptial asperities on thumb and 
second finger; males bearing pectoral plates of 
spines in breeding season (Cei, 1962; Gallardo, 
1970); toes not bearing terminal pads, toes bear- 
ing indistinct to prominent lateral fringes and web 
rudiment (least developed in nodosus); two meta- 
tarsal tubercles, inner much larger than outer; 
plantar surface lacking distinct supernumerary 
tubercles: quadratojugal bone present; nasal bones 
separated (only nodosus studied); frontoparietal 
fontanelle present; occipital condyles narrowly 
separated; terminal phalanges knobbed. 

Two species (quixensis and versus) are included 
in the quixensis group. One occurs in Amazonian 
Ecuador and Peru (quixensis) and the other is 
known from southeastern Brasil. This group is 
defined as follows: larvae not known, develop- 



ment probably direct; eggs of quixensis are un- 
pigmented and about 2.5 mm in diameter (ovarian 
eggs); prevomerine teeth on processes lying pos- 
teromedial to choanae; tympana visible externally; 
fingers not bearing terminal pails, first linger 
longer than second, males lacking nuptial asper- 
ities: no pectoral plates of spines; toes not bearing 
terminal pads, toes lacking lateral fringes and 
webbing: two metatarsal tubercles, equal in size; 
plantar surface bearing numerous conical super- 
numerary tubercles (Fig. 1); quadratojugal bone 
present; nasal bones in broad contact; no fronto- 
parietal fontanelle; occipital condyles widely sepa- 
rated; terminal phalanges knobbed. 

Two Chilean species (roseus and vertebralis) 
make up the roseus group here defined as follows: 
larvae not known; eggs large and unpigmented 
(Cei, 1962); prevomerine teeth on processes lying 
between or slightly posterior to choanae; tympana 
visible externally; fingers not bearing terminal 
pads, first finger shorter than or equal in length 
to second; males bearing nuptial asperities on 
thumb and second finger; no pectoral plates of 
spines; toes not bearing terminal pads, toes lack- 
ing lateral fringes and webbing; two metatarsal 
tubercles, inner much larger than outer; plantar 
surface lacking distinct supernumerary tubercles; 
quadratojugal bone present; nasal bones widely 
separated; frontoparietal fontanelle present; occip- 
ital condyles narrowly separated; terminal pha- 
langes knobbed. 

The Chilean E. taeniatus is distinctive compared 
to the other species and is included in its own 
group: larvae aquatic, not flattened or attenuate, 
labial papillae complete anteriorly (enclosing 
mouth); eggs large (3 mm in diameter) and pig- 
mented (Cei, 1962); prevomerine teeth on pro- 
cesses lying between choanae; tympana visible ex- 
ternally; fingers not bearing terminal pads, first 
finger as long as second; males bearing nuptial 
asperities on thumb and second finger; no pectoral 
plates of spines; toes not bearing terminal pads, 
toes lacking lateral fringes and webbing; two meta- 
tarsal tubercles, inner much larger than outer; 
plantar surface lacking distinct supernumerary 
tubercles: quadratojugal bone absent; nasal bones 
widely separated; frontoparietal fontanelle present; 
occipital condyles widely separated; terminal pha- 
langes T-shaped (comments to the contrary by 
Cei, 1960, 1962, and Grandison, 1961, are in 
error: dissection of the toe as a means of deter- 
mining the shape of the terminal phalanges fre- 
quently results in breakage of the fragile lateral 
expansions). 



1972 



SOU I II AMI.KICAN 1 .1.1' I <)I)A( I) I II > ll«H,\ 




Figure 2. Skulls of Ischnocnema quixensis (left, UIMNH 50643) and Eupsophm roseus 
(right, KU 84731 ). 



The species from the plateau of central Peru 
are rare and poorly known. Four species are in- 
volved (juninensis, peruanus, sylvestris, and wett- 
steini). I have seen preserved examples of three 
(sylvestris is known only from lost type-material, 
fide Peters, 1873a) and have seen skeletons of 
juninensis and wettsteini. No reproductive data 
are available except that juninensis has large un- 
pigmented eggs and none of the specimens known 
have nuptial asperities on the fingers or pectoral 
plates of spines. Prevomcrine teeth are absent in 
juninensis and wettsteini but are present and lie 
between the choanae in the other two species. 
The tympana are visible externally in peruanus, 
sylvestris, and wettsteini but are lost in juninensis. 
The fingers and toes do not bear terminal pads 
and the first finger is shorter than the second. The 
toes lack lateral fringes and webbing; two meta- 
tarsal tubercles are present, the inner is larger in 
juninensis and sylvestris, but the tubercles are 
about of equal size in peruanus and wettsteini. 
Supernumerary plantar tubercles are absent in all 
four species. The quadratojugal bone is present 
in juninensis, peruanus, and wettsteini; the nasal 
bones are moderately separated in juninensis and 
wettsteini: juninensis has a small frontoparietal 
fontanelle. whereas wettsteini does not; the occip- 
ital condyles are narrowly separated in juninensis 
and widely separated in wettsteini; and the ter- 
minal phalanges are knobbed (juninensis and wett- 
steini). The species of this group are difficult to 
deal with because so much data are lackins. For 



purposes of discussion this group is termed the 
peruanus group. 

DISCUSSION 

The six groups defined above are each homoge- 
neous with the exception of the last group 
(juninensis, peruanus, sylvestris. and wettsteini) . 
The major difficulty with that group is the paucity 
of specimens and information. On the basis of 
the group definitions each of the first five groups 
could be recognized as a distinct leptodactylid 
genus. The genus Thoropa is available for the 
miliaris group and is distinctive in tadpole mor- 
phology and osteology. In large measure, the 
genus has been recognized because the terminal 
phalanges are T-shaped (in contrast to the knobbed 
phalanges of the nodosus and roseus groups). Gal- 
lardo (1965) pointed out one skeletal feature 
which readily distinguishes the miliaris group 
(Thoropa) from the nodosus and roseus groups. 
In Thoropa the occipital condyles are widely sepa- 
rated and the atlantal cotyles are widely separated 
producing a gap at the occipito-cervical articula- 
tion, whereas in Eupsophus the condyles (and 
cotyles) are narrowly separated (Fig. 2) and there 
is no gap (Gallardo termed these two character 
states concave and convex atlas, respectively). In- 
clusion of the miliaris group in the genus Eup- 
sophus greatly increases the limits of the genus 
and returns us to the era in which certain genera 
were taxonomic wastebaskets. 



BULLETIN SOI THERh CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



I he taenialus group (taenialus) is distinctive in 
tadpole morphology and osteology (no quadrato- 
jugal hone. T-shaped terminal phalanges, and 
widely separated occipital condyles) when com- 
pared with the nodosus and roseus groups. The 
OSteological data were not known to previous 
workers and thus the distinction of the group 
from other Eupsophus species was not appreciated. 
The definition of the group is identical with that 
for frogs of the genus Batrachyla. Batrachyla and 
Eupsophus have heen separated on the basis of 
T-shaped terminal phalanges in the former by 
earlier authors (Cei, 1962) but are more distinc- 
tive than the single character difference usually 
cited. Loss of the quadratojugal bone is a fre- 
quent occurrence in some groups (e.g., Hylidae) 
but is uncommon among leptodactylid frogs 
(known only in Crossodactylus, Batrachyla. and 
Eupsophus taenialus; Lynch, 1971). The widely 
separated occipital condyles in Batrachyla and 
Eupsophus taenialus strongly contrast to the con- 
dition seen in the nodosus and roseus groups. Two 
solutions are available: (a) to include Batrachyla 
in the genus Eupsophus, or (b) to remove taenialus 
from Eupsophus and include it as a species of 
Batrachyla. In order to maintain some homoge- 
neity within leptodactylid genera, the latter choice 
is most preferable. Accordingly, Eupsophus tae- 
niatus should be called Batrachyla tacniata 
(Girard). new combination. The definition of 
the genus Batrachyla is the same as that listed for 
the taeniatus group. As now constituted, the genus 
Batrachyla contains three species (antarlandica, 
leptopus, and taeniata). 

The quixensis group includes two species. The 
characteristics of the group require its removal 
from Eupsophus. Among the osteological features 
distinguishing this group from Eupsophus (nodosus 
and roseus groups) are the broad contact of the 
nasal bones, absence of a frontoparietal fontanelle, 
the more posterior position of the prevomerine 
dentigerous processes, and the round sacral dia- 
pophyses (slightly expanded in all other groups). 
The osteological features are unlike those found 
in Batrachyla, Eupsophus, Hylorina, Telmatobius, 
or Thoropa but are like those seen in advanced 
telmatobiine genera (Eleutherodactylus and allied 
genera ) . Inclusion of the species of the quixensis 
group in Eupsophus greatly increases the hetero- 
geneity of the genus. In foot structure, the species 
of the quixensis group are distinctive from the 
other groups included in Eupsophus — no other 
groups have the large conical supernumerary 
plantar tubercles; the only argument for inclusion 



of the quixensis group in Eupsophus rests on the 
simple digital tips (i.e., no pads) and ignores the 
other osteological data. The oldest generic name 
for the quixensis group is Ischnocnema Reinhardt 
and Liitkcn. 1X62 (type-species Leiuperus verru- 
cosus Reinhardt and Llitken). Leiuperus verru- 
cosus Reinhardt and Liitken is an older specific 
name for Eupsophus verrucosus Miranda-Ribeiro, 
1937, for which Gorham (1966) proposed a re- 
placement name, Eupsophus versus (see systematic 
account). 

I (Lynch, 1971 ) have previously argued that the 
genus Eupsophus (in the restricted sense) should 
be used for the species of the nodusus and roseus 
groups as well as part of the peruanus group. The 
species of the nodusus and roseus groups and 
juninensis (peruanus group) have narrowly sepa- 
rated occipital condyles (convex atlas of Gallardo. 
1965), a quadratojugal bone, slightly dilated sacral 
diapophyses, knobbed terminal phalanges, sepa- 
rated nasal bones, and a frontoparietal fontanelle. 
The heterogeniety within and between these groups 
involves the secondary sex characteristics (pectoral 
plates of spines in males of the nodosus group but 
not in the roseus group or in juninensis) and ex- 
ternal evidence of the ear (tympanic annulus and 
columella lost in juninensis and monticola, present 
hut concealed beneath the skin in illotus and 
nodosus, and present and visible externally in 
roseus and vertebral is) . On the basis of the con- 
cealed ear, basal webbing of the toe, presence of 
pectoral plates of spines in reproductive males, 
and the enlargement of the forearms in reproduc- 
tive males, Gallardo (1970) proposed generic 
recognition of the nodosus group as Alsodes. He 
included a new species (A. gargola), some species 
usually included in Telmatobius (montanus, prae- 
hasalticus, and reverberii), as well as the three 
nodosus group species (illotus, monticola, and 
nodosus) in the genus Abodes. Barrio (1970) 
followed Gallardo in recognizing Alsodes but did 
not agree with the inclusion of Telmatobius prae- 
basalticus and T. reverberii. Gallardo's evidence 
for removing montanus from Telmatobius and 
placing it near illotus, monticola, and nodosus is 
compelling and appears to be the most correct 
disposition of montanus thus proposed. However, 
generic recognition of the nodosus group should 
be postponed because ( 1 ) the significance of ear 
reduction (or loss) is over-emphasized and does 
not consider the ear loss in juninensis, (2) while 
the pectoral plates of spines of the nodosus group 
are distinctive, the character state is duplicated to 
a lesser degree in some Telmatobius (aemaricus, 



1972 



SOUTH AMERICAN LEPTODAl TYUD FROGS 



hauthali, and jelskii), (3) the enlarged forearms 
in reproductive males of the nodosus group arc 
no more distinctive than the condition seen in 
some Leptodactylus, and (4) the proper generic 
name for Gallardo's gemis is Hammatodactylus, 
not Alsodes, if Fitzinger's (184.3) work is older 
than Bell's (1843) as has been generally assumed, 
The arguments in favor of recognizing the 
nodosus group as generically distinct from the 
roseus group depend in part on the lack of infor- 
mation concerning the species from the central 
plateau of Peru {peruanus group). Complete os- 
leological data are available for only two Peruvian 
species {juninensis and wettsteini). One of these 
(juninensis) has the narrrowly separated occipital 
condyles (convex atlas of Gallardo) like the 
Argentine-Chilean species (nodosus and roseus 
groups), whereas the other (wettsteini) has widely 
separated occipital condyles. Lynch (1968a, 1971) 
used Niceforonia for the Andean leptodactylids 
with widely separated occipital condyles, separated 
nasal bones, lacking a frontoparietal fonlanelle, 
and having knobbed terminal phalanges. Stereo- 
radiographs of paratypes of Eupsophus wettsteini 
reveal that osteologically this species is unlike 
Eupsophus and belongs to Niceforonia. Allocation 
of the other two species (peruanus and sylvestris) 
is not possible at present because data are lacking. 
The types of sylvestris are lost (Peters, 1873a) 
and no specimens matching the original descrip- 
tion were found in Tschudi's Peruvian collections. 
Its inclusion in Eupsophus cannot be justified and 
in view of Peters' (1873a) comments sylvestris 
should be considered a nomen dubiuin. Peters' 
(1873b) Phrynopus peruanus may be a Nice- 
foronia species; determination of its generic posi- 
tion must be postponed pending the availability 
of osteological material. 

SYSTEMATIC ACCOUNTS 

The following is a summary of the generic dis- 
positions of the species included in Eupsophus by 
Gorham (1966) as well as other species referred 
to the genus by other authors but not included in 
Gorham's list. Each generic account includes a 
generic synonymy, definition, content, and when 
appropriate, remarks. 

Batrachyla Bell, 1843 

Batrachyla Bell. 1843:43 [Type-species by monotypy. 
Batrachyla leptopus Bell, 1843]. 
Definition. — see definition of taeniatus group. 



Content. Hircc species /; antariandlco Bai 
1967, /(. leptopus Bell, 1843, and B. tacnlato (Oirard), 
1854. Litorla glandulosa Bell, 1843 
earlier name foi Batrachyla taenlata (sec I 

Remarks. — Lynch (1971) pointed OUl tha 
lenger's (1882) and Myers' (1962) combination 
ol Batrachyla with Eleutherodactylus 



Eupsophus Pitzinger, 1843 

Eupsophus Fitzinger, 1X43:31 [Type-species by orig 
inal designation. Cyslignatlms roseus Dumeril and 
Bibron, 1841], 
Hammatodactylus Fitzinger, 1843:31 |'l ype-specics by 
original designation. Cystignathus nodosus Dumeril 
and Bibron, 1841]. 
Borborocoetes Bell, 1843:34 [Type-species by sub- 
sequent designation (Lynch, 1971), Borborocoetes 
grayi Bell, 1843. Preoccupied by Borboroi 
Schoenherr, 1842 ( Insecta: Coleoptera) ]. 
Alsodes Bell, 1843:41 [Type-species by monotypy, 

Alsodes monticola Bell, 1843]. 
Cacotus Gunlher. 1868:482 [Type-species by mono- 
typy, Cacotus maculatus Giinther, lXf>X|. 
7 'Phrynopus Peters. 1873:416 |Typc-specics by mono- 
typy, Phrynopus peruanus Peters, 1873]. 
Borborocoetea Strand. 1928:55 [Replacement name 
for Borborocoetes Bell. 1843 (preoccupied); there- 
fore taking the same type-species, Borborocoetes 
grayi Bell], 

Definition. — larvae aquatic, not flattened or atten- 
uate, labial papillae incomplete anteriorly: eggs large 
and unpigmented: prevomerine dentigerous processes 
situated between or slightly posterior to choanae or 
prevomerine dentigerous processes absent: tympana 
visible externally, concealed beneath skin, or lost: 
fingers and toes not bearing terminal pads; males 
with nuptial asperities on thumb and sometimes on 
chest; toes lacking webbing or having basal web: two 
metatarsal tubercles, inner much larger than outer: 
plantar surface lacking distinct supernumerary tu- 
bercles; quadratojugal bone present: nasal bones 
widely separated: frontoparietal fontanelle present: 
occipital condyles narrowly separated (convex atlas): 
terminal phalanges knobbed. 

Content. — Eight species, E. gargolus (Gallardo ». 
1970. new combination. E. illotus (Barbour). 1922. 
E. juninensis (Shreve), 1932. E. montanus (Lataste>. 
1902. new combination. E. monticola (Bell). 1843. 
E. nodosus (Dumeril and Bibron). 1 841. E. roseus 
( Dumeril and Bibron). 1841. and E. vertebralis 
Grandison. 1961. Two other taxa may belong here 
but incomplete data preclude generic assignment: 
Cystiganthus sylvestris Tschudi. 1845. and Phrynopus 
peruanus Peters. 1873. 

Remarks. — Gallardo's (1970) recognition of 
A Isodes has not been followed here for the reasons 
cited in the discussion. Before the nodosus group 



in ill n\ ski nil l<\ < iLlFORNIA ACADEMY OF SCIENCES 



VOLUME 7 1 




Figure 3. Dorsum, hand, and side of head of the holotype of Leuipenis verrucosus (ZMK 
1180). {=Ischnocnema verrucosa). 



is placed in a separate genus the relationships of 
the Patagonian Telmatobius must be more fully 
documented. 



lschnocnema Reinhardt and Liitken, 1862 

Isclinocnema Reinhardt and Liitken, 1862:239 [Type- 
species by monotypy, Leuipenis verrucosus Rein- 
hardt and Liitken, 1862]. 

Oreobates Jimenez de la Espada, 1872:87 [Type- 
species by monotypy, Oreobates quixensis Jimenez 
de la Espada, 1872]. 
Definition. — see definition of quixensis group. 



Content. — Two species, /. quixensis and /. verru- 
cosa; both species have acquired synonyms (see re- 
marks). 

Remarks. — The species of the genus bear con- 
siderable resemblance to some of the members of 
the genus Eleutherodactylus of southeastern Brasil 
(binotatus, guentheri, and octavioi). The terminal 
phalanges are knobbed in lschnocnema and the 
tips of the digits lack expansions and the terminal 
transverse groove. In all Eleutherodactylus (in- 
cluding those of southeastern Brasil), there is a 
terminal transverse groove at each digital tip al- 
though there may be no digital expansion and the 



I<)72 



SOUTH .\Mi:i<icAN LEPTODACTYLID FROGS 



terminal phalanges of some lingers knobbed. Ai 
present I am unwilling to place the two species of 
Ischnocnema in the genus Eleutherodactylus, al- 
though the possibility cannot be eliminated thai 
the two species represent a single morphological 
divergence from the binotatus group of Eleuthero- 
dactylus. 

Reinhardt and Lutken (1862) named ;i single 
specimen of a frog from Brasil as Leuiperus ver- 
rucosus in allusion to its warty skin (Fig. 3). In 
an appendix to their paper, the authors proposed 
the generic name Ischnocnema. I hey slated that 
they did so on the advice of Wilhelm Peters, who 
did not consider verrucosus to be a true Leuiperus. 
All subsequent authors, including Boulenger, Nie- 
clen, and Gorham, failed to notice this generic 
name. Apparently, the next author to examine 
the holotypc of Leuiperus verrucosus was Cochran 
(1955), who, while retaining it in the genus 
Pleurodema (following Parker), noted that it bore 
considerable resemblance to the Eleutherodactylus 
guentheri group and little to Physalaemus or 
Pleurodema. Boulenger (1882) had placed the 
species in Paludicola and was uncritically followed 
byNieden (1923). Parker ( 1927), without exam- 
ining the unique holotypc, placed it in Pleuro- 
dema. Much of the synonymy of the frog, as thus 
far presented, has resulted from its transferal 
within a complex of then poorly understood 
genera. The basis for the use of such names as 
Leuiperus, Paludicola, Physalaemus, and Pleuro- 
dema was not clarified until Parker (1927) pointed 
out the osteological bases for the recognition of 
Physalaemus and Pleurodema and the invalidity 
of Paludicola and Leuiperus. 

The only synonyms of Leuiperus verrucosus of 
which I am aware are Eupsophus verrucosus 
Miranda-Ribeiro. 1937, and Eupsophus versus 
Gorham, 1966. Bokermann (1966) suggested that 
this species is an Eleutherodactylus but {in litt.) 
agrees that it is identical with Reinhardt and Li'it- 
ken's specimen; Bokermann has directly compared 
the two holotypes. 

Examination of the holotype of Leuiperus ver- 
rucosus conclusively demonstrates that the species 
is not a leptodactyline (since it lacks a bony style 
in the sternum) and therefore neither a Physalae- 
mus nor a Pleurodema, both of which have prom- 
inent bony styles in their sterna. The absence of 
the terminal transverse groove at the digital tip 
and the knobbed rather than T-shaped terminal 
phalanges do not permit its association with 
Eleutherodactylus. 

Gorham (1966), believing Miranda-Ribeiro's 



name to be valid and com i tl 
ically, proposed a replacement nai ophui 

versus, since verrucosus is preoccupied in the 
genus Eupsophus b\ Borborococles 
I'hilippi, 1902 ( Eupsophu - 1 1 

and Bibron, 1841). Since it now appears thai the 
Brasilian species is valid, a redescription ol the 
type is presented below. Apparently only two 
specimens have been collected, each being the 
holotypc of specific epithets ol verruct 

Ischnocnema verrucosa (Reinhardt and I utken), 

1862 

Leuiperus verrucosus Reinhardt and Lutken, 1862:171 
[Holotype: Zoologiskc Museum Kj0benhavn 
penhagen) 1 1 so. from near Juix de F6ra, Mina 
Gerais, Brasil], 
Ischnocnema verrucosa: Reinhardt and Lutken, 1X62: 

239. 
Paludicola verrucosa: Boulenger. 1882:171: Neidcn, 

1923:507. 
Pleurodema verrucosa: Parker, 1927:476; Cochran. 

1955:355: Gorham. 1966:158. 
Eupsophus verrucosus Miranda-Ribeiro. 1937:67. Fig. 
1 [Holotype: Museu Nacional (Rio dc Janeiro). 
from Rio Mutum, Coluna, Espirito Santo, Brasil. 
not examined]. 
Eleutherodactylus verrucosus [verrucosus of Miranda- 
Ribeiro, 1937. not Reinhardt and I.iitken. 1862] 
(non Hylodes verrucosus Jimenez dc la Espada. 
1875): Bokermann. 1966:40. 
Eupsophus versus Gorham. 1966:120 [Replacement 
name for Eupsophus verrucosus Miranda-Ribeiro. 
1937; preoccupied in Eupsophus by Borborococles 
verrucosus Philippi. 1902 ( = Eupsophus nodosus 
Dumeril and Bibron. 1841)]. 

Redescription of holotype. — (Fig. 3). Snout sub- 
acuminate in dorsal view, rounded in lateral profile: 
lips not flared; canthus rostralis sharp, curved; loreal 
region concave; nostrils directed laterally, much closer 
to tip of snout than to eye; width of eyelid slightly 
greater than interorbital distance: interorbital region 
lacking bony ridges: tympanum small, higher than 
long, partially concealed, its diameter about 20 r <: that 
of eye; maxilla and premaxilla toothed: prevomerine 
dentigerous processes absent but the tissue of the pre- 
vomerine area badly damaged suggesting that denti- 
gerous processes probably were present originally 
(Miranda-Ribeiro (1937:67) recorded them present 
in his specimen]: choanae large, oval, situated lat- 
erally, completely visible when roof of mouth is 
viewed from directly above; tongue large, ovoid, pos- 
terior one-third free, not notched. 

Skin of head, dorsum, flanks, and upper surfaces of 
limbs coarsely tuberculate: skin of venter and region 
about anus areolate; discoidal fold absent: fingers 
long and slender (Fig. 3) with prominent subarticular 



1(1 



BULLETIN SOUTHERN < ILIF0RN1A ACADEMY OF SCIENCES 



VOLUME 7 1 



tubercles; three palmar tubercles, outer smallest; super- 
numerary tubercles present on palm, few in number; 
tips of fingers not expanded and lacking any trace of 
terminal transverse grooves; tarsus lacking folds or 
tubercles on inner and outer edges; metatarsal tuber- 
cles large, subequal in size, inner oval, outer rounded, 
neither compressed; plantar supernumerary tubercles 
large, numerous; subarticular tubercles and lips of 
toes like those of hand. 

Color. — Creamy-brown above spotted with darker 
brown; labial bars present, broad; venter dirty cream, 
chin with scattered pale cream spots; limbs barred 
creamy-brown and dark brown; posterior surface of 
thigh uniform bleached cream; dark brown anal patch. 

Measurements in mm. — Snout-vent length 22.1; 
tibia 12.0; head width 9.3: head length 8.5; tympanum 
length 0.65; eye length 3.2: interorbital distance 2.3: 
eyelid width 2.3 + . 

Variation. — The specimen described and figured 
by Miranda-Ribeiro (1937) had more clearly de- 
fined markings dorsally and the light spots on the 
throat and flanks were better defined. These dif- 
ferences probably best reflect the length of time 
the holotype of Leuiperus verrucosus has been pre- 
served. Miranda-Ribeiro reported the tympanum 
to be 40% the diameter of the eye in his speci- 
men, whereas the tympanum/eye value in the hol- 
otype of L. verrucosus is 0.203. This difference 
may reflect differences in measuring techniques 
and/or variation. An index of variation is obtained 
by examining the same statistic in the only con- 
gener, Ischnocnema quixensis; in 24 adult and 
subadult specimens (sexes pooled) the values have 
a range of 0.48-0.66 and a mean of 0.589. 

Ischnocnema quixensis has at least three syno- 
nyms. Hylodes philippi Jimenez de la Espada, 
1875, and H. verrucosus Jimenez de la Espada, 
1875, are considered synonyms (Lynch and 
Schwartz, 1972) and Leptodactylus tuberculosus 
Anderson, 1945, is here added to the synonymy. 
W. Ronald Heyer and I have examined the holo- 
type. The absence of a bony style in the sternum 
requires removal of tuberculosus from Leptodac- 
tylus. The presence of conical plantar super- 
numerary tubercles, two large metatarsal tubercles, 
smooth skin of the venter, tuberculate skin of the 
dorsum, long first finger, and absence of discoidal 
folds and absence of digital expansion characterize 
Ischnocnema: the size, color pattern, and large 
and prominent tympanum characterize /. quixensis. 



Thoropa Cope, 1865 

Thoropa Cope, 1865:113 [Type-species by monotypy, 
Cystignathus missiessii Eydoux and Souleyet, 18411. 



Definition. — see definition of miliaris group. 

Content. — Three species, T. Ititzi Cochran, 1938, 
T. miliaris (Spi\). 1824, I . petropolitana (Wandol- 
leck). 1907. 

Remarks. — Excellent accounts of the species of 
this genus arc available in the works of Bokcr- 
mann ( 1 965 ) and Cochran ( 1 955 ) . Ololygon Fitz- 
inger is frequently cited as a synonym, whereas 
in fact the generic name Ololygon was proposed 
in 1S43 for Hyla strigilata Spix and is therefore a 
synonym of Hyla. 

The species included in Eupsophus by Gotham 
( 1966) but not allocated to Batrachyla, Eupsophus. 
Ischnocnema, or Thoropa here are listed below 
with their taxonomic disposition. 

Borhorocoetes bolitoglossus Werner, 1897, is 
presently considered a synonym of Craspedoglossa 
sanctaccatherinae (Bokermann, 1966). 

Borhorocoetes columhianus Werner, 1 899, is 
presently included in the genus Niceforonia 
(Lynch, 1968a). 

Cystignathus sylvestris Tschudi, 1845, is so 
poorly known (Peters, 1873a) that it should be 
considered a nomen dubium. 

Eupsophus wettsteini Parker, 1932, is here con- 
sidered a species of Niceforonia. 

Phrynopus peruanus Peters, 1873, cannot be 
assigned to genus with the available data. It is 
either a species of Eupsophus or of the generic 
group now called Niceforonia; that decision must 
be postponed pending acquisition of more material. 

LITERATURE CITED 

Barrio, A. 1970. Insuelophrynus acarpicus.un nuevo 
leptodactilido firmisternio sudamericano (Am- 
phibia. Anura). Physis, 30:331-341. 

Bell. T. 1843. Reptiles. In C. Darwin [ed.] The 
Zoology of the Voyage of H.M.S. Beagle under 
the command of Captain Fitzroy, London, pt. 5, 
51pp. 

Bogart. J. P. 1970. Systematic problems in the am- 
phibian family Leptodactylidae (Anura) as in- 
dicated by karyotypic analysis. Cytogenetics, 9: 
369-383. 

Bokermann, W. C. A. 1965. Notas sobre as especies 
de Thoropa Fitzinger (Amphibia. Leptodactyli- 
dae). Anais da Academia Brasileira de Ciencias, 

37:525-537. 

. 1966. Lista anotada das localidades tipo 

de anfibios brasileiros. Servico documentacao. 
Sao Paulo, RUSP, 181 pp. 



1972 



SOUTH AMERICAN LEPTODAl i > i.m FROGS 



II 



Boulenger, (J. A. IXX2. Catalogue of the Batrachia 
Salientia s. Ecaudata in Hie collections of the 
British Museum. 2rul ed., London, 503 pp. 

Cei, J. M. I960. A survey on the leptodactylid 
frogs, genus Eupsophus in Chile. Breviora, I IX: 
1-13. 

— . 1962. Batracios de Chile. Santiago, I2X 



+ CVIII pp. 

Cochran, D. M. 1955. Frogs of southeastern Brazil. 
Bull. U.S. Nat. Mus., (222). 423 pp. 



— . 196X1.. i he idcntil "i a < h 
Abodes mom/rein Bell, :in<l I 
genus A hod en (Amphibia Leptodactylidae J 
Herpetologii a, 24:255-257. 



— . 1971 . I rolulionai y relation 
and zoogeography of Icplodactyloid fro 

Publ. Mus. Nat. Hist., Univ. Kan 
pp. 



Lynch, J. D., and A. Schwartz. 197.: I 

disposition ol some 19th century leptodactylid 
frog names. J. Herpetol., 5: I'll 114. 



Cope, E. D. 1865. Sketch of the primary groups 
of Batrachia Salientia. Nat. Hist. Rev., 5:97- 
120. 



Miranda-Rihciro. A. 1937. Alguns halrachios novos 
das colleccoes do Mnseu Nacional. O Campo, 

Rio de Janeiro, May 1937:66 69. 



Fitzinger, L. 1X43. Systema Reptilium. Vindobonae. 

106 pp. 

Gallardo, J. M. 1962. Los generos Eupsophus y 
Batrachyla (Anura, l.eptodactylidae) en la Ar- 
gentina, y la verdadera identidad de Paludicola 
illota Barbour. Rev. Mus. Argentino Cs. Nats. 
Cienc. Zool., 8( 10) : 1 13-122. 

. 1965. A proposito de los Leptodactylidae 

(Amphibia, Anura). Papeis Avulsos. 17:77-87. 



. 1970. A proposito de los Telmatobiinae 

(Anura, Leptodactylidae) Patagonicos. Neo- 
tropica, 16:73-85. 

Gorham, S. W. 1966. Liste der rezenten Amphibien 
und Reptilien/Ascaphidae. Leiopelmatidae, Pip- 
idae, Discoglossidae, Pelobatidae, Leptodactyl- 
idae, Rhinophrynidae. Das Tierreich. 85:1-222. 

Grandison, A. G. C. 1961. Chilean species of the 
genus Eupsophus (Anura: Leptodactylidae). Bull. 
British Mus. (Nat. Hist.), Zoology. 8:111-148. 

Giinther, A. C. L. G. 1868. First account of species 
of tailless batrachians added to the collection of 
the British Museum. Proc. Zool. Soc. London. 
1868:478-490. 

Jimenez de la Espada, D. M. 1872. Nuevos ba- 
tracios americanos. Anales de la Sociedad Es- 
panola de Historia Natural, 1:85-88. 

Lynch. J. D. 1968a. Systematic status of some 
Andean leptodactylid frogs with a description of 
a new species of Eleutherodactylus. Herpeto- 
logica, 24:289-300. 



Myers, G. S. 1962. The American Leptodactylid 
frog genera Eleutherodactylus, Hylodes (=Elosia) 
and Caudiverbera {■=Calyptocephalus). Copcia, 
1962:195-202. 



Nieden, F. 
584. 



1923. Anura. I. Das Tierreich, 46:1- 



Parker, H. W. 1927. A revision of the frogs of the 
genera Pseudopaludicola, Physalaemus, and 
Pleurodema. Ann. Mag. Nat. Hist.. (9)20:450- 
478. 

Peters, W. 1873a. tjber die von Dr. J. J. v. Tschudi 
beschriebenen Batrachier aus Peru. Monatsber. 
d. konig. Preuss. Akad. Wiss.. Berlin. 1873:622 
624. 

. 1873b. Zwei Giftschlangen aus Afrika und 

uber neue oder weniger bekannte Galtungen und 
Arten von Batrachiern. Monatsber. d. konig. 
Preuss. Akad. Wiss., Berlin. 1873:411-418. 

Reinhardt. J., and C. F. Lutken. 1862. Bidrag til 
Kundskab om Brasiliens Patter og Krybdyr. 
Medd. Naturh. Foren. Kjobenhavn. 3:143-242. 

Rivero. J. A. 1964. Salientios (Amphibia I en la 
coleccion de la Sociedad de Ciencias Naturales La 
Salle de Venezuela. Caribbean J. Sci.. 4:297- 
305. 

Strand. E. 192S. Miscellanea nomenclatoria zoo- 
logtca et palaeontologia. Arkiv fur Naturge- 
schichte. 92A:30-75. 



Accepted for publication October 21. 1 



FIRST RECORD OF THE BLACK SCABBARDFISH, APHANOPUS CARBO, 

FROM THE PACIFIC OCEAN WITH NOTES ON OTHER 

CAL1FORNIAN TRICHIURID FISHES 

John E. Fitch and Daniel W. Gotshall 1 



Aiistract: Based upon throe specimens collected in the vicinity of Fort Bragg, California, 
Aphanopus curbn I. owe. 1839, is reported from the Pacific Ocean for the first time. Data and 
keys are presented for distinguishing A. carbo and four other species of Californian trichiurids 
(Assurger anzac, Diplospinus mullistriatus, Lepidopus xantusi, and Trichiurus nitens). Key 
characters include shape of caudal fin, shape of dorsal fin. presence or absence of ventral fins, 
body depth into standard length, and counts of dorsal rays, anal rays, vertebrae, and pyloric 
caeca. Otoliths (sagittae) of the five species are illustrated and differentiated on the basis of 
shape, proportions, and size. 



Although the black scabbardfish is relatively com- 
mon in the eastern north Atlantic (Tucker, 1956; 
Sutro, 1967; Forster, 1971), only three individuals 
have been noted from the western north Atlantic 
(Templeman and Squires, 1963), and until now 
only one specimen, from the Gulf of Aden (Nor- 
man, 1939), has been recorded outside the At- 
lantic Ocean. 

On March 18, 1969, a black scabbardfish 
(Aphanopus carbo) was taken in an otter trawl 
being fished from the vessel Pearl Harbor in 270 
fathoms (494 m) of water off Bodega Bay, Cali- 
fornia. This fish, a female, 1038 mm total length 
(TL) and weighing 1950 g (Table 1 ), was turned 
over to the California Department of Fish and 
Game by Kenny Marsh, skipper of the Pearl 
Harbor. Eight months later a smaller individual 
was trawled in 300 fathoms (549 m) west of 
Fort Bragg by the vessel Northern Light, skippered 
by Angelo Urbani. A third specimen was netted 
on February 16, 1971, by Tom Webster, skipper 
of the otter trawler Havana, while fishing in 360 
to 390 fathoms (658-713 m) approximately 20 
miles (32 km) west of Bodega Bay. As nearly as 
we can determine, these are the first and only 
captures of A. carbo in the Pacific Ocean; all 
three specimens have been placed in the fish col- 
lections of the Natural History Museum of Los 
Angeles County. 

Tucker (1956) examined the holotypes of sev- 
eral nominal species of Aphanopus and deter- 
mined that none could be distinguished from the 
earliest described species, A. carbo. Templeman 
and Squires (1963) compared morphometries for 
their three western Atlantic specimens with 
Tucker's data, and aside from having a slightly 



greater number of vertebrae (100-103 vs. 98-99) 
in the western Atlantic fish, there were no obvious 
differences. The three fish from off California 
have an even higher number of vertebrae (105) and 
inasmuch as it does not appear that Tucker (1956) 
checked vertebrae on more than three individuals, 
we asked G. E. Maul (Museu Municipal do 
Funchal. Madeira) to send us vertebral counts 
on additional Madeiran material. He was able to 
obtain counts on 16 individuals as follows [num- 
ber of individuals (precaudal + caudal including 
hypural = total)]; 1 (40+59=99), 2(41+57=98), 
3(41+58=99), 3(42+57=99), 3(42+58=100), 
1(42+59=101), 1(42+60=102), 1(43+57=100), 
1(43+58=101). Total vertebrae (105) for the 
Californian Aphanopus (Table 1) thus exceed the 
highest Atlantic count (103). but one of the 
western Atlantic specimens (Templeman and 
Squires, 1963) matched our 46 precaudal verte- 
brae, and three of the fish examined by Maul 
(data above) had caudal vertebral counts (59 
and 60) that were equal to or exceeded those of 
the Californian specimens. For this reason, we 
believe that the apparent difference in total verte- 
brae merely reflects the effect of colder tem- 
peratures on the developing egg, and would be 
resolved upon examination of additional material 
from higher latitudes in the north Atlantic (e.g., 
Newfoundland, Greenland, Iceland). This hy- 
pothesis assumes of course that the eggs of 
Aphanopus are pelagic and develop in the upper 
surface layers. 



1 California Department of Fish and Game, 
Golden Shore, Long Beach, California 90802. 



350 



12 



1972 



NliW CUTLASSFISH RECORDS FOR < M u ohnia 






Table I. Selected morphometry and mciistic data on three i ol Californian n 

(standard lengths in mm, all other measurements expressed as thousandth ol <i 



Measurements 




Aphanopus carbo 




i 


mrgoi anzm 


' 


Standard length 


955 


965 


990 


15.34 


1612 


1X0 






Head length 


1X7 


196 


190 


074 


0X0 


IX.' 


2 1 1 23 1 




Fleshy orbil 


034 


030 


033 


OKI 


010 


>>■<■ 







Bony interorbital 


031 


032 


029 


OOX 


/ 


02; 






Snout length 


075 


0X0 


074 


02X 


032 


060 069 




071 0X2 


Maxillary length 


0X9 


0X9 


OXX 


024 


024 


05X 072 


072 079 


074 0X4 


Pectoral length 


090 


0K7 


093 


031 


- 


0X1 101) 


091 09 




Snout to D, insert 


155 


165 


I5X 


050 


056 


126 117 


145-157 




Snout to P, insert 


196 


200 


190 


0X5 


072 


1X6 206 


203 223 




Snout to A, insert 


576 


5X0 


586 


391 


422 


529 570 


552 560 




Snout to V, insert 


200 


195 


1X4 


MIX 


II 1 


2I6-22X 


226 247 




Depth 


090 


099 


098 


0.37 


039 


075 096 


097-114 




P to D, insert 


069 


072 


069 


044 


045 


0X2 090 


089 102 


089 102 


V to IX insert 


092 


093 


088 


070 


06X 


117-135 


131-143 


131-146 


Counts: 


















Dorsal rays XI. Ill, 55 


XI. 111,54 


XLII.55 


1 16 


117 




7X-XI 




Anal rays 


[1,47 


il.47 


il,48 


il.74 


il,76 




WM-A1 




Pectoral rays 


12 


12 


12 


12 


12 




12 




Gill rakers 


- 


- 


- 


4 + 9 


4 + 9 


6-8+10-13= 


6-21 


Vertebrae 


- 


46 + 59=105 46+59=105 


- 


43 + 84=127 


32-36+48-55= 


84-87 


Pyloric caeca 


7 


7 


7 


- 


13 




16 




No. of specimens 


1 


1 


1 


1 


1 


10 


3 


12 



* Measurements of L. xantusl are presented by size groups to reveal allometric growth, if present. 



Based upon catch data for 15 specimens of 
Aphanopus carbo taken on longline gear in the 
Biscay area, Forster (1971) suggested that its 
distribution is determined by temperature rather 
than depth. Twelve of these 1 5 specimens were 
studied by Bone (1971 ) and were the basis for his 
report on the behavior, histology, and anatomy 
of A. carbo. It was his view that ambient light is 
more important in determining the distribution of 
A. carbo than either temperature or depth. 

In an excellent narrative of the commercial 
fishery off the island of Madeira, Sutro (1967) 
presents a few details regarding the life history of 
the "black espada." He points out that at one time 
a 4-foot (122 cm) long black scabbardfish would 
sell for five cents (about one cent a pound), but 
it is now Madeira's most popular fish, often selling 
for 30 cents a pound. A typical fishing boat carries 
a crew of four, and utilizes four lines. Each line 
averages about 4600 feet ( 1 402 m ) long and is 
fished vertically with the hooks spaced 10 feet (3 
m) apart on the lowermost 1400 feet (427 m) 
of the line. Although the number of fish cap- 
tured in one night varies considerably, a good 
catch would be about 100 black scabbardfish. 



OTHER CALIFORNIAN TRICHIl RIDS 
Assurger anzac (Alexander. 1916) 

Radovich (1961) noted a razorback scabbard- 
fish caught on May 3. 1958, off the north point 
of North Coronado Island, and mentioned that the 
species previously had been taken off California 
in 1951. The specimen from North Coronado 
Island (Scripps Institution of Oceanography collec- 
tion number SIO 58-75) apparently had been 
snagged, as it bears hook marks near the origin of 
the anal fin: this fish measured 1035 mm standard 
length (SL) and 1054 mm total length (TL). The 
specimen caught in 1951 measured 1640 mm TL 
and was taken on September 23. presumably on 
hook and line, in the surf at Playa del Re\ . This 
fish is deposited in the collection of Scripps 
Institution of Oceanography. La Jolla (SIO 
58-80). 

Since Radovich's report, two additional speci- 
mens of Assurger anzac have been taken in Cali- 
fornian waters. One of these, a female measuring 
1534 mm SL. weighing 555 g (Table 1 >. w^ 
snagged on January 1. 1963. by two fishermen. 
Joe Branin and Guy R. Ludwick. approximately 



14 



in in n\ soi mi i<\ ( mroKM i ,ii,iiu:my of SCIENCES 



VOLUME 71 






Figure 1. Comparison of head and trunk region of 
three adult Californian trichiurids: Aphanopus carbo 
(top), Lepidopits xantusi (center), Assurger anzac 
(bottom). 



1% miles (2.4 km) off Laguna Beach. This speci- 
men was deposited in the fish collection at the 
University of California, Los Angeles. The most 
recently collected individual was picked up by 
Mike McGiffin as it was feebly floundering in the 
surf at Torrance Beach, Santa Monica Bay in 
1965. The entire posterior extremity of this fish 
was missing from a point just behind the anal fin 
insertion (possibly 55% of the length of the fish), 
apparently the result of a shark bite, which may 
also account for its being in the surf. The standard 
length of this fish may have been between 1700 
and 1 800 mm judged by its measurable length. 

Due to the rarity of Assurger anzac the follow- 
ing description is presented. The body color, when 
fresh, is an overall silver that is easily rubbed off. 
The dorsal fin membrane is black between the first 
and third fin spines as is the base of the membrane 
between spines three and four; the remainder of 



the dorsal fill membrane is unpigmented. There 
is one row of sharply pointed teeth along the 
lower jaw that are slightly curved inward and are 
about twice as strong as those in the upper jaw. 
There are one or two lanceolate teeth at the tip 
of the lower jaw that are two to three times as long 
as the other teeth. Two or three teeth, which are 
firmly fixed at the tip of each premaxillary, are 
similar in size to the large lanceolate teeth in the 
lower jaw. 

Each primary gill raker consists of one to three 
bristle-like spines on a slightly raised basal plate. 
Secondary rakers, consisting of fewer and shorter 
bristles on smaller plates, are interspersed with the 
primary rakers. Secondary and rudimentary rakers 
extend to the extremities of both the upper and 
lower limb of the first gill arch. 

At a level directly above the pectoral insertion, 
the lateral line lies approximately one-third of the 
distance between the dorsal and ventral profiles; 
it descends from there in a straight line, reaching 
the mid-side about two head lengths behind the 
pectoral, then runs parallel to the body axis 
(horizontally) until it terminates on the caudal 
peduncle. 

We did not differentiate spines and rays in the 
dorsal fin and counted vertebrae on only one speci- 
men (Table 1 ) . The first 63 to 65 anal rays are 
small, prostrate (lie parallel to the body) and em- 
bedded. 

In addition to the Californian specimens, two 
smaller individuals from the stomachs of Alepi- 
saurus were examined. These two trichiurids are 
deposited in the Scripps Institution of Oceanogra- 
phy collections; one measures 760 mm SL and 
was taken off the coast of Chile (lat. 26°04.9' S, 
long. 101°47.0' W), the other is a much smaller, 
badly mangled, fish from northeast of Midway 
Island. The Chilean specimen had 119 dorsal 
fin rays, but the elements in the anal fin could not 
be counted. 

The stomach of the fish from North Coronado 
Island contained a small squid and an anchovy, 
Engraulis mordax; whereas there were beaks from 
three cephalopods in the stomach of the Assurger 
from off Laguna Beach. An incomplete vertebral 
column from a small Pacific hake, Merluccius 
productus, was found in the stomach of the speci- 
men taken at Torrance Beach in 1965. 

Diplospinus multistriatus Maul, 1948 

In August, 1957, a commercial fisherman ob- 
tained a four-inch long trichiurid that had been 



I<)72 



NEW CUTLASSF1SH RECORDS H>l< < ALU OKNIA 








Figure 2. Left sagittae of Diplospinus multistriatus 
4.2 mm (lop) and Paradiplospinus gracilis 2.7 mm 
(bottom). Photos by Jack W. Schott. 



regurgitated by an albacore which he caught 250 
miles southwest of Morro Bay (lat. 33° 10' N, long. 
125° 10' W). The spinous and soft-rayed portions 
of the dorsal fin were completely separated; the 
soft-rays were much more crowded than the spines. 
This trichiurid appeared to be conspecific with 
Lepidopus gracilis described by Brauer (1906) 
from the west coast of Africa. However, Bussing 
(1965) assigned Brauer's fish to Paradiplospinus 
and presented data to distinguish it from Diplo- 
spinus multistriatus. Judged by Bussing's data, 
our specimen should be assigned to D. multi- 
striatus, but various counts made on 12 additional 
specimens found in stomachs of Alepisaurus in- 
dicate the differences between P. gracilis and D. 
multistriatus are not as distinct as Bussing shows. 
Although the otoliths offer additional proof that 
P. gracilis and D. multistriatus are not conspecific, 
the observed differences in the sagittae of these 
two taxa appear to be of specific rather than 
generic magnitude (Fig. 2). 

Andriashev (1960), in establishing Paradiplo- 
spinus, distinguished it from Diplospinus on the 
basis of a number of substantial characters which 
do not permit both species to be assigned to the 
same genus. Unfortunately, he did not have com- 
parative material of Diplospinus, and he relied 
entirely upon published data. Two of his sub- 
stantial characters were not discussed in the publi- 
cations he consulted, and the variation he at- 
tributed to Diplospinus in these instances may not 
have been based upon fact. Some of his other 



i harai ters should have been dc< i n 

rather than "substantial In fa I il j ■;■ 

Bussing (1965) had reservations rcgardii 
validity ol Paradiplospinus in hi I the 

discovery o) ventral fins in small , 
/'. gracilis docs nol invalidate thi 
diplospinus, although several characters suggi 
similarity with Diplospinus." To the characters 

noticed by Bussing we can now arid the Otoliths 
(sagittae). 

Ahlstrom (1971) records larval Diplospinus 
multistriatus from numerous eastern Pacific lo- 
calities, both north and south ni the equator, and 
recently informed one ol us (Fitch) thai he has 
seen fair numbers ol larval D, multistriatus ofl 
southern California: some within 200 miles (320 
km) of the coast. During several tuna longlinc 
cruises in 1954 and 1955. the California Depart- 
ment of Fish and Game research vessel N. B. 
Scofield caught a number of Alepisaurus fcrox 
that had eaten juvenile Diplospinus. In all, 65 
specimens, ranging from just over two to about 
four inches (50-100 mm) in length, were re- 
covered from the stomachs of Alepisaurus. These 
fish were taken primarily within the area bounded 
by lat. 24° and 35° N, and long. I 24 and 1 37 \V. 
The stomachs of two small individuals of Alepi- 
saurus which had been eaten by a larger Alepi- 
saurus caught on longline gear (lat. 24 03' N, 
long. 137° 12' W) on January 27, 1954, contained 
53 of the 65 juvenile Diplospinus multistriatus 
examined by us. Twelve of these two- to three-inch 
(50 to 75 mm) long fish were cleared and stained 
and yielded the following counts (number of in- 
dividuals noted in parentheses for each count i : 
dorsal spines, 30(1 ). 32(1), 33(7). 34(3 I: dorsal 
rays, 35(1). 36(2). 37(3), 38(1), 4(1(1): anal 
rays, 26(1). 28(1), 29(1), 30(1). 33(1); pre- 
caudal vertebrae. 24(4), 25(6). 27(2): total verte- 
brae, 59(1), 60(7), 61(4). There are two anal 
spines, one pelvic spine with no rays, 1 2 pectoral 
rays, and 9 + 8 principal caudal rays. 

Diplospinus multistriatus previously has not 
been reported from off California, and does not 
appear in the checklist of deep-water fishes given 
in Fitch and Lavenberg (1968). Because of their 
extremely small sizes and generally poor condition, 
none of the specimens we examined from the east- 
ern north Pacific was photographed. 

Lepidopus xantusi Goode and Bean. 1S95 

Tucker (1956) reported that the earliest de- 
scriptions of Lepidopus xantusi are misleading. 



16 



in III il\ SOI FHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



and confusing, yel there is no doubl but what the 
name was applied to a specimen from off Cape 
San Lucas, Baja California. Because of erroneous 
reports, Tucker (1956) stated thai "a new name 
will have to be found for it by the first worker 
able to re-describe it from material." However, 
Jordan ami Starks (1907) described fully and 
illustrated a 2 '--foot (76 cm) long L. xantusi 
collected by C. F. Holder at Santa Catalina Island, 
California. In view of this accurate description, 
in addition to the fact that only one species of 
Lepidopus is known to exist in the eastern north 
Pacific, we prefer to retain the name L. xantusi 
for this species. 

Lepidopus xantusi has been called "black scab- 
bardfish" in California (Fitch and Lavenberg, 
1968) and "scabbardfish" by the American Fish- 
eries Society (Bailey et al., 1970). Inasmuch as 
Aphanopus carbo has historically been known as 
the black scabbardfish. we propose the vernacular 
name "Pacific scabbardfish" for L. xantusi. This 
will circumvent confusion and name changing 
should the closely related Atlantic species L. 
caudatus ever be found to occur within the area 
covered by the American Fisheries Society. 

The distribution of L. xantusi extends from 
Eureka, California, to Mazatlan, Mexico (Fitch 
and Lavenberg, 1968). The silver-colored juve- 
niles up to 12 inches (30 cm) long apparently live 
near the surface and often stray into relatively 
shallow water near shore. The adults, however, 
have always been captured in deep water. Those 
caught on hook and line have been taken from 
350 to 1650 feet (107 to 503 m) beneath the 
surface, usually at or near the bottom. The largest 
measured individual was 900 mm TL (Table 1 ), 
although unpublished records from the California 
Department of Fish and Game show an individual 
102 cm TL that was caught "in 275 fathoms south 
of Santa Catalina Island on November 16, 1938." 
Maximum observed weight was 1120 g for a ripe 
male 835 mm TL. 

The otoliths of numerous individuals were ex- 
amined for evidence of age and from this in- 
vestigation it appears that a specimen 375-400 
mm TL, weighing 70 to 85 g, is approximately 4-5 
years old. Individuals measuring 830-888 mm TL 
(770-1070 g) have 15-18 winter annuli on their 
otoliths. 

Although most captures of L. xantusi have 
involved but one fish, in April 1962, a commercial 
fisherman caught 1% tons (1360 kg) of these fish 
in an otter trawl being towed on the bottom in 140 
fathoms (256 m) off Newport Beach, California. 



On another occasion, a sport fisherman caught five 
large adults while fishing near the bottom in very 
deep water (± 300 m) near San Diego. Ap- 
parently. L. xantusi is a schooling fish at times. 

During the past 25 years, we have examined 
more than 50 individuals ranging in standard 
length from 47 to 850 mm, however, we have 
included counts and morphometries for only 25 
of these in this report (Table 1 ). These specimens 
cover the entire range of variation which we ob- 
served in L. xantusi for the sizes indicated. 

Two badly digested specimens of Lepidopus, 
removed from the stomach of a yellowfin tuna 
(T/utnnus albacares) captured 28 miles (45 km) 
northwest of Zorritos, Peru (lat. 03°22' S, long. 
81°01' VV), probably represent an undescribed 
species and are not included in this account. These 
specimens (deposited in the UCLA fish collec- 
tion) have slightly higher dorsal and anal fin 
counts than any L. xantusi we have examined. 
Furthermore, there are other observable differ- 
ences in spite of their poor condition. A satis- 
factory description of this Peruvian Lepidopus 
must await the collection of less wretched material. 

Trichiurus nitens Garman, 1899 

The Pacific cutlassfish has been recorded from 
a number of localities between San Pedro, Cali- 
fornia, and Paita, Peru, but they were taken in 
commercial quantities only during the early 
1930's. Typically, one or two individuals are 
caught within a period of hours or days, then 
several years pass before the species again appears. 
However, during the early 1930's, small lampara 
boats fishing in relatively shallow water near San 
Pedro, Long Beach, and Seal Beach were capturing 
from a few pounds to a ton or more almost daily 
for short periods each spring. 

The largest specimen that we have observed was 
a 44-inch (112 cm) long female netted in March, 
1962, that weighed approximately 3% pounds 
(1460 g). This fish was 7 years old, judged by 
the growth rings on its otoliths, and was approach- 
ing the onset of spawning. A 33-inch (83 cm) 
male taken a month earlier was 4 years old and 
appeared to have viable milt in its testes. 

Hubbs and Hubbs (1941) pointed out numerous 
characters by which T. nitens can be distinguished 
from the Atlantic T. lepturus. However, Tucker 
(1956), felt that "the satisfactory establishment 
of T. nitens requires, not comparison with Atlantic 
material, but demonstration of a non-cline dis- 
continuity across the Pacific." Since we lack ma- 



I'J72 



NI'AV CUTLASSFISH RECORDS FOR < \l II <>KNIA 









Figure 3. Left sagittae of four Californian tri- 
chiurids: a, Aphanopus carbo 7.5 mm: b, Trichiurus 
nitens 5.0 mm; c, Lepidopus xantusi 5.5 mm; d, As- 
surger anzac 3.0 mm. Photos by Jack W. Schott. 



terial from oceanic areas of the Pacific, we are 
in no position to demonstrate "non-cline discon- 
tinuity," but the material we have examined does 
appear to add strength to the argument for re- 
taining T. nitens. Nine specimens, from various 
localities between southern California and Panama, 
yielded the following fin ray and vertebral counts 
(corresponding counts for T. lepturus taken from 
Tucker [1956] are enclosed in parentheses): dor- 
sal rays, 118-128 (126-137); anal rays. 95-97 
(105-108); vertebrae, 148-155 (162-168). There 
are 36-38 dorsal fin rays to a vertical above the 
anus, and 34-36 of the vertebrae are precaudals. 



OTOL1 I lis 01 ( Al II ORN1 

I RI< llll RIDS 

Trichiurids frequently arc led upon b> ecl 
pinnipeds, tunas, bill-fishes, birds, and squid 
because conventional recognition ch; apidly 

disappear with digestion, otolitl 
identifiable fish remains to be found in the 
stomachs ol these predators ["he otoliths | 
tae) of the live Californian trichiurids (1 
and 3) illustrate close family relationships, es- 
pecially in (he configuration ol the sulcus (i.e., 
groove on the inner lace), yel they are also tcadik 
distinguishable from each olher. I he sagitta "I 
Aphanopus lacks an antirostrum (anterior projec- 
tion above the sulcus) and is broadly rounded 
posteriorly. Sagittae ol Diplospinus also lack an 
antirostrum, but have a squared off notch at the 
posterodorsal corner. In Trichiurus, the greatest 
otolith height occurs near its midlength, and the 
outer lace is deeply concave dorso-vcntralh. In 
both Lepidopus and Assurger, the otolith is 
highest in front of the midpoint, but in Assurger 
the tapered posterior end turns abruptly down- 
ward. Even more spectacular is the relative length 
of the sagitta of these two genera: 3.0 mm in a 
1534 mm SL Assurger and 5.5 mm in a 785 mm 
SL Lepidopus. 



KEY TO CALIFORNIAN TRICHIURIDS 

1. Caudal fin hair-like: ventral fins absent: lateral 

line well below mid-side on posterior three- 
fourths of body - - — Trichiurus nitens 

Caudal fin mackerel-like: ventral fins present, 
sometimes as spine-like scales or rudimentary 
and embedded: lateral line at mid-side on pos- 
terior three-fourths of body _ 

2. Spinous and soft-rayed portions of dorsal fin 

completely separated, or separated by a dis- 
tinct notch — 3 

Spinous and soft-rayed portions of dorsal fin 
continuous, indistinguishable without micro- 
scopic examination 

3. Dorsal spines 30 to 34; dorsal soft-rays 35 to 40: 

vertebrae 59 to 61 Diplospinus mullistriatus 

Dorsal spines 42 to 43 (38 to 41 for Atlantic i: 
dorsal soft-rays 54 to 55 (53 to 56 for 
Atlantic): vertebrae 105 Aphanopus carbo 

4. Body 25 times as long as deep: dorsal rays 116 

to 119; vertebrae 12" Assurger . 

Body less than 15 times as long as deep: dorsal 
ravs 78 to SI: vertebrae S4 to IT . 



Lepidopus xaniusi 



IS 



Bl III ll\ SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 7 'I 



ACKNOWLEDGMENTS 

Mam individuals were helpful at one stage or an- 
other of our investigations concerning the family 
I richiuridae. Over 30 different fishermen con- 
tributed directly by bringing in specimens that 
the\ caught, and although most of these are not 
acknowledged by name in the text we are deeply 
indebted to them. Others who were especially 
helpful were E. H. Ahlstrom. National Marine 
Fisheries Service, La Jolla; Charles R. Clothier 
(deceased): James E. Craddock, Woods Hole 
Occanographic Institution; W. I. Follett, Cali- 
fornia Academy of Sciences: Shelly Johnson and 
Richard McGinnis, University of Southern Cali- 
fornia: Robert J. Lavenberg and Armando Solis, 
Natural History Museum of Los Angeles County; 
G. E. Maul. Municipal Museum, Funchal, 
Madeira: Richard Rosenblatt and Robert L. Wis- 
ner, Scripps Institution of Oceanography; and Jack 
W. Schott, California Department of Fish and 
Game. Research on otoliths has been supported 
by grants (GB-1244 and GB-6490) to Fitch from 
the National Science Foundation. 



Wis*. F.rgeh. Dcutsehen 
Jena, 15(1): 1-432. 



Tiefsee Exp. Valdivia, 



Bussing. W. A. 1965. Studies of the midwater fishes 

of the Peru-Chile Trench. Amcr. Geophysical 
Union. Antarctic Res. Ser., 5:185-227. 

Fitch, J. E., and R. J. Lavenberg. 1 968. Deep-water 
teleostean fishes of California. Univ. California 
Press, Berkeley, 1 55 pp. 

Forster. G. R. 1971. Line-fishing on the continental 
slope III. Mid-water fishing with vertical lines. 
J. Mar. Biol. Assoc. U.K., 51(l):73-77. 

Hubbs, C. L... and C. Hubbs. 1941. Pacific cutlass- 
fish, Trichiurus nitens Garman. California Fish 
and Game, 27(l):29-30. 

Jordan. D. S., and E. C. Starks. 1907. Notes on 
fishes from the island of Santa Catalina, southern 
California. Proc. U. S. Nat. Mus., 32(1510): 
67-77. 

Norman, J. R. 1939. Fishes. Sci. Rept., John Mur- 
ray Exped., 7(1): 1-1 16. 



LITERATURE CITED 

Ahlstrom. E. H. 1971. Kinds and abundance of fish 
larvae in the eastern Tropical Pacific, based on 
collections made on EASTROPAC I. U. S. Bur. 
Commerce, Fish. Bull., 69(l):3-77. 

Andriashev, P. A. 1960. Families of fishes new to 
the Antarctic. I. Paradiplospinus antarcticus, gen. 
et sp. n. (Pisces, Trichiuridae). Zoologicheskii 
Zhurnal, Acad. Sci. USSR, 39(2) :244-249. 

Bailey, R. M., J. E. Fitch, E. S. Herald, E. A. Lachner. 
C. C. Lindsey, C. R. Robins, and W. B. Scott. 
1970. A list of common and scientific names 
of fishes from the United States and Canada. 
Amer. Fish. Soc, Spec. Publ., 6:1-149. 

Bone, Q. 1971. On the scabbardfish Aplumopus 
carbo. J. Mar. Biol. Assoc. U.K., 51( 1 ) :219-225. 



Radovich. J. 1961. Relationships of some marine 
organisms of the northeast Pacific to water tem- 
peratures particularly during 1957 through 1959. 
California Dept. Fish and Game, Fish Bull., 112: 
1-62. 



Sutro, A. G. 1967. 
20(4):27-30. 



Black espada. Pacific Discovery, 



Templeman, W., and H. J. Squires. 1963. Three 
records of the black scabbard-fish, Aphanopus 
carbo Lowe, from the Canadian region of the 
Western Atlantic. J. Fish. Res. Board Canada, 
20(2):273-278. 

Tucker, D. W. 1956. Studies on the trichiuroid 
fishes — 3. A preliminary revision of the family 
Trichiuridae. Bull. British Mus. (Nat. Hist.), 
Zool., 4(3):73-130. 



Brauer, A. 1906. Die Tiefsee Fische, I, Syst. Teil., Accepted for publication September 24, 1971. 



ORGAN WEIGHTS OP NON-CAPTIVE porpoise is// iella spi 

Wll.UAM F. PERRIN 1 AND HdwARU L. Roberi 



Abstract: Weights of heart, lungs, liver, kidney, and spleen ol 68 eastern Pacifii spotted 
porpoise (Stenella graffmani) and 14 eastern Pacific spinnei porpoise is. cf. S. longln 
arc presented and related to total body weight, Values of Huxley's growth coefficient " arc 

presented, Adull female spoiled porpoise have on the average smaller spleens than adllll mall 

a hypothesis of suppression of antibody production in pregnant females is suggested. 



information on organ weights and on other aspects 
of the anatomy of non-captive cetaceans is valuable 
for 3 reasons: 1 ) to establish norms that may be 
used in evaluation of the post-mortem condition 
of animals that die in captivity, 2) to further un- 
derstanding of the physiology and overall biology 
of these animals, which is so radically different 
from that of terrestrial mammals, and 3) to pro- 
vide comparative data for use in the systematics of 
the group. 

Odontocete cetaceans for which various organ 
weights have previously been reported include 
Phocoena phocoena, Tursiops truncatus, Lageno- 
rhynchus acutus, Delphinus delphis, Grampus 
griseus, Delphinapterus latcas, Physeter catodon 
(Slijper, 1959); Globicephala melaena (Cowan, 
1966); and, very recently, Inia geoffrensis (Pilleri 
and Gihr, 1969b), and Stenella styx [= S. coertt- 
leoalba {fide Mitchell, 1970)] (Gihr and Pilleri, 
1969a). Layne (1965) reported weights of heart, 
lungs, liver, and kidneys for 1 specimen of Ste- 
nella longirostris. We report here weights of sev- 
eral organs for 68 eastern Pacific spotted porpoise, 
Stenella graffmani (Lonnberg, 1934), and 14 east- 
ern Pacific spinner porpoise, Stenella cf. S. longi- 
rostris (Gray, 1828). Specimens of the spotted 
Stenella plagiodon and 5. attenuata, both very 



similar to the eastern Pacific spoiled porpoise. 
have been or are on exhibit or held capti 
research purposes at various institutions on the 
southeast coast of North America and in Hawaii, 
respectively. Specimens of Stenella roseiventris, 
very similar to the eastern Pacific spinner porpoise, 
are currently held by aquaria in Hawaii. 

METHODS 

The specimens examined were all from the eastern 
tropical Pacific (Table 1). They were part of a 
large series of animals gathered for extensive 
studies of external morphometries, osteology. 
parasites, and reproduction, which will be reported 
elsewhere. The porpoise were captured and killed 
accidentally during the course of commercial tuna 
seining operations, such as described by Perrin, 
1969. The animals were placed in refrigerated 
holds along with the tuna and upon return to port 
were kept in freezers at -18° C until they could 
be dissected. Before dissection, the specimens 



1 National Marine Fisheries Service. Southwest 
Fisheries Center. La Jolla. California 92037. 

-Animal Hospital of Fountain Valley, 18542 Brook- 
hurst Street, Fountain Valley. California ^2"08. 



Table 1. Collection data. 





Locality 










Number 


of specimens 




Date 


Stenella 
graffmani 


Stenella cf. S. 
longirostris 


Number of 

schools 


9 Apr. 1968 

12 Apr. 1968 

26-27 Mar. 1969 

28 Apr. 1969 

5 Apr. 1969 


12°51' N/93-18' W 
7° 11' N/90-32' W 
7°47'-S° N/106°36'- 
7°40'-8° N/107" W 
8° N/109°45' W 


106' 


50' 


W 


Total 


36 

29 

3 

68 


4 
2 
4 
2 

: 

14 


1 
1 

7 

1 



19 



:o 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



I viii i 2. Total length, total weight, and organ weights of 68 specimens of Stenella graffmani and 14 

specimens of s. cf. S. longirostris. 



Specimen I field 
no.) U. S. N;K. 

Mils. No. 

(skeleton) 


Sex 


rotal 
length 

( cm i 


Total 
weigh) 

(kg) 


Heart 

(gm) 


1. uncs 

(pin) 


Liver 
(gm) 


Kidneys 




1 .ii 
(gm) 


Right 
(gm) 


Spleen 
(gm) 








Stenella graffmani 










CV 259/395276 


M 


168 


44.6 


_ 


_ 


1000 


( 40S 


) 


43 


CV 260/395277 


F 


175 


58.2 


- 


- 


1767 


326 


312 


48 


CV 261/395278 


F 


166 


46.8 


- 


- 


1283 


182 


205 


44 


CV 262/395279 


F 


165 


43.2 


- 


- 


1065 


193 


182 


44 


CV 263/395328 


M 


176 


49.5 


- 


- 


1259 


235 


251 


51 


CV 264/395329 


M 


172 


50.0 


- 


- 


1137 


247 


306 


53 


CV 265/395330 


M 


195 


76.7 


- 


- 


1727 


326 


353 


54 


CV 266/395331 


M 


168 


47.2 


- 


- 


1011 


178 


199 


32 


CV 267/395332 


M 


179 


61.7 


- 


- 


1398 


246 


234 


39 


CV 268/395333t 


F 


190 


76.3 


- 


- 


1480 


250 


240 


46 


CV 269/395334 


F 


184 


59.9 


- 


- 


1476 


217 


209 


31 


CV 270/395336t 


F 


182 


70.8 


- 


- 


1304 


298 


319 


36 


CV 271/395337 


F 


182 


58.2 


- 


- 


1467 


245 


251 


35 


CV 272/395338t 


F 


185 


72.7 


- 


- 


1394 


242 


- 


16 


CV 273/395339 


M 


163 


47.2 


- 


- 


1000 


220 


233 


85 


CV 274/395385 


M 


178 


53.2 


253 


- 


1153 


232 


208 


24 


CV 275/395386 


M 


157 


42.2 


230 


- 


1087 


200 


247 


84 


CV 276/395387 


M 


158 


50.0 


211 


- 


1274 


216 


201 


45 


CV 277/395388 


F 


182 


59.9 


291 


- 


1396 


284 


300 


38 


CV 278/395389 


M 


163 


44.1 


207 


- 


1237 


190 


199 


62 


CV 279/395390 


M 


218 


84.0 


311 


- 


1892 


477 


485 


36 


CV 280/395391 


M 


203 


82.7 


311 


- 


1961 


406 


429 


46 


CV 281/395392 


F 


174 


43.2 


214 


- 


1280 


220 


227 


60 


CV 282/395393 


F 


174 


44.1 


179 


- 


1052 


207 


199 


32 


CV 283/395394 


M 


202 


79.9 


308 


- 


2195 


484 


480 


50 


CV 284/3953951 


F 


193 


71.3 


250 


- 


1371 


309 


290 


30 


CV 286/395397 


M 


165 


47.7 


199 


- 


1112 


190 


233 


100 


CV 287/395417 


M 


173 


53.6 


261 


- 


1325 


260 


296 


60 


CV 289/395594 


M 


177 


48.6 


226 


1576 


1219 


227 


243 


44 


CV 290/395595 


F 


163 


40.4 


184 


1270 


924 


166 


183 


27 


CV 291/395596 


F 


193 


63.6 


293 


- 


1847 


340 


323 


21 


CV 292/395597 


M 


189 


75.8 


289 


2275 


1704 


354 


380 


50 


CV 293/395598 


M 


200 


62.2 


274 


2129 


1281 


291 


316 


65 


CV 295/395526 


F 


139 


29.1 


152 


994 


528 


91 


104 


19 


CV 296/395527 


M 


129 


23.6 


- 


907 


517 


86 


91 


33 


CV 297/395528 


F 


143 


26.8 


129 


824 


470 


88 


100 


30 


CV 298/395529 


M 


154 


40.4 


210 


1323 


1124 


155 


189 


47 


CV 300/395530 


M 


144 


31.8 


169 


1019 


647 


114 


121 


38 


CV 303/395532 


M 


143 


29.5 


141 


1075 


532 


112 


113 


34 


CV 304/395535 


M 


86 


6.8 


52 


382 


135 


35 


35 


6 


CV 305/395458 


F 


78 


5.0 


40 


185 


109 


28 


27 


1 


CV 306/395459 


M 


91 


7.2 


54 


264 


122 


40 


- 


6 


CV 307/395460 


F 


80 


5.5 


43 


254 


122 


32 


35 


6 


CV 308/395461 


F 


79 


5.5 


38 


232 


101 


27 


30 


5 


CV 309/395466 


F 


188 


56.7 


236 


2078 


1798 


281 


330 


26 


CV 310/395463 


M 


180 


58.2 


264 


1642 


1360 


220 


241 


48 


CV 311/395464 


M 


200 


69.9 


286 


2090 


1869 


389 


406 


47 


CV 312/395465 


F 


164 


44.1 


211 


1317 


1087 


233 


198 


42 


CV 313/395462 


M 


160 


41.8 


182 


1291 


999 


177 


179 


62 


CV 314/395467 


F 


184 


55.4 


284 


1789 


1600 


284 


290 


20 


CV 315/395468 


M 


162 


43.6 


227 


1354 


1139 


193 


190 


35 



1972 



Olid AN WEIGHTS OF PORPOISES 
Table 2. (Contlmu d I 






Specimen (field 
no.J/U, S. Nat. 

Mus. No. 

(skeleton) 


.Sex 


lol.il 

length 
(cm) 


Total 
weight 

(km 


|l 


I ung 
(gin J 










liver 


i ell 






CV 316/395603 


M 


168 


45.0 


21 1 


1457 


1013 


187 






CV 317/395604 


M 


138 


25.9 


147 


887 


614 


121 


1 17 




CV 318/395605 


F 


144 


29.6 ' - 


160 


1047 


6X4 


115 


1 11 




CV 319/395606 




168 


46.8 


204 


1681 


974 


200 


222 


72 


CV 320/395607 


M 


167 


46.3 


210 


1672 


1132 


205 


217 


76 


CV 321/395608 


M 


153 


34.6 


175 


1 KM 


774 


130 


140 


34 


CV 322/395609 




142 


26.3 


1 19 


853 


528 


102 


1 10 


26 


CV 323/395610 


M 


162 


46.8 


201 


1700 


1076 


191 


190 


33 


CV 324/395611 




136 


25.4 


126 


959 


523 


82 


94 


11 


CV 325/395612 


M 


135 


29.5 


153 


1090 


608 


94 


108 


43 


CV 326/395613' 


M 


86 


6.6 


42 


246 


104 


33 


31 


3 


CV 327/395614" 


F 


81 


5.9 


44 


245 


84 


29 


30 


3 


CV 328/395615 : ' 


M 


74 


3.4 


29 


156 


62 


21 


28 


2 


CV 329/395616' 




81 


5.7 


- 


- 


- 


26 


26 


4 


PQ 01/395617 


M 


198 


- 


301 


- 


1234 


231 


247 


42 


PQ 02/395618 


M 


170 


- 


170 


- 


657 


161 


149 


27 


PQ 04/395410 


F 


168 


Stenella 


219 
cf. .V. / 


ongirostris 


995 


197 


176 


27 


CV 285/395369 


F 


177 


44.5 


230 


_ 


936 


150 


158 


15 


CV 288/395593 


F 


173 


43.5 


220 


- 


832 


( 300 


) 


23 


CV 294/395526 


F 


149 


33.6 


204 


11 12 


855 


116 


137 


43 


CV 299/395531 


M 


1 19 


18.6 


114 


631 


368 


58 


69 


14 


CV 301/395533 


F 


140 


26.3 


135 


812 


542 


73 


88 


16 


CV 302/395534 


M 


1 18 


18.1 


97 


599 


367 


60 


63 


29 


PQ 03/395409 


F 


157 


39.5 


225 


- 


832 


128 


128 


22 


PQ 05/395411 


F 


168 


46.7 


196 


- 


971 


147 


164 


13 


PQ 06/395412t 


F 


173 


51.3 


245 


- 


776 


125 


136 


21 


PQ 07/395413 


F 


176 


S2 ^ 


191 


- 


871 


145 


144 


- 


PQ 08/395414 


M 


173 


50.3 


240 


- 


972 


152 


147 


20 


PQ 09/395599 


M 


173 


59.0 


272 


- 


997 


192 


201 


31 


PQ 10/395600 


F 


109 


18.1 


97 


- 


323 


55 


58 


22 


PQ 11/395601 


F 


105 


13.2 


72 


- 


282 


46 


51 


10 


t Individual was pregnant. 
> Fetus from CV 270/395336. 
-' Fetus from CV 268/395333. 








' Fetus from CV 
1 Fetus from CV 


272/395338. 
284 595395. 









were weighed on platform scales, accurate to the 
nearest pound. The weights were converted to 
kilogram equivalents lor use here. Some dehydra- 
tion may have taken place during the storage 
periods, which ranged from 5 days to 14 months. 
When each specimen was dissected, the heart, 
lungs, liver, kidneys, and spleen were excised, 
placed in plastic bags, and refrozen. The sample 
size is not the same for all the organs, as these 
data were gathered incidentally to the primary 
purposes of the dissections, and some organs were 
not removed when time was short for other pro- 
cedures. Later, the organs were thawed and 
weighed on a beam balance, accurate to the nearest 



gram. Four of the female specimens of 5. graf- 
fmani collected on April 9. 1958 were gravid 
with near-term fetuses, and the organs of these 
fetuses were included in the sample. Degree of 
sexual development was determined by histological 
examination of gonads. Complete skeletons of all 
the specimens are in the U. S. National Museum. 
Washington. D. C. and museum numbers are 
given in table 2. 

The taxonomy of the spotted and spinner 
porpoises is confused (Perrin and Hunter. 1 
and the use here of Stenella graffmani and 5. 
longirostris is provisional pending the results of 
studies underwav by Perrin and others. 



22 



HI III IIX SOUTHERN CALIFORNIA ACADEMY OF SCIENl I \ 



VOLUME 71 





300 




HEART 










• •• 

.■ ■ ■ • • 


















• 


(f) 


















E 
















o • 


o 


250 


- 












+ 


ka 
















■ 


CP 












°0 


O 




1— 










o 


°& 


o 




C£ 


200 












0o 




< 










D 








LU 










o 




D 






X 








D 










Li_ 


150 








o 










O 








a a 










h- 








a 










X 


100 


_ 












o IMMATURE MALE 


o 
















° IMMATURE FEMALE 


UJ 
















• SEXUALLY MATURE MALE 


£ 


50 


_ cP 

4*- 












■ SEXUALLY MATURE FEMALE 
+ PREGNANT FEMALE 
















A MALE FETUS 




o 


A 


i i 


1 


L 




1 


-0- FEMALE FETUS 

1 1 1 1 



10 20 30 40 50 60 70 

WEIGHT OF PORPOISE (kilograms) 



80 



90 



Figure 1. Stenella graffmani. Weight of heart relative to total body weight for 29 males and 
19 females. 



RESULTS 

The data are presented in table 2. Scatter plots 
of organ weight on total weight are presented 
in figures 1-6 and 8-10. Where sample sizes were 
adequately large, the data were fitted to the equa- 
tion Log Y = a + b Log X by the least squares 
method, where Y = organ weight in grams, X = 
total weight in grams, and b — Huxley's (1932) 
"growth coefficient" n . For the cases where cor- 
relation was statistically significant (at P = 0.05), 
the estimates of a and b are presented in table 3. 
Data for the pregnant females were not included 
in the regression analyses. 



Heart 

S. graffmani (Fig. 1 ). The hearts of 11 sexually 
mature adults (ranging from 180-218 cm total 
length and from 55.5-84.0 kg) weighed from 236- 
311 gm and from 0.37-0.51% (average: 0.43%) 
of total body weight. As is evident in figure 1, 
the relative weight of the heart decreases during 
development; the average relative weight for 28 
calves and sexually immature subadults (ranging 
from 126-178 cm total length and from 25.4-53.6 
kg total weight) was 0.48% (range: 0.41-0.57%) 
and for 8 infants and fetuses was 0.75% (range: 
0.63-0.86%). 



7972 



oi«;an WEIGHTS <>!■ PORPOISES 






3DU 










_ 300 
(/> 

E 
o 

3f 250 


HEART 






• 


2 200 

X 




D 


■ 
■ 

■ 


■ 


° 150 










X 
CD 
jjj 100 


D 
O 

e 






o IMMATURE MALE 


£ 


D 






a IMMATURE FEMALE 

• SEXUALLY MATURE MALE 


50 
n 


1 1 




i 


■ SEXUALLY MATURE FEMALE 
+ PREGNANT FEMALE 

-J 1 1 1 



10 20 30 40 50 60 70 80 

WEIGHT OF PORPOISE (kilograms) 

Figure 2. Stenella cf. 5. longirostris. Weight of heart relative to total body weight for 4 males 
and 10 females. 



90 



Table 3. Log-log regression analyses of relationships 

between organ weights and total body weight in 

S. grafjmani and S. cf. 5. longirostris. 











Sample 




a' 


b 1 


r 1 


size 




s. 


grafjmani 






Heart 


-1.25 


0.769 


0.948 


47 


Lungs 


-0.92 


0.S80 


0.990 


35 


Liver 


-2. IS 


1.114 


0.922 


59 


Kidneys 


-1.87 


0.965 


0.977 


57 


Spleen: 










males 


-3.07 


1.010 


0.874 


36 


females 


-3.40 


1.057 


0.885 


24 




S. cf. 


5. longirostris 




Heart 


-1.48 


0.820 


0.962 


13 


Lungs 


-1.25 


0.947 


0.991 


4 


Liver 


0.01 


0.608 


0.536 


13 


Kidneys 


-1.91 


0.940 


0.986 


13 



1 Values of a, b. and correlation coefficient r in linear 
regression equation Log Y = a 4- b Log X. where Y = organ 
weight and X = total body weight, in gms. 



S. longirostris (Fig. 2). The hearts of 6 sexually 
mature adults (ranging from 168-177 cm total 
length and from 43.5-59.0 kg total weight) 
weighed from 191-272 gm and from 0.37-0.52% 
(average: 0.46 r r ) of total body weight. The 
data are insufficient to determine whether relative 
weight of the heart decreases during development 
in this species. 

Lungs 

S. graffmani (Fig. 3L The combined weight of 
both lungs of 6 adults (from 180-200 cm total 
length and from 55.4-75.8 kg total weight ) ranged 
from 1642-2275 gm and from 2.82-3.66% (aver- 
age: 3.19 c r) of total weight. 

S. longirostris (Fig. 3). The lungs of 4 individ- 
uals, all calves and subadults (from 118-1 
total length and from 18.1-33.6 kg total weight) 
weighed from 599-1 1 12 gm and from 3.0 C 
(average: 3.28%) of total weight. 



24 



Bl III ll\ SOI llllli\ CAI '.I FORM. 4 ACADEMY OF SCIENCES 



VOLUME 71 



2600 



2400 



2200 



2000 



CD 

LU 



1200 



1000 



800 



600 



400 



200 



LUNGS 
S. q raff mani 

o IMMATURE MALE 

o IMMATURE FEMALE 

• SEXUALLY MATURE MALE 

■ SEXUALLY MATURE FEMALE 

-^MALE FETUS 

-(J-FEMALE FETUS 







S. cf. S. lonqirostris 




1800 


A IMMATURE MALE 
IMMATURE FEMALE 


CO 

E 

a 

i_ 






1600 


- 


in 


1400 





-«- 






Oo 



D 



A 
A 



10 20 30 40 50 60 70 

WEIGHT OF PORPOISE (kilograms) 



80 



90 



Figure 3. Stenella graffmani. Weight of lungs relative to total body weight for 22 males and 
15 females. Stenella cf. S. longirosrris. Weight of lungs relative to total body weight for 2 
males and 2 females. 



1972 



<)I«IAN weights <>i porpoises 






2200 



2000 



1800 



1600 

£ 

o 1400 

s 

W 1200 



1000 



X 

& 800 



600- 



400 



200- 







• 
• 


LIVER 




• 


o IMMATURE MALE 


■ 


• 


a IMMATURE FEMALE 


■ 




• SEXUALLY MATURE MALE 


■ 




■ SEXUALLY MATURE FEMALE 




• 


+ PREGNANT FEMALE 






> MALE FETUS 


■ 




A FEMALE FETUS 








■" 


4 


- 


■ • 
• 


♦+ 


- 


o o 

° 8 

o 

o 

o 
o ° o o 


+ 




o <P % 
a 

a 




o 






o 













_ o o 






D 






1 1 1 


.1 ...J 1 


i 



» 10 20 30 40 50 60 70 80 

WEIGHT OF PORPOISE (kilograms) 

Figure 4. Stenella graffmani. Weight of liver in 36 males and 27 females. 



90 



Liver 

S. graffmani (Fig. 4). The livers of 16 adults 
(from 175-2 IS cm total length and from 55.4- 
84.0 kg total weight) weighed from 12S1-2195 
gm and from 2.06-3.179r (average: 2.53%) of 
total weight. Relative weight of the liver appears 



to increase early in development (average for 
fetuses and infants was 1.84%, range: 1.42 
2.22 r 'c ) and remain nearly constant thereafter. 

S. longirostris (Fig. 5 V The livers of 6 adults 
(168-177 cm total length and 43.5-59.0 kg total 
weight) weighed from 832-997 gm and from 1.67 
2.10^ (average: 1.90^) of total weight. Adults 



:o 



HI III ll\ Sorilll-RS CAI. NORMA ACADEMY OF SCIENCES 



VOLUME 71 



2200, 



2000 



1800 



1600 



CO 




E 




o 




k_ 




C7> 


1400 


QC 




UJ 




> 


1200 


_1 




Ll_ 




O 






1000 


h- 




X 




CD 




Ul 


800 


£ 





600 



400 



200 



LIVER 



o IMMATURE MALE 

o IMMATURE FEMALE 

• SEXUALLY MATURE MALE 

■ SEXUALLY MATURE FEMALE 

+ PREGNANT FEMALE 



d a> 
a 



10 20 30 40 50 60 70 

WEIGHT OF PORPOISE (kilograms) 



80 



90 



Figure 5. Stcnella cf. 5. longirostris. Weight of liver relative to total body weight for 4 
males and 10 females. 



of this species have smaller livers than subadults 
of S. graffmani; the livers of 21 calves and sub- 
adults of S. graffmani ranging from 43.2-53.6 kg 
total weight weighed from 974—1457 gm and 
from 2.08-3.00% (average: 2.42%) of total 
weight. 



Kidneys 

S. graffmani (Fig. 6). The relative weight of 
the kidneys drops from a high at birth to a low 
in calves and subadults and then increases again 
to a secondary high in adults. The kidneys of 8 



1972 



ORGAN win ,in s in I'oki'oim.s 






1000 



900 



800 



F 


700 


o 




k_ 




Di 




(/) 


600 


>- 




LU 




z 




Q 


500 


x: 





" 400 

h- 

X 

rr; 300 



200- 











< 


> « 


KIDNEYS 












o IMMATURE MALE 






% 




• 


D IMMATURE FEMALE 












• SEXUALLY MATURE MALE 












■ SEXUALLY MATURE FEMALE 








• 




+ PREGNANT FEMALE 












i A MALE FETUS 








Q 




■^FEMALE FETUS 




■ 

■ 








- 


O 


■ • 

■ 
■ 










6> 


■ 

• 




♦ 




- 


oo ° 


• 

o 

■ 








o 












o o 
o 












a4f 

1 1 i 


i 1 


1 


1 







100 



10 20 30 40 50 60 70 80 

WEIGHT OF PORPOISE (kilograms) 

Figure 6. Stenella graffmani. Combined weight of kidneys relative to total body weight for 
35 males and 27 females. 



90 



neonatals and near-term fetuses (74-86 cm total 
length and 3.4-6.8 kg total weight) ranged from 
49-70 gm (average: 59 gm) and from 0.91- 
1.44% (average: 1.09%) of total weight. The 
kidneys of 31 calves and subadults (129-178 cm 
total length and 23.6-53.2 kg total weight) ranged 



from 129-178 gm and from 0.67-1.1 1% (average: 
O.S4% ) of total weight, and those of 1 5 adults 
(175-2 IS cm total length and 55.4-84.0 kg total 
weight) ranged from 42o-96S gm and from 0.78- 
1.20% (average: 0.98%) of total weight. The 
animal is asymmetrical with respect to weight of 



:,s- 



HI I I I ll\ SOUTHERN CALIFORMA ACADEMY OE SCIEXCES 



VOLUME 71 



LEFT KIDNEY LARGER 
(14) 



RIGHT KIDNEY LARGER 
(43) 



15 10 5 5 10 15 

DEVIATION (per cent) 

Figure 7. Stenella graffmani. Frequency distribution 
of percent deviation of weight of largest kidney from 
average weight of both kidneys, for 59 individuals. 



kidneys (Fig. 7); the right kidney on the average 
is 2-4% heavier than would be expected were 
symmetry to prevail. 

S. longirostris (Fig. 8). The kidneys of 6 adults 
(168-177 cm total length and 43.5-59.0 kg total 
weight) weighed from 289-393 gm and from 
0.55-0.69% (average: 0.65%) of total weight. 
This species also exhibits dextral asymmetry: of 
12 sets of kidneys weighed, in 9 the right kidney 
was heavier, in 2 the left was heavier, and in 1 
set both kidneys weighed the same. 

Spleen 

S. graffmani (Fig. 9). We infer from the 
scatter plot of weight of spleen on total weight 
that the spleen increases in size both absolutely 
and relatively during development to a peak in 
subadulthood, then rapidly decreases in size, again 
both absolutely and relatively, to a relative low 
during adulthood. The spleens of 10 neonatals 
and near-term fetuses averaged 4 gm (0.07%) in 
weight, those of 11 calves (129-153 cm total 
length and 23.6-34.6 kg total weight) ranged 
from 11-43 gm (average: 31 gm) and from 
0.04-0.15% (average: 0.11%) of total weight, 



:\nA those of 25 subadults (154-178 cm total 
length and 40.4-53.6 kg total weight ) ranged 
from 24-100 gm (average: 53 gm) and from 
0.05-0.21% (average: 0.12%) of total weight— 
the variance was greatest for this group, but the 
spleens of 16 adults (175-218 cm total length 
and 55.4-84.0 kg total weight) weighed only 
20-65 gm (average: 41 gm) and from 0.03- 
0.10% (average: 0.06%) of total weight. There 
is also an apparent sexual dimorphism in weight 
of the spleen in adults: the spleens of 9 adult males 
weighed 36-65 gm (average: 48 gm) and 0.04- 
10% (average: 0.06%) of total weight, while 
those of 7 non-pregnant adult females weighed 
only 20-48 gm (average: 31 gm, the same as for 
the calves) and from 0.03-0.08% (average: 0.05%, 
less than for the infants and fetuses). The smallest 
adult spleen (16 gm or 0.02% of total weight) 
was possessed by a pregnant female (near-term). 
S. longirostris (Fig. 10). As is the case for the 
liver and the kidneys, the adults of this species 
possess smaller spleens than do subadults of com- 
parable size of S. graffmani. The spleens of 5 
adults weighed 13-31 gm (average: 30 gm) and 
from 0.03-0.05% (average: 0.04%) of total 
weight. The data conform to the pattern noted 
above for S. graffmani of decreased absolute 
weight of spleen in adulthood but are insufficient 
in number to warrant conclusions. 



DISCUSSION 

The value of the growth coefficient b, the ratio 
of the rate of increase of organ weight to the rate 
of increase of total body weight, for heart weight 
obtained here for S. graffmani (0.769) is lower 
than those obtained by Pilleri (1969) for S. 
coeruleoalba (1.014), Delphinus delphis (0.806), 
and Phocoena phocoena (1.055). That of S. 
longirostris (0.820) falls between those for D. 
delphis and S. coeruleoalba. The value of the 
growth coefficient for lung weight appears to be 
inversely related to maximum body weight. Of 
the 3 species of Stenella, the smallest, S. longiro- 
stris, has the highest coefficient (0.947), and the 
largest, S. coeruleoalba has the lowest (0.684) — 
Pilleri, 1969. The value of b for S. graffmani is 
intermediate (0.880). The value of b for liver 
weight for S. longirostris (0.608) is close to that 
obtained for S. coeruleoalba (0.684) by Pilleri, 
but that for S. graffmani (1.114) is considerably 



1972 



okcan WEIGHTS <>l \'()i(Voi\l.s 






1000 



900 



800 



% 700 

\— 

£ 600 
LU 

| 500 



400 



O 

h- 

X 

o 

W 300 



200 



100 



KIDNEYS 



o IMMATURE MA i 

a IMMATURE FEMALE 

• SEXUALLY MATURE MALE 

■ SEXUALLY MATURE FEMALE 

•♦■PREGNANT FEMALE 



10 20 30 40 50 60 70 

WEIGHT OF PORPOISE (kilograms) 



80 



90 



Figure S. Stciiella cf. S. longirostris. Weight of kidneys relative to total body weight for 4 
males and 9 females. 



higher. The values of b for kidney weight for 
both species dealt with here (0.965 and 0.940) 
are lower than those obtained by Pilleri for 5. 
coeruleoalba (0.977) and D. del phis (1.089). 



Pilleri's (1969) value of b for Phocoena phocoena 
(0.6S5) is very much lower than those for the 4 
delphinids (sensu stricto). 

The odontocete spleen has a relatively large 



to 

E 

o 

i_ 

an 



LU 
Ld 

_l 

Q_ 
CO 

u_ 
o 



X 
CD 



MU 














100 


SPLEEN 

o IMMATURE MALE 


o 










90 


d IMMATURE FEMALE 
• SEXUALLY MATURE MALE 
■ SEXUALLY MATURE FEMALE 
4- PREGNANT FEMALE 


o 
o 










80 


- >MALE FETUS 

■^FEMALE FETUS 


o 

D 










70 




O O 


• 








60 




D O 




o 




• 




50 


- 


o 






• 


> 




o 


o 

o 
d a o 
o 

D 


•a 


• 


+ 


• 


40 


D 





• 
■ 










O O 
O 


o 
o° 


■ 


+ 




• 


30 


— □ 
D 


D 


■ 

■ 
o 


+ 






20 


D 




■ 
■ 


+ 






10 
o 


a 

I" 


1 1 


1 


1 




1 



10 20 30 40 50 60 70 80 

WEIGHT OF PORPOISE (kilograms) 

Figure 9. Stcnclla graffmani. Weight of spleen relative to total weight for 37 males and 
28 females. 



90 



number of lymph nodules (white spleen pulp) 
(Pilleri, 1969). A major function of the lymphatic 
tissue of the spleen is the production of antibodies 
(Jubb and Kennedy, 1963). We offer here the 
tentative hypothesis that smaller relative spleen 
size for adult females of S. graffmani (Fig. 8) is 



tied functionally to suppression of antibody pro- 
duction during pregnancy to eliminate the pos- 
sibility of rejection of the fetus. Obviously his- 
tological investigation of splenic change during 
pregnancy will be necessary if this hypothesis is 
to be critically examined. 



IV72 



ORGAN WEIGHTS <>l PORPOISI : 






110 



100 



90 



_ 80 
id 

E 
o 

ct 70 



LU 

Ll) 60 



Q_ 
CO 



50 



| 40 
LxJ 

30- 



20- 



10- 



SPLEEN 



o IMMATURE MALE 

d IMMATURE FEMALE 

• SEXUALLY MATURE r/ 

■ SEXUALLY MATURE FEMALE 

-f PREGNANT FEMALE 



10 20 30 40 50 60 70 

WEIGHT OF PORPOISE (kilograms) 



80 



90 



Figure 10. Stenella cf. 5. longirostris. Weight of spleen relative to total weight for 4 males 
and 9 females. 



ACKNOWLEDGMENTS 

Specimens were collected through the cooperation of 
the owners, captains, and crews of M V CAROL 
VIRGINIA and M "V PACIFIC QUEEN. Craig J. 
Orange and Susumu Kato helped secure the specimens. 
Support was provided by the National Marine Fish- 



eries Service and the Inter-American Tropical Tuna 
Commission. 



LITERATURE CITED 
Cowan. D. F. 1966. Observations on the pilot 



32 



HI I II ll\ SOI ////.'K.N ( AI.IEORMA ACADEMY OE SCIENCES 



VOLUME 71 



whale Globicephala melaena: organ weight and Renin. W. 1969. Using porpoise to catch tuna. 
growth \nat. Rec, 155:623-628. Work! Fishing, 18(61:42-45. 



Gihr. M.. and G. Pilleri. 1969. On the anatomy and 
biometry of Stenella styx Gray and Delphinus 
delphis L. (Cetacea, Delphinidae) of the Western 
Mediterranean, hi Pilleri. G. |ed.|. Investigations 
on Cetacea. Vol. I. Bentelli A. G.. Berne. 219 pp. 



Pilleri. G.. and M. Gihr. 1969. Zur Anatomic und 
Pathologic von Inia geoffrensis De Blainvillc 
1X17 (Cetacea, Susuidae) aus dem Beni, Bolivien. 
In Pilleri, G. |cd.|. Investigations on Cetacea. 
Vol. I, Bentelli A. C... Berne, 219 pp. 



Jubb, K. V.. and P. C. Kennedy. 1963. Pathology 
of domestic animals. Academic Press. New York, 
2:613 pp. 



Slijpcr, E. I. 1959. Organ weights and symmetry 
problems in porpoises and seals. Arch. Neer. 
Zool. 13 (1st suppl.):97-113. 



Huxley. J. S. 1932. Problems of relative growth. 
Dial Press. New York. 276 pp. 



Accepted for publication November 16, 1971. 



THE EFFECTS OF LOWERED DISSOLVED OXYGEN CONCENTRATIONS AND 

SALINITY ON THE FREE AMINO ACID POOL OF THE POLYCHAETOUS 

ANNELID NEANTHES ARENACEODENTATA 

John L. Abati and Donald J. Reish 1 



Abstract: An inbred strain of the polychaetous annelid Neanthes arenaceodentata was 
subjected to reduced dissolved oxygen and salinity levels under laboratory conditions and 
effects of these treatments on the free amino acid pool studied. Changes in the relative propor- 
tions of free amino acids were observed after exposure to lowered dissolved oxygen concentra- 
tions. It is hypothesized that these relate to changes in total protein levels and conversion to 
energy production under anaerobic conditions. The total free amino acid concentration, 
expressed as total optical density of ninhydrin positive spots, decreased at lowered salinities 
along with alteration in the relative proportions of amino acids. These changes probably 
relate to osmoregulation by the worm under reduced salinities. 



The free amino acid pool is a major constituent 
of the metabolic machinery of an organism, and 
therefore gives a good measure of metabolic ac- 
tivity. It is also unique to each species and has 
been compared to a human fingerprint (Roberts 
and Lowe, 1954). The free amino acid pool has 
been used as a research tool in the study of diseases 
(Roberts and Frankel, 1949; Roberts and Borges, 
1955), in the study of speciation (Ball and Clark, 
1953; Micks, 1954, 1956; Micks and Gibson, 
1957; and Lewallen, 1957), and in measuring en- 
vironmental effects on organisms (Schafer, 1961, 
1963; Lynch and Wood, 1966; Stephens and 
Virkar, 1966; and Clark, 1968a). 

In his review of the free amino acid pool in 
invertebrates, Awapara (1962), noted that high 



concentrations were measured in marine mollusks 
and crustaceans and correspondingly low concen- 
trations in terrestrial and fresh water species of 
these groups. He concluded that free amino acids 
and taurine formed the bulk of the alpha amino 
nitrogen found in the body fluids of most marine 
invertebrates. Clark (1968b) substantiated these 
findings in her study of 14 species of polychaetes. 
She also noted that a great variability occurred 
between related species. Therefore, the free amino 
acid pool in marine invertebrates can be char- 
acterized: 1 ) the pool is large and differs from 
one species to another, 2) the pool has high con- 



1 Department of Biology, California State College, 
Long Beach. California 90801. 



1972 



FREE AMINO ACID fool. OF NEANTHES ARENACEODl VTATA 






centrations of a lew forms (glycine and proline. 
lor example) and low concentrations of a large 
number of amino acids, and 3 ) I he pool is sig- 
nificantly small in terrestrial and fresh water 
species. Awapara (1962) postulated thai the free 
amino acid pool of marine inverlehrales may 
function both as an osmoregulatory device and as 
a possible food reservoir. The work of Stephens 
and Virkar (1966) may be cited as evidence for 
the role of the pool as an osmoregulatory device. 
They found that a decline in salinity lowered the 
total concentration of the free amino acids, the 
uptake of free amino acids from the surrounding 
medium was decreased under lowered salinities, 
and an increase occurred in the rate of assimila- 
tion of amino acids into proteins under lowered 
conditions. The second postulate of Awapara can 
be supported by the uptake of free amino acids 
from an ambient medium and their possible role 
as a nutrient source (Stephens, 1963; Stephens 
and Virkar, 1966; Reish and Stephens, 1969). 

Investigations of the effects of environmental 
stress on the free amino acid pool include studies 
on varying salinities (Lynch and Wood, 1966; 
Stephens and Virkar, 1966; Negus, 1967; Clark, 
1968a) and domestic sewage (Schafer, 1961, 
1963). As discussed above, decreased salinities 
cause a decrease in the total free amino acid pool 
to a specific level (Lynch and Wood, 1966); 
conversely, in hypersalinities, the concentration of 
the pool increases with proline, glycine, and 
alanine accounting for most of the increase. Fur- 
thermore, Negus (1967) found that the oxygen 
consumption increased under decreased salinities 
in the pelecypod Hydrobia ulvae presumably as a 
response to an increase in energy requirements. 
Allen (1961) suggested that an increase in energy 
expenditure would supply short-chain keto acids 
which could be used as substrates in the trans- 
aminase reaction. The amino acids from hydrolyzed 
protein would serve as amino donors. In addition, 
Clark (1968b) pointed out that amino acids may 
be transferred into tissue compartments depending 
upon the ambient salinity which would then act as 
a osmoregulatory mechanism. 

Environmental stresses affect the organism at 
sublethal levels. Boring activity in the marine 
wood borer genus Limnoria was found to be re- 
lated to the salinity; boring activity decreased with 
a decrease or increase in salinity (Eltringham. 
1961; Reish and Hetherington, 1969). Similar re- 
sults were obtained when three species of Limnoria 
were subjected to sublethal levels of reduced 
dissolved oxygen concentrations (Anderson and 



Reish, 1967). Egg producti .•■■■. 

hemoglobin compensation occurred in Neanthei 
arenaceodentata when it was subjected to 
lethal concentral 8 of dissolved oxygct (D 

1969; Raps and Reish, 1971 ). 
The laboratory induced changes in the 

discussed above serve i dicate thai metabolic 

changes may be occurring within the 
.ind thai perhaps changes at the organ ismic level 
and molecular level can be correlated. Therefore, 
it is the purpose of this study to measure the el- 
lecls of lowered dissolved oxygen concentration', 
and lowered salinities on the free ammo acid 
pool of an inbred strain of Neanthei arenaceo- 
dentata and to correlate these results with i (.. 
previously gathered with this species on the 
organismic level. Furthermore, it is hoped that 
we can learn more about the basic effects of en- 
vironmental stress on an organism which may then 
be employed as a monitoring measure of marine 
pollution. 

METHODS 

The Reish strain of Neanthcs arenaceodenlata 
(Moore) was chosen as the experimental animal 
because of its uniform genetic background. The 
stock colony was fed powdered alfalfa and the 
green alga Entcronwrpha crinita during the ex- 
periments. 

The procedure was essentially the same for both 
the dissolved oxygen and salinity experiments with 
only minor differences. For each experiment four 
young males and four females were selected by 
use of the fighting response (Reish and Alosi, 
1957). Each worm was placed into a separate 250 
ml erlenmeyer flask with 100 ml filtered sea water 
and sufficient food (dried, resoaked Entcronwrpha 
crinita). The flasks were sealed with a rubber 
stopper and in the case of the dissolved o\\gen 
experiments each flask was flushed with N. gas 
according to the method of Reish and Richards 
( 1966). Reduced chlorinity conditions were made 
by diluting 100 r r sea water (19.2 ' . chlorinity) to 
90% (16.4 %o) and 70% (12.8 1. All flasks were 
kept in a cold bath at 18° C for seven da>v V 
seven days, in the case with the dissolved ox\gen 
experiments, the rubber stopper was removed and 
the dissolved oxygen measured with an electrode. 
The worms were removed from the flasks and 
placed into separate vials containing SO'""- ethanol. 

Extraction of amino acids was accomplished by 
leaving the worms in the S0 r c ethanol for 4S to 
50 hours. The worms were removed from the 



34 



BULLETIN SOI lllll<\ CAIIEORXIA ACADEMY ()!■ SCIENCES 



VOLUME 71 



I v iii i i. Means, standard deviations, 1 and Student West 8 of the 14 most abundant amino nitrogen compounds 

at varying concentrations of dissolved oxygen. 



Amino Acid Spol 


i onlrol i s s m P 1 . 
Means, S.D. 


2.7 mg/1 
Means, S.D. 


Mcsl 


1.46 mg/1 
Means. S.D. 


/-lest 


II SI in,' 1 

Means. S.D. 


Mcsl 


Aspartic Acid 


2.40 ± 1.27 


2.08 ± 0.61 


1.25 


2.13 ±0.52 


1.19 


0.70 ± 0.33 


4.47** 


Glutamic Acid 


2.77 ± 0.96 


2.73 ±0.74 


[).l 1 


3.72 ± 0.64 


O Sil*< * 


2.87 ± 0.72 


0.29 


L-amino Adipic Acid 


0.47 ± 0.30 


0.23 ± 0.20 


().()() 


0.52 ± 0.20 


0.00 


0.28 ±0.31 


0.00 


Serine 


4.04 ±1.11 


3.8! ±0.87 


0.54 


3.46 ± 0.95 


1.34 


2.67 ± 0.66 


3.60** 


Glycine 


29.03 ± 6.00 


37.64 ± 8.65 


2.71* 


32.14 ± 8.83 


0.96 


22.65 ± 7.61 


2.18* 


I. -threonine 


3.51 ± 1.19 


1.12 ±0.63 


6.28** 


1.77 ± 1.26 


3.48** 


3.96 ± 1.65 


0.75 


Glutamine 


2.65 ± 0.96 


3.17 ± 1.17 


1.18 


2.84 ± 1.04 


0.46 


2.26 ± 0.59 


0.25 


Citrulline 


0.55 ± 0.22 


2.00 ± 1.20 


4.02** 


0.92 ± 0.83 


1.54 


0.83 ± 0.46 


2.00 


L-alanine 


2.92 ± 0.96 


2.05 ± 1.06 


2.07* 


2.14 ± 1.59 


1.41 


5.30 ±2.17 


3.35** 


Tyrosine 


3.14 ±0.89 


1.37 ± 1.07 


4.31** 


2.99 ± 1.40 


0.30 


2.38 ± 0.80 


2.23* 


Valine, norvaline. 
















I -methionine 


5.35 ± 2.61 


3.21 ± 1.97 


2.18* 


4.42 ± 2.51 


0.85 


6.42 ± 3.23 


0.85 


Phenylalanine, 
















leucine, isoleucine 


11.04 ± 4.29 


7.63 ± 5.26 


1.67 


6.97 ± 4.68 


2.08* 


12.79 ±6.85 


0.72 


Asparagine 


3.12 ± 0.93 


2.87 ± 1.15 


0.58 


3.21 ±0.90 


0.24 


2.08 ± 0.95 


2.73* 


Proline 


14.35 ±4.65 


16.64 ± 3.84 


1.26 


15.99 ± 3.05 


0.98 


24.45 ± 4.66 


5.10* 



1 Means and standard deviations recorded as per cent values 
-Compared to control 
* 5% level: 2.07 
•• 1% level: 2.82 



ethanol, weighed, and inspected for possible dam- 
age. Specimens with openings in their body wall 
were discarded. 

Chromatographic Techniques: The chromato- 
graphic techniques employed followed essentially 
the procedure given by Mearns and Reish (1969). 
Dowex 50W, X8, 100-200 mesh ion exchange 
resin was prepared in 4 N NaOH 12 hours prior 
to use. This resin was washed to neutral with glass 
distilled water and activated 30 minutes prior to 
use with 4 N HC1. Each of the eight 200 X 15 mm 
polyethylene drying columns was filled with 6 ml 
of resin and the flow rate adjusted with glass wool. 
The column was washed with 25 ml glass distilled 
water and 2.5 ml extract was pipetted into a tube. 
The salts were eluted with 125 ml of glass distilled 
water in 25 ml aliquots and discarded. Then 12.5 
ml of 4 N NaOH was pipetted into each column 
to elute the amino acids into collecting tubes. 
The contents of each tube was condensed to 0.05 
ml by use of a rotary evaporation. The spotting 
procedure, the chromatography, and development 
of the chromatograms were identical to Mearns 
and Reish (1969). 

The ninhydrin-positive spots were identified 
from a previously prepared chromatographic 
master. After identification, the spots were cut 
out and placed into individual vials containing 10 
ml 50% ethanol for 2 hours. The paper was then 
removed and each eluate was analyzed quantita- 
tively with a spectrophotometer. Optical densities 



were read at 570 m^ wavelength for the purple 
and pink spots and at 300 m^ for the orange and 
yellow spots and adjusted for the 50% ethanol 
blank. The optical densities were recorded and 
converted to relative proportions of the total 
optical density according to the procedure given 
by Hrubant (1959). The relative proportions of 
amino acids or ninhydrin-positive spots from each 
chromatogram were compared using the Student 
/-test. 

RESULTS 

The data are summarized in tables 1 and 2 for 
the dissolved oxygen and salinity experiments, 
respectively, as the means, standard deviations, 
and Student Ntests. The optical densities for each 
amino acid are not included in the tables; they 
have been converted to per cents of total optical 
density. Since nine of the 23 amino acids for the 
dissolved oxygen experiments and ten of the 23 
for the salinity experiments were discernible but 
negligible these data were omitted from the tables. 
Dissolved oxygen experiments: At least 12 and, 
since some spots contained more than one amino 
acid, possibly 16 amino nitrogen compounds were 
significantly different from the controls to at 
least the 5% level on the Student f-test scale after 
lowered concentrations of dissolved oxygen over 
a seven day period. At the mean D.O. level of 
2.71 mg/1, six and possibly eight compounds 



1972 



FREE AMINO ACID I'OOI. Oh NEANTIIES ARENACEODl ' III i 






Table 2. Means, standard deviations 1 and Studcnl West 3 ol the 13 mosl abundant amino nil 

under varying salinities, 



Amino Acid Spot 


Control (normal 

oallnlly > 

Means, S.l ) 


909! Sallnfl 
Mconi, si). 


Mot 


S i< 




Aspartic Acid 


2.40 


± 1.27 


1.62 1: 1.50 


1.34 


1.60 




Glutamic Acid 


2.77 


! 0.96 


2,21 ±0.57 


1.69 


i 60 0.69 


■ 


Serine 


4.04 


! 1.1 1 


4.27 ± 1.25 


0.46 


1 ! 


0.67 


Glycine 


29.03 


i (,.00 


42.18 ± 7.92 


4.39'* 


4 (.58 < 10.73 


1 


1. -(h reonine 


3.51 


' 1.19 


1.74 ± 1.12 


.3.76** 


3.16 -2.13 


0.48 


Glutamine 


2.65 


±0.96 


1 .75 ± 0.57 


2.72* 


1.27 i 0,7", 


.3.83** 


Citrnllinc 


0.55 


± 0.22 


0.80 ± 0.46 


1.78 


0.85 0.92 


1.15 


L-alanine 


2.92 


± 0.96 


2.49 ± 1.40 


0.86 


1.58 ± 1.27 


2.9 1** 


Tyrosine 


3.14 


± 0.89 


1.74 ± 1.30 


3.04** 


1.54 ± 1.14 


3.80** 


Valine, norvaline, L-methionine 


5.35 


±2.61 


2.81 ± 1.41 


2.85* ♦ 


3.96 ± 2.49 


1.28 


Phenylalanine, leucine, isolencine 


1 1.04 


± 4.29 


6.25 ± 3.36 


2.92** 


7.57 ± 5.53 


1 .64 


Asparagine 


3.12 


± 0.93 


3.62 ±0.84 


1.38 


3.25 ± 0.84 


o , 


Proline 


14.35 


± 4.65 


16.61 ± 3.58 


1.28 


13.38 ± 3.30 


II '(, 



1 Mcuns unci standard deviations recorded as per cent values 
- Compared to control 
* 5% level: 2.07 
*» 1% level: 2.82 



changed significantly in their relative propertions 
at either the 5% or 1 % level. These were glycine, 
threonine, citrulline, (Tyrosine, at spot 32 which 
contained valine, norvaline, and methionine. At 
least three compounds differed significantly at 
mean dissolved oxygen levels of 1.46 mg/1; 
namely, glutamic acid, threonine, and spot 35 
which may contain phenylalanine, leucine, and 
isoleucine. At the mean dissolved oxygen level 
of 0.83 mg/1, seven compounds differed signifi- 
cantly from the control: these were, aspartic acid, 
glycine, serine, L-alanine, tyrosine, asparagine, and 
proline. There were no amino nitrogen compounds 
which differed significantly at all three levels of 
experimentally lowered dissolved oxygen con- 
centrations; however, glycine, threonine. L-alanine, 
and tyrosine differed at two of the three dissolved 
oxygen levels. There were a few amino nitrogen 
compounds which showed general trends, although 
they were not significant statistically. Aspartic 
acid and serine both showed drops in their rela- 
tive concentrations with a decrease in the amount 
of dissolved oxygen present. 

Salinity experiments: At the 90% salinity level, 
six spots encompassing ten possible amino nitro- 
gen compounds varied significantly (Table 2). 
These were glycine, glutamine, 1-threonine, tyro- 
sine, spot 32 which may contain valine, norvaline, 
or methionine, and spot 35 which may contain 
phenylalanine, leucine, or isoleucine. At 70% 
level of salinity the concentration of glycine, 
glutamine, L-alanine, and tyrosine differed sig- 
nificantly from the normal salinity control. An 



additional difference was noted in the totals "I the 

optical densities of all ninhydrin-posilivc spots. 
The mean optical density values were 2.326 ± 
0.766, 1.036 ± 0.324, and 1.053 ± 0.300 for nor- 
mal, 90%, and 70% sea water, respectively. The 
latter two values differed significantly at the 1 ' • 
level when compared to the control. 



DISCUSSION 

The free amino acid pool of N. arenaceodentata 
is somewhat similar, in size and composition, to 
those free amino acid pools of marine invertebrates 
which have previously been studied. It is com- 
posed of 32 amino acids and related amino nitro- 
gen compounds of which seven account for over 
60%. These amino acids are glycine, isoleucine. 
leucine, phenylalanine, proline, serine, and valine. 
Simpson, Allen, and Awapara (1959). in their 
survey of 17 different marine invertebrates, found 
that glycine, proline, taurine, and three others to a 
lesser degree, accounted for most of the amino 
nitrogen present. It was not possible to measure 
taurine in this study: therefore, no estimation of 
its presence could be made. Other workers. 
Camien et al. (19511. Kittredge et al. (19621. and 
Clark (1968b) have found similar results in all 
of the major marine invertebrate phyla. 

The free amino acid pool of .V. arenaceodentata 
also showed a large amount of variability from 
one animal to another. Most of the variability was 
found among the amino acids present in the largest 



16 



HI III ll\ SOI ////A'\ < II II OHM. t ACADEMY OF SCIENCES 



VOLUME 71 



concentrations. Clark (1968b), and Mearns and 
Rcish (1969) observed similar results in other 
polychaete species. However, the organisms they 
studied were from natural field populations. The 
specimens of Neanthes arenaceodentata used in 
this studj were from an inbred population which 
had been maintained for some 30 generations. Still, 
a degree of variability remained. To explain this 
variability other factors were examined. First, the 
diet of the experimental animal used in this study 
was controlled. Second, age and maturity were 
controlled to reduce the possibility of variability 
introduced by age differences. Third, the environ- 
mental factors of temperature, salinity, and light 
were controlled. Fourth, seasonal variation was 
controlled by alternating controls and experi- 
mental over a 12 month period. Since all of these 
factors were held to some degree of constancy, 
they then can conceivably be given less importance 
in their role in determining the variability in the 
free amino acid pool of N. arenaceodentata. Per- 
haps, it is as Clark (1968b) has stated, that under 
normal conditions, the amino acids in polychaetes 
are not that closely regulated, and thus vary within 
a given range. 

If the free amino acid pool of N. arenaceo- 
dentata is compared to the free amino acid pool 
of the closely related N. succinea, the same basic 
pattern of distribution of amino acids occurs, but 
differs in relative proportions (Mearns and Reish, 
1969). Glycine, L-alanine, glutamic acid, serine, 
glutamine, and proline accounted for 75% of the 
free amino acid pool in N. succinea but in N. 
arenaceodentata 75% was due to glycine, proline, 
spot 32 which contains valine, norvaline, or 
methionine, and spot 35, which contains phenyla- 
lanine, leucine, or isoleucine. On the basis of the 
chromatograms of these two species, N. arenaceo- 
dentata could easily be distinguished from N. 
succinea. Such comparisons suggest possible 
phylogenetic relationships which probably exist 
between these two organisms, and it may indeed 
be a better method for studying the phylogeny in 
Family Nereidae than morphological characters 
(Clark, 1961). 

Effects of lowered dissolved oxygen on free 
amino acids: It is very difficult to assess with 
certainty the effects of environmental variables 
on the metabolism of a specific organism. How- 
ever, the results obtained in this type of study do 
lend themselves to some general analysis, from 
which it is possible to make further evaluations. 
For example, the major changes in relative 
amounts of amino acids at each level of lowered 



dissolved oxygen were the result of changes in a 
few common amino acids, namely, glycine, isoleu- 
cine, leucine, phenylalanine, proline, and serine. 
These amino acids comprise about 60% of the 
free amino acid pool. This seems to indicate that 
the organism is compensating in some way for 
lowered dissolved oxygen by altering the more 
abundant amino acids while maintaining constant 
levels in the majority. This suggests that those 
amino acids which are maintained at constant 
levels are most essential for maintenance of normal 
metabolic activity. Furthermore, the observed 
fluctuations in free amino acid levels may and 
probably do reflect degrees of protein synthesis 
and degradation, induced by lowered dissolved 
oxygen. Raps and Reish (1971 ), showed that the 
polychaete A', arenaceodentata increased its hemo- 
globin levels in order to compensate for lowered 
dissolved oxygen conditions. R. A. Cripps (pers. 
comm.) also showed that both protein synthesis 
and degradation can be induced by lowered dis- 
solved oxygen. If proteins, as stated, are being 
synthesized and degraded at increased or decreased 
levels under lowered dissolved oxygen conditions, 
the free amino acid pool would necessarily fluctu- 
ate in accordance with supply and demand for 
the required amino acids. 

A number of amino acids showed a trend in 
their response to lowered dissolved oxygen. Among 
these were alanine, aspartic acid, glycine, proline, 
threonine, and serine. An analysis of the known 
synthesis and degradation pathways of these amino 
shows a number of common characteristics which 
each exhibit, namely: 

1 . Alanine, aspartic acid, glycine, proline, ser- 
ine, and threonine, are all closely related and 
interconnected by their known synthesis and deg- 
radation pathways (Mahler and Cordes, 1968), 
and are essentially interchangeable under differ- 
ent conditions. 

2. All except proline are formed by simple one 
or two step synthesis pathways, making them 
susceptible to interconversion to other compounds 
with little or no energy loss (Mahler and Cordes, 
1968). 

3. All are closely related to carbohydrate metab- 
olism. It is conceivable, therefore, that changes 
in these amino acids may be the result of stress 
imposed on the energy production pathways of 
the organism. 

4. All except proline are short chain and simple 
in structure, hence they are easily formed and 
are most susceptible to conversion for energy 
production. 



1972 



FREE AMINO ACID I'OOI. OF NI.ANllll.S ARL l< EODl ITATA 






Thus far, all explanations advanced for changes 
in the free amino acid pool arc based on I ho as- 
sumption dial under low dissolved oxygen condi- 
tions, the organism mnsl compensate in some 
manner lor loss of the terminal electron acceptor, 
oxygen. II indeed this electron acceptor is oxygen, 
then without it, alternative means must be taken 
to insure energy production, Some of these may 
be behavioral. Neanthes arenaceodentata moves 
to the air-surface interphase under lowered dis- 
solved oxygen condition (Reish, 1967). Other 
measures may not be as well pronounced as this 
observed behavior pattern. Compensation by in- 
creasing hemoglobin (Raps and Reish, 1971 ), and 
increasing enzyme protein or enzyme activity 
(Simpson and Awapara, 1966; Cripps, pcrs. 
comm.), are not well clarified. However, each 
of these methods of compensation is related to 
the production of energy; to maintenance of a 
terminal electron acceptor, or to increased de- 
pendence on anaerobic pathways. Changes in the 
free amino acid levels also are not as distinct as 
the observed behavior, but appear to be related to 
compensatory changes in protein metabolism. 

Effects of lowered salinity on free amino acids: 
The most notable change from lower salinities was 
the decrease in the total concentration of amino 
acids, which agrees with the earlier findings of 
Lynch and Wood (1966), Stephens and Virkar 
(1966), and Clark (1968b). Since the total optical 
density, hence concentration, of ninhydrin-positive 
spots was the same at 90 and 70% salinities, it 
appears as if the free amino acid pool is only 
regulated to a certain minimum level below which 
its normal metabolic machinery may be impaired. 
The 28-day TL m for N. arenaceodentata was about 
60% salinity (Reish, 1970) indicating that this 
impairment of metabolic activity occurred over a 
long period of time and may have been the cause 
of death. In short term responses to hyposalinities. 
Clark (1968a) did not find a stable concentration 
of amino acids in several species of polychaetes. 

Very little is known concerning which amino 
nitrogen compounds are responsible for the total 
decreases in concentration. Lynch and Wood 
(1966) found that proline, glycine, and alanine 
accounted for most of the responses to lowered 
salinities in Crassostrea virginica. The results with 
N. arenaceodentata suggest that the major adjust- 
ment is the result of a general decrease in the size 
of the pool. Although the increase in percent 
glycine was significant, the optical density was 
actually less. The other amino acids decreased in 
concentration even greater than glycine, giving an 



increase in the relative amoi Whi 

some Ol the decreases were nol Fit, at 

least eight and possibly 12 (counting the multiples 

Ol One spot) amino acids decreased in rcl 
proportions at 90'; salinity and at l'-asl ten and 

possibly 15 at 70';. Howevct the glycine con- 
centration remained the same from 90$ to 7095 

salinity. All of these results indicate thai ihe total 
free amino acid pool is osmotically diluted by 
decreased salinity and that glycine is m 
at a constant minimum level while the other free 
amino acids decrease. No explanation is advanced 
herein to explain this observed glycine stability. 

From Ihe results of both the dissolved oxygen 
and salinity experiments, it is apparent that Ihe 
free amino acid pool is affected, either directly 
or indirectly by changes in this organism's en- 
vironment. Alterations in dissolved oxygen and 
salinity concentrations accompany some forms ol 
water pollution. Schafer (1961, 1963) has pre- 
viously noted distinct changes in the tree amino 
acid pool of organisms living in a polluted en- 
vironment. Therefore, monitoring the effects ol 
these variables on the free amino acid pool mas 
give a measure of how pollution affects some 
marine organisms. 



LITERATURE CITED 

Allen. K. 1961. The effect of salinity on the amino 
acid concentration in Rangia cuneala (Pelecyp- 
oda). Biol. Bull., 121:419-424. 

Anderson. J. W.. and D. J. Reish. 1967. The effects 
of varied dissolved oxygen concentrations and 
temperature on the wood-boring isopod genus 
Limnoria. Mar. Biol.. 1:56-59. 

Awapara. J. 1962. Free amino acids in inverte- 
brates: a comparative study of this distribution 
and metabolism. Pp. 158-175. In Holden. J. T . 
[ed.]. Amino acid pools. Distribution, formation 
and function of free amino acids. Elsivier. 
Amsterdam. 

Ball. G. H.. and E. W. Clark. 195?. Species dif- 
ferences in amino acids of Culex mosquitos. Syst. 
Zool.. 2:138-141. 

Camien. M. H.. H. Sarlet. G. Duchateau. and M. 
Florkin. 1951. Non-protein amino acids 
muscle and blood of marine and freshwater 
crustaceans. J. Biol. Chem.. 193:881-81 

Clark. M. E. 196Sa. A survey of the effect of 



is 



i;i III I IX SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



osmotic dilution on free amino acids of various 
polychaetes. Biol. Bull., 134:252-260. 

— . 1 968b. Free amino acid levels in the 



coelomic fluid and body wall of polychaetes. 
Biol. Bull., 134:35-47. 

Clark, R. B. 1961. The origin and formation of the 
hetcroncreis. Biol. Rev.. 36:199-236. 



Micks, D. W.. and I-'. .1. Gibson. 1957. The char- 
acterization of insects and licks by their free 
amino acid patterns. Ann. Ent. Soc. Amer., 50: 
500-505. 

Negus. M. R. S. 1967. Oxygen consumption and 
amino acid levels in Hydrobia ulvae (Pennant) 
in relation to salinity and behavior. Comp. Bio- 
chem. Physiol., 24:317-325. 



Dims. D. 1969. The effects of lowered dissolved 
oxygen levels on the reproduction of the poly- 
chaetous annelid Neanlhes arenaceodentata. Un- 
published Master's thesis. California State Col- 
lege. Long Beach. Long Beach, California. 62 pp. 

Eltringham, S. K. 1961. The effect of salinity upon 
the boring activity and survival of Limnoria 
(Isopoda). J. Mar. Biol. Assoc. U. K., 41:785- 
799. 

Hrubant. H. E. 1959. A chromatographic analysis 
of the free amino acids in the blood plasma of 
three inbred strains of the house mouse, Mas 
imtscuhis. Genetics, 44:591-608. 

Kittredge. J. S., D. G. Simonsen, E. Roberts, and B. 
Jelink. 1962. Free amino acids of marine in- 
vertebrates. Pp. 176-186. In Holden, J. T., [ed.] 
Amino acid pools. Distribution, formation and 
function of free amino acids. Elsivier, Amster- 
dam. 

Lewallen, L. L. 1957. Paper chromatography studies 
of the Anopheles maculipennis complex in Cali- 
fornia (Diptera: Culicidea). Ann. Ent. Soc. 
Amer., 50:602. 

Lynch. M. P., and L. J. Wood. 1966. Effects of 
environmental salinity on free amino acids of 
Crassostrea virginica (Gmelin). Comp. Biochem. 
Physiol., 19:783-789. 



Raps, M. E., and D. J. Reish. 1971. The effects of 
dissolved oxygen on the hemoglobin levels of the 
polychaetous annelid, Neanlhes arenaceodentata. 
Mar. Biol., 11:363-368. 

Reish. D. J. 1967. Relationship of polychaetes to 
varying dissolved oxygen concentrations. Adv. 
Water Poll. Res., 3:199-266. 

. 1970. The effects of varying concentrations 



of nutrients, chlorinity, and dissolved oxygen on 
polychaetous annelids. Water Res., 4:721-735. 

Reish, D. J., and M. C. Alosi. 1967. Aggressive be- 
havior in the polychaetous annelid family Nere- 
idae. Bull. So. California Acad. Sci., 67:21-28. 

Reish, D. J., and W. M. Hetherington, III. 1969. 
The effects of hyper- and hypo-chlorinities on 
members of the wood-boring genus Limnoria. 
Mar. Biol.. 2:137-139. 

Reish, D. J., and T. L. Richards. 1966. A technique 
for studying the effect of varying concentrations 
of dissolved oxygen on aquatic organisms. In- 
ternal I. Air Water Poll., 10:69-71. 

Reish, D. J., and G. C. Stephens. 1969. Uptake of 

organic material by aquatic invertebrates. 5. The 
influence of age on the uptake of glycine-C" by 
the polychaete Neanlhes arenaceodentata. Mar. 
Biol., 3:252-255. 



Mahler. H. R., and E. H. Cordes. 1968. Basic 
Biological Chemistry. Harper and Row, New 
York, 509 pp. 



Roberts, E., and P. Borges. 1955. Patterns of free 
amino acids in growing and regressing tumors. 
Canadian Res., 15:697-699. 



Mearns, A. J., and D. J. Reish. 1969. A comparison 
of the free amino acids in two populations of the 
Polychaetous annelid Neanthes succinea. Bull. 
So. California Acad. Sci., 68:43-53. 



Roberts, E., and S. Frankel. 1949. Free amino 
acids in normal and neoplastic tissues of mice as 
studied by paper chromatography. Canadian Res., 
9:645-648. 



Micks, D. W. 1954. Paper chromatography as a 
tool for mosquito taxonomy: the Culex pipens 
complex. Nature. London, 174:217-221. 

. 1956. Paper Chromatography in insect 

taxonomy. Ann. Ent. Soc. Amer., 49:576-581. 



Roberts. E., and T. P. Lowe. 1954. Occurrence of 
the O-phosphodiester of L-serine and ethanol- 
amine in turtle tissue. J. Biol. Chem., 211:1-12. 

Schafer, R. 1961. Effects of pollution on the free 



IV72 



FREE AMINO ACID POOL <>!■ Ni.ANI III.S AKIN |( lain I 1/ • 






amino acid content of two marine invertebrates. 

Pacific Sci., 15:49-55. 

1963, Effects of pollution on the amino 
acid content of Mytilus edulis. Pacific Sci.. 17: 
246-250. 

Simpson, .1. w., K. Allen, and J. Awapara. 1959. 
Free amino acids in some aquatic invertebrates. 
Biol. Bull., 117:371-381. 



by aquatic invertebrates. 2. A' 
amino acids by Ihi bamboo worm ( I 
torquata ( omp. Bioi hem. Ph iol., 10: 191 

Stephens, (<.(.. and R. a, Virkar. 1966 i | 
organic matei ial by aquatii in ■ i l< 
influence of salinity on die briltlcstar, OphL 
arenosa, Biol. Bull., 131:172 185. 



Stephens, CJ. C. 1963. Uptake of Organic material Accepted for publication February x. 1972 



AN ANNOTATED KEY TO THE CERCARIAE THAT DEVELOP IN THE SNAII 

CERITHIDEA CALIFORNIA 

W. E. Martin 1 



Abstract: An annotated key is provided to the 18 species of nematodes found in the 
snail, Cerithidea californica; an intermediate host. 



Cerithidea californica Haldeman, the California 
hornshell snail, is common on mud-sand or mud 
beaches of estuaries and bays of southern Cali- 
fornia. These areas are favorite feeding grounds 
for local and migratory shore birds. Bird feces con- 
taining trematode eggs result in heavy and varied 
infections of this snail. The intensity and types 
of infection vary somewhat with different localities 
because some regions are more commonly used 
by birds. Martin (1955) followed the types and 
infection rates over a one-year period at Newport 
Bay. 

Infected snails are used by biology and zoology 
classes in the study of larval trematodes. An an- 
notated key to these trematodes should be useful 
to such studies. Furthermore, the availability of 
all the stages in a trematode life cycle could be 
advantageous in physiological and biochemical re- 
search. 

An attempt was made to keep the key simple, 
to use distinguishing characteristics that are easily 
recognized, and that do not require observation 
with an oil immersion lens, wherever possible. 

Cerithidea californica is a favorite host for 
many trematodes. Eighteen species of cercariae 



are listed in the following key and it is possible 
that a few more will be found. Perhaps the 
cercaria of Ascocoiyle sexidigita also may use 
this snail as host. The metacercariae of this species 
have been found in Fundulus parvipinnis living in 
waters where this snail is common (Martin and 
Steele, 1970). Judging by the work of Schroeder 
and Leigh (1965) on the life cycle of Ascocoiyle 
pachycystis, the cercaria of A. sexidigita should 
resemble those of Stictodora (Parastictodora) 
hancocki and Euhaplorchis californiensis. 

The figures in this paper include only certain 
details to assist in identification. The size scale 
is based on fixed specimens: fresh material should 
be larger. This study was supported by NSF grant 
GB-6962. 



KEY TO CERCARIAE INFECTING 
CERITHIDEA CALIFORMCA 



a. Tail forked 

b. Tail nonforked __ 



1 Biology Depi.. University of Southern California. 
Los Anceles. California 90007. 



40 



HI ill ll\ OIIII1 l<\ CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 




Figures 1—4. 
Notocotylid. 



2. a. With eyespots — Avian blood fluke. Develops 

in sporocysts (Fig. 1 ). This may be the same 
as Austrobilharzia penneri Short and Holli- 
man, 1961 and Holliman (1961) which uses 
Cerithidea scalariformls as intermediate host 
and chicks, parakeets, and pigeons as experi- 
mental adult hosts. However, we have ex- 
posed chicks and pigeons to large numbers 
of the cercaria from C. californica with nega- 
tive results. Therefore this may be a different 
species of Austrobilharzia. Further work is in 
progress, 
b. Without eyespots 3 

3. a. Flame cells in groups of twos — Strigeid 

cercaria (Fig. 2). Develops in sporocysts, 
cercariae encyst in muscles of Fundulus 
parvipinnis. Life cycle unknown. Smaller 
than 3b. 
b. Flame cells in groups of threes — Meso- 
stephanus appendiculatus (Ciurea, 1916) 
Lutz, 1935 (Fig. 3). Develops in sporocysts, 
encysts in muscles of Fundulus parvipinnis, 
adults experimentally in intestine of chicks, 
naturally in shore birds. Life cycle described 
by Martin (1961). 



Figures 5-11. 5. 10. Echinostomids. 6. Heterophyid. 
7, 8. Microphallids. 9, 1 1 . Philophthalmids. 



4. a. Tail with fins - 14 

b. Tail without fins 5 

5. a. With eyespots 6 

b. Without eyespots 8 

6. a. With a pouch at each posterior-lateral 

margin of body — Catatropis johnstoni Martin, 
1956 (Fig. 4). Develops in rediae, encysts 
on almost any object, adults experimentally 
in ceca of chicks. Life cycle described by 
Martin (1956). 
b. Without a pouch at each posterior-lateral 
margin of body 7 

7. a. Encysts in dish — Himasthla rhigedana Dietz, 

1909 (Fig. 5). Develops in rediae, encysts 
on almost any object, adults in willet and ex- 
perimentally in ileum of chick. Life cycle 
described by Adams and Martin (1963). 
Cercaria has median pigment spot nearly in 
line with eyespots. 
b. Does not encyst in dish — Pygidiopsoides 



7972 



KEY TO THE CERCAR1AE Of CALIFORNIA HOR in U. SNAILS 







Figures 12-18. 12. 17. Echinostomids. 13, 14. Ren- 
icolids. 15. 16. 18. Heterophyids. 



spindalis Martin. 1951 (Fig. 6). Develops in 
rediae, encysts in gills of Fundulus parvipinnis, 
adults experimentally in intestines of chicks 
and cats. Life cycle described by Martin 
(1964). 

a. With stylet (small "spear") near mouth — 
Microphallids 1 and 2. 

1. Develops in sporocysts (Fig. 7). Larger 
than 2. 

2. Develops in sporocysts (Fig. 8). Smaller 
than 1. 

Sarkisian (1959) described Maritrema uca 
from fiddler crabs, Uca crenulata (Locking- 
ton), collected in the same area where the 
two above microphallid cercariae are found. 
Apparently one of the above encysts in this 
crab. Heard and Sikora (1969) placed this 
species in the genus Probolocoryphe. 

b. Without a stylet 9 

a. With pouch opening at end of tail 10 

b. Without pouch opening at end of tail ._ 11 



10. a. Willi spincd collai Parorcl canthu 

(Nicoll, 1906) (Fig n , Devi lo| 
ency i on almo I any object, adull 
ni '.linn- birds, I lb- ■ ■ Ic di 
Stunkard and ' able (19 P 

was declared a synonym '■! /' acuniliu 
< able and Martin (1915). I I,;,.. 
infections in chicks by injecting Ihi m with 
metacercariae per cloaca bul il»- pen enl 
success is low. 'I his cercaria is largci than 10b 
b. Withoui spined collai Cloacltrema n 
ganensis Mcintosh, I91K (Fig, 9). Develop-, 
in rediae, encysts on almost any object, adults 
in cloaca of shore birds. Life cycle outlined 
by Robinson (1952). I have established in- 
fections in chicks by injecting them with 
metacercariae per cloaca but the percentage 
of success is low. 



II. a. With spined collar 
b. Withoui spined collar 



12 
13 



12. a. F.ncysts in radular muscle — Acanthoparyphium 

spinulosum Johnston. 1917 (Fig. 10). De- 
velops in rediae. encysts in Cerilhidea call- 
fornica especially in radular muscles, adults 
experimentally in intestine of chicks. Life 
cycle described by Martin and Adams (I960. 
1961). Cercaria larger than 12b. 
b. Encysts in feces — Acanthoparyphium sp. 
(Fig. 12). Develops in rediae, encysts in feces 
of C. californica and in certain annelids, 
adults experimentally in intestine of chicks, 
Martin and Steele in manuscript. 

13. a. Usually forms clusters by adhesions of 

proximal portions of tails. Y-shaped ex- 
cretory bladder — Renicola hitchanani (Mar- 
tin and Gregory. 1951) (Fig. 13). Develops 
in sporocysts. encysts in liver of Fundulus 
parvipinnis, young adults experimentally in 
kidneys of Larus californicus. Martin (1971). 
Cercaria larger than 13b. 
b. Does not form clusters. Y-shaped excre- 
tory bladder — Renicola cerithidicola Martin. 
1971 (Fig. 14). Develops in sporoc\sts. 
encysts in gills of Fundulus parvipinnis. young 
adults experimentally in kidneys of Larus 
californicus, Martin (1971). 

14. a. With dorso-ventral tail fin only 

b. With dorso-ventral and lateral tail fins _ 16 

15. a. With eyespots — Phocitremoides ovale Martin. 

1950 (Fig. 15). Develops in rediae. encysts 
under scales of Fundulus parvipinnis and 
Alherinopsis californica (Girardl. adults ex- 
perimentally in chicks and cats. Life cycle 
described by Martin (1950c). 
b. Without eyespots — Echinoparypliium sp. (Fig. 
PL Develops in rediae. sometimes forms 
weak-walled cvsts in dish, life cycle unknown. 



BULLETIN SOUTHER* CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 7 1 



Id. a. Flame cells in groups of twos — Euhaplorchis 
calif or niensis Martin. 1950 (Fig. 16). De- 
velops in rediae, encysts on brain of Fundulus 
parvipinnis, adults experimentally in small 
intestines of chicks, turns californicus, cats 
(Bamberger and Martin. 1951 ). monkeys, etc. 
Life cycle described by Martin (1950a). 
b. blame cells in groups of threes — Sliclodora 
[Parastictodora) hancocki (Martin, 1950) 
(Fig. 18). Develops in rediae. encysts in 
various tissues of Fundulus ^parvipinnis and 
Gillichthys mirabilis Cooper, adults experi- 
mentally in small intestines of chicks and 
Lams californicus. Life cycle described by 
Martin (1950b). 



LITERATURE CITED 

Adams, J. F„. and W. E. Martin. 1960. Life history 
of Himasthla sp. an echinostome trematode. J. 
Parasit.. 46: (supple.) : 15. 



— . 1950c. Phocitremoides ovule n.g., n.sp., 
(Trematoda: Opisthorchiidae), with observations 
on its life cycle. J. Parasit.. 36:552-558. 



— . 1951. Pygidiopsoides spindalis n.gen., n.sp., 
(Heterophyidae: Trematoda). and its second 
intermediate host. J. Parasit., 37:297-300. 



— . 1955. Seasonal infections of the snail, 
Cerithidea californica Haldeman, with larval 
trematodes. Essays Nat. Sci. Honor of Capt. 
A. Hancock, 203-210. 



— . 1956. The life cycle of Catatropis johnstoni 
n. sp. (Trematoda: Notocolylidae). Trans. Amer. 
Micros. Soc. 75:117-128. 



. 1961. Life cycle of Mesostephanus ap- 

pendiculatus (Ciurea, 1916) Lutz, 1935 (Trem- 
atoda: Cyathocotylidae). Pacific Sci., 15: 
278-281. 



— . 1963. Life cycle of Himasthla rhigedana 
Dietz, 1909 (Trematoda: Echinostomatidae). 
Trans. Amer. Micros. Soc, 82:1-6. 



. 1964. Life cycle of Pygidiopsoides spin- 
dalis Martin. 1951 (Heterophyidae: Trematoda). 
Trans. Amer. Micros. Soc, 83:270-272. 



Bamberger, J. W., and W. E. Martin. 1951. Effect 
of size of infective dose, partial ileectomy, and 
time on intensity of experimental infections of 
Euhaplorchis calif orniensis Martin. 1950 (Trem- 
atoda) in the cat. J. Parasit., 37:387-391. 

Cable, R. M., and W. E. Martin. 1935. Parorchis 
avitus (Linton, 1914) a synonym of P. acanthus 
(Nicoll. 1906). J. Parasit., 21:436-437. 

Heard. R. W., Ill, and W. B. Sikora. 1969. Pro- 
bolocoryphe Otagaki. 1958 (Trematoda: Micro- 
phallidae). a senior synonym of Mecynophallus 
Cable, Connor, and Balling, 1960. with notes on 
the genus. J. Parasit.. 55:674-675. 

Holliman. R. B. 1961. Larval trematodes from the 
Apalachee Bay area Florida, with a checklist of 
the known marine cercariae arranged in a key 
to their superfamilies. Tulane Studies Zool., 9: 
2-74. 

Martin. W. E. 1950a. Euhaplorchis calif orniensis 
n.g., n.sp., Heterophyidae, Trematoda, with notes 
on its life cycle. Trans. Amer. Micros. Soc, 69: 
194-209. 

. 1950b. Parastictodora hancocki n.gen.. 

n.sp. (Trematoda: Heterophyidae), with observa- 
tions on its life cycle. J. Parasit., 36:360-370. 



Martin, W. E.. and J. E. Adams. 1960. Life cycle 
of Acanthoparyphium spinulosum Johnston, 1917 
(Echinostomatidae). J. Parasit., 46: (Supple.): 35. 

. 1961. Life cycle of Acanthoparyphium 



spinulosum Johnston, 1917 (Echinostomatidae: 
Trematoda). J. Parasit.. 47:777-782. 

Martin, W. E., and V. L. Gregory. 1951. Cercaria 
buchanani n. sp., an aggregating marine trem- 
atode. Trans. Amer. Micros. Soc, 70:359-362. 

Martin, W. E., and D. F. Steele. 1970. Ascocotyle 
sexidigita sp. n. (Trematoda: Heterophyidae) 
with notes on its life cycle. Proc Helm. Soc, 
37:101-104. 

Robinson, H. W. 1952. A preliminary report of 
the life cycle of Cloacitrema michiganensis Mc- 
intosh. 1938 (Trematoda). J. Parasit., 38:368. 

Sarkisian, L. N. 1957. Maritrema ttca, new species 
(Trematoda: Microphallidae), from the fidler 
crab, Uca crenulata (Lockington). Wasmann J. 
Biol.. 15:35-48. 

Schroeder, R. E., and W. H. Leigh. 1965. The life 
history of Ascocotyle pachcystis sp. n., a trema- 
tode (Digenea: Heterophyidae) from the rac- 
coon in South Florida. J. Parasit., 51:594-599. 



1972 



KEY TO THE CERCAR1AE OF < Al.ll OKNIA llt>i< . \iii 1 1 






Short, R. is., and R. B. Holliman. 1961. Auslro- 
bilharzia penneri, a new schistosome from marine 
snails. .1. Parasit., 47:447-452. 



histoi v "I Paron his a\ mi: \ i inton i a m 

from the cloaca "I Ihc cull. Biol. Bull., '.;■ 

12K 338. 



Stunkard, II. W., and R. M. Cable. 1932. The life Accepted lor publication September 25, 



NEW SPECIES OF ROBBER FLIES OF THE GENERA 
WILCOX1A AND METAPOGON (DIPTERA: ASILIDAE) 

J. Wn.cox 1 



Abstract: Four new species of the genus Wilcoxia are described and a key to the species 
occurring in southwestern United States and northern Mexico is presented. In addition, two 
new species of the genus Metapogon are described. 



The genus Wilcoxia has remained monotypic since 
it was described by James ( 1941 ) ; type species W. 
cinerea. These small robber flies, length 5-10 
mm. have the general appearance of Cophura 
Osten Sacken, but lack the twisted spine at the 
apex of the fore tibiae which is characteristic of 
that genus. The middle tibiae at the apex has a 
short straight spine which is usually brown or 
black. Wilcoxia is most closely related to 
Metapogon Coquillett which usually has the 
mesonotum compressed, highly arched, in lateral 
view, and with strong anterior dorsocentral 
bristles, the wings are spotted with brown on the 
crossveins and furcations, the third vein usually 
is branched before the end of the discal cell, and 
the stump vein is long. In Wilcoxia the mesonotum 
is not compressed or highly arched and the strong 
anterior dorsocentral bristles are absent, the wings 
are hyaline, uniformly brown, or brownish, the 
third vein is branched opposite or beyond the 
end of the discal cell, and the stump vein is usually 
absent. Wilcox and Martin (1957) presented a 
key to the related genera. Material received re- 
cently for identification has prompted me to de- 
scribe four new species of Wilcoxia. 

Measurements were made with an ocular 
micrometer; the face and wings at 40X, and the 
antennae at 75 X. 



Wilcoxia marlinorum, new species 

Description: Male. Length 7 mm. Head black, 
densely grayish white pollinose. Mystax composed of 
long sparse oral white hairs becoming slightly shorter 
above and extending to antennae: hairs on frons white: 
four long and several short bristles on ocellar tubercle 
and occipitals. yellowish white: beard, hairs on palpi 
and proboscis white. Face at the antennae 15 28 
(.54) width of one eye. Antennae black, grayish 
pollinose; hairs white; bristles white, two weak bristles 
below on segment 1 and 2. stronger bristles below on 
2; segments 10-8-26-15 in length. 

Mesonotum black, densely grayish white pollinose. 
broadly divided central stripe brown. Sparse hairs 
white, long erect on humeri and anterior and lateral 
margins. Bristles yellowish: 2 presutural. 1 supraalar. 
1 postalar, 6 dorsocentral (3 anterior). Pleura and 
coxae densely grayish pollinose. hairs white. Scutellum 
densely grayish pollinose, sparse discal hairs white, 
six weak and long yellowish marginal bristles. 

Abdomen black, densely gray pollinose with bare 
spots at middle of tergite 1. 2-6 with an anterior 
lateral spot on each side and a central posterior spot, 
and 7 with a small central posterior spot. Hairs white, 
sparse, long on sides of 1-3; five lateral bristle-like 
hairs on 1. Sternites gray pollinose. hairs erect white 
becoming shorter apically. Genitalia brown, hairs 
w r hite. 



1 21171 Mohler Place. Anaheim. California c : 



44 



Hi I I 1 ll\ SOUTHERN < AL1F0RNIA ACADEMY OF SCIENCES 



VOLUME 71 



I egs black, fore and middle femora dorsallj and 
laterally, hind femora dorsally and sides of apical 
third, graj pollinose; hairs and bristles white: claws 
black; empodia and pulvilli, light brown. 

Halteres yellow, base brown. Wings hyaline: veins 
brown; anterior crossvein at 42/60 (.70) length of 
discal cell; posterior and anal cells open. 

Female. Length 7 mm. Frons brownish pollinose. 
Mesonotum with a spot between humeri and central 
stripe and the anterior intermediate spots, golden 
brown pollinose. Abdominal segments 7-8 brownish 
black and bare of pollen, apical spines brown. Wings 
with a faint brown clouding on the crossveins and 
furcations. 

Holotype: male. California. Riverside Co.. White 
Water. 23 December 1947 (J. Wilcox) CAS. Allotype: 
female, same data as holotype. 

Named in honor of the ardent collecting team of 
Dorothy and Charles H. Martin who found this species 
literally swarming on the hillside rocks near the en- 
trance to White Water Canyon on 23 December 1947. 

Prey: A small anthomyiid fly. two borborid flies 
and a small green hymenopteran were taken by P. H. 
Arnaud. Jr. and a small winged termite by A. L. 
Melander. 

Other specimens examined: ARIZONA. Maricopa 
Co., $. 21 mi N Ajo. 11 November 1966 (R. J. 
Hamton) EF; Pima Co.. 8 5 , 7 9. Organ Pipe Cactus 
National Monument. 2 mi N Headquarters, 1 1 No- 
vember 1966 (R. J. Hamton) EF; Yuma Co.. 8 5. 12 9. 
Brenda. 28 October 1967 (S. A. Gorodenski and M. 
A. Cazier) ASU. CALIFORNIA. Imperial Co.. 9. 
Painted Gorge, Coyote Mts., 22 August 1934 (C. D. 
Michener) CIS; Inyo Co.. $, Death Valley National 
Monument, 1.5 mi SW Wildrose Station, 3100 ft. 6 
November 1968 (P. H. Arnaud, Jr.) at flowers of 
Chrysolhamiuts penicillatus (Gray) Hall, CAS; River- 
side Co., 9, Coyote Creek, 6 November 1963 (E. I. 
Schlinger) UCR; 5$, 29, Deep Canyon, P. S. Boyd 
Desert Research Center, 3.5 mi S Palm Desert, marker 
no. 57. 6 November to 22 December 1963, 1969, 31 
January to 10 February 1970 (S. Frommer, M. E. 
Irwin. L. La Pre, E. I. Schlinger, and R. Worley) at 
light and malaise trap. UCR; 9, Desert Hot Springs. 
Los Angeles Aqueduct. 25 January 1947 (G. H. Sperry 
and J. L. Sperry) USNM; 45, 7 9, Joshua Tree Na- 
tional Monument (JTNM); Pleasant Valley, Fried 
Liver Wash, 22 October 1967, 29 October 1965 (E. 
L. Sleeper) part blacklight. CSCLB; 6<j, 109, 5 and 
10 mi S Oasis, 17 November 1947, 10 December 1962 
(A. F. Howland. I. J. Wilcox, and J. Wilcox) JW: 
25, Palm Desert, 10 October 1964 (E. Fisher) EF; 
45, 3 9, Palm Springs. 12 November 1944, 19 No- 
vember 1943 (A. L. Melander) USNM; 77 5, 73 9, 
Palm Springs. 23 December 1950, 29 December 1953 
to 8 January 1954. 21 January 1953 (P. H. Arnaud, 
Jr.) JW; 5 3, 29, Palm Springs. 21 January 1953. 
16 December 1964 (M. W. Stone and J. Wilcox) JW; 
95, 13 9, Palm Springs, Chino Canyon, 22 to 26 



December 1950 (P. H. Arnaud, Jr.) JW; 55, 179, 
Palm Springs, Tahquitz Canyon, 24 November 1944 
(A. L. Melander) USNM; 46 5, 1359, White Water, 
23 November to 27 December 1947. 6 January 1948 
(A. F. Howland. C. H. Martin. D. Martin. I. I. Wilcox, 
and J. Wilcox) CHM, CIS. JW, UCD; 15 5. 12 9, 
White Water, 11 November 1966, 13 November, 16 
December 1962 (E. Fisher and R. J. Hamton) EF. 
NEVADA. Nye Co., 5, Rhyolite, 2 October 1936 
(A. J. Basinger) CAS. MEXICO. Sonora. 3 5, 2 9, 
13 mi SW Sonoyta, 12 November 1966 (E. Fisher) 
EF. 

WUcoxia monae, new species 

Description: Male. Length 6 mm. Head black, densely 
whitish pollinose. Mystax white, composed of weak 
oral bristles and hairs above about half as long; 
hairs on frons white: six short erect bristles on ocellar 
tubercle and occipitals, yellowish: beard, hairs on palpi 
and proboscis white. Face at antennae 20/24 (.83) 
width of one eye. Antennae black, hairs white, seg- 
ment 2 with two longer hairs below; segments 
1 1-11-40-12 in length. 

Mesonotum black, gray pollinose, broadly divided 
central stripe brown. Hairs white, appressed and as 
long as antennae 1. Bristles yellowish, 4 presutural, 
3 supraalar, 3 postalar, 4 posterior dorsocentral. 
Pleura and coxae gray pollinose. hairs white. Disc of 
scutellum gray pollinose with short sparse white hairs, 
broad posterior margin black with four weak, yel- 
lowish bristles. 

Abdomen shining black, sides of tergites 1-6, 
slightly projecting inward on posterior margins of 4-6, 
gray pollinose. Hairs short yellowish white, sparse on 
dorsum; three or four yellowish lateral bristles on 
1. Sternites gray pollinose, hairs erect white, dense 
apically. Genitalia brown, hairs yellowish. 

Femora black, narrow base and about apical third 
yellowish-red; tibiae and tarsi yellowish-red; fore tarsi 
and segment 5 of middle and hind tarsi brown. Hairs 
white; bristles yellowish; claws black; empodia white; 
pulvilli gray. 

Halteres yellow, base brown. Wings brown with 
lighter spots in posterior cells; veins brown, anterior 
crossvein at 38/59 (.64) length of discal cell; third 
vein branched beyond end of discal cell; all posterior 
cells and anal cell narrowly, open. 

Female. Length 7 mm. Oral bristles yellowish. 
Dorsum of abdominal tergite 6, all of 7-8, and apical 
spines brown. Wings light yellowish brown, anterior 
crossvein at 45/67 (.67) length of discal cell. Anterior 
side of fore tibiae and all tarsi brownish black. 

Holotype: male, California, Mono Co., Mammoth. 
9 August 1957 (J. Wilcox) CAS. Allotype: female, 
same data as holotype. 

Other specimens examined: CALIFORNIA. Mono 
Co., 265, 49, Mammoth. 9 August 1957 (I. J. Wilcox 
and J. Wilcox) EF, JW; Nevada Co., 9, 3 mi N 



1972 



NliW SPECIES OF Kuiim-.K FLIES 






Boca, 23 July 1961 (F. I). Parker) UCD; v. I mi 
S Hobart Mills, I September 1957 (E. (J. Linsley) 
Chvysolhamnus viscidiflorus var. typicus, CIS; $ , 
Prosser Dam, 15 July 1966 (I). R. Miller) UCD; 
$, 29, Truckee, 17 August 1955 (K. G. Linsley) Ciim- 
positae, CIS; Tulare Co., 9, Mineral King. 8 Augusl 
1959 (W. E. Simonds) CDA; ,', , 9, 0.5 mi E Smith 
Meadow, Nine Mile Canyon, 7850 ft, 5 Augusl [961 
(C. W, O'Brien) CIS. NEVADA. Douglas Co., S, 
Topaz Lake, 17 August I960 (A. S. Menke) UCD. 



Wilcoxin painteri, new species 

Description: Male. Length 5 mm. Head black den- 
sely whitish pollinose, frons with central golden spot. 
Hairs white, lower third of mystax long, short and 
sparse above; occipital and four erect bristles on ocel- 
lar tubercle white. Face at antennae 16/19 (.84) width 
of one eye. Antennae brownish black; hairs white; 
segment 2 with one weak white bristle below; 3 nearly 
uniform in width and style broad; segments 7-8-25-10 
in length. 

Mesonotum black; grayish pollinose, divided cen- 
tral stripe, anterior and posterior intermediate spots 
and a small spot opposite postalar calli, brown pol- 
linose. Short sparse hairs white. Bristles yellowish, 
2 weak humeral, 2 presutural, 2 supraalar, 1 postalar, 
4 dorsocentral ( I anterior). Pleura and coxae grayish 
pollinose, hairs white. Scutellum grayish pollinose. 
two erect marginal bristles yellowish, fine marginal 
hairs white. 

Abdomen brownish black; narrow lateral margins 
of all tergites and very narrow anterior margin of 
2-6, gray pollinose. Fine hairs white, longer laterally 
on 1-3; three yellowish lateral bristles on 1. Sternites 
grayish pollinose, narrow central line bare of pollen, 
hairs white. Genitalia small and black, hairs white. 

Femora except tips, apices of tibiae and tarsal seg- 
ments black; tips of femora, basal part of tibiae and 
tarsi, yellowish-red. Hairs and bristles white; claws 
black, reddish basally; pulvilli brown. 

Halteres yellowish, base of stem brownish. Wings 
hyaline, veins light brown, anterior crossvein at 28/46 
(.61) length of discal cell: third vein branched op- 
posite end of discal cell; all posterior and anal cells 
open. 

Fetnalc. Length 6 mm. Intermediate spots of 
mesonotum bare of pollen and shining brownish- 
black. Anterior fascia on abdomen slightly broader, 
and absent on tergite 6; apical spines black. Third vein 
branched slightly beyond end of discal cell and in one 
wing with minute stump vein. 

Holotype: male. New Mexico, Catron Co., Datil. 
Continental Divide, 17 July 1930 (T. F. Winburn 
and R. H. Painter) RHP. Allotype: female, same 
data as holotype. 

Named in honor of the late Reginald H. Painter 
who, although not especially interested in Asilidae. 



always collected ran and inti on hia 

many trips. 

Other specimens examined '-I ■'■ Ml • I' I 
imn Co., i . 22, Datil, ( ontincntal 1 1 
July 1930 (I. I Winburn and R H P 
RHP, IW. ARIZONA. Apache I Chin 

I ee (< hinle), 26 July 1935 (Brucs) USNM, i i VH 
Kane Co., 2, 16 mi W Glen I an ■••mbcr 

1969 (P. II. Timberlake) Sutienzia sp., U< R; Millard 
Co., $, 23 mi W Delta, 4900 ft, 4 Scptcmbci 1963 
Ik. II. Painter and I . \1. Painter) RHP; Uinta 
V. 16 mi SW Vernal. 5000 ft, 7 Scplcmhci 1965 

(R. H. Painter :t\u\ E. M. Painter) RHP; county un- 
determined, 2, Showell, 20 August 1932 <<. I 
Knowlton) JW. [anterior fascia on abdomen of this 
specimen much broader and faint fascia on tergite 6|. 



Wilcoxia pollinosa, new species 

Description: Mule. Length 8 mm. Head black, 
densely white pollinose. Hairs and bristles white: 
mystax of about eight oral bristles and short hairs 
extending half way to antennae; two erect bristles on 
ocellar tubercle. Face at antennae 17 23 1.74) width 
of one eye. Antennae black: short sparse hairs white: 
one weak white bristle below on segment I and one 
strong below on 2; segments 9-10-29-14 in length. 

Mesonotum black: white pollinose. divided central 
stripe and small intermediate spots brown. Hairs 
short sparse semierect white. Bristles white. 2 pre- 
sutural. I supraalar. 1 postalar. 4 weak, anterior 
dorsocentral (pin obscures posterior ones). Pleura 
and coxae black; white pollinose: hairs white, short 
on coxae, long on hypopleura. Scutellum black: white 
pollinose; three strong white marginal bristles. 

Abdomen black: tergites 1-8 white pollinose. small 
central and lateral bare spots on 2-6. Hairs short 
white, rucumbent on dorsum, semierect on sides: five 
white lateral bristles on 1 plus a few longer hairs. 
Sternites grayish white pollinose. hairs short semierect 
white. Genitalia yellowish-red basally. brown apically: 
hairs long and white. 

Femora and tibiae yellowish-red: apical half of 
fore femora, apical fourth of middle and hind femora. 
and apical fifth of all tibiae, black; tarsi black. Hairs 
and bristles white; claws black, base reddish: pulvilli 
and empodia yellowish-red. 

Halteres yellowish, lower stem light brown. Wings 
hyaline: veins brown: anterior crossvein at 36 5" 
(.63) length of discal cell: third vein branched slightly 
beyond end of discal cell; posterior cells broadly open: 
anal cell narrowly open. 

Female. Length 10 mm. Weak bristle below only 
on antennal segment 2. Mesonotal pollen with a yel- 
lowish cast; two posterior and four anterior 
central bristles: two scutellar bristles. Abdominal seg- 
ments 1-6 and 7 basally. grayish pollinose. S bare: 
terminalia and spines reddish-brown, hairs yelli 



46 



III III ll\ sol illll<\ i \LIF0RN1A ACADEMY OF SCIENCES 



VOLUME 71 



II. iiis and bristles ol legs white to yellowish. Anterior 
crossvein ;ii 50 76 (.66) length of discal cell; third 
vein branched opposite end of discal cell. 

Holotype: male. New Mexico. Chaves Co.. 6 mi W 
Roswell, 15 September 1965 (P. H. Timbcrlake) 
VCR. Allotype: female. New Mexico. Eddy Co.. 
0.5 mi N Stale I inc. Hwy. 180 and 62. 2 October 
1962 U.S. Papp) VCR. 

Oilier specimens examined: NEW MEXICO. Eddy 
Co.. 9, 16 mi S Artesia. 3000-3500 ft. 24 September 
1950 (W. Gertsch and M. Cazier) AMNH [thorax 
and abdomen greased, ground color black except 
posterior lateral margins, and postalar calli of meso- 
nolum reddish: 3 scutellar bristles]: Torrance Co., 9, 
Gran Quivira, II August 1931 (R. H. Painter) 
RHP [third antennal segment missing and scutellum 
crushed]. TEXAS. Hudspeth Co., 9, 30 August 1940 
(D. I. Knull and J. N. Knull) OSU. 



KEY TO THE SPECIES OF W1LCOXIA 

1. Posterior margin of the scutellum bare of pollen; 

wings of the males brown, females lighter 
brown: abdomen black, lateral margins gray 
pollinose somewhat expanded apically on the 

tergites 2 

Scutellum all pollinose; wings hyaline; abdomen 
with at least some pollen on the dorsum of 
the tergites 3 

2. Legs black, basal fourth of fore, middle, and 

basal half of hind tibiae bright yellow, apices 
of fore and hind femora yellow; mesonotum 
gray pollinose. divided central stripe black, 
and small anterior intermediate spots brown; 
length 6.5-7.0 mm (Colorado) __ cinerea James 
Legs yellowish-red. basal two-thirds of femora, 
except extreme base, black; mesonotum 
densely gray pollinose. divided central stripe 
indistinctly brown; length 6-7 mm (Cali- 
fornia and Nevada) monae Wilcox 

3. Legs black, femora largely white pollinose and 

only the tips reddish; abdomen white pol- 
linose, central part of tergite 1, large central 
posterior spot and small lateral anterior spots 
on 2-6, bare of pollen: length 6-7 mm (Cali- 
fornia, Arizona, and Nevada) 

martinorum Wilcox 

At least the tibiae largely yellowish or reddish .. 4 

4. Femora except tips, tips of tibiae, and tips of 

tarsal segments brown or black, tips of 
femora, and basal part to tibiae and tarsi 
reddish: abdomen black, narrow lateral mar- 
gins of tergites 1-6. narrow anterior margins 
of male 2-6 and female 2-5, gray pollinose; 
length 5-6 mm (Arizona, New Mexico, and 

Utah) painter! Wilcox 

Femora except apical fourth and tibiae except 



tips, yellowish-red. apex of femora and tibiae, 
and all of tarsi, brown or black; abdomen 
black, densely gray pollinose. tergites 2-6 
with small lateral bare spots: length 8-11 mm 
(New Mexico and Texas) pollinosa Wilcox 



Metapogon amargosae, new species 

Description: Male. Length 8 mm. Head black, white 
pollinose, hairs and bristles white. Face at antennae 
22/28 (.79) width of one eye. Antennae black, 
grayish pollinose: hairs white, segments 1-2 each 
with a white bristle below; segments 10-10-35-19 in 
length. 

Mesonotum black; grayish white pollinose; broadly 
divided central stripe, small intermediate spots, and 
dorsocentral stripes broad anteriorly, brown. Hairs 
erect, white, and subequal in length lo antennae 1-2. 
Bristles white, 2-3 presutural, 1 supraalar, 1 postalar; 
bristles brown, four anterior and four posterior dorso- 
central and 12 anterior on the central stripe. Pleura 
and coxae grayish pollinose, hairs white. Scutellum 
white pollinose with a round central bare spot, hairs 
white, two strong, brown and two weak, pale mar- 
ginal bristles. 

Abdomen black; sides of tergites 1-3, anterior mar- 
gins of 2-7 and posterior corners of 2-6 extending 
narrowly inward on posterior margins of 2-4 (entire 
on 4). grayish-white pollinose. Hairs and bristles 
white, long on sides of 1-3. Sternites grayish pol- 
linose. 2-6 with small lateral bare spots, and a median, 
apical, triangular brownish spot, hairs white. Genitalia 
small, black, hairs white. 

Legs black, tips of femora and dorsum of tibiae 
brown. Hairs and bristles white; claws black, base 
reddish; empodia and pulvilli whitish, pulvilli 4/5 
length of claws. 

Halteres yellowish, base and lower part of knob 
brown. Wings hyaline with small, brown spots on the 
crossveins and furcation; veins brown; anterior cross- 
vein at 43/68 (.63) length of discal cell; third vein 
branched before end of discal cell and with short 
stump vein. 

Female. Length 9 mm. One weak, pale scutellar 
bristle (others probably broken off). Anterior fascia 
and posterior pollinose spots on abdominal tergites 
4-6 connected, 7 largely pollinose dorsally. 

Holotype: male. Nevada, Nye Co., 2 mi S Beatty, 
24 December 1964 (D. L. Coates) UI (CAS). 
Allotype: female same data, as holotype. 

This species runs to couplet 4 in the key to the 
species (Wilcox, 1964). In Metapogon pictus Cole 
and M. tricellus Wilcox the white hairs anteriorly on 
the central stripe are subequal in length to antennal 
segment 1. and there are no bristles on the anterior 
central stripe. In M. gibber (Williston) and M. 
carinattts Wilcox, the anterior hairs on the central 
stripe are usually black, but a few M. carinattts have 
white hairs; in M. gibber there are no anterior bristles 



1972 



NliW SPECIES <>!■ KOIiltl.K III/; 



47 



on t lie central stripe whereas, M. carina! us has about 
30 black bristles. 

Oilier specimens examined: NEVADA. Nye Co., 
2c5, 2 mi S Ueatty, 24 December 1964 (I). I .. Coates) 
UI, JW. ARIZONA. Maricopa Co., 7 , Tempo;- 21 
November 1967 (D. Plantz) ASU. CALIFORNIA. 
Riverside Co., 9, BIythe, 28 October 1940 (K. S, 
Snyder) Tamarix, CIS; .San Bernardino Co., $, 9, 
Needles, 20 March 1967 (R. M. Bohart and I). S. 
Horning, Jr.) UCD; 6c?, 3 9, Twentynine Palms, 23 
October, 6 November 1967 (E. Fisher) EF, [taken 
resting on the foliage of low shrubs on the sand dunes 
near the golf course about 2 miles northeast of town]. 



Metapogon obispae, new species 

Description: Male. Length 5 mm. Head black; 
grayish pollinose with a golden tinge on face, and 
central frons brown. Mystax with six brown oral and 
six black bristles above, plus a few short brown and 
black hairs; 3-4 short, black hairs on sides of frons; 
four strong, erect black bristles on ocellar tubercle; 
short occipital bristles black above and brown below; 
sparse beard and hairs on palpi and proboscis white. 
Face at antennae 18/22 (.82) width of one eye. 
Antennae black; short sparse hairs, and one bristle 
below on segment 1, black; segment 3 broader beyond 
middle; segments 9-9-30-9 in length. 

Mesonotum black; central stripe and intermediate 
spots dark brown, lateral margins golden brown, area 
between central stripe and intermediate spots, and a 
narrow median line, silvery pollinose. Sparse hairs 
black and subequal in length to antennae 1. Bristles 
black, 2 presutural, 1 supraalar, 1 postalar, 1 weak 
posterior and three strong anterior dorsocentral. 
Pleura and coxae black, grayish-brown pollinose; 
short hairs on coxae white; 5-6 long hypopleurals 
black. Scutellum black: silvery pollinose with golden 
tinge; four strong, black marginal bristles. 

Abdomen black; grayish pollinose with golden 
tinge, at some angles lateral, anterior black spots are 
apparent on tergites 2-5 and small round central spots 
on 2-6. Hairs short, sparse, recumbent, and black, 
white laterally with a few longer hairs on 1-2; four 
to five brown lateral bristles on 1. Sternites black, 
grayish-golden pollinose. short, sparse erect hairs 
white. Genitalia black, hairs white. 

Legs black, tips of femora and dorsum of tibiae 
dark reddish; hairs white; bristles brown; claws black; 
pulvilli dark brown, 4/5 length of claws; empodia 
brown. 

Halteres yellow, stem brown. Wings light brown, 
the brown intensified on the crossveins, furcations, 
and apex; veins black; anterior crossvein at 31/45 
(.69) length of discal cell; third vein branched slightly 
before end of discal cell and with long stump vein: 
posterior cells broadly open; anal cell narrowly open. 



Female. I ength 6 mm Mi dian line ■■! mi 

golden Abdi :ii i gmi ntt I gra i»h . ollinoi i 

K shining black; apical spines black, ihorl hairs while 

Holotype: male, California, San I I 

Haywood Park. 24 Octobei 1970 (I. Wilcox. 
Allotype: female, same data ;e. holotypc, 25 
1970. 

In the key lo the species (Wilco> 1964 i thi pecics 
runs lo couplet 10; with its lonj,' pulvilli it would I": 
close to M. tarsalus Wilcox from which it differs by 
having the abdomen all pollinose and is without ban; 
spots on the mesonotum. 

Oilier specimens examined: CALIFORNIA. San 
Luis Obispo Co., 41 $, 2X9. Baywood Park. 23 to 2^ 
October 1970 (J. Wilcox) CAS. [collected on Ifal 
ground and low weeds on the dunes overgrown with 
trees, brush, and weeds about two miles east of Morro 
Bay (Baywood Park P.O.) near the intersection of 
Nipomo Ave. and Willow Drive], 



ACKNOWLEDGMENTS 

1 am indebted to the following persons for the loan of 
specimens: J. G. Rozen. American Museum of Nat- 
ural History. AMNH: Mont A. Cazier, Arizona State 
University, ASU; Paul H. Arnaud. Jr.. California 
Academy of Sciences, CAS; M. S. Wasbauer. Cali- 
fornia Department of Agriculture. CDA: Charles H. 
Martin, Tucson, Arizona, CHM: J. A. Powell, Cali- 
fornia Insect Survey. University of California. Berke- 
ley, CIS; Eric Fisher, California State College. Long 
Beach, CSCLB, EF; Charles A. Triplehorn. Ohio 
State University. OSU; Elizabeth M. Painter. Kansas 
State University, RHP; Robert O. Schuster. Univer- 
sity of California, Davis. UCD; Saul I. Frommer. P. 
H. Timberlake, University of California. Riverside. 
UCR; William F. Barr. University of Idaho. UI: 
Willis W. Wirth. Entomology Research Division. Agr. 
Res. Serv., U.S.D.A., USNM. 



LITERATURE CITED 

James, M. T. 1941. The Robber Flies of Colorado 
(Diptera. Asilidae). J. Kansas Ent. Soc. 14: 

27-53. 

Wilcox, J. 1964. The Genus Metapogon (Diptera: 
Asilidae). Pan-Pacific Ent.. 40:191-200. 

Wilcox. J., and C. H. Martin. 1957. Backomyi; 
(Dipteria-Asilidae). a New Genus. J. K. 
Ent. Soc, 30:1-5. 



Accepted for publication October 21. 1 



RESEARCH NOTES 



A FIRST REPORT OF 

SELF-FERTILIZATION IN THE 

WOOD-BORING FAMILY TEREDINIDAE 

(MOLLUSCA: BIVALVIA) 



Three mechanisms for fertilization have been rec- 
ognized in the family Teredinidae: (1) Sex products 
are released into sea water where fertilization occurs 
as in Teredo (Nototcredo) norvagica (Lebour, 1938, 
1946; Nair. 1962). Teredo (Psiloteredo) megotara 
(Sigerfoos, 1908: Nair. 1962), Bankia sctacea (Coe, 
1941: Quayle. 1953). B. indica (Nair. 1956a, b). 
Nausitora dunlopei (Smith. 1963). B. australis 
(Turner. 1971), and perhaps B. goiddi (Sigerfoos. 
1908). (2) Sperm is released into sea water by the 
male and drawn into the incurrent siphon of a 
neighboring female where fertilization occurs in the 
mantle cavity as in Lyrodus pedicellatus (Roch, 
1940; Becker. 1959). L. diegensis ( = pedicellatus) 
(Kofoid and Miller, 1927), L. medilobata (Edmonson, 
1942). Teredo navalis (Grave, 1928), T. poculifer 
(Smith. 1963), and L. pedicellatus (cf. T. bartschi) 
(Isham and Tierney, 1953). Turner (1966) also re- 
ported T. furcifera, T. parksi (= sexually mature but 
small, young T. furcifera. Turner, 1966), T. somersi, 
T. clappi, L. affinis, and L. massa with brooded young 
from test panels, which proved that fertilization had 
occurred in the epibranchial cavities in these species. 
(3) Sperm possibly being transferred from one male 
to a neighboring female by the insertion of the male 
excurrent siphon into the female incurrent siphon as 
observed by Clapp (1951) in Bankia gouldi and by 
Townsley, Richy. and Trussell (1966) in B. setacea. 

No report of self-fertilization in the family has 
been made although Coe (1941) demonstrated that 
the sexual phases in Lyrodus and Teredo were not 
sharply demarcated and functional hermaphroditism 
in which the male and female stages occurred simul- 
taneously was common. However, there has been 
no laboratory evidence to suggest that self-fertilization 
in brooding forms is indeed a fact. 

During laboratory studies on the effects of tem- 
perature and reduced salinity on larvae and adults of 
L. pedicellatus (Eckelbarger and Reish. 1972), adults 
isolated from the veliger stage to an age of five months 
produced apparently normal veligers. Although the 
purpose of the experiment was to assess the effects of 
temperature and reduced salinity on growth and 
survival, the fact that all test animals were com- 
pletely isolated, one per flask from the pediveliger 
stage, yet produced larvae proves the existence of a 
mechanism for self-fertilization in this species. 



METHODS 

Lyrodus pedicellatus larvae were obtained by placing 
infested wood blocks brought from the field in room 
temperature sea water as a stimulant for the release 
of larvae. Actively swimming pediveligers were 
pipetted individually into separate petri dishes con- 
taining 75 ml of normal chlorinity (19.2 '/,.„) sea water 
and a presoaked block of Douglas fir measuring 
5 X 25 X 42 mm. All dishes were left undisturbed 
for a week in reduced light at room temperature. Only 
those boring larvae with normally appearing calcareous 
caps and visible siphons were selected for the experi- 
ment. 

A series of eight concentrations of sea water was 
prepared by dilution with appropriate volumes of 
distilled water; these included 19.3, 16.0, 13.0, 11.0, 
9.0, 7.0, 5.0, and 3.0 % . Chlorinities were determined 
by titration with silver nitrate (Barnes, 1959). A wood 
block containing one Lyrodus was placed in each of 
ten 500 ml erlenmeyer flasks with 100 ml of sea 
water at each chlorinity level. The flasks were sealed 
with a number seven rubber stopper. The experiment 
was conducted at two temperatures, one at room 
temperature (22-24 C) and the second at 14-16 C. 
Each wood block was removed from its flask and 
examined weekly under a dissection microscope. Ob- 
servations were noted on the appearance of the pal- 
lets, the calcareous cap, and the behavior of the ani- 
mal. The blocks were then returned to the flasks 
with fresh filtered sea water. At no time was sea 
water from one flask transferred to another in order 
to prevent the exchange of sperm between organisms. 

The experiment was terminated at the end of 5 
months at which time all animals were removed from 
their blocks. The mantle cavity and gill filaments were 
examined for the presence of mature veligers and were 
counted if present. 

RESULTS 

Table 1 indicates that larvae were found in 40, 25, 
and 11% of the animals in chlorinities of 19.3, 16.0, 
and 13.0 % , respectively, at 14—16 C. Larvae were 
found in 30 and 20% of the animals at 19.3 and 
16.0 % c , respectively, at 22-24 C. Animals raised at 
14—16 C contained 49 larvae as compared to 25 at 
22-24 C. 



DISCUSSION 

Protandry in teredinids was proved by Yonge ( 1926) 
from histological work on Teredo (Nototcredo) 
norvagica although it had been earlier postulated by 
Sigerfoos (1908) in Bankia gouldi. Coe (1933-41) 



48 



1972 



RESEARCH NOII-.S 



19 



' I a it 1 1 I. The effects of temperature and reduced 

chlorinity on survival and reproduction of Lyroclus 

pedicellatus. 



Chlorinity 


No. of Sur- 
viving I est 
Animals 


No 
Vc 


Willi 
Igers 


No. of Siir- 

vivni" 1 ,' II 

Animals 


No'. With 

Vcligcrs 


14- 


li, i 






22-24 C 




19.3 


II) 




4 




10 




3 


16.0 


8 




2 




10 




2 


13.0 


9 




1 




8 







11.0 


4 









4 







9.0 












1 








found Teredo /nivalis, li. setacea, and L. diegensis 
(— L. pedicellatus) all' to be protandrous hermaph- 
rodites. He found the sexual phases not sharply 
demarcated in L. diegensis, and was able to achieve 
artificial fertilization and development through gastru- 
lation. In the experiments presented herein, direct 
evidence is given proving that at least one species of 
teredinid is capable of self-fertilization. 

The reduction in animals producing larvae at lower 
chlorinity levels was an expected result since the 
species is sensitive to reduced chlorinities (Eckel- 
barger and Reish, 1972). Kinne (1963) confirmed 
that embryonic and larval stages are almost universally 
more sensitive than adults of the same species. 

The greater number of larvae produced at the lower 
temperature level may reflect a lower sensitivity to 
reduced chlorinity conditions as a result of the lower 
temperature. It maybe that there is increased maternal 
retention of larvae at lower temperatures. Coe (1941) 
stated that this species could retain its larvae within 
the maternal gill for several months until more favor- 
able temperatures. Since the 14-16 C range corre- 
sponded to the seasonal low reported by Moore and 
Reish (1969) for Alamitos Bay, Long Beach, and 
for Los Angeles-Long Beach Harbors by Menzies. 
Mohr, and Wakeman (1963), it is possible that the 
higher larval count reflected this maternal retention. 



ACKNOWLEDGMENTS 

The authors wish to thank Ruth D. Turner, Harvard 
University, for her critical review of the manuscript 
and many helpful suggestions, and John L. Culliney 
for his constructive criticisms. 



LITERATURE CITED 

Barnes, H. 1959. Apparatus and methods of 
oceanography. Part one: Chemical. Intersci. 
Publ. Inc., New York. 341 pp. 

Becker, G. 1959. Biological investigations on marine 



borers in Berlin Dahlcm, In D i I 

Murine boring and foil 

Washington Prci . Seattle pp. 62 76. 

( lapp, w. I . 195 1. Observations on I 

dinidae. Fourth Progn Report 'Kept. no. 
7550), William I. Clapp Laboratori i 

Duxbury, Massachusetts. 9 pp. 

Coe, W. K. 1933. Sexual phases in Tcrcdi 
Bull., 65:293-303. 

. 1934. Sexual rhythm in the pclccypod 

mollusk Teredo. Science, 80:192. 

. 1936. Sequence of functional sexual phases 

in Teredo. Biol. Bull., 71:122-133. 

. 1941. Sexual phases in wood-boring mol- 



lusks. Biol. Bull., 81:168-176. 

Eckelbarger. K. L. and D. J. Reish. 1972. The ef- 
fects of temperature and reduced salinity on the 
settlement, growth, and reproduction of the wood- 
boring pelecypod, Lyrodus pedicellatus Quatre- 
fages. Submitted manuscript. 



Edmonson, C. H. 1942. Teredinidae in 
Occas. Pap. Bishop Mus.. 17:97-150. 



Hawaii. 



Grave. B. H. 1928. Natural history of the ship- 
worm, Teredo naralis. at Wood's Hole. Massachu- 
setts. Biol. Bull.. 55:260-282. 

Isham, L. B.. and J. Q. Tierney. 1953. Some aspects 
of larval development and metamorphosis of 
Teredo {Lyrodus) pediccllata de Quatrefages. 
Bull. Mar. Sci. Gulf and Caribbean. 2:574-589. 

Kinne. O. 1963. The effects of temperature and 
salinity on marine and brackish water animals. I. 
Temperature. Oceanogr. Mar. Biol., Ann. Rev.. 
2:281-339. 

Kofoid. C. A., and R. C. Miller. 1927. Biological 
Section. In: C. L. Hill and C. A. Kofoid. [Eds.]. 
Marine borers and their relation to marine con- 
struction on the Pacific coast. San Francisco Bay 
Marine Piling Comm.. San Francisco, pp. 188— 
343. 

Lebour. M. W. 193S. Notes on the breeding of 
some lamellibranchs from Plymouth and their 
larvae. J. Mar. Biol. Assoc. U.K. 23:135-138. 

. 1946. The species of Teredo from Plym- 



50 



Hi III il\ \c>l llli:i<\ (AIIIORNIA ACADEMY Or SCIENCES 



VOLUME 71 



ouili waters. J. Mar. Biol. Assoc. U.K.. 26: 
381 389. 

Menzies, R. J.. I. Mohr, and C. M. Wakeman. 1963. 
I ho seasonal settlement of wood-borers in Los 
Angeles-Long Bench Harbors. Wasmann J. Biol., 
21:97-120. 

Moore, D. R., and D. J. Reish. 1969. Studies on 
the Mytilus edulis community in Alnmitos Bay. 
California. IV. Seasonal variation in gametes 
from different regions of the bay. The Veliger. 
11:250-255. 

Nair, N. B. 1956a. Sex changes in the wood-boring 
pelecypod, Bankia indica Nair. J. Madras Univ.. 
Sect. 13. 26:277-280. 

— . 1956b. The development of the wood- 



boring pelecypod. Bankia indica Nair. J. Madras 
Univ.. Sect. 13. 26:303-318. 

— . 1962. Ecology of marine fouling and 



wood-boring organisms of western Norway. 
Sarsia. No. 8:1-88. 

Quayle, D. B. 1953. The larvae of Bankia setacea 
Tryon. British Columbia Dept. Fish. Rept., 
1951:88-91. 

Roch. F. 1940. Die Terediniden des Mittelmeeres. 
Thalassia, 4 (3), 147 pp. 

Sigerfoos, C. P. 1908. Natural history, organiza- 
tion, and late development of the Teredinidae, 
or shipworms. Bull. U. S. Bureau Fish., 37: 
191-231. 

Smith. M. L. 1963. The Teredinidae of the Queens- 
land coast from Cairns to Brisbane, (unpublished 
thesis). Zoology, Univ. Queensland. Brisbane, 
206 pp. 

Townsley, P. M.. R. A. Richy, and P. C. Trussell. 
1966. The laboratory rearing of the shipworm, 
Bankia setacea (Tryon). Proc. Nat. Shellfish. 
Assoc, 56:49-52. 

Turner. R. D. 1966. A survey and illustrated 
catalogue of the teredinidae (Molluscs: Bivalvia), 
Mus. Comp. Zool., Harvard Univ., 265 pp. 

. Australian shipworms. Australian Nat. Hist. 



Kevin J. E( KEl iiakgek. Marine Science Institute, 
Northeastern University. Nahant, Massachusetts 
0190$. and Donald J. Reish. Dept. Biology, Cali- 
fornia State College, Long Beach, California, 
90801. 

Accepted for publication December 1, 1971. 



(In press.) 

Yonge. C. M. 1926. Protandry in Teredo norvegica. 
Quart. J. Micros. Sci.. 70:391-394. 



RHADINAEA GODMANI, AN ADDITION 
TO THE SNAKE FAUNA OF HONDURAS 



Rhad'maca godmani (Giinther) has been recorded 
from Guatemala (Slevin, 1939; Stuart, 1951, 1963), 
El Salvador (Mertens, 1952a. 1952b; Uzzell and 
Starrett. 1958). and Costa Rica (Scott, 1969). Re- 
cent field work by us has revealed its presence in 
Honduras. On the afternoon of 9 April 1971 we 
secured one male specimen (LSUMZ 24398) from 
underneath a rock near a log slide at an elevation of 
1740 m in Piiuts pseitdoslrobus forest (pinabetal of 
Carr, 1950; Lower Montane Moist Forest formation 
of Holdridge, 1962) on Cerro Uyuca, Depto. de 
Francisco Morazan. On 29 April 1971 an additional 
specimen (LACM 72070) was found by Eric and 
Susan Festin at an elevation of 1450 m in Pinus 
oocarpa forest (ocotal of Carr, 1950; Subtropical 
Moist Forest formation of Holdridge, 1962) at El 
Hatillo, Depto. Francisco Morazan. 

Both specimens are males exhibiting the following 
measurements and features of scutellation (data for 
LSUMZ 24398 listed first): ventrals 159, 156; sub- 
caudals 92, 92; supralabials 8-8 in both; infralabials 
9-9 in both; dorsal scale rows 19-21-21 in both; 
total length 405, 407 mm; tail length 122, 129. Other 
scutellational data are typical for the species. 

In life LACM 72070 exhibited the following color 
and pattern; rows 1 through 3 yellowish-orange, adja- 
cent portions of rows 1 and 2 and rows 2 and 3 with 
black flecking giving impression of narrow, diffuse 
stripes; lower portion of row 4 light yellow; upper 
portion of row 4. most of row 5, and lower portion 
of row 6 black; upper portion of row 6 and lower 
edge of row 7 light yellow; ground color of upper two- 
thirds of row 7 through row 10 medium brown, 
edges of these scales black giving impression of nar- 
row, irregular, diffuse stripes on adjacent portions of 
scale rows; middorsal row black lightly mottled with 
brown. 

Dorsum of head dark brown with faint light yellow 
mottling and a tiny, but distinct, light yellow dot in 
middle of each parietal; light yellow band across 
snout on posterior portion of rostral and anterior por- 
tion of internasals, in connection with light pigment 



1972 



RESEARCH NOTl S 






on supralabials; supralabials mostly ivory, supralabials 
I through 4 wiih dark brown upper and lower edges, 
dark blotch on upper portion of supralabial 6 cx- 
tending diagonally across lower portion of supralabial 
6, upper edge of Supralabial 6. upper half of 7. and 
all but anteroventral corner of 8 dark brown (the 
dark diagonal subocular blotch and the dark pigment 
on supralabials 6 through 8 appear to outline a pule 
diagonal stripe from eye lo lip); infralabials ivory 
with dark brown mottling on anterior ones; chin 
ivory; iris dark brown. The oilier specimen (LSUMZ 
24398) agrees in essential aspects with the above 
description. 

Keels are present on the dorsal scales above the 
vent in l.ACM 72070 but not in LSUMZ 24398. 
Mertens (1952a) described Rhadinaea zilchi on the 
basis of a single specimen from HI Salvador, distin- 
guishing this taxon from Rhadinaea godmani on the 
basis of differences in dorsal color pattern and Ihe 
presence of supra-anal keels in the former. In a 
later paper Mertens (1952b) pointed out that there 
was actually no difference in color pattern between 
godmani and zilchi. The differences given in the 
original description of zilchi were based on an error 
in the pattern representation of godmani given by 
Stuart and Bailey ( 1941 ). an error later corrected by 
Stuart (1951). Mertens (1952b) suggested that since 
the presence of supra-anal keels in zilchi and their 
absence in godmani served to distinguished the two. 
that zilchi be regarded as a subspecies of godmani. 
Two additional adult male R. godmani (from El 
Salvador) discussed by Uzzell and Starrett (1958). 
possess well-developed supra-anal keels, whereas, a 
topotypic Guatemalan specimen of the nominate sub- 
species has supra-anal keels that are less well-de- 
veloped. They suggested that the differences between 
godmani and zilchi "seem too slight to be of taxonomic 
importance." However. Merten's view was accepted 
by Peters and Orejas-Miranda I 1970) along with 
the erroneous conception that pattern differences exist 
between the two. Inasmuch as the name zilchi is 
based upon only two specimens and the character of 
supra-anal keeling appears to vary intrapopulationally. 
we suggest that Rhadinaea zilchi Mertens be regarded 
as a synonym of Rhadinaea godmani (Giinther ) 
1865. 

Scott (1969) recorded Rhadinaea godmani from 
several localities in Costa Rica, apparently the first 
records for the species in that country. Taylor (1951. 
1954) did not list R. godmani as a member of the 
Costa Rican snake fauna, but he (Taylor, 1954) did 
describe a new species. Rhadinaea altamontana, on 
the basis of one specimen from the Cordillera de 
Talamanca in Costa Rica. In this paper, he regarded 
altamontana as most closely related to godmani but 
stated that the two could be distinguished on the basis 
of differences in color pattern, head length, and 
supra-anal keeling (keels present in altamontana. keels 
not present in godmani). We have not examined the 
holotype of altamontana but we discount the impor- 



i; ■ of supra anal I •■• ling and poinl i 

description ol the coloi pattern ol Ihc holi 

very similar (bm nol identical, ■ rding 

description i io the coloi pattern Wc 

suggest thai Rhadinaea altamontana I prob- 

ably a synonym ol H qodmani and thus support the 
implication ol Scotl (1969) lhal altamontana 
recognizable. 

Presently, Rhadinaea godmani ii known ; 
at moderate and intermediate elevations (1450 
in) ol both versants from southwestern Guatemala lo 
Costa Rica; il has thus far nol been recorded from 
Nicaragua. 

LITERATURE (NED 

Carr, A. F.. Jr. 1950. Outline for ;i classification 
of animal habitats in Honduras. Bull. Arncr M 
Nat. Hist.. 94:567-594. 

Holdridge. L. R. 1962. Mapa ecologico dc Hon- 
duras. Organizacion de los Estados Americanos. 
San Jose, Costa Rica. 

Mertens. R. 1952a. Neues iibcr die Rcptilicnfauna 
von El Salvador. Zoologische Anzciger. Leipzig, 
148:87-93. 

. 1952b. Die Amphibien und Reptilien von 



El Salvador. Ahhandlungen der Senckenber- 
gischen Naturforschenden Gesellschaft. Frank- 
furt am Main. 487:1-120. 

Peters. J. A., and B. Orejas-Miranda. 1970. Cata- 
logue of the Neotropical Squamata: Part I. 
Snakes. Bull. U. S. Nat. Mus.. 297:1-347. 

Scott. N. J.. Jr. 1969. A zoogeographic analysis of 
the snakes of Costa Rica. Ph.D. Thesis. Univ. 
Southern California. 390 pp. 

Slevin. J. R. 1939. Notes on a collection of reptiles 
and amphibians from Guatemala. I. Snakes. Proc. 
California Acad. Sci.. ser. 4. 23:393-114. 

Stuart. L. C. 1951. The herpetofauna of the Guate- 
malan Plateau, with special reference to its 
distribution on the southwestern highlands. Con- 
trib. Lab. Vert. Biol.. Univ. Michigan. 45:1-71. 

. 1963. A checklist of the herpetofauna of 

Guatemala. Misc. Publ. Mus. Zool., Univ. Michi- 
gan. 122:1-150. 

Stuart. L. C. and J. R. Bailey. 1941. Three new- 
species of the genus Rhadinaea from Guatemala. 
Occas. Pap. Mus. Zool.. Univ. Mich 
1-11. 



5? 



BULLETIN SOUTHERN CALIFORNIA ACADEMY (>E SCIENCES 



VOLUME 71 



raylor, III 1951. A brief review of the snakes 

Of Costa Rica. Univ. Kansas Sci. Bull., .14:3-188. 
. 1954. Further studies on the serpents of 



Costa Rica. Univ. Kansas Sci. Bull.. 36:673-800. 

Uzzell, T. M.. Jr.. and P. Starrett. 1958. Snakes 
from 11 Salvador. Copeia, 1958:339-342. 

Larry David Wilson, Horizon School, 9025 
Sunset Drive. Miami, Florida 33143, and John R. 
Meyer. The Research Ranch. Elgin. Arizona 
85611. 

Accepted for publication October 21, 1971. 



A NEW SPECIES OF 

BAETODES FROM ARIZONA 

(EPHEMEROPTERA: BAETIDAE) 

The first record of Baclodes in North America, north 
of Mexico, was reported by Edmunds (Ent. News, 
61:203-205, 1950) from nymphs collected by J. G. 
Needham in the Rio Frio, Garner State Park, Uvalde 
Co.. Texas. An assemblage of mayfly nymphs from 
Arizona were found to belong to an undescribed species 
of Bactodes, representing the second record of the 
genus for North America, north of Mexico. The 
holotype is deposited in the California Academy of 
Sciences. We thank Jerry Battagliotti for preparing the 
included figures. The research upon which this report 
is based was supported by the National Science 
Foundation. 



reddish-brown; tibiae brown; tarsi brown to dark 
brown: coxae without gills: tarsal claws with 6-7 
denticles. Abdominal terga light brown with reddish- 
brown markings; abdominal terga 1-9 with a reddish- 
brown transverse band: abdominal terga 6-10 with a 
median pale stripe, and often with submedian pale 
spots; abdominal tergum 10 light brown; abdominal 
terga 1-6 with small median tubercles, tubercle on 
tergum 6 barely discernable (Fig. 1); abdominal terga 
without setae; abdominal gills pale; abdominal sterna 
light brown with reddish brown markings; abdominal 
sterna 1-7 with an anterior transverse reddish brown 
band (band often disjunct on terga 5-8); abdominal 
sternum 9 pale (Fig. 2). Caudal filaments light brown. 

Holotype: mature male nymph, Tonto Creek at 
Kohl's Ranch. Gila County, Arizona, 19 July 1970, 
R. K. Allen. Paratopotypes: 1 male and 1 female 
nymph, same data as holotype, in collection of Cali- 
fornia State College, Los Angeles. 

Remarks: The only described species in the genus 
are known from South America. Baetodes serratus 
Needham and Murphy and B. itatiayanus Demoulin 
were described from Brazil, and B. spiniferum Traver 
is known only from Venezuela. Baetodes sigillatus 
would seem to be distinguished from the other de- 
scribed species by distribution alone as there are no 
records of North American species of mayflies oc- 
curring in South America. The nymphs of this species 
are further distinguished by possessing small dorsal 
median tubercles on abdominal segments 1-6 (Fig. 
1), and by the presence of transverse reddish-brown 
bands on abdominal sterna 1-7 (Fig. 2). 




Baetodes sigillatus, new species 
Figures 1 and 2 

Description: Nymph. — Length: body 5.0-6.0 mm; 
caudal filaments 4.0-5.0 mm. General color light 
brown with reddish-brown markings. Head light 
brown with brown and reddish-brown markings; 
occiput brown, frons and genae reddish-brown; head 
without setae. Thoracic nota light brown with 
brown and reddish-brown markings; pronotum light 
brown with intricate brown and reddish-brown mark- 
ings; mesonotum light brown with reddish-brown an- 
terior transverse band, and with submedian longitudi- 
nal brown streaks; pronotum with a dorsal, median, 
posterior elevation; mesonotum without a dorsal 
elevation; thoracic nota without setae; thoracic sterna 
pale: legs light brown to brown with reddish-brown 
markings: femora light brown often margined with 




Figures 1-2. The nymph of Baetodes sigillatus, new 
species: Fig. 1, abdominal terga, lateral view; Fig. 2, 
abdominal sterna, ventral view. 



Richard K. Allen and Esther S. M. Chao, Dept. 
Biology, California State College, Los Angeles, 
California 90032. 

Accepted for publication October 21, 1971. 



1972 



RESEARCH NOTl S 






AN INI RODUCLI) POPULATION OP 

SOCIAL WASPS, POLISTES APACHUS, 

THAT HAS PERSISTED FOR 10 YEARS 

(HYMENOPTERA: VESPIDAE) 



The large social wusp Polistes apachus Saussure often 
builds its nests under the eaves of houses. This speeies 
has a typical life cycle for wasps of the genus. Over- 
wintering queens, usually in groups of two to four, 
establish nests timing April. Workers are produced 
until about midsummer, and then increasing numbers 
of sexual forms are produced. Observations made in 
this study indicate that nests terminate in September 
and October. When a nest terminates, reproduction 
ceases, the workers and males die, and the current 
season's queens eventually leave the comb. Polistes 
apachus, like all social wasps in North America, has 
an annual life cycle, and only the newly produced 
generation of queens survives the winter. There is 
some evidence that queens emerging from hiberna- 
tion have a strong tendency to initiate nests in the 
area of the nest of the previous season. Using marked 
queens, Eberhard (Misc. Pub. Mus. Zool., Univ. Michi- 
gan, 140, 1969) found no movement of marked in- 
dividuals of P. fuscatus (Fabricius) between neigh- 
boring buildings over a period of two years. This 
note reports the artificial establishment of a popula- 
tion of P. apachus that has persisted in the same gen- 
eral locality for 10 summers. 

Ten nests of P. apachus were removed from the 
eaves of a house in Chatsworth. California during 
the middle of June, 1962. They were removed during 
the period when most nests have a few capped cells 
containing pupae, but very few adult workers. One 
nest appeared to have produced two workers, as two 
wasps were noticeably smaller than the others and 
two of the central cells were being reused. Each of 
the nests had at least one, and as many as five, wasps 
in attendance. 

The wasp nests were transferred to wooden boxes. 
Each box was 14 X 18 X 30 cm. had a large plastic 
window and a hinged screen door, and was provided 
with a small petri dish of sucrose and water. Each 
nest was fixed to the roof of a box by a drop of 
polyvinyl chloride glue placed on the nest pedicel. 
The wasps were introduced into the boxes after the 
glue had dried. 

A method similar to that described by Rabb and 
Lawson (J. Econ. Ent., 50:778-784. 1957) was used 
to relocate the wasp nests. The boxes were placed 
in the desired location and left there for three days. 
After this period, the screen door of each box was 
opened, allowing the wasps to resume normal for- 
aging activities. In the present study, the new nest 
site was the roof of the science building of San 
Fernando Valley State College in Northridge. Cali- 



I ia. I his site wa 

nestS of any species had been found in lh<- I 

overhangs ol ;my of the buildii 

because nests on the root of the science building 

nol be disturbed. 'Ihe ncsl boxes wci <■ in a 

row to the top oi a safety wall on the pi 
the roofs. Each box was about io cm from an) 

box. The new ncsl site was 4 miles east and 1.3 miles 
south of the original location in ' hatsworth, 

After the Hi nests were relocated, they wen 
served through the summer and fall. All ten nests 
increased in si/c. produced sexual forms, and ter- 
minated in the fall. During the following spring 
( 1963), /'. apachus queens were observed flying about 
in the vicinity of the science building, and a number 
of new nests were started under an eavc-likc overhang 
immediately below the old nest boxes, this was the 
first time that nests of any Polistes species had been 
found on any building on campus. 

Annual checks for active P. apachus nests on the 
science building and on two nearby buildings of simi- 
lar structure were made each summer from 1962 to 
1969. A final check, made in 1971. included the 
science building, the two nearby buildings, and four 
other similar buildings in the area. A tract of bunga- 
lows near the science building, included in the annual 
check until 1965. was also checked in 1971. Each 
year at least one nest was located in the immediate 
vicinity of the old nest boxes. More nests were found 
suspended from overhangs on other parts of the 
building, usually on the south side at the first floor 
level. For ten seasons, no P. apachus nests have been 
observed on the two nearby buildings. However, the 
bungalows, which had been free of Polistes nests in 
1965. had a large P. apachus population in 1971. 

This study confirms Eberhard"s (1969) observations 
on the strong tendency of Polistes queens to start nests 
in the vicinity of the nest of the previous season. 
After ten seasons on the San Fernando Valley Slate 
campus. P. apachus is still restricted to the science 
building and to a nearby tract of bungalows. Con- 
sequently, some Polistes species are able to form pop- 
ulations that can persist for many generations in a 
very restricted locality, despite the annual termination 
of individual nests. 



ACKNOWLEDGMENTS 

I wish to thank Philip J. Pointing for criticizing the 
manuscript. I would also like to thank Peter Bellinger 
for encouraging the research and for obtaining ap- 
proval from the college for placement of the nest 
boxes. 

David L. Gibo. Erindale College. Universal 
Toronto. Clarkson. Ontario. 

Accepted for publication February S. 1 



.w 



in 1 1 1 1 1\ sot I ui:i<\ / < u.ii-'ORNIA l< i/»/ Wl OF SCIENCES 



VOLUME 71 



I III I'm I I OSIOMAI1D BAT. 

VAMPYKI-.SS . I BROCKI, 

IN COLOMBIA 



I ho species Vampyressa brocki was described from 
Guyana by Peterson (Life Sci. Contrib.. Royal On- 
tario Mus.. 73:1-17. 1968) on the basis of a single 
female. Subsequently, a male was obtained in Gu- 
yana (Peterson. Canadian Jour. Zool., 50:in press, 
1972) and the species presently is known only from 
these two specimens. Between 28 June and 1 July 
1969. while conducting studies on the karyotypes of 
phyllostomatid bats, we collected three additional 
specimens of this species in mature tropical rainforest 
at Leticia. Amazonas. Colombia. These bats were 
taken in mist nets at a height of no more than 3.5 
meters over footpaths through the forest. 

The measurements of our specimens (all adult 
females) TTU 8827, TTU 8832. and TTU 9047. re- 
spectively, as follows: Length of forearm, 35.4, 32.1, 
33.2: length of hind foot, 8.5, 8.5, 9.5: length of ear, 
I 1.0. 11.0, 11.5: greatest length of skull (including 
incisors). 18.4, 18.3. 18.4; condylobasal length. 16.0, 
15.8. 16.2: zygomatic breadth. 10.9, 10.8. 10.7: 
breadth of braincase, 8.4. 8.4, 8.0; mastoid breadth. 
9.4, 9.4. 9.5: interorbital constriction, 4.6, 4.6, 4.9; 
postorbital breadth, 4.9, 4.7, 5.1; breadth across 
molars (Ml-Ml). 7.9. 7.6, 7.8: breadth across ca- 
nines (C-C). 4.2. 4.2. 4.3: length of maxillary tooth- 
row (C-M2). 5.7. 5.7, 5.7: palatal length (including 
incisors), 8.0, 8.2, 8.0; length of mandibles (con- 
dylo-incisive), 11.5, 11.5, 11.0; height of ramus, 3.7, 
3.6, 3.7. 

In all measurements, our females closely resemble 
those given for Guyanan specimens by Peterson 
(1968. 1972). However, in the key provided by 
Peterson (1968:13). bats with a forearm greater 
than 34 mm would be identified as Vampyressa 
nymphaea. One of our specimens has a forearm 
length of 35.4 but its greatest length of skull (18.4) 
identifies it as V. brocki, which is distinctly smaller 
than V. nymphaea in cranial dimensions. 

In other characters, such as number of lower 
incisors, size and shape of lower premolars and m2. 
and absence of m3, our specimens are identical with 
those reported by Peterson. A dorsal stripe is pres- 
ent, but faint, in our specimens. The facial markings 
appear to be somewhat less distinct than those of 
the holotype as illustrated by Peterson (1968). 

The karyotype of V. brocki is shown in figure 1. 
The diploid number is 24 and the fundamental num- 
ber is 44. This karytoype is similar to that of V. 
nymphaea from Nicaragua and Honduras, which 
have a diploid number of 26 and a fundamental 
number of 48. Vampyressa pusilla from Leticia, Co- 
lombia, have a diploid number of 23 (males) or 
24 (females), with a fundamental number of 24. 
Although the degree of chromosomal divergence be- 



nxHmxvxx 

Aft hh II Aft 

Figure I . Representative karyotype of a female Vam- 
pyressa brocki from Leticia, Colombia. 



tween V. brocki and V. nymphaea does not neces- 
sarily imply that the two taxa are specifically distinct, 
such divergence is typical of populations representing 
different species in the family Phyllostomatidae. Fur- 
ther, these chromosomal data suggest that V. brocki 
and V. nymphaea are more closely related to each 
other than either is to V . pusilla. These relationships 
were suggested by Peterson (1972). 

Two females were pregnant when taken on 30 June 
and 1 July. The single embryo in each was minute. 
None of the three specimens evinced molt. 

We thank C. J. Marinkelle of the Universidad de 
Los Andes for assistance during this study. 

Robert J. Baker and Hugh H. Genoways, Dept. 
Biology and The Museum. Texas Tech University, 
Lubbock, Texas 79409, and Alberto Cadena, Mu- 
seum of Natural History, The University of Kansas, 
Lawrence, Kansas 66044. 

Accepted for publication April 1. 1972. 



TEMPERATURE CHANGES IN HEAT 
PRODUCING PLANTS 



Heat production in aroid plant flowers was first 
noticed by Lamarck (Thomas, 1960) and at present 
the details of the biochemical and physiological 
mechanisms involved are well documented (Dormer, 
1960; Fischer, 1960; Forward, I960; Hackett, 1957; 
James and Beevers. 1950; James and Clapham, 1935; 
Meeuse. 1966). There are no records of the daily 
cycle of temperature within the spadix. Although a 
few temperatures have been recorded, there is no in- 
dication as to the magnitude of heat production, nor 
has there been much work reported on tropical aroids. 
The cycle of temperature changes during flowering 
of Philodendron selloum on the campus of the Uni- 
versity of California, Los Angeles, and in my own 
yard, Anaheim, California were recorded. All plants 
were in the shade for most or all of the day. Tem- 
peratures of 28 spadixes were taken with a Schult- 



1972 



KICSIiAHCIl NOTl 5 






'♦-♦SPATHE CL03ED-* 



"•DROsWhILA 




7 12 IB 24 


6 12 IB 24 


6 12 18 


JULY 10 


JULY 1 1 


JULY 12 


DAYI 


DAY 2 


DAY 3 



Figure I. Spadix, leaf, and ambient temperatures of 
a single flower (17) of Philodendron selloum. 



45 1 



40' 



35' 



^ 25 J 



15 



-.i-/i r hi m-iic 



M 



M«'xi 

'.< ON D 



MU / MM ' 




t . ■ . 
[UOHCI 



MMWPH » 



!< 



DAY I 



Figure 2. Summary of spadix and ambient temper- 
atures of 28 Philodendron selloum flowering cycles. 
Vertical lines indicate range and dots indicate means. 



heis quick recording thermometer or with a YSI 
Telethermometer thermistor Model 44TD using vinyl 
402 or banjo probes. Temperature readings were 
taken as follows: spadix tip. within center of spadix, 
1" below tip; spadix base, within spadix 2" below 
beginning of female flowers; air temperature, within 
spathe but 1" away from and 2" above bottom of 
spadix; and leaf, 3" inside leaf base and 3" below 
spadix. Morning temperatures were taken between 
0830-0930, afternoon: 1300-1700, night: 2000- 
2200 his. 

The cycle for a typical flower is shown in figure 
1. A summary of all temperatures is shown in fig- 
ure 2. At no time was the ambient temperature, 
measured at spathe base, above 26.7°C. Air temper- 
atures at night at UCLA were usually below 20°C 
with fog present, yet a maximum temperature of 
42.5" (AT: 18.5°C) was recorded within the spadix 
base, a figure 24°C above ambient. Thus (Fig. 2) 
the spadix temperature is rising while the ambient is 
falling. The tip of the spadix begins to rise in tem- 
perature before the base, but the latter reaches a 
higher maximum temperature. 

As noted by others (Dormer, 1960: James and 
Clapham, 1935) the emergence of the flowers from 
the solid spadix gives a shaggy appearance to the 
spadix. Flower emergence follows the night after 
maximum temperature is reached (night of day 2). 
This is followed by the beginning of a strong odor 
and the appearance of Drosopliila. The flies crawl 
about the flower, presumably pollinating the flowers. 
The spathe subsequently closes accompanied by a 
drop in flower temperature, an increase in the odor, 
and the decomposition of the flower. The Drosopli- 
ila are present within the closed spathe for several 
days after closing and presumably lay eggs in the 
decaying flower. 

I wish to thank Allen Strickler. Beckman Instru- 



ment Co., for calling my attention to this problem. 
Portions of this work were supported by the Na- 
tional Science Foundation (GB-2307, and a Senior 
Postdoctoral Fellowship, 56017). 



LITERATURE CITED 

Dormer, K. J. 1960. The Truth about Pollination 
in Arum. New Phytol.. 59:298-301. 

Fischer, H. 1960. Atmung von Bliiten und Bliiten- 
standen. Encyc. Plant Physiol.. 12:521-535. 

Forward, D. F. 1960. Effect of Temperature on 
Respiration. Encyc. Plant Physiol.. 12:235-258. 

Hackett. D. P. 1957. Respiratory Mechanisms in 
the Aroid Spadix. J. Exp. Bot.. 8:157-171. 

James, W. O., and H. Beevers. 1950. The respira- 
tion of Arum spadix: a rapid respiration, re- 
sistant to cyanide. New Phytol.. 49:353-374. 

James. W. O.. and A. R. Clapham. 1935. The 
Biology of Flowers. Clarendon Press. Oxford. 
1-115 pp. 

Meeuse. B. J. D. 1966. The Voodoo Lily. Sci. 
Amer.. 215:S0-88. 

Thomas. M. 1960. History of Plant Respiration. 
Encyc. Plant Phvsiol.. 12:1-46. 



Bayard H. Brattstrom, Dept. Biology 
State University. Fullerton. California 9263 1. 

Accepted for publication April 5. ' 72 



56 



!:i II.ET1N SOI THERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



RANGE EXTENSIONS OF PORCELAIN 

AND HERMIT CRABS IN THE 

GULF OF CALIFORNIA 



During 1970-1972 the senior author made numerous 
collections of interticlal invertebrates throughout the 
Gulf of California. From examination of these col- 
lections and one by Alex Kerstitch (University of 
Arizona) the following range extensions have been 
recorded. 



Section Anomura 

Superfamily Galatheidea 

Family Porcellanidae 

Petrolisthes edwardsii (Saussure) 

Former northernmost records: Bahia de Santa 
Maria and Bahia de la Magdalena, west coast of Baja 
California, and Los Frailes. near the mouth of the 
Gulf of California (Haig. Zoologica, 53:57-74, 1968). 

New record: Puerto Penasco, Sonora, Mexico, near 
the head of the Gulf of California: under stones in the 
mid-intertidal region: November 14. 1970; R. C. 
Brusca. 

Petrolisthes sanfelipensis Glassell 

Former southernmost record for west coast of 
mainland Mexico: Guaymas, Sonora, Mexico (Haig, 
Hopkins, and Scanland, Trans. San Diego Soc. Nat. 
Hist., 16(2):13-32, 1970). 



New record: Cabo San Lucas, Baja California Sur, 
Mexico; November 28. 1968: Alex Kerstitch. 

Petrolisthes hirtispinosus Lockington 

Former northernmost record: Bahia de Tepoca, 
Sonora, Mexico (Haig, Allan Hancock Pac. Exp. 24: 
1-440, 1960). 

New record: Puerto Penasco, Sonora, Mexico; 
under stones in the mid-intertidal region; November 
14. 1970; R. C. Brusca. 

Superfamily Paguridea 

Family Lithodidae 

Subfamily Hapalogastrinae 

Hapalogaster cavicauda Stimpson 

Former southernmost records: Punta Santo Tomas 
and San Geronimo Island on the west coast of Baja 
California (Garth. Syst. Zool., 9(3) : 105-123, 1961). 

New record: Guaymas, Sonora, Mexico; under 
stones in the low-intertidal region; January 3, 1969; 
R. C. Brusca. We believe that this is the first record 
of the subfamily Hapalogastrinae from the Gulf of 
California. 



Richard C. Brusca, Dept. Biology, University of Ari- 
zona, Tucson, Arizona 85721 and Janet Haig, Curator 
of Crustacea, Allan Hancock Foundation, Los Angeles, 
California 90007. 

Accepted for publication February 29, 1972. 



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McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii -f 326 pp. 
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Brattstrom, B. H. 1969. The condor in California. Pp. 369-382 in Vertebrates of California. 
Payne, ed.), Univ. California Press, xii + 635 pp. 

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CONTENTS 

Editorial: A bil of History 1 

Generic partitioning of the South American leptodactylid frog genus Eupsophus Fitzinger, 1843 
( Sensu Lato) . By John D. Lynch 2 

record ot the black scabbardfish, Aphanopus carbo, from the Pacific Ocean with notes on 
other Californian trichiurid fishes. By John E. Filch and Daniel W. Gotshall 12 

Organ weights of non-captive porpoise (Stenella spp.). By William F. Perrin and Edward L. Roberts 19 

The effects of lowered dissolved oxygen concentrations and salinity on the free amino acid pool of 
the polychaetous annelid Neanthes arenaceodentata. By John L. Abati and Donald J. Reish 32 

An annotated key to the cercariae that develop in the snail Cerithidea californica. By W. E. Martin 39 

New species of robber flies of the genera Wilcoxia and Metapogon (Diptera: Asilidae). By J. Wilcox 43 

RESEARCH NOTES 

A first report of self-fertilization in the wood-boring family Teredinidae (Mollusca: Bivalvia). By 
Kevin J. Eckelbarger and Donald J. Reish 48 

Rhadinaea godmani, an addition to the snake fauna of Honduras. By Larry David Wilson and 

John R. Meyer . 50 

A new species of Baetodes from Arizona (Ephemeroptera: Baetidae). By Richard K. Allen and 

Esther S. M. Chao 52 

An introduced population of social wasps, Polistes apachus, that has persisted for ten years (Hy- 

menoptera: Vespidae) . By David L. Gibo 53 

The phyllostomatid bat, V ampyressa brocki, in Colombia. By Robert J. Baker, Hugh H. Genoways, 

and Alberto Cadena 54 

Temperature changes in heat producing plants. By Bayard H. Brattstrom 54 

Range extensions of porcelain and hermit crabs in the Gulf of California. By Richard C. Brusca 
and Janet Haig 56 



cover: Aeolian erosion and contorted bedding in a Cretaceous stream deposit, El Gallo Formation, 6 miles 
west of El Rosario, Baja California Del Norte. Specimens of hadrosaurian dinosaurs related to Lambeosaurus 
have been collected from this formation. The large brown mass in the center of the photograph is a fossil tree, 
lorn loose, and washed downstream in the flood that produced the sandstone strata. 

Photograph by William Morris, Occidental College. 



SOUTHERN CALIFORNIA ACADEMY OF S( I 



Volume 71 



BULLETIN 



JOHN ADAMS COMSTOCK MEMORIAL ISSUE 





IBRARY 

NOV 10 1972 

NEW YORK 
!>TAS\ICAL GARDEN 






Number 2 



BCAS-A71(2) 57-112 (1972) 



AUGUST 1 



Southern California Academy of Sciences 

Founded (> November 1891. incorporated 17 May 1907 

OFFICERS 

, Andrew Starrett, President 

Elbert Sleeper. First Vice President 

Jules Crane. Jr.. Second Vice President 

Stuart L. Warter, Secretary 

Donald R. Patten. Treasurer 

Patrick II. Wells. Technical Editor 

James Dale Smith, Managing Editor 

Roberta S. Greenwood, Index Editor 






1970-1972 

Donald Bright 
Jules Crane. Jr. 

John E. Fitch 
William J. Morris 
Charles R. Weston 



BOARD OF DIRECTORS 

1971-1973 

John J. Baird 

Takashi Hoshizaki 

Elbert Sleeper 

David L. Walkington 

Stuart L. Warter 



1972-1974 

Wayland D. Hand 

Donald R. Patten 

Donald J. Reish 

Andrew Starrett 

Patrick H. Wells 



Membership is open to scholars in the fields of natural and social sciences, and to any per- 
son interested in the advancement of science. Dues for membership, changes of address, and requests 
for missing numbers or numbers lost in shipment should be addressed to the Treasurer, Donald R. 
Patten, the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, Califor- 
nia 90007. 

Annual Members $ S.00 

Student Members 5.00 

Life Members 150.00 

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



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



BULLETIN OF THE SOUTHERN CALIFORNIA 

ACAD E MY () F SC I I. N( ES 



Volume 71 



October 20, 1972 



Nl Mill R .' 



IOIIN ADAMS COMSTOCK. I8K3 1970 

Lloyd m. Martin 1 



At his home in Del M;ir, California, the long and 
productive life of Dr. John Adams Comsiock came 
to an end on December 26, 1 970. 

Born in Evanston, Illinois on January 30, 1883. 
he was a descendant of William Comstock, who 
arrived in Massachusetts, from England, in 1635. 
His father was John Adams, Sr., born October 2, 
1845 in Franklin, Oakland Co., Michigan. His 
mother was Cornelia Hurd; daughter of Judge 
Harvey B. Hurd, one of the founders of North- 
western University. John Comstock married 
Caroline Townsend in 1905. They had three 
children: John Sterling, Jean, and Bette; all three 
now reside in California. 

At an early age Dr. Comstock became interested 
in natural history, especially in entomology. Soon 
he narrowed his studies to the Lepidoptera. He 
was elected Secretary of the Chicago Entomologi- 
cal Society and served from 1898 to 1903. John 
was elected Recorder of the Entomological 
Section of the Chicago Academy of Sciences in 
1903-1904. Because of his innate talent for illus- 
tration and design, he was given ( 1904) the posi- 
tion of Architectural Designer with Myron Hunt 
in Chicago. During the summers, he instructed 
in nature study at Vrilie Heights Summer School, 
Lake Geneva, Wisconsin. 

In 1901, he and his brother Hurd travelled to 
Lake Josephine and Avon Park, Florida, to collect 
butterflies. Many fine specimens were obtained 
and some are still in the collection of the Natural 
History Museum of Los Angeles County. Later, 
in 1902, accompanied by Hurd, and a Professor 
Snyder, he went to Hall Valley, Colorado ( in the 
Rocky Mountains). This party travelled all the 
way to Hayden where they caught many rare and 
little-known butterflies. 

In the fall of 1905 he went to East Aurora. New 
York, to further his natural ability as a designer 
and writer. His furniture and book plate designing, 
book binding and illustrations, metal work. 
jewelry design, and associated creative work were 



continued in the Roycrofl Shop under Ihe gin 
of Elberl Hubbard. As a natural consequence o) 
his love for the outdoors, he conducted nature 
study classes and brought nature-inspired designs 
into the Roycroft work. 

"The Doctor" moved with his family to Santa 
Rosa, California late in 1906. In 1 90S. he and 
his sister, Catherine Comstock Seideneck estab- 
lished the Craft-Camarata arts and crafts shop. 
He also worked for Luther Burbank, as a designer 
for Burbank"s publications. In 1910, another 
Craft-Camarata was established in Santa Barbara. 
California. 

Two years later John Comstock attended medi- 
cal college in Los Angeles. During his summer 
vacations he taught the arts and crafts of jewelry 
and metal work at Carmel summer schools. In 
1915, he was licensed to practice medicine and 
surgery in California and from 1928 to 1930 he 
was an honorary member of the attending staff 
of the Los Angeles County General Hospital. 

During these years, his interest in entomology 
never wavered and he collected whenever time per- 
mitted. In 1917, he helped found and was elected 
the first president of the Lorquin Entomological 
Club (formerly a section of the Lorquin Natural 
History Club). The first meetings of the Club 
were held at the Los Angeles Public Library. In 
October of 1919, the club"s meetings were held in 
the tower of the Southwest Museum. The club 
moved from the Southwest Museum in January. 
1927 to the Natural History Museum of Los 
Angeles County where it became the Lorquin 
Entomological Society. These progressive changes 
were made possible by the constant work of Dr. 
Comstock. 

The Royal Entomological Society of London 
elected Dr. Comstock as a Fellow in 1919. The 
same year he was appointed Assistant Director of 



; 2063 Kachina Drive. Prescott, Arizona 8 



57 



58 



BULLETIN SOUTHERN ( IL1F0RN1A U IDEMY OF SCIENCES VOLUME 71 



JOHN ADAMS COMSTOCK 



1972 



JOHN ADAMS COMSHX K, 1883 1970 






the Southwest Museum of Los Angeles and in 
1921 was appointed Director of the Museum; a 
position which he held until I 926. Al lliis lime 
the fields of the Museum were restricted to 
anthropology, archeology, and history ol (he 
Southwest. The natural hisiory exhibits and 
specimens of the Museum were turned over as an 
indefinite loan to the Natural History Museum of 
Los Angeles County. Dr. Comslock's entomologi- 
cal collections were transferred as a definite gilt. 

In 1922, while at the Southwest Museum, Dr. 
Comstock organized the Indian Welfare league 
and served as Executive Chairman, helping to 
secure beneficial legislation for Indians. In rec- 
ognition of his efforts he was appointed to the 
Advisory Committee on Indian Affairs under 
Hubert Work, Secretary of the Interior. He served 
as Chairman of a Joint Parks Committee from 
1925 to 1928 and worked to establish parks 
throughout the state of California. 

John Comstock returned to private medical 
practice in 1926 at the Edwards-Wildey Building 
in Los Angeles. His medical skills, along with 
his kindness, gentleness, and warmth toward his 
patients, made him an outstanding physician. 

On March 28, 1928, he married Ruth Mary 
Gard, a descendant of an early California pioneer 
family. She now resides at their home in Del Mar, 
California. 

The Doctor's interest in Lepidoptera took him. 
in 1924, to Moran, Wyoming, in the Jackson Hole 
region of the Teton Mountains", and to Mt. Wash- 
burn, in Yellowstone National Park. Many 
museums throughout the world have specimens 
from this extensive collecting trip. In addition to 
collecting specimens, he took many photographs 
on glass plates which he later tinted to bring out 
the natural beauties of the country. 

Dr. Comstock became interested in the South- 
ern California Academy of Sciences shortly after 
he arrived in Los Angeles. He was elected Secre- 
tary and editor of the Bulletin in 1921. He served 
as President of the Academy 1926-27. He con- 
tinued as editor until 1962. In 1929 he was elected 
Fellow of the Academy. The financial assets of the 
Academy increased greatly under his guidance and 
the welfare and progress of this organization has 
benefited greatly from the many years of his close 
association. 

An ambition to publish a book on the butter- 
flies of California was realized on April 14, 1927. 
Dr. Comstock spent many years corresponding 
with outstanding lepidopteran collectors in Cali- 
fornia and in securing authentic records of Cali- 



fornian species from various muscumt and privati 
collections. The Butterflies •</ California -till re- 
mains a classic ami 1 6 COll >] pi U ■'• hit h 

are exceptionally clear and precise. 

I .uly in I92K John Adam ' "in tocl 
pointed Act mi' Director ol tin-- I < 
County Museum ol History, Science, ami An on 
October l, be was made Assistant Dircctoi Hi 
was Director of Science from 1939 i" 1942. Al 
that time his title was changed in ( hicl < Uralor 
of the Science Division and after his retirement 
in November of 1948 he was given the position 
of (uralor Emeritus of Entomology. Alter his 
retirement he remained closely associated with 
the Museum and on February 16, 1970 he was 
appointed Research Associate in Entomolog 
the fifth three-year period. 

During his twenty years al the Natural History 
Museum John Comstock became a dearly loved 
member of the staff. His understanding of per- 
sonal problems, his financial help, and his kind. 
gracious, and warm concern for his fellow man. 
helped many employees through the trying times 
of the "great depression" of the 1930s. 

The Doctor's wide range of interests extended 
to genealogy and in 1930. he was elected President 
of the Los Angeles Genealogical Society. He was 
elected Life Fellow and given a Certificate of Merit 
by the Institute of American Genealogy in 1939. 
His book. A History and genealogy of the Coin- 
stock family in America, was published in 1949 
and the enormous amount of information com- 
piled in this book is extremely valuable. 

During World War II. Dr. Comstock was active 
in the Station Wagon Evacuation Corps. Company 
D. First Evacuation Regiment of Los Angelev As 
a medical doctor, his services were invaluable and 
greatly needed. 

In addition to the above-mentioned societies. 
Dr. John Comstock was awarded many honorary 
memberships in Greek letter societies. These in- 
cluded Pi Gamma Mu (social sciences). Phi 
Sigma (biological sciences), and Sigma \i 
(science). 

After retiring from the Natural History Museum 
of Los Angeles County in 1948. John and his 
wife Ruth Comstock moved to Del Mar. Cali- 
fornia. He was unfailingly interested and active 
in civic affairs and in 1952. he served as President 
of the Del Mar Civic Association. 

Dr. Comstock's continuing responsibility to pre- 
serving important California natural areas is 
exemplified by his efforts and those of his staunch 
conservation companion. Guy Fleming. Together 



Ml 



HI III IIS sol llll l<s ( ILIFORNIA ACADl \IY OF SCIENCES 



VOLUME 71 



thej foughl to preserve the I ones Pines in a 
natural state. The Doctor served as President of 
the Torres Pines Association from 1955 to 1962. 

His work al the San Diego Museum of Natural 
History was recognized with highest esteem. He 
was elected as President of the Fellows. San Diego 
Society of Natural History, for 1958 and served 
as a member of the Board of Directors of the San 
Diego Museum from 1957 to 1970 when he re- 
signed i\vw to age and poor health. His resignation 
was not accepted however, and instead the mem- 
bers of the Board voted him a Member Emeritus. 
He had been named Honorary Curator of Ento- 
mology for the San Diego Museum in 1969. 

The Lepidopterists' Society elected him Presi- 
dent in 1962 in recognition of the vast work that 
he had accomplished in working with young 
entomologists on the life cycles of butterflies and 
moths. In 1968, the Executive Council unan- 
imously voted him an Honorary Life Member: 
an appointment which was confirmed by more 
than 80 percent of the membership. 

During the period from 1932 to 1964 he made 
many trips to Arizona collecting large numbers of 
moths and butterflies, which he gave to the Nat- 
ural History Museum of Los Angeles County. 
At the same time, he illustrated the early stages 
of many Lepidoptera that had not previously been 
described. 

In November of 1952. Dr. Comstock ac- 
companied by his wife Ruth, drove from the 
United States to the Guatemalan border. In the 



spring of 1957 they went to Puerto Vallarta, 
Mexico, where they spent four months (June- 
October) collecting specimens. Much of the re- 
search from these expeditions to Latin America 
were published with Dr. Leonila Vasqucz by 
Mexican foundation grants. Later he and his wife 
spent six months (May-October 1961) working 
in American Samoa where they resided in Mulinuu 
in the country home of the former governor. The 
scientific knowledge accumulated in this area was 
published by the Bishop Museum of Hawaii. 

His contributions to entomology, published in 
numerous scientific journals, number 220. He 
helped many entomologists who, through the 
years, have expressed great affection for him as 
an excellent worker in the field, and a delightful 
companion for fellow collectors. It would take 
many pages to record their reactions. His honors, 
positions, accomplishments, talents, personality, 
and affection for his fellow man will long be re- 
membered by those who were fortunate to know 
him. His talents in art, science, medicine, and 
genealogy, and his qualities for leadership, have 
been widely recognized. In the words of Robert 
F. Heilbron, President of the Board of the San 
Diego Society of Natural History, "He was wise, 
gentle, always kind. His life touched ours and we 
were enriched. Now he is gone and we are 
diminished." 

I wish to express my appreciation to Mrs. Ruth 
Comstock for information and her assistance in 
preparing this manuscript. 



BIBLIOGRAPHY OF JOHN ADAMS COMSTOCK, 1883-1970 
Deborah Birnie and James Dale Smith 1 



Between the years 1902 and 1969, Dr. John 
Adams Comstock, produced 236 scientific papers: 
the majority of which dealt with Lepidoptera of 
the southwest and Pacific coast. These papers in- 
clude many excellent illustrations which attest to 
the Doctor's ability with pen and ink as well as 
water colors. In preparing Dr. Comstock's 
bibliography which follows, we have made several 
annotations, in brackets, in order to clarify the 
content of certain papers. 



1902 

A trip to Lake Josephine, Fla. [with Hurd Comstock]. 
Ent. News, 13(3):75-77. 

An incident of Prof. Synder's trip in Colorado. Ent. 
News, 13(8) :258. 



1 Dept. Biology. California State University, Fuller- 
ton, California 92634. 



1972 BIBLIOGRAPHY 01 lolIN ADAMS COMSTO 

1918 

A new aberration, The Lepidopterist, 2(2): 13, I p] 



61 



Butterflies ol ' alifornia ' oni i I (Tic Parna 

Hull. s.,, ( alifornia .' 



Melitaea anicia: two new aberrations. I Me I epidop- 
terist, 2(5):34-37, I pi. 

Notes on the pola-minuta group of Melilaeas, wiih 
description of a new species. The Lepidopterist, 

2(7):55. 

Noies on the pola-minuta group of Melilaeas, wiih 
description of ;i new species (com.). The 
Lepidopterist, 2(9-10) :70-73, I pi. 

I 920 

A new species or race of Argynnis from California. 

Southwest Sci. Bull., pp. 4-8, I pi. 

Butterflies of California: Preliminary announcement. 
Bull. So. California Acad. Sci., 19(3):4-8. 

1921 

Butterflies of California; the Swallowtails and allies. 
Bull. So. California Acad. Sci.. 20(l):5-6. 
[Plate 3 published BSCAS. 21(1): 18]. 

Butterflies of California, (conl.) |The swallowtails 
and allies]. Bull. So. California Acad. Sci., 20(2) : 
45-46, I pi. 

Studies in Pacific Coast Lepidoptera [Early stages of 
Etiphydryas sierra Wright]. Bull. So. California 
Acad. Sci., 20(2):46-47. I pi. 

A giant Palm-boring beetle — Dinapate wrightii. Bull. 
So. California Acad. Sci., 21(1):5-17. 

Butterflies of California, (cont.) [Plate 3 — Swallow- 
tails and their allies]. Bull. So. California Acad. 
Sci., 21(1): 18. [Text for plate published earlier 
BSCAS. 20(1) :5-6]. 

The Mission Indians of California. The Golden West, 
Nov. 1, p. 7, and Nov. 15, 4-5. 

1922 

Studies in Pacific Coast Lepidoptera, (cont.): Notes 
on the acmon-neurona group Lycaenids. with de- 
scription of a new species. Bull. So. California 
Acad. Sci.. 21(2):43-48. 

1923 

Butterflies of California, (cont.): The Parnassians. 
Bull. So. California Acad. Sci.. 22(1): 15-16. 
1 Pi- 

Studies in Pacific Coast Lepidoptera [Early stages of 
Melitaea neumoegeni]. Bull. So. California 
Acad. Sci, 22(2):68-69. 



192 1 

Butterflj migration, i he ' luzel, 1(1) 
Studies in Pacifii Coast I epidoptcra i 

cil a "lost species." Bull. So. ( alifornia 

Sci., 23(1):13 16, I pi. 

Butterflies of California, (cont.): The Whiti 

allies. Bull. So. California Acad. Sci.. 23(1) 
18-20, I pi. 

Studies in Pacific Coasl lepidoptera: New r;i 

California butterflies. Bull. So. < alifornia Acad. 
Sci.. 23(2):51-52. 

Butterflies of (alifornia. (cont.): The Whites ami 
allies. Bull. So. California Acad. Sci, 23(2):5 

Butterflies of California, (cont.): The Whites and 
allies. Bull. So. California Acad. Sci.. 23(4): 
124-125. 

Butterflies of California, (cont): The Whites and 
allies. Bull. So. California Acad. Sci, 23(5): 
157. 



A new record for California [Papilio polydamas in 
California]. Bull. So. California Acad. Sci, 

23(5): 157. 

Studies in Pacific Coast Lepidoptera. (cont.): A new 
Melitaea from Oregon. Bull. So. California Acad. 
Sci, 23(6): 173-174. 

Studies in Pacific Coast Lepidoptera: Notes on the 
genus Cercyonis. Bull. So. California Acad. Sci, 
23(6) : 174— 177. [Includes notes on the genera 
Picris and Eurymus]. 

Butterflies of California, (cont.) [The Dwarf Yel- 
lows, Marbles, and Southern Marble]. Bull. So. 
California Acad. Sci, 23(6): 177-178. 1 pi. 

Larva and pupa of Desmocerus californicus (Horn) 
[with Alonzo Davis]. Bull. So. California Acad. 
Sci, 23(6): 178-181. 

1925 

Studies in Pacific Coast Lepidoptera. (cont.l: A new 
variety and two new 7 aberrant forms of California 
butterflies. Bull. So. California Acad. Sci.. 
24(11:3-4. 

Butterflies of California, (cont.) [The Marbles 
Orange Tips]. Bull. So. California Acad 

24(11:4. 1 pi. 



Ill III IIS sol llll A'.V CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



Studies in Pacific Coast Lepidoptera: A new race of 
Mitoura siva Edw. in California. Bull. So. Cali- 
fornia Acad. Sci., 24(2):36-38. 

Butterflies of ( allien ma. (cont.) [The Marbles]. Bull. 
So. California Acad. Sci., 24(2) :38. l pi. 

Butterflies of California, (cont.) [The Sulphurs and 
Dog-faces]. Bull. So. California Acad. Sci., 
24(3):61-62, I pi. 

Studies in Pacific Coast Lepidoptera, (cont.): Eight 
new races, forms or aberrations of California 
butterflies. Bull. So. California Acad. Sci.. 
24(3):62-68. 

1926 

Butterflies of California, (cont.) [The Sulphurs). Bull. 
So. California Acad. Sci.. 25(1) :28, 2 pis. 

Studies in Pacific Coast Lepidoptera. (cont.): Thirteen 
new species or aberrations of California butter- 
flies. Bull. So. California Acad. Sci.. 25(1): 
29-34. 

Studies in Pacific Coast Lepidoptera. (cont.): A new 
aberrant butterfly from southern California. Bull. 
So. California Acad. Sci., 25(2) :48. 

Butterflies of California, (cont.) [The Sulphurs]. Bull. 
So. California Acad. Sci., 25(2): 62-65, 2 pis. 

Butterflies of California, (cont.) [The Milkweed but- 
terflies]. Bull. So. California Acad. Sci.. 25(3): 
84-87. 1 pi. 

1927 

Butterflies of California. McBride Printing Co., 
(privately printed by John Adams Comstock), 
1-334, 63 pis. 

Protective coloration and mimicry. Bull. So. Cali- 
fornia Acad. Sci., 26(1): 1-4. 

1928 

Studies in Pacific Coast Lepidoptera. (cont.) [Life 
history studies (eggs, larvae, and pupae) of six 
California Lepidoptera]. Bull. So. California 
Acad. Sci.. 26(3):67-70. 

Studies in Pacific Coast Lepidoptera, (cont.) [Life 
history studies of four California Lepidoptera]. 
Bull. So. California Acad. Sci., 27(2):63-67. 

Studies in Pacific Coast Lepidoptera. (cont.): Early 
stages of Calephelis australis Edw. Bull. So. Cali- 
fornia Acad. Sci., 27(3): 80-89. 

1929 

The egg of Glaucopsyche lygdamus australis Grinnell. 
Bull. So. California Acad. Sci., 28(1 ):6, pi. 5 
on page 1 1 . 



Studies in Pacific Coast Lepidoptera. (cont.) [Life 
his tor j studies of eight California Lepidoptera]. 
Bull. So. California Acad. Sci., 28(2):22-32. 

A new species or form of Anthocharis from Cali- 
fornia. Bull. So. California Acad. Sci., 28(2): 
32-33. 



1930 

Studies in Pacific Coast Lepidoptera, (cont.) [Life 
history studies of eight California Lepidoptera], 
Bull. So. California Acad. Sci., 28(3):50-58. 

The life history of Philotes sonorensis Felder [with 
Carl Coolidge], Bull. So. California Acad. Sci., 
29(1): 16-21. 

Studies in Pacific Coast Lepidoptera. (cont.) [Life 
history studies of eight California Lepidoptera], 
Bull. So. California Acad. Sci., 29(1):21-31. 



1931 

Studies in Pacific Coast Lepidoptera, (cont.) [Life 
history studies of four California Lepidoptera]. 
Bull. So. California Acad. Sci., 29(3) :135-142. 

Studies in Pacific Coast Lepidoptera, (cont.) [Life 
history studies of three California Lepidoptera]. 
Bull. So. California Acad. Sci., 30(1): 15-20. 

Notes on the life history of Poanes melane Edw. 
(Lepid.) [with Charles M. Dammers]. Bull. So. 
California Acad. Sci., 30( l):20-22. 

Notes on the early stages of four California argynnids 
[with Charles M. Dammers]. Bull. So. California 
Acad. Sci., 30(2):40-44. 



1932 

The metamorphosis of Heterochroa bredowii cali- 
fornica Butl. (Lepid.) [with Charles M. Dam- 
mers]. Bull. So. California Acad. Sci., 30(3): 
83-87. 

Studies in Pacific Coast Lepidoptera, (cont.) [Life 
history studies of five California Lepidoptera], 
Bull. So. California Acad. Sci., 30(3):88-92. 

Early stages of Melitaea leanira wrightii Edw. and 
Calephelis nemesis Edw. (Lepidoptera) [with 
Charles M. Dammers], Bull. So. California Acad. 
Sci., 31(1):9-15. 

Studies in Pacific Coast Lepidoptera, (cont.) [Life 
history studies of three California Lepidoptera], 
Bull. So. California Acad. Sci., 31(1): 16-18. 

Metamorphoses of five California diurnals (Lepi- 
doptera) [with Charles M. Dammers]. Bull. So. 
California Acad. Sci., 3 1(2): 33-45. 



1972 



BIBLIOGRAPHY OF JOHN ADAMS COM.STOCI 



' 



Metamorphoses of six California Lepidoptera. Bui 
So. California Acad. Sci., 31 ( 3 > :KK 99. 



Miscellaneous notes on ■■■■ Icrn Lcpidoplcra Hull 
So, ' alifornia Acad S( i., 34(2) 



1 933 

A new lycaenid from Southern California I with Chris- 
topher llennel. Bull. So. California Acad. Sci.. 
32(l):23-26. 

Notes on the life histories of two California lepidop- 
terous insects | with Charles M. Dammers]. Bull. 
So. California Acad. Sci., 32(l):27-37. 

Notes on the life histories of four Californian lepidop- 
terous insects (with Charles M. Dammers]. Bull, 

So. California Acad. Sci., 32(2):77-83. 

Notes on the life histories of two Arizona butter- 
flies [with Grace H. and John L. Sperry]. Bull. 
So. California Acad. Sci., 32(3) : 99-1 04. 

Rarly stages of three California diurnals (Lepidop- 
tera) [with Charles M. Dammers], Bull. So. 
California Acad. Sci., 32(3): 105-1 13. 

Studies in Pacific Coast Lepidoptera. (cont. ) [Life 
history studies of four California Lepidoptera]. 
Bull. So. California Acad. Sci., 32(3) : 1 13-120. 



19.34 

Additional notes on the early stages of California 
Lepidoptera [with Charles M. Dammers]. Bull. 
So. California Acad. Sci.. 33(1) :25-34. 

Studies in Pacific Coast Lepidoptera. (cont.) [Life 
history studies of four California Lepidoptera]. 
Bull. So. California Acad. Sci., 33(l):34-38. 

The metamorphoses of three California diurnals [with 
Charles M. Dammers]. Bull. So. California Acad. 
Sci., 33(2):79-92. 

1935 

Notes on the early stages of three butterflies and 
five moths from California [with Charles M. 
Dammers]. Bull. So. California Acad. Sci., 
33(3): 136-151. 

Early stages of Papilio polydamas lucayus R. and J. 
[with Florence Moore Grimshawe], Bull. So. Cali- 
fornia Acad. Sci.. 34(11:76-80. 

Notes on the early stages of two butterflies and one 
moth [with Charles M. Dammers], Bull. So. 
California Acad. Sci., 34(l):81-87. 

Notes on the early stages of three butterflies and six 
moths from California [with Charles M. Dam- 
mers]. Bull. So. California Acad. Sci., 34(2): 
120-142. 



I •>:,(, 

Notes cm the life histories ol three bullcrflii and 
three moths from California [with Charlc M 
Dammers], Bull So California Acad. Sci 
34(3 ):21 1-225. 

Metamorphosis ol Stivnxm leda I dw. il.cpid. 

Rhopal.) | witii < harlcs \i. Dammers] Bull So 
California Acad Sci., 35< I I I 

Notes on the early slaves ol I rebus odora I 

(Lepidopt.). Bull, So California Acad 

35(2):95 98. 

Notes on the early stages of five moths from southern 
California [with (harlcs m. Dammers]. Bull, So 
( alifornia Acad. Sci.. 35(2):99-107. 



1937 

Notes on the life history of Oencis daura. Stir 
(Lepidoptera) |with John L. and Grace H. 
Sperry], Bull. So. California Acad. Sci.. 35(3) 
165-169. 

A new record and a life history (Lepidoptera). Bull. 
So. California Acad. Sci.. 35(3 1:169-174. 

Life history of Papilio rudkini Comst. |with Charles 
M. Dammers]. Bull. So. California Acad. Sci.. 
36(1): 13-18. 

Miscellaneous notes on western Lepidoptera. Bull. 
So. California Acad. Sci.. 36(11:19-23. 

Notes on the early stages of three California moths 
[with Charles M. Dammers]. Bull. So. California 
Acad. Sci., 36(2):68-78. 



19.38 

Miscellaneous notes on western Lepidoptera. Bull. 
So. California Acad. Sci.. 36(3) : I 1 1-1 24. 

Notes on the metamorphosis of Mitoura spinetorum 
Hew. (Lepidoptera. Theclinae) [with Charles M 
Dammers]. Bull. So. California Acad. Sci.. 
37(l):30-32. 

A new race of Euproserpinus phaeton from the 
Mojave Desert (Lepidoptera: Sphinigidae). Bui 
So. California Acad. Sci.. 37(l):33-42. 

Notes on the life history of a noctuid moth 

Charles M. Dammers]. Bull. So. California Acad. 
Sci.. 37(2):78-79. 



64 



HI III I IS sol I ill l<s < ALIFORN1 I i< ADl M) OF SCIENCES 
1 939 



VOLUME 71 



Studies in the metamorphoses of six California moths 
[with Charles M. Dammers]. Bull. So. California 
Vcad. Sci., 371 J):105 128. 

A new Apodemia from California. (Lepidopt.). Bull. 

So. California Acad. Sci.. 37(3 ): 129-1 32. 

Historical sketch oi the Academy. Bull. So. California 
Vcad. Sci.. 37(31:146-153. 

Notes on the life histories of two moths from Cali- 
fornia (with Charles M. Dammers]. Bull. So. 
California Acad. Sci.. 38(l):31-33. 

Studies in Pacific Coast Lepidoptera [Larva and pupa 
of Callopistria floridensis Gn.]. Bull. So. Cali- 
fornia Acad. Sci.. 38(11:34-35. 

[The fauna and flora of the El Segnndo sand dunes — 
cont.] 7. Two new phaloniid moths. Bull. So. 
California Acad. Sci.. 38(2): 112-119. 

[The fauna and flora of the El Segundo sand dunes — 
cont.] 8. Notes on the larva of Lorita abornana 
Busck. (Lepidopt.). Bull. So. California Acad. 
Sci.. 38(2): 119. 

A new Sparganothis with notes on early stages 
(Lepidoptera, Tortricidae). Bull. So. California 
Acad. Sci., 38(2) : 120-121. 

Notes on the early stages of Mammifrontia rileyi 
Benj. (Lepid.). Bull. So. California Acad. Sci., 
38(2) -122-123. 

1940 

Contributions from the Los Angeles Museum — 

Channel Islands Biological Survey: Introductory 

note. Bull. So. California Acad. Sci., 38(3): 
133-134. 

Four new California moths with notes on early stages. 
Bull. So. California Acad. Sci., 38(3) : 172-182. 

Notes on the early stages of three California moths 
[with Charles M. Dammers]. Bull. So. California 
Acad. Sci., 38(3) : 183-187. 

[The fauna and flora of the El Segundo sand dunes — 
cont.] 10. Notes on the early stages of Euphyia 
implicata. Gn. Bull. So. California Acad. Sci., 
38(3): 198-200. 

Argynnid notes. Bull. So. California Acad. Sci., 
39(l):75-77. 

Notes on the early stages of Nemoria pistaciaria 
Pack, [with Christopher Henne]. Bull. So. Cali- 
fornia Acad. Sci., 39(11:78-80. 

Notes on the life history of Aseptis perfumosa Hamps. 
Bull. So. California Acad. Sci., 39(2): 104-105. 



rhe early stages of Trachea ftimeola Hamps. Bull. So. 
California Acad. Sci.. 39(2) : 106-107. 

Notes on the early stages of Euphydryas gilletli 
Haines. Bull. So. California Acad. Sci., 39(2): 
111-1 13. 

1911 

[Contributions from the Los Angeles Museum — 
Channel Islands Biological Survey | 15. A com- 
parison of the larva of Arachnis picta insularis 
and Arachnis picta maia [with Christopher 
Henne]. Bull. So. California Acad. Sci., 39(3): 
189-190. 

Notes on the early stages of Xanthothrix ranunculi 
[with Christopher Henne]. Bull. So. California 
Acad. Sci., 39(3) : 198-199. 

Notes on the life history of Sparganothis larreana 
Comst. [with Charles M. Dammers]. Bull. So. 
California Acad. Sci., 39(3) : 199-200. 

Additional notes on Carolella willettana, Comst. Bull. 
So. California Acad. Sci., 39(3) : 20 1-202. 

[Contributions from the Los Angeles Museum — 
Channel Islands Biological Survey] No. 19. 
Ctenucha brunnea Stretch, on Santa Rosa Is- 
land [with Charles M. Dammers]. Bull. So. Cali- 
fornia Acad. Sci.. 40(1 ):1 1-12. 

Life history of Paphia Cinderella Sm. Bull. So. Cali- 
fornia Acad. Sci., 40(2) : 111-1 13. 

1942 

Notes on Loxophora dammersi Van Duzee. Bull. So. 
California Acad. Sci.. 40(3): 160. 

[Contributions from the Los Angeles Museum — 
Channel Islands Biological Survey] No. 24. The 
larva and pupa of Trichoclea edwardsi Sm. [with 
Christopher Henne]. Bull. So. California Acad. 
Sci., 40(3): 165-166. 

Notes on the life histories of two California moths 
[with Charles M. Dammers], Bull. So. Cali- 
fornia Acad. Sci., 41(l):44-47. 

[Contributions from the Los Angeles Museum — 
Channel Islands Biological Survey] No. 26. A new 
race of Arachnis picta from Santa Catalina Is- 
land. Bull. So. California Acad. Sci., 41(2): 
83-85. 

Notes on the life history of Tolype glenwoodii 
Barnes [with Christopher Henne]. Bull. So. Cali- 
fornia Acad. Sci.. 41(2):86-90. 

Notes on the metamorphoses of two California moths 
[with Charles M. Dammers]. Bull. So. California 
Acad. Sci.. 41(2):91-96. 



1972 



B1BL10GRAPHI OF JOHN ADAMS COMSTOCI 



• 



[Obituary] William Allanson Bryan Dec. 23, 1X75 

June IK, 1942. Hull So. California Acad. Sci., 
41(2): 115 1 17. 

i 94 :t 

The early stages of Arctonotus lucidus Bdv. 
(Lepidopt.) |wiih Christopher Henne], Hull. So. 
California Acad. Sci.. 41(3) :167— 171. 

Noics on the life histories of iwo common mollis 
[with Charles M. Dammers]. Bull. So. California 
Acad. Sci., 41(3): 172 178. 

Western Poplar Sphinx | with Charles M. Dammers]. 
Bull. So. California Acad. Sci.. 42(1):39-41. 

California species of Smeiinlhus with notes on lin- 
early stages of S. ccrisyi ophthalmicus |wiih 
Charles M. Dammers]. Bull. So. California Acad. 
Sci., 42(l):42-45. 

The larva of Copicucullia basipuncta B. and McD. 
(Lepidoptera) [with Christopher Henne], Bull. 
So. California Acad. Sci.. 42(1 ):45-46. 

Life history of Pseudalypia crolchi Hy. F.dw. Bull. So. 
California Acad. Sci., 42(2):59-66. 

Notes on the metamorphosis of Leptarctia californiae 
Wlk. (Lepidopt.) |wilh Charles M. Dammers]. 
Bull. So. California Acad. Sci., 42(2):67-68. 

Mature larva of Graptolitha longior Sm. [with 
Christopher Henne). Bull. So. California Acad. 
Sci., 42(3): 132. 

Additional notes on Erinnyis ello L. (Lepidoptera; 
Sphingidae). Bull. So. California Acad. Sci., 
42(3): 133-135. 

1944 

Four California moths associated with Cat-tails. Bull. 
So. California Acad. Sci., 43(2):81-83. 

The larva and pupa of Arzama gargantua Dyar [with 
Charles M. Dammers], Bull. So. California Acad. 
Sci.. 43(2): 84-85. 

Brief notes on the early stages of three California 
moths [with Charles M. Dammers). Bull. So. 
California Acad. Sci.. 43(3) :1 13-1 18. 

1945 

Notes on the early stages of Nemoria delicataria 
Dyar (Lepidoptera). Bull. So. California Acad. 
Sci., 44(1):20-21. 

1946 

[Obituary] George Willett 1879-1945. Bull. So. Cali- 
fornia Acad. Sci., 44(3) : 140-142. 



Noti on Hi' iii' in lor) ol Anai an 

fragilaria ( Iroi il <\ < nidopu ra I [with < hi 
M. Dammers] Bull So ' uliforni 
45(1 ):I7 20. 

I' ontributions from Ihi Lo Vi 

' hanncl Island Bii ill ii al Survc) i B 

on the expedition - ondu led bctwcci ' i I 

1940 and Deccmbci 14, 1941. Bull So. ( alifi 

Acad. Sci.. 45(21:94 107 

A few pesis of Sunflowci m Califon I Bull So 
( alifornia Acad. Sci., 45(3) Ml 144 

Phlegethontius rustica Fabr. in California (Lcpidop 
tera; Sphingidae). Bull, So. California Acad 
Sci.. 45(3): I 15-147. 

19 I 7 
[The fauna and flora ol the I I Si gundo sand dunes 

com. I 17. Notes on thi of I "sma 

hehnei Clarke. Bull. So. ( alifornia Acad. 
46(l):53-54. 

Notes on the early stages of Adelocephala hciligl 

f. hubbardi Dyar. Bull. So. California Acad. 

Sci., 46(2):72-77. 

Notes on the life history of Orthodes accurata Hy. 
Edwards. Bull. So. California Acad. Sci.. 46(3) 
124-126. 

1918 

The larva and pupa of Eumaeus debora Hbn. Bull. 
So. California Acad. Sci.. 4 7 { I 1:3-5. 

The mature larva and pupa of Arctonotus tcrlootii 
Hy. Edw. Bull. So. California Acad. Sci.. J~ 
49-5 1 . 

1949 

[Obituary] Dr. Robert Dudley Emery. Bull. So. Cali- 
fornia Acad. Sci.. 48 ( 21:94-95. 

A history and genealogy of the Comstock famil\ in 
America. Commonwealth Press. Los Angeles 
(privately printed for John Adams Comstock i. 

xvi + 715 pp. 

1952 

[Obituary] Dr. John Herman 1878-1951 [with Homer 
P. King). Bull. So. California Acad. Sci.. 51(1): 
21. 

1953 

Collecting butterflies in the coastal area of Mexico 
near Manzanillo. Colima with notes on the life 
history of a rare skipper. Bull. So. California 
Acad. Sci., 521.21:43-15. 



BULLETIN sol IIIIR\ i ILIFOKMt KM/)/. \/> Ol- SCII-.NCK.S 



VOLUME 71 



Life history notes on four southern Arizona butter- 
flies. Hull So. California Acad. Sci., 52(3): 
127-136. 

1951 

1 ife history notes on Ascia monuste crameri. Hull. 
So. California Acad. Sci.. 53( I 1:46-49. 

Beauty anil the hectic. Bull. So. California Acad. Sci., 
53(21:118-121. 

Notes on the early stages of Forsebia perlaeta (Hy. 
F.dw. ) Lepidoptera: Phalaenidae. Bull. So. Cali- 
fornia Acad. Sci.. 53(3): 166-168. 

1955 

Miscellaneous notes on North American Lepidoptera. 
Bull. So. California Acad. Sci.. 54(l):30-36. 

A partial life history of a California butterfly. Bull. 
So. California Acad. Sci.. 54(2):57-60. 

Brief notes on the life histories of six California 
moths. Bull. So. California Acad. Sci., 54(2): 
60-66. 

The egg. and first larval instar of a geometrid moth 
from Arizona. Bull. So. California Acad. Sci., 
54(2): 105. 

The egg and young larva of a geometrid moth from 
California. Bull. So. California Acad. Sci., 54(2): 
105. 

A new Papilio from California [with Lloyd M. Mar- 
tin]. Bull. So. California Acad. Sci., 54(3): 
142-150. 

A surmise confirmed. Bull. So. California Acad. Sci., 
54(3):167. 

1956 

Notes on metamorphoses of the giant skippers 
(Lepidoptera; Megathyminae) and the life his- 
tory of an Arizona species. Bull. So. California 
Acad. Sci., 55(1): 19-27. 

Is this a new, and giant clothes moth? Bull. So. 
California Acad. Sci., 55(l):51-53. 

Notes on the life history of a rare Arizona Sphinx 
moth, Xyophanes falco Walker. Bull. So. Cali- 
fornia Acad. Sci., 55(2) :102-106. 

The larva of Euclidimera diagonalis Dyar, (Lepidop- 
tera; Catocalinae). Bull. So. California Acad. 
Sci., 55(2): 115-1 16. 

The larva of Stenoporpia grisearia (Grt.) (Lepidop- 
tera: Geometridae). Bull. So. California Acad. 
Sci., 55(2): 116-1 17. 



Notes on the life histories of two southern Arizona 
butterflies. Bull. So. California Acad. Sci., 
55(3): 171-179. 

Brief notes on the life histories of three Arizona 
moths. Bull. So. California Acad. Sci., 55(3): 
180-183. 



1957 

Notes on the metamorphosis of an Agave-boring but- 
terfly from Baja California, Mexico. Trans. San 
Diego Soc. Nat. Hist., 12( 13) :263-276. 

Notes on the early stages of two western American 
moths. Bull. So. California Acad. Sci., 56(1): 
42-47. 

[Obituary] Commander Charles Montagu Dammers 
1878-1956. Bull. So. California Acad. Sci., 
56(1 ):50. 

Life histories of two southern Arizona moths of the 
genus Caripeta. Bull. So. California Acad. Sci., 
56(2):88-96. 

Brief notes on the life histories of two Arizona 
geometrid moths. Bull. So. California Acad. Sci., 
56(2):99-100. 

The habits and life histories of Coehisea sinuaria B. 
and McD. and Coehisea sonomensis B. and McD. 
[with Christopher Henne and Frank Sala]. Bull. 
So. California Acad. Sci., 56(3) : 169-177. 

Scientific notes. [On eggs and first instar larvae of 
Aemilia ambigua Stkr. and Dasylophia seriata 
Druce.]. Bull. So. California Acad. Sci., 56(3): 
178-179. 

1958 

On the early stages of the Lepidoptera (Abstract). 
The Lepidopterists' News, 1 1(1-3) :4. 

Early stages of Eutachyptera psidii (Lasiocampidae), 
a rare moth from southern Arizona. The Lepi- 
dopterists' News, 11 (4-5): 99-102. [title in- 
correct, see Comstock (1964. J. Lepid. Soc. 
18(3):164)]. 

The ova and first larval instars of three southwestern 
moths. Bull. So. California Acad. Sci., 57(1): 
34-38. 

The early stages of Oncocnemis perscripta (Guenee) 
(Lepidoptera: Phalaenidae). Bull. So. California 
Acad. Sci., 57(2):81-84. 

Random notes on early stages of Lepidoptera. Bull. 
So. California Acad. Sci., 57(2) : 114-1 15. 

Notes on the early stages of Feralia fehrualis Grote. 
Bull. So. California Acad. Sci., 57(3): 140-143. 



IV72 



BIBLIOGRAPHY OF Jims ADAMS COMSTOCK 






A brief nolo on die pupa of Melitaea pola Boisduval 
(Lepidoptera: Nymphalidae), Bull. So. ( lali 
fornia Acad. Sci., 57(3): 143 14-4. 

Random notes on early stages of Lepidoptera (cont.). 
Bull. So. California Acad. Sci., 57(3): 179. 

Butterfly collecting in the Mexican tropics. The 
Lepidopterists' News, 12(3-4): 127 129. 

L959 

Brief notes on four Arizona moths. Hull. So. Cali- 
fornia Acad. Sci., 58(1 ):38-44. 

Scientific notes. |On Dalcerides ingenita (Hy. F.dw. )]. 
Bull. So. California Acad. Sci.. 58(1):53. 

Notes on the early stages of three southwestern 
moths. Bull. So. California Acad. Sci., 58(2): 
95-100. 

Notes on the early stages of Phaeoura cristifera 
Hulst (Lepidoptera: Geometridae). Bull. So. 
California Acad. Sci., 58(2) : 101-106. 

Rare or common! With notes on the life histories of 
two southwestern moths. Bull. So. California 
Acad. Sci.. 58(3) : 155-161. 

The life history of Herculia phoezalis (Pyralidinae ). 
J. Lepid. Soc. 13(2):67-69. 

Entomological briefs [Comments on life histories, 
food plants, and new records of nine California 
Lepidoptera]. Bull. So. California Acad. Sci., 
58(3): 169-171. 

1960 

Notes on the life histories of three western phalaenid 
moths. Bull. So. California Acad. Sci., 59(1): 
22-29. 

Scientific notes. [On larva and pupa of Nasusina 
vaporata (Pears.)]. Bull. So. California Acad. 
Sci.. 59(1):56. 

Notes on the early stages of two phalaenid moths 
from southern California. Bull. So. California 
Acad. Sci., 59(2) : 95-1 02. 

Inherent and applied camouflage in the Subfamily 
Geometrinae (Lepidoptera). including three new 
life history studies. Trans. San Diego Soc. Nat. 
Hist., 12(26) :421-440. 

Life history notes on a saturniid and two lasiocampid 
moths from California. Bull. So. California 
Acad. Sci.. 59(3) : 170-181. 



1961 

Bricl notes on the life histories <4 tl western 

moths. Bull, So. ' alifornia 60(1) 

40-44. 
Estudios dc los ciclos biologii 

mexicanos [with I . \ Oai it del 

Instituto dc Biologia, 11(1 and I) 

Notes on the life history ol Palthis angulalis Hub 

(Lepidoptera; llerminiinac ) [will 

land]. Bull. So. ' alifornia 

112-116. 

Notes on the early stares nl two Texas butterflies. 
Bull. So. California Acad. Sci.. 60(3) : 147-155 

Scientific notes. Bull. So. ( alifornia Acad. Sci.. 60(3): 
196. [Corrects reference from Fernaldclla 
fimetaria (Comstock. Bull. So. (alifornia Acad 
Sci.. 60(1 ):40-44, 1961 ) to I . parlilaria (Grote) 
and comments on distribution of Sterrha 
dimidiata Hufn.]. 

1962 

Life history notes on Epiplatymetra vroiearia Packard. 
Bull. So. California Acad. Sci.. 61(1): 3-6. 

1963 

A fruit-piercing moth of Samoa and the South Pacific 
Islands. Canad. Ent.. 95(2 ) :2 18-222. 

Notes on the life histories of two southwestern 
phalaenid moths. Bull. So. California Acad. Sci.. 

62(l):33-40. 

The early stages of Pcro macdunnoughi Cassino and 
Swett (Lepidoptera. Geometridae). Trans. San 
Diego Soc. Nat. Hist.. 13(2):45-48. 

Notes on the early stages of two California geometrids. 
J. Res. Lepid.. l(3):195-200. 

The early stages of Stenapilates apapinaria Dyar 
(Lepidoptera: Geometridae). Trans. San Diego 
Soc. Nat. Hist.. 13(2):45-48. 

Presidential address to the twelfth annual meeting of 
the Lepidopterists' Society. J. Lepid. Soc. 16i 4 i : 
247-248. 

Early stages of a southern California geometrid 
moth. Drepanulatrix hulsti hulsti (Dyar». J. Res. 
Lepid.. 1(41:245-248. 

The life history of Hulstina inconspicua. a ceometri 
moth from southern California. J. Lepid. Soc 
17(l):28-30. 

Notes on the ova of six California moths. I 
California Acad. Sci.. 62(4): 202-208. 



M 



Hi I I I n\ sol llll.KX CALIFORNIA ACADEMY OF SCIENCES 



I OI.I'MF 71 



Motes on the earlj stages oi Drepanulatrix monicaria 
(Ouenee) (Geometridae). I. Res. Lepid., 2(3): 
201-203. 

Hstudios de los ciclos biologicos en lepidopteros 
mexicanos: II Heterocera [with L. V. Garcia]. 
Analcs de [nstituto del Biologia, 34(1 and 2): 
217-273. 

I <)(. I 

The larva and pupa of Agathymus dawsoni (Lepidop- 
tera: Megathymidae). Trans. San Diego Soc. 
Nat. Hist.. 13(8): 169-172. 

Additional notes on the life history of Neoterpes 
edwardsata (Geometridae). J. Lepid. Soc., 
18(l):32-34. 

The early stages of Didaematiphorus phaceliae McD. 
(Lepidoptera: Pterophoridae). Canad. Ent., 96: 
538-540. 

Life history notes on Catocala euphemia. Bull. So. 
California Acad. Sci.. 63(3) : 151-153. 

Early stages of Dichogaster (Lasiocampidae) : A cor- 
rection. J. Lepid. Soc, 18(3): 164. 

Life history notes on Eupithecia macrocarpata (Lepi- 
doptera: Geometridae). Bull. So. California 
Acad. Sci., 63(4): 171-174. 

Resignation of Dr. William Dwight Pierce. Bull. So. 
California Acad. Sci., 63(4) :204. 

1965 

Studies in life histories of North American Lepidop- 
tera; California Annaphilas [with Christopher 
Henne]. J. Res. Lepid., 3(3) : 173-191. 

[Obituary] Edwin Ray Hulbirt, (1886-1965). J. 
Lepid. Soc, 19(4): 243-244. 

Notes on the life history of Philotes enoptes dammersi 
[with Christopher Henne]. Bull. So. California 
Acad. Sci., 64(3) : 153-156. 

Ciclo biologico de Prodenia ornithogalli Guenee 
I Lepidoptera: Noctuidae). Anales del Instituto 
de Biologia, 36(1 and 2): 199-202. 



1966 

Life history of Philotes mojave (Lepidoptera: 
Lycaeninae). Trans. San Diego Soc. Nat. Hist., 
14(10): 133-136. 

Life history of Philotes enoptes bayensis (Lepidoptera: 
Lycaenidae) [with R. L. Langston]. Pan-Pacific 
Ent., 42(2): 102-108. 



The early stages of Graptolitha (Lithophane) 
dilatocula Smith (Lepidoptera: Phalaenae) [with 
Christopher HenncJ. Bull. So. California Acad. 
Sci.. 65(3): 155-159. 

The larva ami pupa of Orthosia hibisci quinquefasciata 
(Noctuidae) [with Christopher Henne]. J. Lepid. 
Soc, 20(4):213-215. 

Lepidoptera of American Somoa with particular refer- 
ence to biology and ecology. Pacific Insects 
Monograph, 11:1-74. 

The eggs and first instar larvae of three California 
moths. J. Res. Lepid., 5(4) :215-21 9. 



1967 

Studies in life histories of North American Lepidop- 
tera; California Annaphila II [with Christopher 
Henne]. J. Res. Lepid., 5(1): 15-26. 

Notes on the life history of Philotes rita elvirae 

(Lepidoptera; Lycaenidae) [with Christopher 

Henne]. Bull. So. California Acad. Sci., 66(2): 
99-102. 

Early stages of Sphinx sequoiae engelhardti (Sphingi- 
dae) [with Christopher Henne]. J. Lepid. Soc, 
21(1) :27-31. 

Notes on the early stages of the Barberry geometrid 
moths. Genus Coryphisla, and the description of 
a new subspecies of C. meadii (Lepidoptera). 
Bull. So. California Acad. Sci., 66(2):92-98. 

Life history of Gesta gesta invisus (Lepidoptera: 
Hesperiidae) [with Roy O. Kendall]. Trans. San 
Diego Soc. Nat. Hist., 14(18):233-236. 

An additional food plant record for Papilio thoas 
autocles R. and J. J. Res. Lepid., 5(4):220. 

Early stages of Lycomorpha regains Grinnel, with 
notes on the imago. (Lepidoptera: Amatidae) 
[with Christopher Henne]. J. Res. Lepid., 6(4): 
275-280. 



1969 

Notes on the life history of Fishia evelina hanhami 
(Lepidoptera). Bull. So. California Acad. Sci., 
68(l):54-56. 

Life history notes on Lithophane snhtilis (Noctuidae) 
[with Christopher Henne]. J. Lepid. Soc, 23(1): 
15-18. 



Accepted for publication June 30, 1972. 



THRHi: NI'W SYMI'ATKK PLEOCOMA FROM Mil. s< >1 Mil R1 .1! RR 
NEVADA MOUNTAINS OF CALIFORNIA KOI I OF] I RA: S( ARABA1 ID i 

Frank I. H0VORE 1 



Abstract: Two new species and one new subspecies iii Pleocoma an d< cribctl from 
sympatric populations in the Greenhorn Mountains of the southern Siena Nevada range, 
[ntraspecific character variation, comparison to closely related forms and biology arc di 
following each description, An ecological description of the lype locality is also presented. 



Beetles of the genus Pleocoma have long been 
prized and much sought alter by Pacific coast 
coleopterists, yet our knowledge of the habits and 
distribution of the genus is still rather fragmentary. 
The three new forms described herein help I ill a 
large distributional gap between the central Sierra 
Nevada and Tehachapi Mountains. They also 
present the first recorded instance of three 
Pleocoma species occurring sympatrically. Refine- 
ments in collecting apparatus, easier access to re- 
mote areas, and increased knowledge of male 
Pleocoma flight habits will undoubtedly lead to 
the discovery of more such areas of species 
sympatry. 

As in most populations of Pleocoma where 
series have been available for study, the three 
new taxa presented exhibit considerable minor 
morphological character variation. 1 have there- 
fore selected as diagnostic characters those which 
appear most constant, both quantitatively and 
qualitatively. 

Holotypes of new taxa described herein are on 
deposit in the Natural History Museum of Los 
Angeles County collection (LACM). Other in- 
stitutions or persons receiving paratypes will be 
abbreviated in the text as follows: California 
Academy of Sciences, CAS; California Insect Sur- 
vey, Berkeley, CIS: U. S. National Museum, 
USNM: F. T. Hovore, FTH: D. G. Marqua, 
DGM; C. E. Langston, CEL: T. W. Taylor, TWT: 
H. F. Howden, HFH; B. D. Streit, BDS: G. C. 
Walters, GCW; M. T. Gannon. MTG: R. L. 
Westcott, RLW. 

Pleocoma marquai, new species 
Figure 1 

Description: Male. Form robust, oblong-oval, 
moderately convex, dorsum slightly flattened; in- 
tegument mostly dark brown to piceous; pubescence 
reddish-golden. Head black, vertical horn and an- 
terior process of clypeus thinly clothed with reddish- 



golden hairs; dorsal surface closely, irregularly 
punctate, with broad smooth area extending from 
lateral base of vertical horn anteriorly to base of 
ocular canthus; clypeal process small, constricted at 
base, only moderately rcflcxcd. anterior face slightly 
concave medially, apex with moderately deep, obtuse 
notch, apical angles of notch acute, rounded; vertical 
horn elongate, sides subparallel, apex with shallow, 
obtuse notch, apical angles of notch rounded, anterior 
face of horn evenly convex, surface coarsely punctate, 
punctures elongate, sparsely clothed with long reddish- 
golden hairs; ocular canthi projecting forward from a 
right angle, anterior edge sinuate, apex acutely 
rounded, dorsal surface concave, smooth except for 
few elongate punctures apically: palpi and antennae 
dark reddish-brown, scape and lamellae darker, scape 
stout, subcorneal. second segment moniliform. 
strongly flattened, third segment elongate, much 
shorter than scape, slightly reflexed. fourth segment 
cylindrical, slightly more than one-half as long as 
third segment, fifth segment strongly transverse with 
acute process, segments six to eleven distinctly lamel- 
late, sixth segment with lamella about two-thirds as 
long as that of seventh segment, lamella of seventh 
segment slightly shorter than that of eighth segment, 
that of eighth distinctly shorter than that of ninth. 
lamellae of ninth and tenth segments subequal in 
length, that of eleventh only slightly shorter, ratios of 
segments six to eleven in holotype 20:30:32:36:36:35. 
Pronotum piceous, less than twice as wide as long 
(length-to-width ratio in holotype 5.85:10.8). widest 
at posterior angles, posterior angles broadly rounded, 
lateral discal impressions feeble, with indistinct 
reddish macula; disc evenly convex, anterior median 
impression lacking, pubescence entirely absent, surface 
shining, finely, moderately densely punctate, punc- 
tures denser anteriorly, less distinct and more widely 
spaced laterally. Legs dark reddish-brown to piceous. 
clothed with long reddish-golden hairs. Scutellum 
finely, sparsely punctate centrally, nearly glabrous. 
Elytra black, shining, finely, shallowly, irregulari 
punctate, sutural striae moderately impressed, coarsely. 



1 Placerita Canyon Nature Study Center. 1 
Placerita Canvon Road. Newhall, Ca 



69 



70 



in III ll\ sol mi i<\ < ILIFORNIA ACADEMY OF SCIENCES VOLUME 71 




Figure I. Pleocoma marquai, new species, left lateral view of head and pronotum, holotype 
(male). Scale equals 2 mm. 



irregularly punctate, geminate striae at margins of 
costae feeble, indicated only by single row of fine, 
shallow punctures, costae with few minute punctures, 
not elevated. Abdomen dark reddish-brown, sternites 
moderately densely, irregularly punctate, punctures 
moderate in size, thinly clothed with reddish-golden 
pubescence. Length 22-28 mm. 

Female. Form ovate, robust; color reddish-brown; 
pubescence pale reddish. Head with clypeus coarsely, 
densely punctate, expanded apically, apical angles 
acute, rounded, anterior margin feebly sinuate near 
median notch, median notch small, shallow, narrowly- 
rounded; vertical horn very short, stout, apical notch 
broadly obtuse, rounded, apices rounded; antennae 
castaneous. lamellae slightly darker, third segment 
short, subcylindrical, about one and one-half times 



longer than second segment, fourth segment slightly 
transverse, fifth segment with acute process, segments 
six to eleven distinctly lamellate, forming club, sixth 
segment with short lamella, lamellae of segments 
seven and eight subequal, only about three-fourths 
as long as that of segment nine, lamella of segment 
nine longest, those of segments ten and eleven slightly 
shorter than nine and of decreasing length. Pronotum 
convex, shining, dark reddish-brown, lighter laterally, 
less than twice as wide as long (length-to-width ratio 
in allotype 9:16.8), widest at posterior angles, pos- 
terior angles obtusely rounded, disc moderately, ir- 
regularly punctate, punctures larger and denser 
laterally and anteriorly, interrupted medially by vague 
longitudinal impunctate line. Sculellum finely, sparsely 
punctate anteriorly, few punctures with short re- 



I'}/ ' 



THREE NEW SYMPATR1C PLEOCOMA FROM ( \LIFOR I 







Figure 2. Pleocoma fimbriata Lee, left lateral view of head and pronohim. male. Scale 
equals 2 mm. 



cumbent hairs. Elytra widest behind middle, trans- 
parent, surface shining, finely, irregularly punctate, 
costae elevated, nearly attaining elytral apices, sutural 
striae distinct, impressed, coarsely punctate, geminate 
striae at costae, feebly impressed, finely, irregularly 
punctate. Length 31-40 mm. 

Holotype: Male, California, Tulare Co., Posey, 16 
October 1971 (B. D. Streit) LACM. Allotype: Fe- 
male, California. Tulare Co.. Posev. 12 November 
1971 (T. W. Taylor) LACM. 

Paratopes (241): CALIFORNIA. Tulare Co., 144 $ 
15 9, Posey, 8 November 1969 to 20 December 1969, 
23 October 1970 to 26 October 1970, 15 October 
1971 to 29 November 1971 (D. G. Marqua. T. W. 
Taylor, B. D. Streit. G. C. Walters. F. T. Hovore, 
C. E. Langston, Dawn, Rod, and Blaine Bryan. M. T. 
Gannon) DGM 17, TWT 31, BDS 42. FTH 12, CEL 



7, GCW 21, LACM 4. USNM 3. HFH 4. RLW 4, 
CIS 5, CAS 4, MTG 5: 14 a 5 9. .6 mi E Posey. 15 
October 1971 to 25 October 1971 (F. T. Hovore) 
FTH: 23 i 7 9 . 1 mi N and 2 mi N Posey. 16 October 
1971 to 1 December 1971 (C. E. Langston l CEL: 
24 • . Balance Rock Resort, 16 October 1971 to 12 
November 1971 (F. T. Hovore. B. D. Streit) FTH 4. 
BDS 20. Kern Co.. 9 ; . Glennville. 14 November 
1968, 15 November 1969. 26 November 1970. 11. 12 
November 1971 (D. G. Marqua. C. E. Langston. 
B. D. Streit) DGM 3. BDS 2. CEL 4. 

Other specimens examined: CALIFORNIA. Tidarc 
Co., 17$, California Hot Springs. 13 October 196S. 
6 November 1966 (David Fields. 4 with no collector 
listed). 

Discussion and diagnosis: Both sexes 
marquai can be immediately distinguished from 



72 



Hi III n\ sol THERh ( IL1F0RN1 I i< ADEMY OF SCIENCES VOLUME 71 




Figure 3. Pleocoma tularensis Leach, left lateral view of head and pronotum, 
(male). Scale equals 2 mm. 



paratype 



their closest congeners, P. fimbriata Leconte and 
P. tularensis Leach, by the presence of distinct 
lamellae on antennal segments six and seven, with 
that of segment six two-thirds or more as long 
as that of segment seven. In both tularensis and 
fimbriata, segment six is at most angulate, seg- 
ment seven sometimes with a short projection. 
Males of P. marquai may be further differentiated 
from fimbriata and tularensis by the fifth antennal 
segment which is strongly transverse with at least 
a short projection (at most slightly transverse in 
typical fimbriata and tularensis) , and by the 
shorter, less strongly reflexed clypeal process 
(Figs. 1-3). In all specimens examined of 
marquai the pronotum is less than twice as wide 



as long, while in the 59 males of tularensis ex- 
amined, the pronotum is at least twice as wide as 
long in all but two specimens, or about 3 percent. 

Pleocoma marquai represents the southernmost 
form of the fimbriata— tularensis species group. 
Thus far, the northernmost known locality for 
marquai is about 30 miles south of the southern- 
most recorded point of collection of tularensis, 
and it is highly probable that somewhere in the 
intervening area the two populations overlap. 

Biology: Adults of both sexes and larvae of P. 
marquai have been collected from burrows 
beneath Ceanothus cuneatus Nutt., and it is prob- 
able that this plant serves as the primary larval 
host. Pleocoma tularensis males and females have 



1972 



THREE NEW SY MP ATRH PLEOCOMA FROM CAL1FOR IA 







Figure 4. PIcocoma rubiginosa, new species, left lateral view of head and pronotum. holotxpe 
(male). Scale equals 2 mm. 



been collected from pupal cells beneath Ceanothus 
cuneatus in Fresno County, and I have also col- 
lected female P. fimbriate! beneath Ceanothus sp. 
(prob. integerrimus H. and A.) in El Dorado 
County. Although it appears that Pleocoma 
larvae are not necessarily oligophagous (Fellin. 
1966), the fact that the three species in this group 
show an apparent preference for the genus 
Ceanothus as a primary host seems to substantiate 
their close biological relationship. 

Adult activity of P. marquai in the vicinity of 
the type locality begins during or shortly after the 



first precipitation of the fall season, usually in 
early October, and may continue until early De- 
cember in years of relatively low rainfall. Males 
will fly throughout the night while it is raining 
or snowing, and at dusk for several days there- 
after, during weather conditions varying from 
heavy overcast to bright late afternoon sunlight. 
I take pleasure in naming this species after 
David G. Marqua. who first brought specimens of 
Pleocoma from the Posey-Glennville area to i 
attention, and who has made numeroi - 
tions to this studv. 



74 



in III ll\ SOI nil l<\ < ILIF0RN1A ACADEMY OF SCIENCES VOLVMEll 




Figure 5. Pleocoma hoppingi Fall, left lateral view of head and pronotum, male, 
equals 2 mm. 



Scale 



Pleocoma rubiginosa, new species 
Figure 4 

Description: Male. Form robust, broadly oblong- 
oval, moderately convex, dorsum slightly flattened; 
integument reddish-brown, pubescence pale reddish. 
Head dark reddish-brown, narrowly margined an- 
teriorly with piceous, very densely clothed with long 
reddish hairs; dorsal surface irregularly, shallowly 
rugoso-punctate, with vague smooth area extending 
from anterior base of vertical horn laterally to apex 
of ocular canthus; clypeal process moderately re- 
flexed, anterior face distinctly impressed medially, 
apex deeply, obtusely notched, apical angles produced, 
acute, rounded; vertical horn elongate, sides sub- 
parallel, apex with moderate rounded notch, apical 
angles acutely rounded, anterior face of horn only 
slightly concave medially, coarsely, irregularly 
punctate, punctures elliptical, surface clothed with 



long reddish hairs; ocular canthi stout, subquadrate, 
projecting forward slightly from a right angle, an- 
terior edge sinuate, dorsal surface slightly concave, 
nearly glabrous; oblique supra-orbital carina distinct, 
extending medially on ocular canthus; palpi and 
antennae light reddish-brown, lamellae of antennae 
slightly darker, scape stout, subconical, slightly flat- 
tened dorsally, second segment moniliform, strongly 
flattened, third segment elongate, subequal to scape 
in length, slightly angulated anterior at apex, fourth 
segment only slightly transverse, angulated, fifth 
segment strongly transverse with acute process, seg- 
ments six to eleven distinctly lamellate, sixth segment 
with lamella about two-thirds as long as that of seventh 
segment, lamella of seventh segment more than four- 
fifths as long as that of eighth segment, that of eighth 
distinctly shorter than that of ninth, lamella of ninth 
segment longest, those of tenth and eleventh slightly 
shorter than ninth, and of decreasing length, ratios 



1972 



THREE NEW SYMPA1 me PLEOl OMA I ROM ' \LIFO) 






of segments six lo eleven in holotype 20:33:40:43: 
42:40. Pronotum less than Iwiee as wide as long, 
barely widest at middle, sides broadly rounded, pus 
tenor angles broadly, obtusely rounded, lateral diseal 
impressions feeble, maculate with piceous; dise eon- 
vex, anterior median impression moderate, flattened, 
surface shining, densely, coarsely punctate except for 
narrow, median longitudinal impunctate line extend- 
ing from anterior lo posterior margins, entire surface 
of disc densely clothed with long, erect reddish hairs. 
Scutellum irregularly, moderately coarsely punctate, 
moderately densely clothed with long snbreenmbent 
reddish hairs. Elvira rich reddish-brown, transparent, 
shining, fairly uniformly, finely, shallowly punctate, 
sutural striae moderately impressed, irregularly 
punctate, geminate striae at margins of costac feeble, 
indicated only by single row of fine, shallow punc- 
tures, costac impunctate, not elevated. Abdomen 
dark reddish-brown, stcrnites finely, moderately 
densely punctate, clothed with long reddish pubescence. 
Length 24-29 mm. 

Holotype: Male, California, Tulare Co.. 1 mi W 
Posey, 16 January 1972 (C. E. Langston) LACM. 

Paratypes: 41 i , same locality as holotype. 2 
January 1972 to 1 April 1972 (C. E. Langston) at 
blacklight trap. CEL 20. LACM 2. CAS 2, CIS 2, 
USNM 1, FTH 15, TWT 1, BDS 1, DGM I, HFH 1. 
RLW 1. 

Discussion and diagnosis: Pleocoma rubiginosa 
is perhaps only subspecifically related to P. hop- 
pingi Fall, which it closely resembles in form and 
color. Males differ from hoppingi by the presence 
of only five long lamellae in the antennal club 
(seven in typical hoppingi) , the absence of a 
flattened anteroventral process on the third an- 
tennal segment, the conspicuous median longitudi- 
nal impunctate line on the pronotal disc, the much 
more produced, more strongly reflexed clypeal 
process, and the longer vertical horn (Figs. 4-5). 
From the other similarly colored species, badia 
Fall and linsleyi Hovore, rubiginosa can be im- 
mediately distinguished by the fewer antennal 
lamellae (seven long lamellae in badia and 
linsleyi), densely hairy, heavily punctate pro- 
notal disc, and subquadrate ocular canthi. 

As in most of the other species of Pleocoma, 
males of rubiginosa exhibit some variation in ex- 
ternal morphology. The relative lengths of the 
first few antennal lamellae vary slightly within 
the type series, but the basic diagnostic formula 
is sufficiently constant to distinguish rubiginosa 
from other closely related species. The absolute 
shape and size of the clypeal process is also some- 
what variable, but in all specimens it is much more 
produced and more strongly reflexed than in any 
specimen examined of hoppingi. 



Mi, lmn : Nothing is known "i tl i lai ■ 4 habit 
this specie. The specimens of thi typi crii 
captured with blacklighl traps from lanuar) I 

April, considerably later in thi on than the 

normal majoi flighl periods ol othci Pleocoma in 

the area. Although the traps were I I 
various limes during any given night, spci 
were usually found in the Iraps in the morning, 
indicating that males ol rubiginosa, like tin 
several other species, fly most numerousi) at 01 
slighily before dawn (Hazeltine, 1950, 1952: 
Hovore, 1971). Most of the specimens were col- 
lected during heavily overcast skies and log. with 
light rain or snow. A few specimens were found 
in the traps during a week of warming weather 
following a major snowstorm, flighl probably 
having been initiated by melting snow or ground 
ice late season flight habits have also been 
recorded for /\ hoppingi on the South Fork of the 
Kaweah River (Fall, 1906): in the vicinity ol 
Yosemite National Park (Linslcy. 1941. 1942. 
1943); and at Dry Meadow, Kern County 
(I.insley, 1957). It is probable that the last record 
refers to a population more closely related to 
rubiginosa than to typical hoppingi. Dry Meadow 
is in the Piute Mountains, about 24 miles south- 
east of Posey, while the type locality for hoppingi 
(South Fork of the Kaweah River. Tulare County) 
is approximately 48 miles north of Posey. Linsley 
(1938b) lists hoppingi from Kernville. approxi- 
mately 8 miles east of Posey, and this record un- 
doubtedly refers to a specimen of rubiginosa 
rather than typical hoppingi. 

Pleocoma hirticollis rcflcxa. new subspecies 
Figure 6 

Description: Male. Form robust, oblong-oval, con- 
vex, dorsum only slightly flattened: integument 
reddish-brown to piceous: pubescence pale golden. 
Head black, vertical horn and anterior process of 
clypeus moderately densely clothed with long, pale 
golden hairs; dorsal surface irregularly punctate: 
clypeal process pronounced, strongly reflexed. nearly 
parallel with vertical horn, anterior face deeply con- 
cave medially, apex with deep, acutely rounded notch, 
apical angles acute, rounded: vertical horn elongate. 
sides subparallel, apex with shallow, obtusely rounded 
notch, anterior face of horn evenly convex, surface 
coarsely punctate, punctures elongate: ocular canth 
nearly right angular to longitudinal midline of head, 
anterior edge sinuate, apex broadly, acutely rounded, 
dorsal surface concave, irregularly punctate: pah 
antennae dark reddish-brown, scape and lame 
slightly darker, scape stout, subcorneal, second sec- 



76 



i;i iii n\ SOI ////a'\ c.u.iroiiMA academy of sciences voevmeu 




Figure 6. Pleocoma hirticollis refle.xa, new subspecies, left lateral view of head and pronotum, 
holotype (male). Scale equals 2 mm. 



ment moniliform, flattened, third segment elongate, 
subcylindrical, slightly reflexed, fourth segment 
strongly transverse with short projection, segments 
five to eleven distinctly lamellate, fifth segment with 
lamella more than four-fifths as long as lamella of 
sixth segment, lamella of sixth segment only slightly 
shorter than lamella of seventh segment, lamella of 
seventh segment subequal to that of eighth segment, 
that of eighth segment slightly shorter than that of 
ninth segment, ninth longest, lamellae of tenth and 
eleventh segments subequal, slightly shorter than that 
of ninth. Pronotum piceous, more than twice as wide 
as long, widest at posterior angles, posterior angles 
rounded, lateral discal impressions distinct, maculate 
with pale reddish-brown; disc strongly convex, ir- 
regularly clothed with long, suberect pale hairs, an- 
terior median impression indicated by narrow flatten- 
ing of anterior surface, surface shining, densely, 
irregularly punctate, punctures small to moderate in 



size on disc, larger anteriorly. Scutellum finely, 
moderately densely punctate, moderately clothed with 
long recumbent hairs. Elytra rich chestnut-brown, 
transparent, shining, finely, shallowly, irregularly 
punctate, sutural striae moderately impressed, ir- 
regularly punctate, geminate striae at margins of 
costae feeble, indicated only by single row of fine, 
shallow punctures, costae impunctate, not elevated. 
Abdomen reddish-brown, sparsely punctate, thinly 
clothed with long, pale golden hairs. Length 20-25 
mm. 

Female. Form robust, convex; color reddish-brown, 
pubescence pale golden. Head with dorsal surface 
densely, coarsely rugoso-punctate; clypeus expanded 
apically. apical angles obtuse, anterior margin 
bisinuate. median notch small, very shallow, rounded; 
vertical horn very short, stout, apical notch shallow, 
rounded, anterior face deeply impressed medially; 
ocular canthi subtriangular, anterior edge feebly 



1972 



THREE NEW SYMPATRIC PLEOCOM I FROM ' ALlFOi 







Figure 7. Pleocoma hirticollis vandykei Linsley. left lateral view of head and pronotum. 
male. Scale equals 2 mm. 



sinuate, apices acutely rounded; antennae reddish- 
brown, scape and lamellae slightly darker, third seg- 
ment elongate, subequal to scape in length, fourth 
segment with acute projection, about one-third as 
long as lamella of fifth segment, segments five to 
eleven lamellate, forming club. Pronotum dark reddish- 
brown, lighter laterally: slightly less than twice as 
wide as long, widest at posterior angles, posterior 
angles obtuse, rounded; disc convex, shining, surface 
coarsely rugoso-punctate, punctures forming in- 
distinct transverse rows, denser and larger anteriorly, 
lateral discal impression feeble, maculate with 
piceous. Scutellum sparsely punctate medially, few 
punctures with short erect hairs. Elytra pale reddish- 
brown, transparent, surface shining, finely, irregularly 
punctate, sutural striae feebly impressed, indicated 
by single row of punctures, geminate striae at margins 
of costae indistinct, finely, shallowly punctate, costae 
only slightly elevated, nearly attaining elytral apices. 
Length 29-31 mm. 



Holotype: Male. California. Tulare Co.. 1 mi N 
Posey, 18 December 1971 (C. E. Langston) at black- 
light trap. LACM. Allotype: Female. Tulare Co.. I 
mi W Posey. 3500 ft. 27 February 1972 (F. T. 
Hovore) dug from pupal cell. FTH. 

Paratypes (17): CALIFORNIA. Tulare Co.. 12 $, 
1 mi N Posey. 18 December 1971. 21 December 1971 
(C. E. Langston) at blacklight trap. CEL 5. FTH 3, 
CAS 1. CIS 1. LACM 1. USNM 1. Kern Co.. 2 
Glennville and vicinity. 14 January 1970. IS January 
1970 (D. G. Marqua). DGM: 1 ' . Glennville. 26 
November 1970 (F. T. Hovore) FTH: 1 ' Glennville. 
26 November 1970 (C. E. Langston | TWT; 19, 
Glennville. 25 November 1970 (Dawn Bryant TWT. 

Discussion and diagnosis: Three apparent!) 
distinct forms of Pleocoma hirticollis Schaufuss 
can be recognized: P. hirticollis vandykei Linsley 
(Fig. 7) occurs in the grassland regions stir 
ing San Francisco Bay: the nominate 
P. h. hirticollis Schaufuss. has been c 



78 



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VOLUME 71 



the Sierra Nevada foothills of Yuba and Nevada 
Counties: /'. h. reflexa occurs in the foothills of 
the Greenhorn Mountains along the Kern-Tulare 
Count) border. Males of the three subspecies may 
be distinguished as follows: 

Third segment o( antenna usually angulate an- 
teriorly, lamella of fourth segment one-third or 
more as long as that of fifth segment, lamella of 
eighth segment usually longest: clypeal process 
moderate, slightly reflexed: elytra chestnut-brown. 
Length 25-28 mm. Yuba and Nevada Counties, 
California h irticollis Schaufuss (1870). 

Third segment of antenna usually not angulate 
anteriorly, lamella of fourth segment less than one- 
third as long as that of fifth segment, lamella of 
ninth segment usually longest: clypeal process re- 
duced, only feebly reflexed; elytra piceous to 
black. Length 17-24 mm. Alameda, Sonoma, and 

Yolo Counties, California — 

_ vandykei Linsley (1938). 

Third segment of antenna usually not angulate 
anteriorly, fourth segment of antenna transversely 
angulate or with vestigial lamella, much less than 
one-third as long as that of fifth segment, lamella 
of ninth segment usually longest: clypeal process 
very pronounced, strongly reflexed; elytra chest- 
nut-brown. Length 20-25 mm. Kern and Tulare 
Counties, California reflexa Hovore. 

In the type series the specific diagnostic char- 
acters are remarkably constant. The anterior 
clypeal process exhibits slight variation in size and 
in the depth of the median notch, but in all speci- 
mens the clypeus is more pronounced and much 
more strongly reflexed than in any specimen seen 
of either vandykei or hirticollis. The color of the 
pronotum in the male may vary from reddish- 
brown to piceous. 

Biology: Males of reflexa in the vicinity of the 
type locality have been collected only after the 
area has received several inches of precipitation, 
with dates ranging from late November to mid- 
January. Most specimens were collected at night 
or at dawn, during or shortly after a drizzling rain. 
I collected a male at incandescent light shortly 
after dusk following a day of clear cold weather, 
and it is probable that male activity continues for 
several days subsequent to a major flight. 

The allotypic female and two larvae were dug 
from their burrows in a grassy clearing, at depths 



of from one to four inches. The paratype female 
was found in a schoolyard baseball field. It is 
possible that the larvae of reflexa feed upon grass 
rootlets, as do those vandykei (Smith and Potts, 
1945; Ritcher. 1947). but the soil from which 
they were dug is interlaced with numerous rootlets 
of Ceanothus cuneatus and Quercus Douglasii H. 
and A., and a definite feeding preference has not 
vet been observed. 



ECOLOGY OF THE TYPE LOCALITY 

The sympatric occurrancc of three distinctly dif- 
ferent species of Pleocoma is unique in the 
recorded distribution of the genus. There are 
several localities known where two species are 
sympatric, and three species {fimbriata, hirticollis, 
and edwardsi Leconte) have been recorded from 
the vicinity of Nevada City, Nevada County, Cali- 
fornia. However, the exact localities of their col- 
lection have not been documented in sufficient de- 
tail to establish a definite area of sympatry. 

The site near Posey where P. rubiginosa, P. 
marquai, and P. h. reflexa are sympatric is in a 
rather poorly defined ecotonal area. The grassy hill- 
sides surrounding the site are dotted with deciduous 
Blue Oaks (Quercus Douglasii H. and A.) and 
Buckbrush (Ceanothus cuneatus Nutt.), charac- 
teristic of the Foothill Woodland plant community, 
while the shaded canyon bottoms hold mature 
stands of Yellow Pine (Pinus ponderosa Dougl.), 
Incense Cedar (Libocedrus decurrens Torr.), 
Alder (Alnus sp.), Maple (Acer macrophyllum 
Pursh.), and Willow (Salix sp.). Yellow Pine and 
Incense Cedar indicate the lowest influence in 
this area of the Yellow Pine Forest plant com- 
munity (Munz and Keck, 1970), which pre- 
dominates at higher elevations. 

The blacklight traps from which the specimens 
were collected were placed near private dwellings 
along the bottom of a well-drained west-facing 
canyon at an elevation of approximately 3500 ft. 
The north-facing canyonside is dominated by large 
Black Oaks (Quercus Kelloggii Newb.), with 
brush cover primarily composed of low-growing 
stands of Gooseberry (Ribes sp.). Blackberry 
(Rubus sp.), Rose (Rosa calif ornica Cham, and 
Schlect.). Buckbrush. and Mountain Mahogany 
(Cercocarpus betuloides Nutt.). A thick carpet 
of leaves covers much of the slope, and the soil 
beneath is a poorly consolidated clay loam. On 
the drier, south-facing canyonsides the dominant 
tree is the Blue Oak, with brush cover widely 
spaced, composed of Buckbrush, Coffeeberry 



I')72 



THREE NEW SYMPATR1C PLEOCOMA I hum < \LIFORI I I 






(Rhamnus calif ornica Ksch.), Hlderbcrry (Sam- 
bucus mexicana Presl.), Mountain Mahogany and 
Honeysuckle (Lonicera interrupta Benth.). The 
groundcover consists of numerous grasses and 
small annuals, with Mariposa Lilies (C'alocboi tits 
sp.) and Buckwheat (Eriogonum sp.) abundant 
in May and June. The soil varies from hard- 
packed decomposing granite to a loose clay loam. 

Pleocoma marquai shows no evident preference 
for a particular soil type, and is extremely abun- 
dant beneath most stands of Buckbrush. The eleva- 
tion range observed for marquai is from approxi- 
mately 2900 ft to 4500 ft, and roughly follows 
the irregular lower margins of the Woodland- 
Forest ecotonal area. The vegetation mix is simi- 
lar to that of the recorded localities for fimbriata 
and tularensis. These areas usually receive snow 
only during the colder winter storms, often not 
until late in the season. The flight periods of 
marquai, tularensis, and fimbriata are somewhat 
earlier than those of most other Pleocoma species 
in the Sierras, and in normal winters they will have 
experienced their major period of adult activity 
before the first significant snowfall. 

Pleocoma rubiginosa has only been collected 
at the type locality, at an elevation of 3500 ft. 
although its affinities are to species which favor 
higher elevations. Its closest congener, hoppingi, 
has been recorded from 4000 ft to above 6000 ft, 
and specimens of an undescribed species in the 
"hoppingi group" have recently been collected on 
the eastern side of the Sierras at elevations up to 
10,000 ft. It is therefore probable that rubiginosa 
is more widely distributed in the Yellow Pine 
Forest community above the Posey site, and prob- 
ably does not occur much lower than 3400 ft. 

The affinities of P. h. reflexa are to Pleocoma 
populations primarily found in foothill grassland 
areas, and at the Posey site specimens have been 
dug from the soil on the south-facing slope in an 
open grassy clearing. The other localities for 
reflexa are also in the grassy Foothill Woodland 
community, at 3150 ft and 3700 ft elevations. 
It is probable that reflexa is the most widely dis- 
tributed Pleocoma in this area, as it would appear 
to have the widest potential area of suitable habitat. 
If the larva is in fact a mobile rootlet feeder in 
grassy areas, reflexa could conceivably occur in 
several floral communities at elevations both 
higher and lower than presently recorded. The 
hirticollis species group is found in both the 
Coastal and Sierra Nevada Ranges, and may ulti- 
mately prove to be the most widespread and 
evenly distributed species in the genus. 



ACKNOW1 I DGM1 I S 

I he authoi wishes lo extend Ins gratitude i" Ihc follow 
ing persona and institutions i Gorton Linili 
John A. ( hemsa) oi thi Univi ity of < alifi 
Berkeley, foi ihcir generously given time nd d 
sistance during the initial research tagi "I thii 
manuscript; Paul Arnaud and Hugh Leech ol Ihc 
( alifornia Academy of Sciences foi making availabli 
the type specimens oi Pleocoma undei theii can 
Charles Hogue of the Natural Histor) Museum "i I o 
Angeles County for reviewing the text ol the inanu 
script. To the following individuals special gratitude 
is extended for having loaned their fine collection! 
of Pleocoma to the author: D. G. Marqua. I W 
Taylor. C. E. Langston. B. I). Streit. and \l I 
Gannon. 



LITERATURE CITED 

Fall. H. C. 1906. A new Platycerus and a new 
Pleocoma. Km. News. 16:393-395. 

Fellin. D. G. 1966. Biology and feeding habits of 
Pleocoma larvae in western Oregon coniferous 
forests. Oregon State University, unpublished 
Ph.D. dissertation. 156 pp. 

Hazeltine. W. 1950. Observations on flights of 
Pleocoma conjungens. Pan-Pacific Ent.. 26:188- 
189. 



. 1952. Notes on flights and food plants 

of Pleocoma. Pan-Pacific Ent.. 28:202. 

Hovore. F. T. 1971. A new Pleocoma from South- 
ern California with notes on additional species. 
Pan-Pacific Ent.. 47:193-201. 

Linsley. E. G. 1938a. Notes on the habits, distribu- 
tion and status of some species of Pleocoma. 
Pan-Pacific Ent.. 14:49-58. 



— . 1938b. Notes on the habits, distribution 
and status of some species of Pleocoma. Pan- 
Pacific Ent.. 14:97-104. 



. 1941. Additional observations and descrip- 
tions of some species of Pleocoma. Pan-Pacific 
Ent.. 17:145-152. 



. 1942. Notes on the habits of some beetles 

from the vicinit\ of "i osemite National Park. 
Bull. So. California Acad. Sei.. 41:164-166. 



— . 1943. Pacific Coast Ent. S -dines, 

in Pan-Pacific Ent.. 19 : 5o. 



S7J 



/(( / / / //\ SOI I III R\ ( II II ORXIA ACADEMY Of- SCIENCES 



VOLUME 71 



. [957, Distributional records for some Schaufuss, L. W. 1870. Pleocoma staff. Nunquam 

species of Pleocoma. Pan-Pacific I'm.. 33:102- Otiosus, 2:50-59. 



104. 



Munz, P. v. and D D Keck 1970. A California 

Flora. Univ. California Press. 5th Ed. 1550 pp. 

Ritcher, P. O. 1947. Description of the larva of 
Pleocoma hirticollis vandykei 'Linsley. Pan- 
Pacific Ent., 23:11-20. 



Smith, R. F., and R. W. L. Potts. 1945. Biological 
notes on Pleocoma hirticollis vandykei Linsley. 
Pan-Pacific Ent.. 21: 115-1 IK. 



Accepted for publication May 8, 1972. 



(COLEOPTERA: 



A REVIEW OF EUCYLLUS HORN 
CURCULIONIDAE, BRACHYRHININAE, PERITELINI) 



Frank W. Pelsue and Elbert L. Sleeper 1 



Abstract: The genus Eucyllus Horn is reviewed with the three species: vagans Horn. 
unicolor Van Dyke, and echinus Van Dyke, redescribed and discussed. Three additional 
species are described as new: saesariatus from Sonora, Mexico, and Arizona, carinarostris 
from California, and cinereus from California. Eucyllus tinkhami Tanner is placed in the 
genus Eucilinns Buchanan. Distributional and biological information is given for each. 



While we were reviewing the Peritelini it became 
evident to us that the whole tribe was in need of 
study. Many new species were encountered in 
fieldwork in western North America. Because an 
abundance of material exists and several new 
species required description, the genus, Eucyllus 
Horn was chosen as a starting point. 

The following standard abbreviations ( Arnett. 
ct al., 1969) are used: BYUC — Entomological 
collections. Brigham Young University: CASC — 
Entomological collections. California Academy of 
Sciences: CIAN — Centro de Investigaciones 
Agricolas del Noroeste; CSCLB — Entomological 
collections, California State University, Long 
Beach: ELS — E. L. Sleeper collector; ELSC— 
E. L. Sleeper collection: FWPC — Frank W. Pelsue 
collection: INIA — Instituto Nacional de Investiga- 
tiones: LACM — Natural History Museum of Los 
Angeles County: and USNM — United States Na- 
tional Museum. 

Field studies in Baja California were supported 
by the Association for Biological Studies in Baja 
California. Studies in the desert regions of South- 
western United States and the Joshua Tree Na- 



tional Monument were supported by grants from 
the Long Beach California State College Founda- 
tion. 

EUCYLLUS HORN 

Eucyllus Horn. 1876:74. 
Encyllus Van Dyke, 1936:31. (lapsus). 
Type species: Eucyllus vagans Horn, by monotypy. 
Description: Rostrum slightly longer than broad, con- 
tinuous with head; epistoma smooth, devested of 
scales, triangular in shape with carinate margins, and 
a fringe of 10-12 setae bordering the carinate margins; 
head not as long as broad, with smaller and more 
abundant punctures than rostrum, little or no sculp- 
turing; antennae inserted in the basal third of rostrum; 
scrobe a flat-bottomed channel with no pronounced 
convex area distad of the eye. Funiculus with seven 
segments, vestiture of scales and setae similar to re- 
mainder of body; scape feebly arcuate attaining an- 
terior margin of prothorax; club oval-acuminate, 
clothed with short fine setae. Eyes nearly round, 



1 Dept. Biology, California State University. Long 
Beach, California 90840. 



1972 



REVIEW <)E THE snoiii BEETLES, GENl S / ( < YLU 



HI 












Figures l-10a. Spermathecae of Eucyllus. 1. Eucyllus vagans Horn. Mexico. Baja Cali- 
fornia Norte. Puertecitos. V-31-58. 2. E. vagans, California. San Bernardino Co.. Upper 
Lucerne Valley. X-2-64. 3. E. vagans, California. San Diego Co.. Pamitas Spring (3 mi W 
Carrizo Springs. Ill- 1-58. 4. E. vagans, Mexico. Baja California Norte. Laguna Salada. 11-20-59. 
5. E. saesariatus. new species, Mexico. Sonora. 51 mi E San Luis. XII-30-60. paratype. 6. E. 
carinarostris, new species. California. Riverside Co.. Pinto Wash Well. II-l 7 -62. holotype. 
7. E. unicolor Van Dyke. Nevada. Nye Co.. 19 mi SE Lathrop Wells. III-24- _ 0. 8. E. echinus 
Van Dyke. Nevada. Lincoln Co.. 10 mi N Hwy. -91 on Elgin Road. V-1S-62. 9. E. echinus. 
Mexico. Sonora, 51 mi E San Luis. XII-30-60. 10. E. cinereus. new species. California. Inyo 
Co.. Grandview P.C., VIII- 10-65. paratype. 10a. E. cinereus. a different individual. Line 
represents ] 2 mm. 



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VOLUME 71 



strongl] convex, and coarse!} faceted. Prothorax not 
.is long as broad, sides feebly arcuate, anterior margin 
truncate, posterior margin rounded. Sctitellum small. 
obscured by elytra. Elytra nearly twice as long as 
broad; humeri rounded, feebly prominent, sides 
parallel, tapering to apex: striae defined by round 
deep punctures, setae usually arranged in two longi- 
tudinal rows alternately spaced on the interval. Venter 
of body with vestiture similar to dorsum; however, 
the recurved setae are slender, and truncate at apices; 
pro- and mesocoxae contiguous with metacoxae, 
widely separated. Femora one-third longer than 
tibiae, beginning rather slender and gradually be- 
coming clubbed distally. no teeth or definite spines 
present, vestiture of scales and recurved setae similar 
to other parts of the body; tibiae slender, broadening 
distally. vestiture similar to femora, apex with teeth, 
tibiae 1 and 2 with a short mucro. tibia 3 without 
mucro: tarsus with first segment as long as second 
and third combined, fourth segment almost as long 
as first three segments combined, dorsal vestiture of 
segments 1-3 recurved setae and scales, fourth segment 
with fine setae. 

Remarks: Horn described Eucyllus in 1876 
placing it between Dysticheus Horn and Thin- 
oxenus Horn, a placement retained by all sub- 
sequent workers through Kissinger (1964). Eucyl- 
lus Horn, Thinoxenus Horn, Dysticheus Horn, 
Rhypoclillus Cockerell, Eucillinus Buchanan, and 
two undescribed genera form a group of mor- 
phologically very similar genera. The described 
genera can be separated easily in Kissinger (1964). 

Eucyllus, for the most part, inhabits arid habi- 
tats of the desert (Fig. 32). With the exception 
of E. cinereus, members of this genus are restricted 
mostly to the Creosote Bush Scrub plant com- 
munity, with E. echinus and saesariatus barely 
reaching into the Mesquite-Grassland plant com- 
munity. Eucyllus cinereus is found in Pinyon- 
Juniper communities. The other genera of the 
group are coastal or montane. 

Eucyllus remained monotypic until 1936 when 
Van Dyke added two species; E. echinus and 
unicolor. In this paper we are describing three 
new species of Eucyllus, bringing the total number 
of species to six. 

In 1959, Tanner described Eucyllus tinkhami. 
Upon re-examination of paratypes of this species, 
particularly the proventriculus and tibial apices, we 
conclude that it does not belong in Eucyllus and 
is in fact a subspecies of Eucilinus (Thysano- 
chorhinus) aridus Van Dyke. It should then be 
known as Eucilinus (Thysanoehorhinus) aridus 
tinkhami (Tanner), new combination. The type 
locality for E. aridus aridus is Cronise Lake, Cali- 



fornia, approximately 35 miles northwest of the 
Kelso Dunes; the type locality of E. aridus 
tinkhami. 

The paucity of specimens of Eucyllus in collec- 
tions results from the fact that members of the 
genus are nocturnal and seek refuge about the 
bases of food plants during the daylight hours 
when most insect collectors are active. Eucyllus 
vagans and saesariatus have been taken at night 
throughout the year. 

Species of Eucyllus are largely parthenogenetic; 
males being less frequently encountered than fe- 
males. Populations from south and east of the 
Colorado River have a higher frequency of males 
than those encountered elsewhere. In the material 
available to us, E. echinus from Arizona has a 
male-female ratio of 1 :2.5; whereas, in saesariatus 
it is 1:1.25. Males were not found in the popula- 
tion of E. echinus from Nevada and Utah, E. 
unicolor, or E. carinarostris. 

Eucyllus vagans has an interesting change in sex 
ratio from south to north in its range. In Baja 
California del Norte, Mexico (the extreme south- 
ern part of the range) the male-female ratio ap- 
proaches 1:1 from below sea level to about 1000 
ft elevation. No males are known from above 
1000 ft. In the vicinity of Borrego Valley and the 
Salton Sea in California the ratio is 1:1.5. From 
north of the Riverside-Imperial County line or the 
vicinity of the northern end of the Salton Sea, only 
one male is known. It is a very old specimen, 
presented to Sleeper by J. G. Shanafelt who veri- 
fied that the data were correct. It bears the 
locality label "Taquitz Cyn. nr. Palm Sprs." From 
the remainder of the range northward and north- 
eastward for more than 260 miles, no males are 
known. 

In all species, except cinereus, the number of 
mature ova per female at the time of peak pro- 
duction varied from 5-10, with a mean of 8.25 
per female. There seemed to be no significant dif- 
ference between species as to ova number. Ova 
averaged 0.5 mm in echinus and 0.65 mm in 
vagans. Eucyllus vagans deposits approximately 
250 eggs over a one to two month period after 
which ova production nearly ceases. Ova are de- 
posited in the soil about the bases of the food 
plant. 

In the four species in which males are known, 
there are three distinct types of aedeagi. In the 
larger species the aedeagi are very long, slender 
and twisted at the apical third, terminating apically 
in a more slender, elongate, blunt knob (Fig. 20). 
In echinus the aedeagus is rather short, parallel- 



1972 



REVIEW <)l- I III. SNotiT BEETLES, GENl ; i I ( YLU 






sided and tapering apically lo an abrupt, hluni 
apex, not curved or twisted (Fig. 21). ["he 
aedeagus of cinereus is short, narrowed, and blunt 
at the apex (Fig. 19). 

The spermathecae are useful for separating the 
species. Both fresh and dried specimens were 
dissected; no delectable differences were noted be- 
tween either. 



KEY TO THE SPECIES OF EUCYLLUS 

1. Larger species, 4.5-7.7 mm in length; second 

segment of funiculus almost twice as long as 
third .. 2 

I'. Smaller species, 3.7-5.3 mm in length; second 

funicular segment slightly longer than third 3 

2. Setae of dorsal surface of body 4-7 times as 

long as broad, a few only 2-3 times as long 
as broad, acute to blunt apices, with a 
strongly arcuate concavity on posterior side: 
feeble indication of a median sulcus on 

rostrum (Fig. 28) . vagans Horn 

2'. Setae of dorsal surface of body 10-15 times 
as long as broad with a few 4-7 times as long 
as broad, acute apices, with feebly arcuate 
concavity on posterior side: rostrum with a 
feeble indication of a median carina (Fig. 
31) saesariatus, new species 

3. Setae of dorsal surface as much as three times 

as long as broad, blunt at apices, erect and 

appearing club-like 4 

3'. Setae of dorsal surface not more than twice as 
long as broad, recurved and peg-like in ap- 
pearance 5 

4. A rather robust species with many dark patches 

on the elytra and prothorax. overall color 
straw; setae robust and erect; feeble indica- 
tion of a median carina on dorsum of rostrum 
posterior to the epistoma (Fig. 23): sper- 
matheca as in figures 8-9 .... echinus Van Dyke 
4'. A smaller species, uniformly gray in color with 
a few darker patches; setae finer than pre- 
ceding species, prominently erect; sper- 

matheca as in figures 10-10a 

.. cinereus, new species. 
[Note: the recently described species Eucyl- 
lus horridus Hatch (1971:265) will key to 
cinereus. It appears that horridus is not a 
Eucyllus, but belongs to an as yet unde- 
scribed genus of Peritelini. Although this 
casts some doubt on the placement of 
cinereus in Eucyllus. it seems wise to leave 
all species in their present status until the 
new genera of this group are described.] 

5. Strong median carina posterior to epistoma 

(Fig. 22); spermatheca as in figure 6 

carinarostris. new species 

5'. Feeble indication of a median sulcus posterior 



to epistoma p u in figi 

nun ill, ,r Van 

/ iii i Uus agan I lot n 
Figures 14. 15, 28, JO 

Eucyllus vagans Horn 1876 6 I pe local it) 
/una and California adjacent Hen r< I icted in 
C'arri/.o Springs. Imperial ' o., ( alifornia i he <>nl 
specimen in the Museum of Comparative Zoology 
Harvard University was examined and is dcsigi 
lectotype. It bears the following data I > gold di • 
(= California), 2) 1937. 3) Type, 4) Eucyllus 

vagans Horn. 5) lectotype label- (A female). 

Description: Body length 4.5-7.7 mm (mean 
5.85; 9 6.77 mm); width 2.3-3.5 mm (mean i 2.56; 
9 3.03 mm); elongate oval; vestiture brownish gray 
with silver patches on the head, prothorax and ab- 
domen. Body densely clothed with white and silver 
triangular scales; moderately clothed with erect to 
suberect and recurved seta-like scales, set in deep 
punctures, four to seven times longer than broad at 
base with acute to nearly blunt apices. Shape of 
elongate setae, in cross section, indicates a strongly 
arcuate concavity. Rostrum clothed with white scales 
with a small silver patch dorsad of each eye and a 
median patch extending from posterior margin of the 
head to just past the anterior margin of the eyes: 
strongly sculptured with large punctures, this evident 
when denuded: with feeble indication of dorsal sulcus 
extending from just posterior of epistoma almost lo 
head. Head feebly sculptured with smaller and more 
abundant punctures than rostrum. Funiculus with 
first and second segment same length, second twice 
length of third. Prothorax with vestiture of while 
scales with three longitudinal stripes of brown to 
silver scales; disc coarsely punctured with nearly 
round deep punctures with a mean of 17.1 per .25 
mm- and rather uniform in distribution. Elytra with 
dirty white scales and several irregular patches of 
silvery to brown scales forming no definite pattern. 
Femora with band of silvery scales al distal third: 
apical comb of tibia with anterior teeth rather short 
and thick, becoming slightly longer and thinner 
towards posterior. Aedeagus long and rather slender, 
sides almost parallel for basal third of length tapering 
for another third then broadening again twisting down, 
tapering to a blunt knobbed apex: spermatheca as in 
figures 1-4. 

Distribution: California. In\o Co., 8.5 mi SW West- 
gard Pass. 6000 ft. eastward (below 6000 ft) to 
Nevada. Nye Co.. Cane Springs, thence to the 
Mormon Mountains in Lincoln Co., Washington Co.. 
Utah, southeastward to near Colorado City. Arizona, 
westward to Littlefield, Arizona, south to the Colo- 
rado River, west of the Colorado to the delta in 
Baja California del Norte then south lo Miss 



84 



BULLETIN SOUTHERN t IL1FORNIA H [DEMY OF SCIENCES VOLUME 71 




I')72 



REVIEW OF THE SN0U7 BEETLES GENl LLVS 




Figures 22-31. 22. Rostrum of Eucyllus carinarostris, 2. California. Riverside Co.. Pinto 
Wash Well, 111-17-62. holotype. 23. Same of E. echinus, 2. Mexico. Sonora. 51 mi E San 
Luis, XII-30-60. 24. Same of E. unicolor, 2, Utah. St. George. 25. Apical setae of E. echinus. 
26. Same of E. cinercus. 27. Same of E. unicolor. 28. Rostrum of E. vagans. 29. Apical 
setae of elytra of E. saesariatus. 30. Same of E. vagans. 3 1 . Rostrum of E. saesariatus. 
Figures of various scales. 



Figures 11—21. 11. Eucyllus unicolor. 2, St. George. Utah. 12. E. echinus. 8, Arizona. 
Pinal Co.. Picacho. V-l-42. 13. E. echinus. 2. Mexico. Sonora. 51 mi E San Luis. XII-30-60. 
14. E. carinarostris. 2. California. Riverside Co.. Pinto Wash Well. 11-17-62. paratype. 15. E 
vagans. 2. California. San Diego Co.. 3 mi W Carrizo Springs. XI-15-58. 16. E. 
2, Mexico, Sonora. 51 mi E San Luis. XII-30-60. holotype. 17. Eucilinus {Thysan 
aridus tinkhami, 2. California. San Bernardino Co.. Kelso Dunes. V-25-58, ELS. IS. 1 
cinercus. 2, California, Inyo Co.. White Mts.. Grandview Public Camp. YIII-1( 
19. E. cinercus aedeagus. 20. E. saesariatus aedeagus. 21. E. echinus aedeagus. f 
the line equal to 1 mm. Figures 19-21 no scale. 



86 



HI I II ll\ SOI THERN CALIFORNIA ACADEMY Ol '-SCIENCES 



VOLUME 71 



I mii i I. si/o of Ettcyltiis vagans Horn from south 
to north. All measurements in millimeters. Each 
sample contained 30 random!} selected individuals. 



l ength 



W Kill, 



Mean ( range) ± SD 



Mean (range) -i SD 



Mission Calamiijne, 900 ft.. Baja California del Norte 
■ 5.45 (4.5-5.8) ±.012 2.38 (2.0-2.6) ±.013 

9 6.23 (5.5-6.9) ± .014 2.78 (2.5-3.1 ) ± .026 

Vicinity of Carrizo Springs. 250 ft.. Imperial Co., 

California 
' 5.'H) (5.3-6.4) ± .01 I 2.51 (2.4-2.6) ± .0006 

2 6.31 (5.5-6.8) ± .012 2.76 (2.4-3.0) ± .002 

Indian Cove. Joshua Tree National Monument, 

Riverside Co.. California* 

2 6.17 (5.0-6.5) ± .012 2.77 (2.2-3.0) ± .025 

Lucerne Valley. 3000 ft., San Bernardino Co., 

California 1 '-' 

2 6.66 (6.2-7.7) ± .013 3.01 (2.8-3.6) ± .003 

Wildrose Ranger Station. Pane/mint Mts.. 4200 ft.. 

Inyo Co., California* 
2 5.19 (4.5-5.7) ± .012 2.39 (2.1-2.6) ± .020 

Lathrop Wells. 2900 ft., Nye Co., Nevada* 
2 6.12 (5.5-6.7) ± .014 2.74 (2.5-3.1) ± .002 

* No males found in sample. 



Calamujue. northwest to El Rosario. eastward around 
the south end of the Sierra San Pedro Martir. thence 
north along the eastern slopes of this mountain range 
(below 2000 ft) over San Matias Pass, south to La 
Parra on the northwestern slope of the Sierra San 
Pedro Martir, north to Valle de la Trinidad to San 
Matias Pass northward along the eastern flanks of 
the Sierra de Juarez into California, throughout the 
Colorado Desert parts of Southern California to 
Banning and Morongo Valley, around and along 
the north flank of the San Bernardino and San 
Gabriel Mountains north to Mojave and throughout 
the Owens Valley wherever Larrea occurs (Fig. 32). 
A total of 4286 specimens were examined. 

Remarks: Eucyllus vagans is the most plastic 
member of the genus and may represent a com- 
plex of species. Morphological variation presented 
several problems in this respect. Isolated pockets 
of this species appeared, superficially, to be 
distinct. Certain of these populations (for ex- 
ample, those from Valle de la Trinidad, Baja 
California del Norte, and Wildrose Ranger Station 
in the Panamint Mountains, Inyo County, Cali- 
fornia) are so different in scalation as to be easily 
distinguished from the remainder of the specimens 
examined. These could be separable as subspecies. 



However, the authors do not feel it wise to name 
subspecies in parthenogenetic species. No sig- 
nificant differences were found in the sper- 
mathecae, although body proportions varied in the 
different populations. Table 1 indicates the range 
in lengths and widths and their respective means. 

As can be seen from the means in table 1 , 
specimens from Wildrose Ranger Station are rela- 
tively smaller and broader than the other samples. 
The means of leg lengths, antcnnal proportions, 
and other body proportions follow a similar pat- 
tern. The sample from Valle de la Trinidad was 
not large enough to analyze (only 15 specimens). 

In vagans there is a relationship between setal 
length and age. Newly emerged specimens have 
rather elongate pointed setae (not approaching 
length of saesariatus) . The tips of these setae 
break or wear off shortly after emergence in most 
individuals resulting in the broader, blunter scales 
normally considered typical of vagans. 

The selection of Carrizo Spring as a type lo- 
cality was based on two factors. First, other 
species of snout beetles collected and described 
at about the same time have this as a type locality 
(= Carisa Creek). These species have been re- 
cently collected with E. vagans at this location. 
This location was a stage and rest stop where the 
travelling collectors, of that time, would have 
felt safe to wander around collecting specimens. 
Second, material taken there seemingly represents 
the typical species (as currently recognized) and 
this population has a sex ratio of males to females 
of about 1:1. 

Biology: Eucyllus vagans has been encountered 
under or feeding upon a larger variety of plant 
species than any of the other species. These 
include Larrea tridentata. Franseria dumosa, 
Hymenoclea salsola. H. monogyra, and A triplex 
spp. The species also has been taken in pitfall 
traps and walking over blacklite and white light 
sheets. 

Newly emerged forms appear on vegetation in 
May or June depending upon the latitude. The 
newly emerged individuals feed briefly, then most 
become dormant until October. Occasional 
stragglers are taken year around on the host 
plants, particularly Hymenoclea and Larrea. 

Eucyllus saesariatus, new species 
Figures 5, 16, 20, 29, 31 

Holotype: Female, Mexico, Sonora, 51 mi E San 
Luis, XII-30-60, ELS, in ELSC #80. Allotype: Male, 
same data as holotype. Length 6.3 mm; width 2.7 mm. 



1972 



REVIEW OF THE snout BEETLES, GENUS EU( YLU 






120 



115 



110 



35 



30 



\ 






^"^ 


1 


i 


\ 






i 




i 

i 


\ 






i 




■ 


\ 


\ 




/ 




i 




4^r 


ill 


rfl@fli 










w^Mm^®^ — " 






"^C^V- 










i 

i 






Wm 


f A % 

3 ■ ' 






• carinarostris 
. O cinereus 




) r~ s 


@-V a "a 












■ echinus 
A saesariatus 






( 100 

L, 11 I 1 




200 


▲ unicolor 






&\ p. V Scale of mil 


es 




j^vagans 




\ 







30 



no 



Figure 32. Distribution of Eucyllus species in the United States and Mexico. Symbol enclosed 
in a circle indicates the type locality for the species. 



Differing little from the holotype. except for con- 
cavity in sternites 1 and 2. Aedeagus similar to 
vagans (Fig. 20). 

Description: Length 7.1 mm; width 3.4 mm; 
elongate oval; vestiture dirty gray with silver to 
brownish patches on the head, prothorax. and ab- 
domen; seta-like scales erect, ten to fifteen times 
longer than broad with some four to seven times as 
long as broad, all setae with acute apices: in cross 
section, setae with a feebly arcuate concavity. Rostrum 
slightly longer than broad; posterior to apex of 



epistoma a feeble indication of a median carina with 
a teardrop-shaped fovea on each side of the carina 
forming a "V". distad to the carina a deep oval pit. 
this evident only when denuded: rostrum strongly 
sculptured when denuded. Funiculus with firs 
ment as long as second, second segment almost twice 
as long as third. Prothorax dirty white will 
longitudinal stripes of silver scales: dia 
irregular punctures with a mean of 13.5 per 
Elytra dirty white, with irregular patchc- 
brown scales. Femora with a bar 



\,s- 



m ill li\ sol nil i;\ < IL1FORN1 i « ADl M) OF SCIENCES 



VOLUME 7/ 



neaj distal end; apical comb with short thick teeth 
anteriorl) gradually becoming longer and thinner 
toward posterior margin. Spermatheca as shown in 
figure 5. 

Paralypes: 2 . 109 same data as holotype, 9(5, 
209 52 mi I San I uis, XH-27-60, LO '. 209. IX-30- 
68, I I S. Size of paratypes: length, 5.6-8.0 mm 
(moan 6.45; 9 7.13 mm); width 2.5-3.6 mm 

(mean 2. 84: 9 3.31 mm). Paratypes deposited 
in the following collections: CASC. CIAN. CSLB, 
CWOB, IINC. FWPC, 1NIA. LACM, and USNM. 

Other specimens examined: MEXICO. Sonora, 
72 mi E San Luis. VIII-22-60 (T. Schowalter); 57.9 
mi W Sonoyta, IV-14-62: Sa. Pinacate, 2 mi S Mexico 
#2. 1-26-62. CA1 IFORNIA. Imperial Co.. Imperial 
Dam. V-3-58. ARIZONA. Pima Co.. (the follow- 
ing locations in Organ Pipe Cactus National Monu- 
ment) Alamo Wash Yi mi E Hwy. #85. 11-23-62. 
V-26-62: Bonito Well, 11-12-61: Dos Lomitas XII- 
29-60: 15.8 mi N Headquarters, X-29-62: Quito- 
baquito, XII-28-60. X-28-61. 1-24-62. VIII-5-62; Ajo 
Mlns.. X-16-35, Bryant. CASC. Maricopa Co.. 
Picacho. V-l-42 and VI-28-41. Bryant. CASC. 
'.'UTAH. Washington Co.. St. George, Wickham, 
CASC. (probably a labeling error). A total of 432 
specimens examined. Unless noted otherwise, material 
was collected by ELS and is in ELSC. 

Remarks: This species is very similar to E. 
vagans; however, it can easily be differentiated 
by the length of the seta-like scales on the dorsum, 
morphology of the spermatheca. and the feeble 
indication of a median carina on the rostrum. 
Greater morphological continuity of specimens 
from different locations (Fig. 32) is seen in this 
species than in vagans. This may be due to the 
existence of males in all the locations recorded. 

Biology: All of the material was collected from 
Larrea tridentata at night. The adults feeding on 
vegetation are most numerous in late December 
and early January. A few stragglers are present 
year around. 



Eitcyllus echinus Van Dyke 
Figures 8, 9, 12, 13, 21, 23, 25 

Eucyllus echinus Van Dyke 1936, 31. Type locality — 
"Cave Creek, Maricopa Co., Arizona." (CASC 
4144 Ent.) a female. Holotype examined. 

Description: Body length 4.0-5.3 mm (mean 4.9): 
width 1.8-2.2 mm (mean 2.1); elongate oval; vestiture 
brown to dirty gray; derm black, with irregular 
patches of brownish silver scales; seta-like scales twice 
as long as broad, truncate at apices or slightly rounded, 
appearing club-like and erect. Rostrum having a 
feeble indication of a median carina extending from 
epistoma to posterior margin of head; strongly sculp- 



tured when denuded. Funiculus with first, second, and 
third segments nearly same length, slightly longer 
than scape. Head feebly sculptured with a few rather 
small, oval punctures, which arc most evident when 
denuded. Prothorax with irregular dark patches; disc 
with large deep punctures with a mean of 24.3 per 
.25 mm 3 . Elytral vestiture brownish in color with 
irregular dark patches. Femora uniformly brown 
in color without distinguishing markings or spurs; 
apical comb of tibia 3 with rather short thick teeth 
from anterior to posterior all about the same length. 
Aedeagus rather wide with sides almost parallel, 
tapering to a rather abrupt, blunt apex, not curved or 
twisted, however, slightly arcuate dorsoventrally (Fig. 
21); spermatheca as in figures 8-9. 

Distribution (Fig. 32): MEXICO. Sonora, 52 mi 
E San Luis, XII-27-60; 51 mi E San Luis, XII-30-60. 
ARIZONA. Pima Co., (the following locations in 
Organ Pipe Cactus National Monument) Alamo 
Wash y 2 mi E Hwy. #85, 11-23-62; Dos Lomitas, 
XII-29-60; Headquarters. IX-29-61; 14 mi N Head- 
quarters, 11-24-62; 15 mi N Headquarters, X-29-61; 
Quitobaquito, XII-28-60, X-28-62. Pinal Co., 3 mi 
NE Apache Junction, XII-4-62. Maricopa Co., Gila 
Bend, Wickham coll., Picacho, V-l-42, Bryant, and the 
type locality. UTAH. Washington Co., 15 mi SW 
Shivwits, VIII-13-62, Ross Hardy coll.; St. George, 
A. M. Woodbury. NEVADA. Lincoln Co., 10 mi 
N US #91 on Elgin Road in the Mormon Mountains, 
V-18-62, Ross Hardy coll.; Nye Co., 19 mi SE 
Lathrop Wells, 111-24-70, and 2 mi SW Mercury, 
V-25-71. Unless otherwise noted, material was col- 
lected by ELS and is in ELSC. Material will be 
distributed to appropriate institutions. One hundred 
four specimens were examined. 

Intensive collecting has not turned up specimens of 
echinus between Cave Creek, Maricopa County, 
Arizona and the Washington-Lincoln Counties area of 
Utah-Nevada, a distance of 240 airline miles. 

Remarks: This species is closely related to 
unicolor but can be distinguished by its dorsal 
seta-like scales and a feeble median carina on the 
rostrum. The spermatheca is very distinct and 
shows only a slight variability in form throughout 
the specimens examined. Parthenogenicity seems 
to exist in the population in Nevada and Utah 
as no males have been taken there. 

Biology: The specimens from Quitobaquito and 
15 mi N Headquarters (Organ Pipe Cactus Na- 
tional Monument) were taken on Franseria dum- 
osa and Franseria sp. The material from Nye 
County, Nevada was beaten from an undetermined 
species of Franseria and Larrea divaricata. All 
other material collected by ELS and that collected 
by Ross Hardy were from Larrea tridentata. In 
the Arizona-Sonoran portion of the range, the 
adult population is present in largest numbers on 



1972 



REVIEW (>!■ THE SN0U7 BEETLES Gl ' EU( YLLl 






the vegetation in December at which time the 
females were found to be carrying an average of 
lour mature ova. 

Eucyllus cinereus, new species 
Figures 10, 10a, IS. 19, 26 

Holotype: Female, California, Inyo County, While 
Mountains, Grandview Public Camp. 8400 ft, VIII 
13-64, ELS, ELSC #82. Allotype: same data as 
hololype. Length 3.8 mm. width 1.7 mm; differing 
in narrow form and with scattered brown scales on 
dorsum. 

Description: Length 4.4 mm: width 2.2 mm: 
elongate oval; derm mahogany brown; scales and 
seta-like scales uniformly white giving the body a 
gray color, no irregular dark patches on dorsum; 
setae three times as long as broad, erect, truncate at 
apices, appearing club-like. Rostrum as long as 
broad, shorter than in other species; feebly sculp- 
tured; a feeble indication of a median sulcus extending 
posteriorly from the epistoma to the anterior margin 
of the head; punctures on the rostrum moderate 
in size, oval in shape, rather shallow compared to the 
other species; scrobes with dorsal and lateral margins 
reaching orbital of eye channel of scrobe with a 
feeble median carina converging with orbital of eye. 
this evident when denuded. Funiculus with second 
segment one and one-half times longer than third. 
Prothorax with disc feebly flattened with large deep 
punctures separated by their diameter and numbering 
43.2 per .25 mm'-'; elytra uniformly gray in color; 
length of second abdominal sternite not quite as long 
as three plus four combined, and shape slightly dif- 
ferent than occurs in other species; tibia has apical 
comb with anterior teeth short and thick gradually 
becoming longer posteriorly until posterior teeth are 
one-half again as long as anterior teeth while main- 
taining same thickness. Spermatheca as in figures 10 
and 10a. 

Paratypes: All 21 are from California. Inyo Co., 
White Mountains, 9 5 same data as holotype (4 
ELSC, 2 FWPC. 1 CASC. 1 LACM. 1 CSLB), 9 9 
from type locality. VIII- 10-65 (4 ELSC. 1 CASC. 1 
LACM, 2 USNM, 1 BYUC); 1 9 type locality. VIII- 
15-65, M. L. West coll.; 12. 7.0 mi S Schulman 
Grove, 9000 ft. VI-25-63. M. L. West coll.: 12. 3.0 
mi N Westgard Pass, 7300 ft, VIII- 15-64. Length 
3.7-4.6 mm (mean 4.0 mm): width 1.5-1.9 mm 
(mean 1.62 mm). A total of 23 specimens were 
examined. Unless otherwise noted, the material was 
taken by ELS and is in ELSC. A few of the paratypes 
have faint indications of mottling with brown. 

Distribution: The range of this species is at present 
limited to the White Mountains, Inyo County. Cali- 
fornia (Fig. 32). It would seem probable that it will 
be encountered in the drier mountain ranges to the 
east in Nevada, such as the Silver Peak Mountains. 

Remarks: This species is generally smaller than 



the othei spi cii ithin 

i inereus can be easily scparati d from 

species by size, coloration, shortci ro 

like scales, the spermatheca (Figs. 10 10a), and 
the aedeagus. Other minoi - harai aid in 

defining this species. These are the 
length of the second abdominal sternite, an 

length of the pOSterioi teeth on the apical 
of the tibia 3. 

Biology: The specimen-, from the type locality 
were taken on Artemesia tridenlata and Purshia 
tridentata, The other specimens were taken dur- 
ing sweeping of various vegetation types. v - 
ture ova were encountered in the material 
from the period of late June through mid August. 
The habitat of this weevil is the high I'inyon- 
Juniper region. The apparent altitudinal range 
of this species is from 7300-9000 ft: the highest 
of any in the genus. 

Eucyllus carinarostris, new species 
Figures 6. 14. 22 

Holotype: Female. California. Riverside Co., Pinto 
Wash Well, 11-17-62. 1000 ft. ELS. in I 1 S< 

Description: Length 4.6 mm: width Z.O mm: 
elongate oval; derm black, scales dirty white giving 
body a brownish color; seta-like scales twice as long 
as broad, recurved, peg-like, and brown in color. 
Rostrum with strong indication of a median carina 
extending from epistoma to anterior margin of head: 
this evident when denuded: a rather deep fova on 
each side of the carina with large shallow punctures; 
rostrum rather strongly sculptured. Funiculus with 
second segment longer than third, but not twice the 
length of the third. Disc of prothorax feebly convex 
with round deep punctures with a mean of 29.7 per 
.25 mm" spaced approximately their diameter apart. 
Elytra with a few irregular dark patches, however. 
mostly uniformly straw colored. Femora with no 
distinctive markings: tibiae having the apical comb 
with anterior teeth short and thick, gradually be- 
coming longer towards the posterior margin and 
closer together while maintaining same thickness. 
Spermatheca as in figure 6. 

Paratypes: All five from California. Riverside Co.. 
1 2 . same data as holotype: 1 2 . Joshua Tree 
National Monument. 7 mi NW Old Dale Junction. 
1-28-61; 3 2 . 5.4 mi NW Old Dale Junction. Ill 
All material taken by ELS. Length 4.1-5.0 
(mean 4.4 mm); width 2.0-2.4 mm (mean 2.14 mm). 
Distribution of paratypes 2 ELSC. 
LACM. 1 USNM. Only six specimens were examined. 

Distribution: Apparently limited to the 
Basin and the old Pinto Lake drainage 
Riverside County. California (Fig. 3 



90 



in I 1 II IX SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



Remarks: This species closelj resembles E. 
unicoloi bul can be readily distinguished bj the 
median rostral carina and iho shape of the 
spermatheca. 

Biology: All material was taken on Larrea 
tridentata. Northwest of Old Dale Junction it oc- 
curred with E. vagans. At Pinto Wash Well it was 
taken on the same plants with E. vagans and 
\4iloderes sp. 

Eucyllus unicolor Van Dyke 
Figures 7. 11. 24, 27 

Eucyllus unicolor Van Dyke. 1936. 32. Type lo- 
cality— "Utah in 1921." (CASC 4145 Ent.). Holo- 
type examined. 

Description: Length 4.5-5.0 mm (mean 4.65 mm); 
width 2.0-2.4 mm (mean 2.09 mm); derm brown 
to black; scales white, seta-like scales dirty white, no 
distinct color markings on dorsum; seta-like scales 
generally twice as long as broad, recurved, rounded 
at apices. Rostrum with feeble indication of a 
median fova approximately midway up the rostrum 
from the epistoma. no indication of a median carina; 
feebly sculptured, this evident when denuded: punc- 
tures on rostrum larger than those on head, hut not 
as large as those found on related species. First seg- 
ment of funiculus approximately same length as 
second, third segment slightly shorter than preceding 
segments. Punctures on prothorax round and rather 
deep with a mean of 32.4 per .25 mm-; prothorax 
without distinct marking or patches; the sides not as 
rounded as in most species. Elytra with no distin- 
guishing marks, uniformly straw color; femora with 
no distinguishing marks, tibia with apical comb of an- 
terior teeth short, gradually becoming longer pos- 
teriorly while maintaining same thickness, posterior 
teeth not more than one-half again as long as anterior 
teeth. Spermatheca as in figure 7. 

Other type material: One paratype from Westgard 
Pass, Inyo Co., California, not examined. 

Distribution: UTAH. Washington Co., St. George 
(and vicinity), no other data; Dixie State Park, VII- 
3-63. NEVADA. Nye Co.. 19 mi SE Lathrop Wells, 
111-24-70. Tanner (1959) reports this species from 
"St. George, Washington Co.. Utah, A. M. Wood- 
bury; Virgin River. Washington Co., Utah, C. J. 
Weidt, 1892; SW end Cedar Mountains, Toole Co., 
Utah, W. J. Thomas, VIII-25-53; Peach Springs, Ariz., 
Ulke. 1896; 'Ariz.'" and again Tanner (1966) from 
the Nevada Test Site "Area 28 (vicinity of Rock 
Valley) and . . . Cane Springs." Both of these loca- 
tions are in Nye County. It is our opinion that this 
species is confined to western Utah and south central 
Nevada (Fig. 32). This is based on extensive collect- 
ing in southwestern United States and the absence of 
recently collected, correctly labeled material from 
other than the locations cited. Numerous snout beetle 



specimens have been encountered bearing the label 
"Peach Springs, Ariz.. Ulke" that were apparently mis- 
labeled. Several of these species have been searched 
for, in vain, in the Peach Springs area. They have been 
encountered in southwestern Utah and southeastern 
Nevada. The "Ariz." labeled material gives no real 
information. This leaves the paratype from "West- 
gard Pass." This paratype has not been located but 
it is probably E. cinereus which we have previously 
mentioned resembles unicolor. Nineteen specimens 
were examined. 

Discussion: Eucyllus unicolor resembles car- 
inarostris and echinus, but can be readily separated 
by the length and shape of the setae and the lack of 
a median carina on the rostrum. The spermatheca 
is very different from all of the species of the 
genus (Fig. 7). 

Biology: The material examined from St. 
George was captured on Larrea tridentata. Most 
of the specimens from 19 mi SE Lathrop Wells 
were on Franseria sp. 

ACKNOWLEDGMENTS 

We wish to thank J. F. Lawrence, Museum of Com- 
parative Zoology. Harvard University, for informa- 
tion on the lectotype of E. vagans Horn, and aid in 
studying the LeConte Collection; Hugh B. Leech, 
California Academy of Sciences, for use of Academy 
facilities while working on that collection and for 
lending material; Ross Hardy, Alan Hardy, Eric M. 
Fisher, John A. Gruwell. and Sandra L. Jenkins for 
additional material. 

Special thanks are extended to the Superintendents 
of Joshua Tree National Monument and Organ Pipe 
Cactus National Monument and their Park Nat- 
uralists and Rangers who significantly aided the 
efforts of the collectors. 



LITERATURE CITED 

Arnett, R. H., Jr., el al. 1969. Directory of Coleop- 
tera Collections of North America. Center for 
the Study of Coleoptera, Lafayette, Indiana, vii -f- 
123 pp. 

Hatch, M. H. 1971. The beetles of the Pacific 
Northwest. Part V: Rhipiceroidea, Sternoxi, 
Phytophaga, Rhynchophora, and Lamellicornia. 
Univ. Washington Press, Seattle, xiii 4- 662. 

Horn. G. H. 1876. In LeConte and Horn. The 
Rhynchophora of America north of Mexico. 
Proc. Amer. Philos. Soc, 15:1-455. 

Kissinger, D. G. 1964. Curculionidae of America 



1972 



REVIEW OF Till. SNoiri mi ills. GENUS EUt I / I ' 






north ol Mexico. Taxonomic Publications, South 
Lancaster, Massachusetts. 143 pp. 



rada iiM Site Brigham Voun 
Biol. Sen, 8: 1 



Tanner, V. M. 1959. Studies in the weevils of the v '' ,n Dyke, I. C. 1936. New ipecica ol ' 



western United States No. IX: Description of a 
new species of Eucyllus (Colcoptera: Curculioni 
dac). Great Basin Nat., 19:53-55. 



American weevils in the family ' urculionidsu 
subfamily Brachj rhininac IV. Pan P I 

12:19-32. 



1966. Rhynchophora Beetles of t he Ne- Accepted for publication August 31, 1971. 



TWO NEW SPECIES OF NORTH AMERICAN FLAT BUGS 
(HEMIPTERA: ARADIDAE) 

Nicholas A. Kormilev 1 



Abstract: Two new species of Aradidae are described. 



Among unidentified material in the entomological 
collections of the Natural History Museum of Los 
Angeles County (LACM). 1 discovered specimens 
of the following two undescribed species of 
Aradidae. 



SUBFAMILY ARADINAE 
Genus Aradus Fabricius, 1803 

Aradus nevadensis, new species 
Figures 1-2 

Description: Male. Elongate ovate: head, pronotum, 
and scutellum finely granulate. Head longer than its 
width across eyes (1.27:1): anterior process strong. 
with parallel sides, reaching basal % of antennal seg- 
ment II; antenniferous tubercles acute, diverging, 
reaching apical Va of antennal segment I; lateral tooth 
small, but distinct: preocular tubercles acute, postocu- 
lar also acute. Eyes reniform, strongly protruding. 
Depressions of vertex deep, converging backward in 
an arc. Antennae slender, almost twice as long as 
width of head across eyes (1.93:1); antennal segment 
II narrower than fore femora, gradually dilating 
toward the tip; relative length of antennal segments 
I to IV: 1:4.28:1.86:1.14. Labium reaching meso- 
sternum. Pronotum less than half as long as its 
maximum width across middle of lateral borders 
(1:2.22): the latter strongly convex, rounded: straight 
and converging anteriorly, and bearing a few small 



teeth. Disc raised before and behind deep, transverse 
depression; inner carinae parallel, very slightly di- 
verging backward. Scutellum triangular, longer than 
its basal width (1.23:1): lateral borders slightly con- 
vex at base, then straight; raised. Tip narrowl) 
rounded. Disc raised at basal -.-,. transversely rugose 
on apical %. Hemelytra reaching apical ' :I of genital 
lobes; corium expanded and rounded laterally at base. 
reaching 1 i> of connexivum V. Abdomen ovate: pos- 
teroexterior angles of connexiva II to IV not pro- 
truding. V and VI slightly protruding. VII forming 
rounded lobes; inner border of genital lobes slightly 
diverging behind middle. Sternum VI longer than 
VII medially (1.2:1). Color black to piceous on head, 
pronotum and scutellum. with exception of latero- 
posterior borders of pronotum and tip of scutellum. 
which are white or whitish. Antennal segment'- I. II. 
and basal % of III. dark brown: apical x 3 of III 
whitish. IV black. Corium of hemelytra ochraceous 
mottled with whitish, infuscate on apical -.-,: membrane 
fuscus. whitish at base. Connexivum dark brown with 
whitish posteroexterior angles and hind borders, the 
latter with a slight, reddish tinge in middle. Ventral 
side of body reddish brown, with whitish postero- 
exterior angles of connexiva. Legs dark brown: coxae 
and tips of tibiae, whitish. Size — total length 



1 Natural History Museum of Los Angeles Co 
Los Angeles. California. 90007 
102-34 93rd Avenue. Richmond Hill. N 
11418). 



«: 



/.'/ /// //.\ SOI I III l<\ ( till OHM. I l( I/)/ W> Ol \(ll.i\( I \ 



VOLUME 71 




Figures 1-5. Fig. 1, Aradus nevadeiisis. n.sp., S, head and pronotum. Fig. 2, tip of abdomen 
from below. Fig. 3, Mezira tropicalis, n.sp., <$ . head and pronotum. Fig. 4. tip of abdomen 
from above; 9 . Fig. 5, tip of abdomen from above. 



mm; width of pronotum 2.50 mm; width of abdomen 
3.25 mm. 

Diagnosis: Aradus nevadensis, n.sp. is similar to A. 
cincticornis Bergroth (Canadian Ent., 38:198-202. 
1906). but it is larger, the head is longer than its width 
across eyes, antennal segment II is as long as the 
width of head including both eyes, sternum VI (V 
visible) is longer than VII (VI visible), and the color 
is different. 

Holotypc: Male. California. Nevada Co.. Sagehen 
Creek. 18 June 1962 (R. L. Westcott) LACM. 

SUBFAMILY MEZIRINAE 
Genus Mezira Amyot and Serville, 1843 

Mezira tropicalis, new species 
Figures 3-5 

Description: Male (Female identical except for dif- 
ferences where indicated). Elongate ovate, covered 



with a fine, setigerous granulation, with setae ex- 
tremely short and erect. Head slightly shorter than 
its width across eyes (male 1:1.07. female 1:1.15); 
anterior process slender, slightly constricted in middle 
and incised anteriorly, reaching apical % of antennal 
segment I; antenniferous tubercles acute, diverging, 
reaching basal Vs of antennal segment I; postocular 
small, acute, reaching outer border of eyes: the latter 
semiglobose. protruding. Vertex with V-form rows 
of setigerous granules. Antennae slender, less than 
twice as long as width of head across eyes (male 
1:79:1, female 1.72:1); relative length of antennal 
segment I to IV: male 1:0.8:1.35:0.7, female 1:0.8: 
1.4:0.75. Labium reaching hind border of labial 
groove, the latter closed posteriorly. Pronotum 
trapezoidal, shorter than its maximum width (male 
1:1.92, female 1:2.12); fore lobe is narrower than hind 
lobe (male 1 : 1.35, female 1 : 1.42); collar thin, straight 
anteriorly; anterolateral angles rounded, slightly ex- 
panded and reflexed, produced forward as far as collar; 



I<>72 



TWO NEW FLAT BUGS I ROM NoHl II I Ml ri< \ 






lateral border parallel ;ii humeri, converging and barely 
sinuate anteriorly; hind holder almost straight, Fore 
disc with 4(2-|-2) high, granulate ridges; hind disc 
granulate; interlobal depression deep, Scutellum 
shorter than its h;is;il width (male 1:1.30, female 
1:1.42); lateral borders thinly carinate and slightly 
sinuate on apical half; lip rounded; disc with a thin 
median carina, areas lalerad of latter granulate. 
Hemelytra reaching beyond fore bolder of tergum VII 
( i? ), or reaching hind border of tergum VI (?); 
apical angle of corium blunt, apical border convex, 
rounded. Abdomen ovale, longer than its maximum 
width across segment IV (male 1.33:1, female 
1.41:1); connexivum wide and slightly raised laterally; 
postcroexterior angles of the connexiva II lo VI 
slightly protruding, blunt; those of VII produced 
backward as rounded lobes, reaching 'j of paratergites 
(c5), or rounded, reaching '/■; of tergum IX (9). 
Paratergites ( $ ) thin, clavate, reaching apical 1 /i of 
hypopygium; the latter cordate, with a thin median 
ridge, slightly shorter than disc of hypopygium. 
Paratergites ( 9 ) large, rounded, reaching apical Vi 
of slightly tricuspidate segment IX. Spiracles II to VII 
ventral, placed far from margin, VIII also ventral, but 
placed closer to margin and not visible from above. 
Legs unarmed. Color dark ferrugineous. connexivum 



and viniii fcrrugincoui 

7.7H, female K.f)7 mm; width ol i lotum: mail . 

female 2.K3 mm; width ol al di i 
male 3.56 mm 

Diagnosis: Mezlra tropicalls, n. sp i 
to A/, mexicana Kormilcv (Proi ' nilcd State 
Mms„ 1 19:245 258, 1964) from Vera ' ruz M 
but is larger, with the anterioi process ol 
longer and more slender; antennal segment III 
lively longer, almost twice as long as IV; and paia 
tergites ( ? ) longer, reaching to the apical 
segment IX. 

Holotype: Male. Mexico. Jalisco. M mi W 

Atenquique, 7800 ft, 13 July 1966 (J, R. Dixon and 

W. K. Meyer) I A( M. Allotype: Female, I female 
paratype and 5 nymphs, same data as holotype. 



ACKNOWl FIX, Ml \| 

For the privilege of studying the Aradidac in the 
collections of the Natural History Museum of I OS 

Angeles County. I express my sincere thanks to 
Charles L. Hogue. Senior Curator of Entomology. 

Accepted for publication November 1(S. 1971. 



A NEW SPECIES OF AMBUSH BUG FROM ARIZONA 
(HEMIPTERA: PHYMATIDAE) 

Nicholas A. Kormilev 1 



Abstract; A new species of macrocephaline ambush bug from Arizona is described. 



The ambush bugs are represented in the con- 
tinental United States by two subfamilies, Phy- 
matinae and Macrocephalinae. The first is com- 
mon throughout the country, but the second is 
rather rare, being distributed mainly in the south 
and southwest, although Macrocephalus pre- 
hensilis (Fabricius), 1803, has been recorded as 
far north as Kentucky and Kansas (Evans, Ann. 
Ent. Soc. America, 24:711-736, 1931). While 
examining specimens of the latter subfamily in 
the collection of the Natural History Museum of 
Los Angeles County (LACM) and in a lot sent 
to me by T. Halstead, I found a few specimens of 



an undescribed Macrocephalus, collected in 
Arizona. 

In the description all measurements are given in 
millimeters. The first figure in a ratio represents 
the length and the second the width of the mea- 
sured part. The length of the abdomen was mea- 
sured from the anteroexterior angles of con- 
nexivum II to the tip of abdomen. 



1 Natural History Museum of Los Angeles C 
Los Angeles. California. 
102-34 93rd Avenue. Richmond Hi!:. !• 
L1418). 



94 



HI I 1 1 ll\ sol THERN ( II IIORNIA ACADEMY OF SCIENCES 



VOLUME 71 



SUBFAMILY MACROCI -THAI INAE 
Genus Macrocephalus Swederus, 1787 

Macrocephalus similis, new species 
Figure I 

Description: Female. Elongate ovate; bead, fore lobe 
of pronotum and corium, finely granulate; connexiva 

II roughly granulate. Head almost twice as long as 
its width across eyes (1.60:0.92); anteocular portion 
narrower than postocular (0.64:0.80): ocelli placed 
nearer hind border of head than eyes (0.24:0.28); 
upper lobe of genae rounded, lower angular; bucculae 
evenly rounded and granulate on border; lateral 
borders of labial groove also granulate. Antennae 
almost as long as head (1.56:1.60); antennal segment 
I subtriangular, flattened laterally: II almost globose, 

III evenly enlarged toward tip. IV fusiform; relative 
length and width of antennal segments I to IV: 
0.40:0.32 — 0.24:0.20 — 0.28:0.16 -- 0.64:0.36. 
Labium reaching middle of prosternal cavity. 
Pronotum shorter than its maximum width across 
humeri (2.28:3.04): fore lobe narrower than hind 
lobe (1.52:3.04). Anterior angles dentiform, acute, 
directed forward; anterior border deeply sinuate; 
anterolateral-anterior borders slightly convex, diverg- 
ing backward: interlobal notch sinuate; anterolateral- 
posterior borders convex, rounded; lateral angles 
incised as in M. cimicoides Swederus. 1787; postero- 
lateral borders firstly convex, then sinuate: hind 
border convex in middle. Fore disc convex, finely 
granulate, more densely granulate along lateral bor- 
ders: pronotal pit elongate. Hind disc roughly punc- 
tured, punctures forming a net-like surface and are 
larger anteriorly and laterally. Pronotal carinae short, 
stout anteriorly, diverging, tapering and evanescent 
posteriorly, without any knob or ridge anteriorly. 
ScuteUum twice as long as its maximum width 
(110:55), with a thin median carina, which is en- 
larged on basal 1/7. Disc roughly punctured at base 
laterally, finely punctured elsewhere on disc, the basal 
portion moderately inflated and with a weak, ivory- 
like elevation in the form of a 3-pronged spear-head 
on basal half medially, more distinct in the males. 
Hemelytra reaching tip of abdomen; corium finely 
granulate, granules arranged in groups. Abdomen 
cordate, slightly longer than its maximum width 
across segment III (4.12:4.00); posteroexterior 
angles of connexiva barely protruding; connexiva II 
covered with a rough and dense granulation; other 
connexiva sparsely and finely granulate along ex- 
terior border. Legs with fore coxae cylindrical, with- 
out knob or tooth, but roughly granulate inferiorly. 
Fore femora longer than wide (2.16:0.88), convex 
exteriorly and with a row of rough, setigerous granules 
on upper side. Fore tibiae without tarsi. Middle and 
hind femora with rough, setigerous granules on upper 
and lower sides, with smaller granules laterally; 
middle and hind tibiae densely granulate. Color 




Figure 1. Macrocephalus similis, n.sp., holotype (fe- 
male), dorsal aspect; A — paratype (male), right 
paramere. 



grayish brown; head on upper side medially, fore lobe 
of pronotum in middle, and exterior borders of con- 
nexiva III and IV, black or blackish; rest of fore lobe 
of pronotum and connexiva III and IV, reddish 
brown; antennae, pronotal carinae. scutellar carina 



1972 



NEW AM HUSH HI Id I ROM AKI/o \ 






and inflated portion of disc, and connexiva II. 
ochraceous or slightly brownish: anterolateral portions 
of scutellum (roughly punctured) brown; ventral side 
of hotly ochraceous, brownish on fore femora, coxae, 
middle and hind femora, and venter laterally and 
posteriorly; middle tarsi black, hind tarsi greenish- 
yellow. Size — total length 8.16 mm; width ol 
pronotum 3.00 mm; width of abdomen 4.00 mm. 

Male. Smaller anil more slender than female, more 
densely granulate and with color generally darker; 
upper surface of head black; antennae, pronotum 
and scutellum, with exception of a three-pronged, 
spear-shaped ivory spot, black mottled somewhat with 
yellow; connexiva 111, IV and most of V, black; 
ventral side of the body yellow with some brown 
stripes on fore femora and venter laterally; reddish 
brown near lateral angles of venter. One male is 
colored as the females. 

Measurements: Head 1.60:0.90; relative length 
and width of antennal segments 1 to IV: 0.48:0.22 - 
0.22:0.20 — 0.32:0.20 — 0.80:0.32; pronotum 2.20: 
3.08, ratio width of fore lobe: width of hind lobe 
as 1.60:3.08; scutellum 4.20:2.20; abdomen 4.16:3.60 
(across segment III); fore femora 2.0:0.9. Paramere 
is of Macrocephalus type. 

Size: Total length 7.92 mm; width of pronotum 
3.08 mm; width of abdomen 3.60 mm. 

Diagnosis: Macrocephalus similis, n.sp. is very 
similar to M. barberi Evans. 1931, from California, 



and probably was confu cd latti i 
he separated from it by: largi i i otal ci 

without knob oi ridge anterior!) granulation 
pronotum and connexivum generally 

dense, head relative!', hortCI pronol relative!) 

longer, and abdomen slightly longct than il ma imum 

width 

Holotype: Female. Arizona Cocl ' mi w 

Portal. 25 June 1959 (I.. A. StangC i I V M lllol pt 
Male. Arizona. Santa ' ruz Co., Pena Blanca Laki 

22 May 1970 (I. H.iKlead) I A( \t 

Paratypes: 5 ' and 7 7. some were collected with 
the allotype, and others in the same locality on 15 
November 1969. I. Ilalslcad coll. Deposited in the 
Natural History Museum of l.os Angeles fount', 
and in the collections of Department ol I ntomi 
University of Arizona. I ucson, and of the author. 



ACKNOWLEDGMENTS 

For the privilege of studying Phymatidae in the col- 
lection of the Natural History Museum of Los Angeles 
County. I express my sincere gratitude to Charles I 
Hogue. Senior Curator of Entomology. I am also 
indebted to T. Halstead. Phoenix. Arizona, for the 
gift of Phymatidae collected by him in Arizona. 

Accepted for publication November 16. 1971. 



THE ASCTDIANS STYELA BARNHARTI, S. PLICATA, S. CLAVA. AND 
5. MONTEREYENSIS IN CALIFORNIAN WATERS 

Donald P. Abbott 1 and Jeffrey V. Johnson- 



Abstract: Reexamination of the holotype of Styela bamharti Ritter and Forsyth. 191". 
shows that it is Styela plicata (Lesueur. 1823). The species designated Styela barnharti by 
Van Name (1945) and subsequent authors is Styela clava Herdman. 1881. an asiatic form 
probably introduced into Californian waters in the late 1920"s. The latter species is distinct 
from the native west coast form Styela montereyensis (Dall. 1872). 



The stalked simple ascidian usually referred to as 
Styela barnharti is common on floats and pilings 
in protected coastal waters of southern California 
(Ricketts and Calvin, 1968; MacGinitie and Mac- 
Ginitie, 196S). In some harbor areas it is found 
with another stalked form, Styela montereyensis: 
the two are not always easily distinguished, es- 
pecially when overgrown by hydroid and bryozoan 
colonies. 



Van Name (1945) expressed the opinion that 
the original description of 5. barnharti by Ritter 
and Forsyth (1917) gave ""a very poor idea of the 
size and usual character of the species." and pro- 



1 Hopkins Marine Station of Stanford 1 
Pacific Grove. California 93950. 

2 Dept. Zoology. University of Nebraska. L 
Nebraska 6S50S. 



w, 



Bl III ll\ SOI THERh ( ILIFOKMA ACADEMY OF SCIENCES 



VOLUME 71 




Figure I. Holotypc of Styela barnharti (= S. 

plicata); positions of apertures marked in right hand 
view. 



vided a new description. Subsequent investigators 
have found it difficult to reconcile Ritter and 
Forsyth's original description of S. barnharti with 
the features of the species called S. barnharti by 
Van Name. J. R. Whittaker (pers. comm., 1971) 
called our attention to the fact that Ritter and 
Forsyth's figure of the intact holotype actually 
resembles another Californian ascidian (not 
named), and suggested that the S. barnharti of 
Van Name (1945) might be merely a shorter- 
stalked variety of S. montereyensis. 

In the present paper we show: (1) the holotype 
of Styela barnharti Ritter and Forsyth, 1917. is a 
specimen of S. plicata ( Lesueur, 1823); (2) the 
entity called S. barnharti by Van Name (1945) 
and subsequent authors is actually S. clava Herd- 
man, 1881: and (3) the latter species is quite 
distinct from the Californian stalked ascidian S. 
montereyensis (Dall, 1872). 

THE HOLOTYPE OF 

STYELA BARNHARTI 

RITTER AND FORSYTH, 1917, 

AND COMPARISON WITH 5. PLICATA 

(LESUEUR, 1823) 

The holotype of 5. barnharti was discovered re- 
cently at Stanford University, and is now deposited 
in the Department of Invertebrate Zoology, Cali- 
fornia Academy of Sciences (CASIZ Type Series 
No. 552). It has been reexamined by the senior 
author. In external features the tunic (Fig. 1) 
closely resembles Ritter and Forsyth's fig. 2. pi. 
3S. The body within the tunic exists in several 
pieces but most of the parts are present and it has 
been possible to verify nearly all of Ritter and 



Figure 2. Styela plicata from different localities. 
1-4, San Diego. California. 5, holotype of S. barnharti, 
San Diego, California. 6, Long Beach, California. 
7, Newport Bay, California. 8-9, Cedar Key, Levy 
Co., Florida. 



Forsyth's recorded observations. The following 
account corrects a few details and adds others 
not included in the generally adequate original 
description. 

Branchial tentacles. — "Of several sizes, about 
forty," according to Ritter and Forsyth. Reex- 
amination shows 64, of 3 orders (but of 4 sizes, 
as some third order tentacles are very small), 
fairly regularly arranged in the pattern 1-3-2-3-1. 

Internal longitudinal vessels. — A recount on the 
type, taken across the middle of the pharynx, 
shows the following distribution (DL = dorsal 
lamina, vessels on branchial folds in parentheses) : 



Right 4(19)8(21)7(16)8(14)4 



DL 

Left 5(20)6(20)7(21)4(21)2. 

Ritter and Forsyth found 94 and 102 vessels, on 
right and left, respectively; we found 101 and 
106. The discrepancies found probably reflect 
a difference in the level on the pharynx at which 
the counts were taken, and differences in assign- 
ment of vessels near the bases of folds to branchial 
folds or interfold areas: they are not considered 
significant. 

Gut. — In cross section the stomach showed 33 
folds; Ritter and Forsyth found about 35. The 
presence of numerous endocarps on the intestine 
is confirmed. 

Testes. — These are considerably more numerous 
and the larger testes more complexly branched 
(Fig. 5) than is indicated in Ritter and Forsyth's 
illustrations (their figs. 39 and 40, pi. 42), though 
no testes exhibit such attenuated branches as those 



1072 



CAL1F0RNIAN Sir I.I. ID ASC1D1ANS 







Figure 3. A, Styela clava, San Diego, California. 
B, Styela montereyensis, Monterey, California. 



shown for an old and large individual of 5. pliccita 
by Van Name (1945:295, fig. 192). 

The holotype has been compared directly with 
individuals of Styela plicata from several localities 
(see below). In external features (Figs. 1-2) and 
in all internal qualitative features, the holotype is a 
"typical" specimen of S. plicata (compare Fig. 
8C, this paper, with Ritter and Forsyth's figs. 39 
and 40, pi. 42). The quantitative characteristics, 
both those which increase more or less regularly 
with body size (e.g., number of internal longitudi- 
nal pharyngeal vessels) and those which do not 
(e.g., number of ovaries, number of plications in 
the stomach wall), are found in figure 6. In most 
of these features the holotype of 5. barnharti is not 
an exceptional specimen of S. plicata. The num- 
ber of gonads (9 on the right side. 3 on the left) 
is rather large (Van Name, 1945, records 4-7 on 
the right and 2 on the left as more usual), but it is 



wiihin ilic known rangi riation 

species; Tokioka (19 6 I 

righl and I 3 "i! the li fl in lap 

and Tucker (1942) recorded 11 on Ihe 

ami I -3 on the leli in m.lr. iduals from ' 

< arolina. 

The first correct published record ol the oc- 
currence of S. plicata, undei thai name, "n the 
west coast of North America is thai ol Keish 
(1963), based on material collected in the spring 
of I960 at Alamitos Bay, Long Beach, < alifornia 
and determined by one of us ( Abbott ) . An earlier 
report of S. plicata from Point Richmond s. • 
Francisco Bay (Abbott. 1954) is erroneous: re- 
examination of the specimens slums they are S. 
clava. New records of S. plicata in southern ( ali- 
fornia are given by Fay and Johnson (1971). It is 
now clear that .S'. plicata was present at San Diego 
at least as early as July 1915, when Hitter and 
Forsyth's material was collected. Its present known 
distribution on this coast extends only from San 
Diego to Santa Barbara. California, at depths ol 
0-3 m. Elsewhere it is widely distributed in the 
Atlantic. Pacific, and Indian Oceans, and has been 
the subject of numerous field and laboratory in- 
vestigations. 

Specimens examined: Point Loma. San Diego. 
California, 24 November 1971. coll. G. I. ingle: 
Mission Bay, San Diego. California. 7 August 
1966, coll. K. M. Taylor: Newport Bay. California. 
autumn 1966, coll. G. Bane: Alamitos Bay, Long 
Beach, California, 18 August 1960. coll. D. J. 
Reish; Cedar Key, Levy Co., Florida, 21 August 
1950, coll. K. C. Strawn: Boca Ciega Bay. Pinellas 
Co., Florida. 3-28 November 1955. coll. R F. 
Hutton: Beaufort, North Carolina. 13 December 
1915, coll. unknown: and Wakanoura. Kii. Japan, 
no date, coll. Jordan and Snyder. 

REEXAMINATION OF THE STYELA 
BARNHARTI OF VAN NAME (1945) 

Van Name (1945) presents the first unequivocal 
published account of this species on the west 
coast. We have examined specimens from eleven 
different localities (see below). An especially 
large series from Venice. California was examined 
in connection with studies of reproduction and 
growth (J. V. Johnson. 1971). Most animals were 
taken intertidally or in the shallow subtidal zone 
on floats, pilings, or rocks: only the specimen 
the Hyperion sewer outfall comes from 
depths (50 ft). 

Van Name's description of 5. b 



98 



III 111 ll\ sol ////A'\ ( tLIEORNIA ACADEMY OF SCIENCES 



VOLUME 7/ 




Figure 4. Styelid ascidians found growing together, Santa Barbara harbor, California. 
1-6, Styela montcreyensis (specimen 4 bearing an 5. plicata). 7-11, Stycla clava. 



where it deals with the placement of the stomach 
and intestine, is generally accurate, though it does 
not always convey the range of variation to be 
expected. The following account augments or 
modifies only those portions of the description 
which require this. 

External features. — Our largest specimens 
reached a total length of about 200 mm. Length 
of the stalk seldom approaches half the total 
length, and may be less than one tenth the total 
length, or virtually absent in small specimens. 
Division between body and stalk ranges from 
sharp to very gradual and indistinct. Number and 
distribution of tubercles on the test varies greatly; 
at the extremes they may cover as much as the 
anterior two thirds of the body proper, or they 
may be confined to a few small bumps about the 
apertures. Development of longitudinal wrinkles 
likewise varies greatly, but the longitudinal ridges 
and grooves, when present, represent folds in the 
relatively thin and leathery tunic, and not al- 
ternating areas of thick and thin tunic. The oral 
siphon may point anteriorly or be canted at a 
slight angle, but it never recurves to point pos- 
teriorly as it usually does in S. montereyensis. 



Dorsal tubercle. — The aperture often shows 
greater inrolling of the horns, and greater ir- 
regularity, than indicated by Van Name (1945); 
specimens exhibit the same sorts of variations de- 
picted by Millar (1960, fig. 1 B-D) for 5. clava. 




Figure 5. Outlines of testes from holotype of Styela 
barnharti (=S. plicata), showing nature of branch- 
ing. 



I>)72 



CAUI'OKNIAN STYEL1D ASCID1ANS 








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HOLOTYPE OF S. barnharti 



• SAN DIEGO, CALIF. 
■ NEWPORT BAY, CALIF. 
« LONG BEACH , CALIF. 







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a BOCA CIEGA BAY, FLORIDA 

A CEDAR KEY, LEVY CO., FLORIDA 

O BEAUFORT, NORTH CAROLINA 

□ WAKANOURA KM, JAPAN 



Figure 6. Styela plicala: relation of total length (body and stalk) to numbers of oral 
tentacles, internal longitudinal vessels, gonads (ovaries), and plications on stomach wall. 



Tentacles. — Average number 55 in 7 individuals bear closely crowded internal longitudin 

counted; range 44 (in a specimen 135 mm long) whose number increases with body size (Fie 

to 76 (in a 90 mm individual ) . Total number of internal longitudin 

Pharynx. — The pharyngeal folds are low but sides) ranged from 137inananii 



100 



350 



300 



CO 

w 250 

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Z> 

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2 150 



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50 



B( / / / TIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 

-i 1 1 1 1 r- 



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i i — r - 1 — i 1 — <— — i — r 



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STYELA clava 

■ VENICE, CALIF. 

O SANTA BARBARA, CALIF. 
STYELA montereyensis 

▲ MONTEREY, CALIF. 

• SANTA BARBARA CALIF. 



-I L 



:; I ' 



-I 1_ 



10 15 

LENGTH (cm) 



-i i i i i_j_ 



20 



25 



Figure 7. Styela clava and S. montereyensis; relation between total length (body and stalk) 
and number of internal longitudinal vessels on pharynx. 



1072 



CALIFORNIA^ STYELID ASCIDIANS 
Table I. Distinguishing features of Styela clava and Siycla montert 



Feature 



S. clava 



Tunic 



Siphons 
Pharynx 

Gut 



Atrial tcnlacles 
Ovaries 

Testes 

West coast dis- 
tribution and 
habitat 



Usually with conspicuous tubercles, at least 
anteriorly (Figs. 3 A, 4). 



Longitudinal folds absent to well-developed, 
confined mainly to posterior body and stalk, 
consisting of wrinkles in tunic rather than 
structural folds (Figs. 3A, 4). 

Both siphons straight, directed anteriorly 
(Fig. 4). 

Esophageal aperture located halfway down 
pharynx; branchial folds with numerous 
internal longitudinal vessels (Fig. 7). 

Stomach in descending position (Fig. 8A); 
first intestinal loop near posterior end of 
body: intestinal wall bearing numerous 
endocarps. 

Forming more than one row. 

Tubular, typically 4-9 on the right and 2-5 
on the left, one or more often branched 
posteriorly (Fig. 8A). 

Clustered into compact lobes (Fig. 8A). 

San Diego Bay to San Francisco Bay, Cali- 
fornia; in areas not exposed to strong surf: 
low intertidal to at least 75 ft depth. 



5 monii 



Tubercles l;' 1 1 in 

I. ii get animals with a few loo • .varl 

tunic matei ial, casilj di Lai bi d 

near the siphons (Figs, 3B, 4). 

Longitudinal I idr<-\ and irrOOVl on bod) and 

stalk extending antcriorlj near); to iphi 
and representing respectively thickci and 

thinner regions of the test (lies 1|i, 4). 

Oral siphon curved, directed vcnlrally 01 
posteriorly digs. 3B, 4 ). 

Esophageal aperture located near posterior 
end of pharynx: branchial folds with fewer 
internal longitudinal vessels (Fig. 7). 

Stomach in ascending position (Fig. 8B); 
first intestinal loop about halfway up body; 
intestinal wall smooth, without endocarps. 

Forming only a single file. 

Tubular, typically 2 on each side (rarch I 
or 3). seldom branched posteriorly il 
8B). 

Individual testes separate. simple or 
branched, not clustered into lobes (Fig. 8B). 

San Diego Bay. California to British Colum- 
bia: on outer coastal rocks and pilings but 
also invading calmer harbors: low intertidal 
to about 100 ft depth: coexists with S. clava 
at Santa Barbara and Mission Bay. San 
Diego. 



to 327 in a specimen 140 mm long. Distribution 
of vessels in 3 animals exhibiting low, intermediate, 
and high vessel counts were as follows: Total 

Length 
-Right 2(16)1(16)2(14)4(10)3 



DL 



DL 



-Left 4(16)3(12)3(14)3(11)3 
-Right 6(35)4(31)3(31)5(21)4 

-Left 6(30)4(29)7(26)7(17)4 



25 mm 



100 mm 



-Right 4(44)6(36)4(42)5(24)3 

DL 140 mm 

-Left 6(35)4(36)4(43)5(22)4. 

The esophageal opening occurs about halfway 
down the pharynx. One would never infer this 
from Van Name's figure 205, though he correctly 
notes that the dorsal lamina "extends far behind 
the esophageal opening." 



Gut (Fig. S A). — The esophagus and stomach 
descend posteriorly from the esophageal opening. 
and the stomach is always in a descending posi- 
tion. Beyond its pyloric end the intestine curves 
sharply anteriorly again and extends toward the 
base of the atrial siphon. In relaxed specimens the 
intestine shows little curvature: in contracted ani- 
mals it may bend, even rather sharply, where it 
passes the esophageal region. In none of the 
specimens examined by us was the placement of 
esophagus and stomach and the course of the 
intestinal loop at all similar to the situation 
portrayed by Van Name (1945:310. fig. 

Gonads (Fig. S A).— Van Name ( 1945) found 
5-9 ovaries on the right and 3-5 on the left. We 
find 4-9 (average 6) on the right and 2 
age 4) on the left in a series of 26 individuals 
25-160 mm in total length. Most specimens 
at least one ovary which is branched po: 



102 



HI III IIS SOI I III l<\ CAI.II OKMA AC ADhMY Ol SCIENCES 



VOLUME 71 





Figure 8. Medial views of right and left halves of body with pharynx removed, showing gut 
and gonads. A, Styela clava, body as shown 70 mm long. B, Styela montereyensis. C, Styela 
plicata. 



such ovaries were counted as single ovaries. Testes 
are grouped in tight clusters, as described and 
figured by Van Name. 

Atrial tentacles. — Minute, forming a narrow 
band several tentacles in width, circling the in- 
terior base of the atrial siphon. The condition is 
similar to that shown by Tokioka (1955b, fig. 3) 
for Styela clava from Japan, but instead of two 



distinct rings of tentacles separated by a bare 
strip, the area between the rings is also occupied 
by tentacles. 

Through the courtesy of David Pawson and 
Maureen Downey it was possible to examine sev- 
eral specimens of Styela clava Herdman, 1881 
in the collections of the U. S. National Museum of 
Natural History, taken near the type locality 



1>>72 



CAI.IIOKNIAN SI Villi) ,ISC//</i 






(Kobe, Honshu, Japan; USNM I 1 703 > ; these ani- 
mals had been examined and identified previously 
by Tokioka (1967:191). Test tubercles in these 
Japanese specimens are somewhat more sparsely 
and regularly placed than is typical of ( aliform, m 
individuals, but this is a character in which both 
American and Asiatic populations show enormous 
and widely overlapping variation. A slighl differ- 
ence exists in the atrial tentacles, as noted earlier. 
In all other respects the species dealt with by Van 
Name under the name Styela barnharti agrees 
with the Japanese specimens and with recent de- 
scriptions of Styela clava (Millar I960: Tokioka 
1953, 1959, 1967; Rho 1971 ). 

The source of Van Name's curious figure of the 
gut (1945:310, fig. 205) remains a mystery. 
Harold S. Fcinberg placed at our disposal the 
earliest Californian collection of .V. clava in the 
American Museum of Natural History, New York 
(AMNH 931, taken in Newport Bay on 15 No- 
vember 1933 by G. E. MacGinitic, and noted on 
the label to "occur abundantly on piles, etc."). 
The specimens, mostly dissected already, had 
been identified by Van Name as S. barnharti. 
All are typical S. clava as regards the gut and 
other features. A search through the other collec- 
tions of 5. barnharti in the American Museum 
which had been examined and identified by Van 
Name, failed to reveal a specimen designated as 
the source of the figure in question. There is a 
possibility that in making the drawing Van Name 
removed the gut and sketched it by itself, in an 
unnatural position: one specimen in lot AMNH 
931 has the gut wholly separated from the body, 
and it can be twisted to resemble that in Van 
Name's figure. At any rate, the figure is either 
inaccurate or portrays an individual with a grossly 
distorted or aberrant gut. 

Styela clava is widely distributed on Asiatic 
shores, ranging from the Kurile Strait, Sea of 
Okhotsk, through southern Siberia, Japan, Korea, 
and the China coast at least as far south as 
Shanghai. It was discovered in the vicinity of 
Plymouth, England in 1953, where it was described 
as a new species (Styela mammiculata Carlisle, 
1954). Carlisle noted its similarity to the Styela 
barnharti of Van Name (1945). and its identity 
with S. clava was shown by Tokioka ( 1955a) and 
Millar ( 1960). Clearly a recent introduction here, 
it has now spread to other parts of southern Eng- 
land (Millar. 1970), and the French channel coast 
(Monniot, 1970). 

In Californian waters 5. clava is regularly abun- 
dant in sheltered areas from San Diego to Santa 



Barbara ( Fay and loh 

Point ' onception its occurrcm 

ll is recorded once li MoiTO Ba and II 

most years from Monterey Bay It ... I 

in most years in San Francisco Ba) though pop 

ulations in some localitic: 'ii appi at afti i , 

nl heavy rains (W I li' rg, ( harlcs N| 

pers. comm.). The species has never been 

in either Tomales Hay or Bodega Bay i<> the north 

of San Francisco (W. B. Gladfeltei and ( n 

Hand, pers. comm.), despite the seeming suit., 

hilily of the habitat .ind extensive collecting by 

field biologists. 

Specimens examined from: Point I Oma S 
Diego. California. 24 November 1971, coll. G. 
1. ingle: Mission Bay, San Diego. California, 15 
June 1959. coll. J. H. McLean, and 10 August 
1966, coll. K. M. Taylor: Newport Bay. Cali- 
fornia, 15 November 1933. coll. G. I '•' 
Ginitie: Hyperion sewer outfall. Palos Vcrdes 
Peninsula, Los Angeles, California. 10 April 1971. 
coll. W. J. North; Venice, Santa Monica Bay. 
California, various dates in 1971, coll. R. C. Fay; 
Santa Barbara, California, June 1971. coll. K ( 
Fay; Morro Bay, California, 4 February 1972. 
coll. D. H. Montgomery: Monterey, California, 15 
November 1961. coll. D. P. Abbott: Oakland. San 
Francisco Bay, California. 26 April 1955. coll. 
R. Rinaldi; Berkeley. San Francisco Bay. Cali- 
fornia. 20 April 1955. coll. R. Rinaldi: Point 
Richmond. San Francisco Bay. California. 12 June 
1949. coll. R. Shaw. 



COMPARISON OF 5. CLAVA AND S. 
MONTEREYENSIS. AND THE IN- 
TRODUCTION OF 5. CLAVA 
TO CALIFORNIA 

Styela clava can usually be distinguished from S. 
montereyensis by its shorter stalk, plumper body, 
and possession of tubercles anteriorly. Variation 
in all of these features is great, and occasional 
specimens are found which are difficult to identify 
in the field. Internally the species are un- 
questionably quite different, and no specimens 
have been examined by the authors, or reported 
by others, whose features suggest hybrid origin. 
The most useful characters distinguishing the two 
species are shown in table 1. 

Information now available permits a reas 
guess as to the date S. clava was introduced 
Californian waters. Ritter collected asciJ: 
tensively over a period of two d< 



104 



i:i 111 IIS sol iiiiks CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



1917, especially in southern California, and ii 
seems unlikel) he could have missed this species 
had il been common in liis lime. 1 1 is also un- 
likelj thai Ritter mistook ii for S. montereyensis. 
Me mentions taking the latter "from main points 
on the coast from San Diego to Mendocino" 
i Kiuer and Forsyth, 1917:451), and the accom- 
panying description and figures of S. montereyen- 
sis clearly apply to that species alone. We con- 
clude that S. clava was absent (or at least extra- 
ordinarily scarce) in Californian waters prior to 
I 1 ) 17. 

No species resembling 5. clava is reported by 
Johnson and Snook ( 1927). though these authors, 
too. collected extensively in areas where S. clava 
is now abundant, and were familiar with S. 
montereyensis. Johnson had worked with ascidians 
under Ritter (Ritter. 1909:66) and would prob- 
ably have spotted the difference between S. clava 
and S. montereyensis. Ricketts and Calvin (1939) 
mention only 5. montereyensis, but their figure 107 
almost certainly depicts S. clava (see also Van 
Name, 1945:310). In any event. S. clava was 
on the coast by that time, as documented by the 
collections of MacGinitie in 1933 at Newport Bay 
(cited above). George and Nettie MacGinitie, 
who arrived at the Kerckhoff Marine Laboratory 
in 1932. inform us (pers. comm.) that S. clava 
also was abundant in Newport Bay in 1932 and 
all subsequent years, and that S. montereyensis 
has never been common there. We conclude that 
S. clava probably gained its foothold on the Cali- 
fornian coast in the late 1920's. 

ACKNOWLEDGMENTS 

Thanks are due to J. R. Whittaker. Wistar Institute, 
whose letter to the junior author provided the im- 
mediate stimulus for the present investigation; to 
Rimmon C. Fay, Pacific Bio-Marine Supply Co., and 
to many other individuals cited in the text, for pro- 
viding ascidian collections and distributional informa- 
tion; to David Pawson and Maureen Downey, Smith- 
sonian Institution, U.S. National Museum of Natural 
History, and Harold S. Feinberg, American Museum 
of Natural History, for the loan of specimens; to 
William E. Berg. Cadet Hand, and Charles Spowart, 
University of California, Berkeley, William B. Gladfel- 
ter, University of the Pacific, and George and Nettie 
MacGinitie. formerly of the California Institute of 
Technology, for providing information on the occur- 
rence of styelid ascidians in San Francisco, Bodega, 
Tomales. and Newport Bays; to Mitsuaki Nakauchi, 
Kochi University, Japan, for information on Japanese 
styelids: and to Lois E. Kruschwitz, University of Ne- 
braska, for the photograph in figure 4. 



LITERATURE CITED 

Abbott, D, P, 1954. Key to the littoral ascidians of 
the central California coast, p. 301-310. //; 
Light, S. F., R. I. Smith. F. A. Pitelka. 15. 1'. 
Abbott, and F. M. Weesner, Intertidal inverte- 
brates of the central California coast. Univ. 
California Press. Berkeley. 446 pp. 

Carlisle, D. B. 1954. Styela mammiculata n. sp., 
a new species of ascidian from the Plymouth 
area. J. Mar. Biol. Assoc. United Kingdom, 33: 
329-334. 

Fay, R. C, and J. V. Johnson. 1971. Observations 
on the distribution and ecology of the littoral 
ascidians of the mainland coast of southern 
California. Bull. So. California Acad. Sci., 70: 

114-124. 

Johnson. J. V. 1971. The annual growth and re- 
productive cycle of Styela sp. in the Marina del 
Rey, Venice, California. M.S. thesis, Dept. 
Zoology. Univ. Nebraska. 

Johnson, M. E., and H. J. Snook. 1927. Seashore 
animals of the Pacific coast. Macmillan Co., 
New York. 659 pp. 

MacGinitie, G. E., and N. MacGinitie. 1968. Nat- 
ural history of marine animals. 2nd ed. Mc- 
Graw-Hill. New York. 523 pp. 

Millar, R. H. 1960. The identity of the ascidians 
Styela mammiculata Carlisle and S. clava Herd- 
man. J. Mar. Biol. Assoc. United Kingdom, 39: 
509-511. 

. 1970. British ascidians. Synopses of the 

British Fauna No. 1, Linn. Soc. London. Aca- 
demic Press, London, 92 pp. 

Monniot, C. 1970. Sur quatre Ascidies rares ou mal 
connues des cotes de la Manche. Cahiers de 
Biologie Marine, 11:145-152. 

Reish. D. L. 1963. Mass mortality of marine or- 
ganisms attributed to the "red tide" in southern 
California. California Fish and Game, 49:265- 
270. 

Rho, B. J. 1971. A study on the classification and 
the distribution of the Korean ascidians. J. 
Korean Research Institute for Better Living, 6: 
103-166. 



1972 



CAI.II-OKNIAN yiYII.ll> ASCIDIA 






Ricketts, E. F., and J. Calvin. L939. Between Pacific 
tides, Stanford Univ. Press, 320 pp, 

. 1968. Between Pacific tides. 4th eil. (J. 



W. Hedgpeth, rev.). Stanford Univ. Press, 614 pp. 

Kilter, W. E. 1909. Halocynthia johnsoni n, sp., a 
comprehensive inquiry as to the extent of law 
and order that prevails in a single animal speeies. 
Univ. California Publ. Zool., 6:65- 114. 

Ritter, W. E., and R. A. Forsyth. 1917. Aseidians 
of the littoral zone of southern California. Univ. 
California Publ. Zool., 16:439-512. 

Tokioka, T. 1953. Aseidians of Sagami Bay col- 
lected by His Majesty the Emperor of Japan. 
Iwanami Shoten, Tokyo, 315 pp.. 79 pi. 

. 1955a. Record of Styela clava Herdman 

from the European waters. Zool. Mag. Tokyo, 
64:200. 



1955b ' onlt ibutiot 

ft a XI. Sporadii mi mot u P 

Mar. Biol, I ab I 10 218. 

— . 1959, ( ontributions to lapi 
fauna XV. Sporadic memorandun 
clava var. symmetrica no* a new pedum 
styelid from the Inland Sea. Publ 
Biol. Lab., 7:457-463. 

1967. Pacific Tunicata ol thi I i 



National Museum. Bull. U.S. Nat. Mus., 251 

1-247. 

Tucker, G. H. 1942. The histology ol the 

and development of the egg envelopes of an 
ascidian {Styela plicata Lesueur). I. Morph.. 
70:81-113. 

Van Name. W. G. 1945. The North and South 
American aseidians, Bull, Amer. Mus. Nat. II 
84:1-476. 31 pi. 

Accepted for publication June 10. 1972. 



KI-SI ARCH NOTES 



A NEW SPECIES OF PUGNUS FROM 

COCOS-KEEL1NC. ISLANDS. INDIAN 

OCEAN (GASTROPODA) 



The littoral molluscan fauna of Cocos-Keeling Islands, 
isolated atoll group of the eastern Indian Ocean, was 
catalogued by Maes (Proc. Acad. Nat. Sci. Phil- 
adelphia. 119(41:93-217. 1967). and its geographic 

affinities assessed. Included in the collections which 
formed the basis of that report were several speci- 
mens of an undescribed species of Pugnus, a peculiar 
gastropod which may be an aberrant member of 
the Margtnellidae, a family under study by the 
author. Through the kindness of Virginia Orr Maes, 
Academy of Natural Sciences. Philadelphia, and 
Winston F. Ponder. The Australian Museum. I have 
had the opportunity to compare the Cocos-Keeling 
specimens with specimens of the Australian Pugnus 
parvus Hedley (Rec. Australian Mus., 2:105-106, 
1896). the type and only species of the genus 
previously described. 



Sciences. Philadelphia. No. 288324. Fight paratypes. 
ANSP No. 32474, from same locality as holotypc. 

Referred material: ANSP No. 288311, one speci- 
men, from two miles east of Ujong Tanjong. West 
Island. Cocos-Keeling Islands, in 4 fathoms (7.32 
m). hard sand and weed; Ostheimcr, Orr, Ross, 
collectors, 1 1 February 1963. 

Discussion: From Pugnus parvus Hedley, originally 
described from Manly, near Sydney, New South 
Wales, the new species differs in several shell details: 
P. maesae has four nearly equal columellar folds: 
P. parvus has two large folds and a smaller, deeply 
seated fold posterior to them. Pugnus maesae has 
dentition on the outer lip; P. parvus has none. The 
specimens of P. parvus examined by the author, 
from Mallacoota. Victoria (nearly 300 miles south of 
Sydney), were rather narrower, in proportion to their 
length, than the new species. They showed a curious 
surface sculpture of shallow hexagonal pits arranged 
"honeycomb" fashion, elongating behind the outer 
lip into spiral sculpture like that originally described 



Pugnus maesae, new species 
Figure 1 

Pupmis sp. Maes. 1967. p. 141. pi. 14. fig. D. 

Description of holotype: Shell minute, solid, white, 
short-cylindrical, slightly broader posteriorly; surface 
polished: involute, spire immersed, covered by a pad 
of callus: aperture as long as shell, nearly straight, 
narrow, widening somewhat anteriorly, arching above 
summit of body whorl; outer lip varicose externally, 
thickened and denticulate within, the denticles very 
fine and close together on central portion of lip. 
larger and more widely spaced anteriorly and pos- 
teriorly: body whorl slightly contracted medially, 
sculptured with approximately 40 wavy spiral grooves 
which are crossed by very fine raised axial threads: 
parietal wall covered by thin, transparent, well- 
defined layer of smooth callus which bears a longi- 
tudinal row of about seven weak, irregularly spaced 
tubercles: anterior portion of columella having four 
evenly spaced, oblique folds (including fold at base 
of columella), most anterior fold the largest; folds 
decreasing in size and becoming more horizontal 
posteriorly; anterior margin of shell produced, evenly 
rounded, without a siphonal notch. Length 1.5 mm, 
breadth 1.1 mm. 

Type locality: One mile north of Tanjong Puji. 
West Island, Cocos-Keeling Islands, Indian Ocean; 
dredged in 6 feet (1.83 m). coral mud and sand, 
with some Caulerpa algae: collected by R. Ostheimer 
and V. Orr (Maes). 30 January 1963. 

Type material: Holotype. Academy of Natural 




Figure 1. Pugnus maesae, new species. Ventral view 
of holotype. Copy, Maes, 1967, pi. 14, fig. D. 



106 



I')72 



RESEARCH NOTES 






for the species hy llcdlcy. These specimens ;il •-.< > 
had perforate apices. 

Familial placemen! of the genus Pugnus is un- 
certain, li was described by Hedley (IK96) as a 
member of Ringiculidae, and Zilch (Handbuch dcr 
Palaozoologie, 6: 1-835. 1959) placed ii in Cephalas 
pidca. Maes (1967) transferred it to Marginellidae, 
remarking on its resemblance to the [?marginellid] 
genus Marginellopsis Bavay, 1911. The character ol 
the columellar folds and the lip denticulation appear 
typically marginellid. The wavy, incised spiral sculp- 
line recalls that of some cephalaspidcan genera, 
such as Acteon Montfort. 1X10. Knowledge of its 
true relationships awaits an anatomical study. 



Barry Roth, 1217 
California 94117. 



Waller Street. San Francisco, 



Accepted for publication May 24. 1972 



A NOTOPHYCID POLYCHAETE FROM 
CALIFORNIA 



Phyllodocella bodegae, 

I igun I and .' 

Material examined Mason's Marina Bodega Hai 
bor, California, luly 20, 1971, from •■ gclatinou 

sac attached to a colons ol Obi lm »p I 

on a floating boal docl - inc ipccimcn 1 1 

deposited in the collection , ol the Allan M. 

Foundation. 
Description'. The holotype is a complete, sexually 

mature female with 24 segments that is H nun long 
and 2 mm wide without setae. It is white with 
reddish pigment spots ovci the antcrioi end in 
alcohol preservation. 

The pygidium is a small, rounded cushion with- 
out anal cirri; the anus is dorsal. 

The prostomium ( Fig, I ) is pentagonal and has 
two pairs of long, slender antennae near the antcnor 
margin. A pair ol small, distinct frontal lobes arc 
present on the anterovcntral margin. Two pan- ol 
eyes are present at the middle and mid-posterior part 
of the prostomium: the anterior pair is lensed; 
the posterior pair is semi-lunular in shape. The 
peristomial segment is a complete ring, forming 



While collecting at a floating boat dock in Bodega 
Harbor, California, one of us (Belman) found a 
thick gelatinous sac attached to a colony of a hydroid 
(Obelia sp.). A polychaete was moving around 
inside the sac; this worm could not be identified 
as any polychaete reported from California (Hart- 
man, Atlas of enantiate polychaetous annelids from 
California. Allan Hancock Foundation. Los Angeles. 
1968; Atlas of the sedentariate polychaetous an- 
nelids from California. Allan Hancock Foundation. 
Los Angeles, 1969) and turned out to belong to the 
family Notophycidae. recently described from New 
Zealand (Knox and Cameron, Trans. Roy. Soc. New 
Zealand. Biol. Sci.. 12:73-85. 1970). The Cal- 
ifornian specimen differs from the other known 
specimens in several respects and is described as a 
new species in a new genus. 

The relationship between the family Notophycidae 
and related polychaetes was discussed in detail by 
Knox and Cameron (1970). 



Phyllodocella, new genus 

Notophycids with a muscular proboscis, but with- 
out jaws. 

The other known genus in the family. Nolophyeus 
Knox and Cameron (1970) has a pair of lateral 
jaws in the proboscis. Such jaws are absent in the 
present specimen. 

The generic name refers to the resemblance be- 
tween this notophycid and members of the Phyl- 
lodocidae. 




Figure 1. Phyllodocella bodegae, new spc- 
anterior end. dorsal view, right dorsal tent 
cirrus broken. \ 50. B. anterior end. vent! 
X50. 



108 



111 I I I 1 1\ sol I 111 A'\ CA1 II 0RN1A ACADEMY <>l SCIENCES 



VOLUME 71 




Figure 2. 
seta from 
anterior v 



Phyllodocella bodegae, new species. A. 
parapodium 7. x 950. B. parapodium 7. 
iew, X 52. 



phycus milium Kno\ anil Cameron (|y70) from 
Snares Island. New Zealand, in that it has two pairs 
of antennae and a single peristomial segment with 
two pairs of tentacular cirri and in the shape and 
structure of the parapodia. Notophycus minuta has 
a pair of lateral jaws in the proboscis; jaws are 
absent in I', bodegae. This character is here con- 
sidered of generic rather than specific value. The 
two species further differ in the shape and detailed 
equipment of the parapodial lobes and in the devel- 
opment of the antennae and tentacular cirri. 

Both species described in the Nolophycidae are 
quite small and arc probably easily overlooked by 
collectors: it is however, rather remarkable that 
these worms have not been seen more frequently, 
considering the apparent wide geographical distri- 
bution of the family. 

Kristian Fauchald, Allan Hancock Foundation, 
University of Southern California, Los Angeles, 
California 90007 and Bruce W. Belman, Department 
of Biological Sciences, University of California, Santa 
Barbara, California 93106. 

Accepted for publication June 2, 1972. 



the lateral and posterior lips ventrally; it has 
two pairs of long, subdistally slightly inflated ten- 
tacular cirri. 

The proboscis, which is strongly muscular, stretches 
through the four first setigers; jaws are absent. 

Notopodia are absent in the first two setigers. 
The first neuropodia (Fig. IB) project strongly 
ventrally with the setae pointing anteriorly and 
ventrally: the second neuropodia are slightly more 
lateral in position. Noto- and neuropodia project 
dorsolateral^ and ventrolateral^ in all setigers from 
setiger 3. Each parapodium (Fig. 2B), where fully 
developed, has similar, uni-acicular noto- and neuro- 
podia. Each ramus has a cylindrical base and is 
distally expanded into a large, bulbous pad. The 
single fascicle of setae forms a straight line along 
the distal side of this pad. The notopodial pad is 
expanded to form a very large, bulbous lobe over- 
hanging the dorsum in all setigers posterior to 
setiger 4: this development is absent in the two 
first pairs of notopodia. The dorsal cirrus is lateral 
to the notopodial bulbous lobe. The ventral cirrus, 
which is of the same size and shape as the dorsal 
one, is near the ventrolateral corner of the neuro- 
podial pad. 

All setae are composite spinigers. Each seta 
(Fig. 2D) has a long, slender shaft that is distally 
crenulated: it has a series of poorly defined trans- 
verse ridges subdistally. The appendage of each seta 
is long, evenly tapering and has series of fine teeth 
along the margin. 

Discussion: Phyllodocella bodegae resembles Noto- 



A KEY TO THE FISHES OF THE FAMILY 

GOBIIDAE (TELEOSTOMI) OF 

CALIFORNIA 

Gobies comprise an interesting group of fishes, oc- 
cupying diverse habitats along the Californian coast. 
They are frequently encountered in fish collections, 
yet a complete key to all species has never been 
published and identification, especially of preserved 
sub-adults, is often difficult. 

All species are native to California except Acan- 
thogobius flavimanus (Temminck and Schlegel) and 
Tridentiger trigonocephalus (Gill). Both of the 
above were introduced to the San Francisco area 
from the Orient. Listings for the occurrence of 
Evermannia longipinnis (Steindachner ) in Californian 
waters are the result of an error by Fowler (1923), 
copied by Ulrey and Greely (1928) and Barnhart 
(1936) (Robert J. Lavenberg, pers. comm.) and thus 
this Gulf of California species is excluded from the 
key. 

Several characters are taken from Jordan and 
Evermann (1896) and Norman (1957). Meristic 
counts that deviate from the above sources are the 
result of a study of the collections in the Natural 
History Museum of Los Angeles County and Cali- 
fornia State University, Long Beach, verified where 
possible by Clothier (1950). 



19 7 2 



RESEARCH Null \ 






FAMILY (iOHIIDAi: 

Dorsal fins separate, the pelvic Tins 
united to form a Flaring cone-shaped 
sucking disc, free from the body 
(Fig. I). 




Figure I. 



I. a. Without five dark bands on leading edge 
of both the first and second dorsal fins; 
teeth in jaws simple, neither clubbed nor 
incised at their extremities (Fig. 2a) 2 



row ol teeth trii u pi 

dilated 1 1 ig 2b) 

goby- Trielenllgei Irlgonoi 




I ruin- 4. 



2. a. Byes normal; no specialized folds and flap 1 - 

in front and below the eyes 
b. Eyes very small, vestigial or nearly so in the 
adults; cheeks much enlarged, tumid; skin 
of head with tactile organs well developed 
(I ig. 4) 

3. a. Dorsal spines VI 



Halfblind goby — Leihops connectens Huhhv 

b. Dorsal spines III Blind goby 

— Typhlogobius californiensis Steindachner. 

4. a. Dorsal spines VII or less _ 
b. Dorsal spines VIII 

Yellowfin goby — Acaniho- 
gobius flavimanus (Temminck and Schlegel). 

5. a. Caudal fin normal, not greatly elongate: 

broad and rounded (Fig. 5a) 




Figure 2. 



b. Five dark bands on leading edge of both the 
first and second dorsal fins (Fig. 3); outer 





Figure 3. 



Figur, 5. 



b. Caudal fin greatly elongate: pointed 

5b) Longtail goby— 

his longicaudus (Jenkins and Ever 



Ill) 



HI I I I I IX SOI I HERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 




Figure 6. 



6. a. Inner margin of shoulder girdle without 

dermal flaps - _ - — 7 

b. Inner margin of shoulder girdle with 1 to 3 
dermal flaps (Fig. 6) _ 11 

7. a. Maxillary not extending beyond the posterior 

margin of orbit (Fig. 7a) 9 

b. Maxillary extending beyond the posterior 
margin of orbit (Fig. 7b) 8 




9. 




Figure 7. 

a. Dorsal V to VI-I, 8 to 13; anal fin elements 
9 to 14 Long- 
jaw mudsucker — Gillichthys mirabilis Cooper. 

b. Dorsal IV to V-O to I, 14 to 17; anal fin 

elements 15 to 18 Arrow 

goby — Clevelandia ios (Jordan and Gilbert). 

a. Anal fin elements 13 or less 10 

b. Anal fin elements 14 or more _ _ Blue- 
banded goby — Lythrypnus dalli (Gilbert). 



10. a. Anal I, 11(10 to 12); head and sides with- 
out crossbars encircling posterior part of 



body Black- 

eye goby — Coryphopterus nicholsi (Bean), 
b. Anal I, 8(7 to 9); head and sides with 15 
crossbars encircling posterior part of body 
_ Zebra goby — Lythrypnus zebra (Gilbert). 

11. a. Anal fin elements 13 or more _ 12 
b. Anal fin elements 12 or less ... Tidewater 

goby — Eucyclogobius new berry i ( Girai d ) . 

12. a. Maxillary not prolonged beyond the posterior 

margin of orbit 13 

b. Maxillary much prolonged beyond the pos- 
terior margin of orbit Shadow goby — 

Quietula y-cauda (Jenkins and Evermann). 

13. a. Dorsal spines V; conspicuous blue-black 

metallic spot on opercle Cheekspot goby — 
llypnus gilberli (Eigenmann and Eigenmann). 
b. Dorsal spines VI or VII; no conspicuous 
blue-black metallic spot on opercle .. 
Bay goby — Lepidogobius lepidus (Girard). 



COMMENT 

Provisional species distributions given below were 
obtained from the card catalogues for fish col- 
lections at the Natural History Museum of Los 
Angeles County and California State University, 
Long Beach. Other sources included Barnhart 
(1936) and Clemens and Wilby (1961). 

Tridentiger trigonocephalus. — Color variable. A 
distinctive black band followed by a pale yellow or 
white band encircling the base of the pectoral fins. 
Maxillary short not reaching to the middle of the 
eye. Habitat: sloughs and estuaries. Range: San 
Francisco Bay, California (L. Dempster, pers. 
comm.). 

Lethops connectens. — Dorsal fins almost connected. 
Eye is functional in young, degenerate in adults. 
Specialized folds and flaps on head are apparent 
even in specimens less than 40 mm standard length. 
Habitat: tide pools. Range: Point Fermin to Carmel, 
California. 

Typhlogobius californiensis. — Head covered with 
numerous folds and flaps. Eyes rudimentary in adults. 
Respiration almost entirely through pink-colored 
integument. Arrangement of free neuromast organs 
of the cephalic-lateralis system suggests that the 
Blind goby is not a simple derivative of the Half- 
blind goby. Adults live in pairs in the burrows of the 
Ghost shrimp Callianassa af finis. Habitat: rocky 
intertidal. Range: Cerros Island. Baja California to 
Point Vincente. California. 



l')72 



RESEARCH Noll \ 






Acanthogobius flavimanus. Color variable. Four- 
teen soft rays in dorsal Tin with eight tlnsk y spols, 
each somewhat larger than ilic eye-diameter, ar- 
ranged in a nearly evenly-spaced scries down each 
side, the first three being concealed beneath 'he 
pectoral fin; the lasl forming a more prominent 
spot at the base of the caudal fin ( lirillan. Alhrechl. 
and Hopkirk, 1963). Habitat: lagoons and estuaries 
ascending rivers to fresh water. Range: l.lklioin 
Slough to Tomales Hay, California (I.. Dempster, 
pers. comni.). The range of this introduced species 
is expanding. 

Gobionellus longicaudus. — Distinctive dark brown 
blotch on each shoulder above the origin of pectoral 
fin. A row of five dark blotches extending to the 
base of tail on both sides of body. Not closely 
related to other Calilornian gobies at least on the 
basis of the cephalic-lateralis system. Habitat: la- 
goons and estuaries. Range: San Diego Bay, Cali- 
fornia to Guayaquil, F.quador. 

Gillichthys mirabilis. — Maxillary extending pos- 
teriorly to base of pectorals in adult males. Ca- 
pable of remaining alive out of water for several 
days, breathing aerially by means of heavily vas- 
cularized buccopharynx. Often found burrowed in 
the banks of mudflats at a level exposed by low tides. 
Habitat: channels of tidal mudflats and lagoons at 
the mouths of dry rivers. Range: Bahia Magdalena. 
Baja California to Tomales Bay. California; north- 
ern Gulf of California southward to Mulege on 
west side and Bahia Agiabampo on the east of Baja 
California (Barlow, 1961 ). Introduced into Salton 
Sea, California. 

Clevelandia ios. — Capable of making rapid color 
changes to match varying substrates and light in- 
tensities. Black band on anal fin of mature males. 
During breeding season, both sexes develop yellow 
pigments on the ventral surfaces. At low tide or 
when threatened. Arrow gobies retreat to burrows of 
echiuroid worms Urechis caupo, Ghost shrimp Cal- 
lianassa californiensis, and Blue mud shrimp Upogebia 
pugettensis. Habitat: channels of tidal mudflats 
and estuaries. Range: Baja California to Vancouver 
Island, British Columbia. 

Lythrypnus dalli. — First dorsal spines greatly ex- 
tended. Colored orange-red with four to six dorso- 
ventrally oriented blue bands. Lives in crevices 
on the sides or upper surfaces of reefs, frequently 
clinging to the rocks with its sucker. Habitat: reefs 
down to a depth of 100 m, commonly between 3 
and 30 m. Range: northern Gulf of California: 
Cedros Island, Baja California to Mono Bay, Cali- 
fornia. I believe two specimens of this species, 
collected 10 November 1962 in Morro Bay and 
deposited in the fish collection of California State 



( fniversily, I ong Beai h (CSC I B I 
:i northern rangi 

Coryphopterus nichol.il Pale ycllo 
a distinctive black margin on Hie fn. I dorsal tin 
Large black eyes. A fleshy cresl on lop id I 

tends posteriorly from behind eyi 
first dorsal fin. Habitat: commonlj found bclwcc 
5 and 50 m, rcsiing on sand bottom lo "■ tand 
rock interfaces; i il has been captured al depths 
greater than 700 m. I his gobi seldom venturing 
further than a lew meters from protci I 
Range: San Martin Island. Baja California fl.bcri 
and Turner, 1962) to the Queen ' harlotlc Islands. 

British Columbia, 

Lythrypnus zebra. — Cherry red with 15 blue bars 
oriented dorsoventrally. Bars alternating with a thin 
blue line along the length of Ihe body. Habitat: 
caves and under rocks or well within deep crevices: 
commonly found to depths of 30 m. Range: Cedros 
Island. Baja California to Ventura County, Cali- 
fornia. 

Eucyclogobius newberryi. — Maxillary reaching to 
or slightly beyond the posterior margin of orbit. 
Olivaceous, with dorsals mottled. Live specimens 
quite transparent showing much of the internal organs. 
Habitat: shallow protected bays, estuaries, and clear 
freshwater streams which flow into the sea. Range: 
San Luis Obispo. Santa Barbara and Marin counties, 
California. 

Quietula y-cauda. — The length of the maxillarv 
increases with age and may not reach to posterior 
margin of orbit in specimens less than 30 mm 
standard length. Color variable. In young, black 
streak traversing dorsal portion of opercle at an angle, 
becoming indistinct in adults. Lower lip black. 
Black band on anal fin of mature males. A row 
of nine or ten dark blotches along the sides, the one 
at the base of the tail often shaped like the letter 
"Y". Enters burrows of invertebrates during low 
tide or when threatened. Habitat: channels of tidal 
mudflats. Range: Lower California to Morro Bay. 
California: northern Gulf of California. Baja Cali- 
fornia. 

llypnus gilberti. — Maxillary extending to below- 
middle of the eye. Conspicuous metallic blue-black 
spot on opercle. Habitat: shallow ba\s and channels 
of tidal mudflats. Range: northern Gulf of Cali- 
fornia: Bahia Magdalena. Baja California to Tomales 
Bay. California (L. Dempster, pers. coram.). 

Lepidogobius lepidus. — Distinctive black on ventrs 
surface of head and tips of all fins. Pale i 
green, with rust colored blotches on sides. ! 
Muddy bottoms to 100 m depth. 
California to Vancouver Island. B 



//: 



/;( /// //\ sin llll.l<\ CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



ACKNOWLEDGMENTS 

I wish to ill. ink Camm Swifl and Robert J. Laven- 
berg for their assistance and permission to examine 
the collections of the Natural History Museum of 
Los Angeles County, Dan Odenweller and Peter 
Haaker, California Department of Fish and Game. 
for obtaining data on the introduced Oriental gobies, 
and I ilian Dempster. California Academy of Sciences. 
San Francisco for distributional information and 
specimens of Tridentiger trigonocephaly. 



Clothier. C. R. 1950. A key to some Southern 

California fishes based on vertebral characters. 

California Dept. Fish and Game, Fish Bull., 
(79):l-83. 

Ebert. E. E., and C. H. Turner. 1962. The nesting 
behaviour, eggs and larvae of the Bluespot goby. 
California Fish and Game, 48(4) :249-252. 

Fowler, H. W. 1923. Records of West Coast Fishes. 
Proc. Acad. Nat. Sci. Philadelphia, 75:239. 



LITERATURE CITED 

Barlow. G. W. 1961. Gobies of the Genus gillichtliy s, 
with comments on the sensory canals as a 
taxonomic tool. Copeia. 1961 (4) :423-437. 

Barnhart. P. S. 1936. Marine Fishes of Southern 
California. Univ. California Press, Berkeley, 
209 pp. 

Brittan. M. R., A. B. Albrecht. and J. B. Hopkirk. 
1963. An Oriental goby collected in the San 
Joaquin River delta near Stockton, California. 
California Fish and Game. 49(4) : 302-304. 

Clemens. W. A., and G. V. Wilby. 1961. Fishes of 
the Pacific coast of Canada. Fish. Res. Bd. 
Canada Bull., 68(2nd ed.), 443 pp. 



Jordan. D. S., and B. W. Evermann. 1896 (Reprint 
ed. 1963). The Fishes of North and Middle 
America. Bull. U.S. Nat. Mus., (47), 3313 pp. 

Norman. J. R. 1957. A draft synopsis of the orders 
and genera of recent fishes and fish like verte- 
brates. Unpublished photo offset copies distrib- 
uted by British Mus. (Nat. Hist). 






Ulrey, A. B., and P. O. Greely. 1928. A list of 
the Marine Fishes (Teleostei) of Southern Cali- 
fornia with their distribution. Bull. So. California 
Acad. Sci., 27:1-53. 

Craig K. Macdonald, Dept. Biology, California 
State University, Long Beach, California 90801. 

Accepted for publication February 16, 1972. 



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CONTENTS 

John Adams Comstock, 1883-1970. By Lloyd M. Martin 57 

Bibliography of John Adams Comstock, 1883-1970. By Deborah Birnie and James Dale Smith .... 60 

Three new sympatric Pleocoma from the southern Sierra Nevada Mountains of California (Coleop- 
tera : Scarabaeidae) . By Frank T. Hovore 69 

A review of Eucy Hits Horn (Coleoptera: Curculionidae, Brachyrhininae, Peritclini). By Frank W. 
Pebue and Elbert L. Sleeper 80 

Two new species of North American flat bugs. By Nicholas A. Kormilev 91 

A new species of Ambush bug from Arizona. By Nicholas A. Kormilev 93 

The ascidians Styela barnharti, S. plicata, S. clava, and S. montereyensis in Californian waters. By 
Donald P. Abbott and Jeffrey V. Johnson 95 

RESEARCH NOTES 

A new species of Pugmts from Cocos-Keeling Islands, Indian Ocean (Gastropoda). By 

Barry Roth 106 

A notophycid polychaete from California. By Kristian Fauchald and Bruce W. Belman 107 

A key to the fishes of the family Gobiidae (Teleostomi) of California. By Craig K. Macdonald . . 108 



cover: Larva of Adelocephala heiligbrodti hubbardi. Reproduced from a painting by Dr. John Adams Comstock. 



OUTHRRN CALIFORNIA ACADEMY OF S < II N < IS 



Volume 71 



brarY 

JAN 22 W\ 



BULLETIN 



Number 3 




BCAS-A71(3) 113-164 (1972) 



November 1°72 



Southern California Academy of Sciences 

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90007. 



BULLETIN OF THE SOUTHERN CALIFOR 

AC A DI'M Y ()l ; SC I liNC] S 



Volume 71 



December 30, 1972 



Ni Mi". i R ! 



ORNITHOPHILY AND EXTRAFLORAL COLOR PATTERNS IN COLUM i I 
FLORIDA MORION (GESNERIACEAE) 

C. Eugene Jones, Jr. 1 and Pat Vickers Ru h 



Abstract: Hummingbirds, such as the Little Hermit (Phaethornis longuemareus) and the 
Green-Crowned Brilliant (Heliodoxa jacula) apparentlj are attracted to the hiddi n Hi 
Columned f loridd Morton (Gesneriaceae) by two conspicuous red spots located on the upper 
surface near the apex of the large leaves. This extrafloral attraction mechanism, in which the 
attracting unit, or leaf, does not simply simulate a portion of the flower, seems to be a new 
and unique type of hummingbird pollination system. 



Most species of Columned (Gesneriaceae) possess 
a group of characteristics which tends to delineate 
ornithophilous plants (Motley, 1966; 1971). Al- 
though most hummingbird-pollinated species have 
bright red flowers, (Faegri and van der Pijl, 1966; 
Grant and Grant, 1968; Meeuse, 1961; Percival, 
1965), some species of Cohmmea lack prominent 
red flowers, and have small yellowish-orange 
blooms completely concealed by large leaves with 
red markings near the apices. We suggest that the 
red color patterns on the leaves serve as stimuli 
which increase the probability of visitation by 
hummingbirds. 

Leaf and flower size, as well as flower color 
and the presence or absence of foliar color pat- 
terns, are quite variable in the genus Columnca. 
The opposite leaves range from small (less than 
1 cm long) and isophyllous to rather large (40 
cm long) and anisophyllous. A detailed analysis 
of the nature of the extrafloral color patterns in 
the genus Cohmmea, which is presently being 
completed, has revealed that, in general, species 
with small, isophyllous leaves tend to have large, 
conspicuous red flowers and no foliar pattern, 
whereas species with larger, anisophyllous leaves 
tend to have small yellowish-orange flowers con- 
cealed under the larger leaf. This leaf usually has 
a conspicuous red color pattern near its apex. The 
fact that flower size and color is related to leaf 
size and the presence or absence of red foliar pat- 
terns seems, in turn, to be directly associated with 
the mode of pollination. 



One species with an unusual foliar pattern. 
Columnca florida Morton, an understory, epi- 
phyte, is conspicuous in the flora of the Prcmon- 
tane Wet Forest (Holdridge, 1967) ol southeast- 
ern Costa Rica. The leaves of this species .ire 
strongly anisophyllous; the larger leaf may be 40 
cm long and 1 1 cm wide, whereas the smaller is 
less than I cm long. The larger leaf complete!) 
conceals the relatively small (about \'j-2 cm 
long), yellowish-orange axillan flowers (Fig. la). 
A large blotch of red pigment is present on the 
lower (abaxial) surface near the apex of the leaf. 
but during most of the year the upper (adaxial I 
surface is a uniform dark green. Prior to the on- 
set of flowering two conspicuous red spots (cover 
illustration) appear on the upper surface. Since 
C. florida is pollinated by hummingbirds, these 
spots may serve as flags advertising the concealed 
flowers to these birds. 



METHODS 

Field observations on hummingbird visitation of 
C. florida were conducted from 30 July to 1 Au- 
gust 1970, and from 31 January to 1 February 
1972 at Finca Las Cruces. near San Vito, Puntare- 
nas. Costa Rica. Visitations by pollinators were 



1 Dept. Biology. California State University, Ful 
ton, California 926?4. 

- Dept. of Vertebrate Paleontology 
seum of Natural History. New x . 

1 0o:4. 



113 



IN 



m in 1 1\ SOI I in A'\ ( 



ALIFORNIA ACADEMY OE SCIEKCES 



VOLUME 71 






Figure 1. Flowers of Columnea florida depicting front and side views (a) and lateral exposed 
views of the protandrous condition with the anthers extended and stigma closed (b) and the 
anthers withdrawn and stigma receptive (c). 



observed primarily during the morning hours, 
from 0530 to 1200, since hummingbird activity 
was greatest during that time. 

In 1970, a survey of C. florida in the area re- 
vealed that flowering had not begun. The plants 
chosen for observation were, therefore, somewhat 
modified to simulate natural morphological char- 
acteristics during the peak flowering period, as 
determined from the Flora of Costa Rica (Stand- 
ley, 1937). On 30 July, two plants were selected 
and a pair of red spots, cut from orange-red sur- 
veyor's tape, was glued onto the leaves in the 
position where they normally appear. Yellow flow- 
ers from Heliconia sp. were trimmed to simulate 
the flower size of C. florida and inserted in the 
axils of the leaves. In 1972, observations were 
made on individuals of C. florida which were in 
bloom so modifications were unnecessary. 



RESULTS AND DISCUSSION 

During the observation period in 1970, only two 
species of hummingbirds, the Little Hermit (Phae- 
thornis longuemareus — eight observations) and 
the Green-Crowned Brilliant (Heiiodoxa jacula — 
four observations) visited C. florida. No other 
pollinators were noted on or near the flowers. 
Two behavioral patterns emerged when the birds 
spotted the conspicuous red pattern on the leaves. 
Two Little Hermits hovered directly above the 
two red spots and after 2-4 seconds dipped 
quickly under the large leaves and approached the 
axillary flowers. At this point the bill was inserted 
into the "dummy" flowers and the visitation com- 
pleted. All other birds of both species followed a 
similar behavioral pattern but did not fly directly 
to the flowers. Instead these individuals appeared 
to discern the red cue, but did not perceive the 



1972 



0RNITH0PH1LY AND EXTRAFLORAL <<>l.oi< PATTER 






location of the food source. Consequently, these 
birds spent several seconds inspecting various por- 
tions of the plant before finding the flowers. Ii is 
our suggestion that the former hummingbirds 
were older birds, that had previously experienced 
this exceptional pollination system. The latter 
birds would then be younger, inexperienced 
individuals. 

In 1970, actual visitations by these humming- 
birds occurred only on 1 August 1970. Prior to 
that day 27 hummingbirds were observed in the 
immediate vicinity, but none actually visited the 
experimental plants. Perhaps, hummingbirds re- 
quire some time interval before notice is made of 
signals regarding new food sources in the environ- 
ment. 

The Little Hermit (eight observations) was the 
only hummingbird seen visiting flowering plants 
of C. florida in 1972. The Little Hermit is a rela- 
tively small hummingbird, measuring only 3%-4 
inches from bill tip to tip of the tail. Its bill is 
slightly curved (Cover illustration) and varies 
from 2-2.5 cm in length as compared to the floral 
tube of C. florida which ranges from 2.5-3 cm 
long. 

In the San Vito area of Costa Rica, C. florida 
flowers from late November until April (R. Wil- 
son, pers. comm.). The flowers are protandrous. 
The dehiscing anthers are extended on elongated 
filaments positioned near the orifice of the corolla 
on the first day, followed the next day by a coiling 
of the stamen filaments removing the anthers and 
any remaining pollen from the opening of the 
flowers and from possible contact with visiting 
hummingbirds. Pollen is released into the wedge 
or "V" formed by the connation of the lateral 
axes of the four anthers. (Fig. lb). Although the 
stigmatic surface also is positioned near the en- 
trance of the flower, it is receptive only on the 
second day, thus decreasing the opportunity for 
self-pollination. When receptive, the stigmatic sur- 
faces also form a "V" (Fig. lc). The angle of the 
"V" formed by the connate anthers and the stig- 
matic surfaces is approximately the same as the 
angle of the upper bill of the Little Hermit. Upon 
insertion of the bill into the flower opening, pollen 
is deposited on the upper bill and transported from 
flower to flower in that way. 

As Grant (1950) and Grant and Grant (1968) 
pointed out, flowers pollinated by hummingbirds 
usually have some means of ovule protection. In 
the case of C. florida it appears that the "V- 
shaped" nature of the connate anthers and the 
stigmatic surfaces serve to direct the bill awav 



i mm the Btiperioi i >■ ary and i > with ihc 

nectar which collects in •> ilighl depn in On; 

lower portion <>i the corolla. Since On.- n 
are horizontal in the ground na tai i olio U in On: 
depression in the corolla by gravity. An cxamina 
lion of 25 flowers from 10 separate plants, re- 
vealed no damage i<> Oil- ovaries from the probing 
bills oi ihc Little Hermit hummingbirds. 

Behavioral responses ol ihc Little Hermits were 
similar in 1972 to those noted in 1970. exo . 
birds flew directly to the flowers ol ( . florida 
without first hovering above the red spots. This 
suggests that these hummingbirds may have had 
their feeding sites memorized and no longer relied 
on the extrafloral red spots as orientation cues. 
Thus, it would appear that the red spots on the 
leaves of C. florida are of prime importance as 
initial visual signals to hummingbirds denoting the 
presence of a new food source in the immediate 
area, but lose at least some of their importance as 
visual orientation cues after flowering begins. 

Position and shape of the red spots on the leaf 
is of interest. Because this species is epiphytic on 
the trunks of large trees, the plants extend hori- 
zontally about 2 to 3 feet. As a result the most 
noticeable portion of the large nodding leaves is 
the upper surface near the tip. hence the location 
of visual cues there should be advantageous. Con- 
centration of the red pigment into two distinct 
spots, rather than a single larger, red blotch, prob- 
ably increases the effectiveness of the visual cue. 
since any consistently repeated pattern, associated 
with a food source, would be a more effective 
stimulus than randomly associated red blotches. 
The latter blotches would only provide a red flash, 
in the immediate environment, which might re- 
semble those of other origins, and which would 
not predictably indicate the presence of a food 
source. 

Our field observations of hummingbird behavior 
lead us to suggest that the red foliar spots are 
indeed functioning as visual cues, attracting hum- 
mingbirds, such as the Little Hermit and the 
Green-Crowned Brilliant, to the flowers of C. 
florida, thus insuring successful pollination. Al- 
though the attraction of hummingbirds to flowers 
by extrafloral mechanisms is well documented ii 
the case of Castilleja and others. (Faegri and van 
der Pijl, 1966: Grant and Grant. 196S). the mech- 
anism for C. florida is unique in that the leaf does 
not simulate a portion of the flower, but : 
simply acts as a visual signal dene 
ence of nectar in the immediate vicin 



116 



IU III ll\ SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



ACKNOWLEDGMENTS 

This work was supported by a Sigma Xi Grant-in-Aid 
of Research and by a grant from California State Uni- 
versity. Fullerton Foundation. We thank U. H. Bralt- 
strom, T. L. Hanes. and C. B. Heiscr for their com- 
ments on the manuscript. The first part of this study 
was conducted during tenure as faculty member 
(Jones) and participant (Rich), in the Organization 
for Tropical Studies program in 1970. Special thanks 
goes to Susan E. Payne who did the cover drawing 
and figure 1 and to James F. Jackson and Stephen L. 
Buchmann for their field assistance. 

LITERATURE CITED 

Faegri, K., and L. van der Pijl. 1966. The Prin- 
ciples of Pollination Ecology. Pergamon Press, 
London, 248 pp. 

Grant. K. A., and V. Grant. 1968. Hummingbirds 
and Their Flowers. Columbia Univ. Press, New 
York. 115 pp. 



Holdridge. L. R. 1967. Life Zone Ecology. Tropi- 
cal Science Center, San Jose, Costa Rica, Revised 
Ed., 206 pp. 

Meeuse. B. J. D. 1961. The Story of Pollination. 
Ronald Press, New York, 243 pp. 

Morley. B. D. 1966. Columnea and aspects of its 
evolution. Sci. Notes and News, Jamaica, 2: 13— 

14. 

. 1971. A hybrid swarm between two hum- 



mingbird-pollinated species of Columnea (Ges- 
neriaceae) in Jamaica. Bot. J. Linn. Soc, 64: 81— 
96. 

Percival, M. S. 1965. Floral Biology. Pergamon 
Press, London, 243 pp. 

Standley, P. C. 1937. Flora of Costa Rica. Field 
Mus. Nat. Hist., Bot. Ser. Publ. No. 391 and 392, 
Chicago, 1616 pp. 



Grant, V. 1950. The Protection of the ovules of 
flowering plants. Evolution. 4:179-201. 



Accepted for publication June 14, 1972. 



EFFECTS OF VARYING TEMPERATURES AND SALINITIES ON SETTLEMENT, 
GROWTH, AND REPRODUCTION OF THE WOOD-BORING PELECYPOD, 

LYRODUS PEDICELLATUS 

Kevin J. Eckelbarger 1 and Donald J. Reish 2 



Abstract: The larvae and adults of the wood-boring pelecypod, Lyrodus pedicellatus, 
were subjected to various conditions of temperature and reduced salinity under laboratory 
conditions. The effects of reduced salinity on the larvae were studied at 9-11 C, 14—16 C, and 
22-24 C, and for the adults at 14-16 C and 22-24 C. Optimum activity for the larvae was 
found to be above 25% c and from 14 to 24 C. The minimum temperature and salinity levels 
for larval boring were 12-14 C and 20%«. The long term minimum requirements for survival 
of adult L. pedicellatus was near 22-25%„ and about 11 C. Reproduction occurred above 



28.8%,; in the temperature range of 14 to 24 C. 



Lyrodus pedicellatus Quatrefages is one of three 
teredinids present on the Pacific Coast of North 
America. It is abundant in southern California 
and is reported north to San Francisco Bay 
(Miller, 1926). Many of the earlier reports re- 
ferred to L. pedicellatus as Teredo diegensis 
(Turner, 1966). This species is frequently found 
associated with Teredo navalis and the colder 



water shipworm, Bankia setacea Tryon. Both L. 
pedicellatus and B. setacea occur in Los Angeles- 
Long Beach Harbors with the former species the 



1 Marine Science Institute, Northeastern University, 
Nahant, Massachusetts 01908. 

- Dept. Biology, California State University, Long 
Beach, California 90801. 



1972 



EFIECIS <)!■' I l:MI'l:KA I IIKIi AND S A I. INI I Y ON I VI /</< ///,!/' 






more prevalenl (Menzies, Mohr, and Wakeman, 
1963). 

Little data arc available on the tolerances of L. 
pedicellatus from ihe Pacific Coast. Marrows 
(1917) found this species able to temporarily 
withstand salinities as low as \(Y/<< in San Fran- 
cisco Bay. Menzies et al. ( 1 963 ) reported that 
larvae displayed maximal settlement during the 
warmer months of the year in Los Angeles-Long 
Beach Harbors. Roch (1940) determined that 
spawning occurred between 10 and 19 C, boring 
was reduced at 25#o and death occurred below 
20%» in the Adriatic Sea. Edmonson (1942) 
found that the larvae were able to survive 10 
days at 15%» and adults survived 12 days in Yl%„. 
Watson (1957) reported L. pedicellatus in Queens- 
land, Australia only in salinities above 25%c. 

Table 1 reviews the literature on temperature- 
salinity effects on teredinids. Species are arranged 
according to larval type {i.e., oviparous, short or 
long-term larviparous). Oviparous forms are 
those that release their sex products directly into 
sea water where fertilization occurs; whereas, 
larviparous species undergo fertilization in the 
mantle cavity and brood the young to the straight 
hinge stage (short-term) or to the mature pedi- 
veliger (long-term), before releasing them (Tur- 
ner, 1966). 

The majority of the previous studies with L. 
pedicellatus have been based on field observations 
and have largely been based on short-term obser- 
vations. The purpose of this study, therefore, was 
to define under laboratory conditions the tempera- 
ture-salinity tolerances of the different stages, in- 
cluding settlement, growth, and reproduction, in 
the life cycle of this wood-boring pelecypod. 

METHODS 

Adult Lyrodus pedicellatus were obtained from 
small Douglas fir blocks which had been suspended 
from ropes at the surface to a depth of 3 m in the 
Cerritos Channel region of Alamitos Bay, Long 
Beach, California. The wood blocks were scraped 
free of fouling organisms bimonthly. The blocks 
were removed as required but generally they were 
suspended for about 6 months. Wood blocks were 
brought to the laboratory, placed in a gallon jar 
with aerated, filtered sea water, and placed in a 
cold bath at 14-16 C. To obtain larvae, the blocks 
were placed in a container with sea water at room 
temperature. This change in temperature gener- 
ally caused the adult to release pediveliger larvae 
present in its brood chamber. The larvae were 



pipetted from the containci and Ihi 
were returned t<> the cold bath, ["hi I nun, 

ber cil larvae were obtained bj using the wood 
blocks the day they were removed from ihe field 
l)iil they could be used as a furthci "urec ol 

larvae in the luimc with decreasing iuco 

obtaining large numbers. 
Larval Salinity Experiments', An experimcnl 

was conducted lo determine the ellecls ol tem- 
perature and reduced salinity on the swimming, 

boring, and survival ol ihe pediveliger stage. 
Eleven concentrations ol sea water weie used. 
35.0, 31.5, 28.8, 25.2, 21.6, 18.0, 14.4, 10.8, 7.2, 
3.6, and 1.8'.. salinity. Solutions were prepared 
by dilution with appropriate volumes of distilled 
water and the salinity determined (Barnes, 19 ■ 
A single 350 ml finger bowl containing a pre- 
soaked block of Douglas fir measuring 5 / 25 
42 mm, was used at each salinity. Thirty swim- 
ming pediveligers were pipetted into each bowl 
and kept in reduced light at 22-24 C, 14-16 C, 
and 9-11 C. Cultures were examined every (i 
hours and larval swimming, crawling, and boring 
activity including the formation ol the calcareous 
cap was noted. The experiment was conducted 
for 10 days which was well beyond the potential 
period for larval boring. 

Survival and Growth Experiments: The long- 
term effects of reduced concentrations of salinity 
on survival, growth, and reproduction in L. pedi- 
cellatus were conducted utilizing an experimental 
procedure similar to that described above. Indi- 
vidual swimming pediveligers were pipetted into 
petri dishes each containing a wood block and 
normal salinity sea water. Cultures were main- 
tained at room temperature in reduced light. 
Larvae were allowed to bore for 14 days to allow 
sufficient time for the calcareous cap and siphons 
to become well developed. This experiment was 
designed to determine long-term salinity and tem- 
perature effects on adults. Eleven concentrations 
of sea water were prepared as before, namely. 
35.0. 31.5. 28.8. 25.2. 21.6. 18.0. 14.4. 10.8. 7 2, 
3.6, and 1.8& salinity. A wood block containing 
a single, healthy L. pedicellatus from the above 
cultures was placed in each of ten 500 ml stop- 
pered erlenmeyer flasks with 100 ml of sea water 
at each salinity level. The experiment was con- 
ducted at 22-24 C and at 14-16 C giving a total 
of 110 specimens at each temperature. The 9 
C temperature bath was unavailable for this ex- 
periment. Sea water in each flask was replaced 
by a fresh solution even' five days. The e 
ment was terminated at five months a 



118 



BULLETIN SOUTHERN CALIFORMA ACADEMY OF SCIENCES 



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8.2 



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122 



HI III ll\ sor////A'\ CAlllORNIA ACADEMY OF SCIENCES 



VOLUME 71 



IMui 2. A comparison of the survival of Lyrodus 
pedicellaius larvae and adults under reduced salinity 
conditions ai the end of seven days al two temperature 

levels 



Table 3. The effects of temperature and reduced 

salinity on the larval swimming activity of Lyrodus 

pedicellaius 24 hours after release. 















Salinity 
(So) 


Number 


of Swimming L 


irvac* 






4-16°C 




22- 


-24°C 


9-1 rc 




14-16°C 


22-24° C 




a 






I 


% 












Salinity 


1 .11 V.l 




Adult 


Larva) 


Adult 


35.0 


6 




6 


3 


('.. ■) ' 


Surviva 


* Si 


rvival** 


Survival* 


Survival** 


31.5 
28.8 


9 

21 




3 
9 





35.0 


90 




100 


.70 


100 





31.5 


65 




100 


70 


100 


25.2 


18 




3 





2S.S 


70 




100 


50 


100 


21.6 


15 




15 





25.2 


60 




100 


10 


90 


18.0 


6 




3 





21.6 


50 




100 


8 


50 


14.4 


3 










18.0 


50 




100 


3 


4(1 


10.8 













14.4 


35 




100 





40 


7.2 













10.8 







60 





10 


3.6 













7.2 








20 











1.8 













3.6 


* Based upon 30 larvae tested 


at 


each salinity 


level. 


1.8 



























* Based upon 30 larvae tested at each salinity level. 
** Based upon 10 adults tested at each salinity level. 



all living animals were removed from their bur- 
rows. Measurements were made of the burrow and 
pallet length. The mantle cavity and gill filaments 
were examined for the presence of veligers which 
were counted if present. 

RESULTS 

The results of the larval and adult experiments of 
reduced salinity at the two temperatures are sum- 
marized in table 2. These data indicate that the 
adults are much more tolerant of reduced salini- 
ties than larvae and, furthermore, both stages are 
tolerant of reduced salinities at the lower experi- 
mental temperature. The seven-day median toler- 
ance level (TL ni ) values for larvae and adults were 
28.2 and 21.6&., respectively, at 22-24 C and 18.0 
and approximately 9.9'^, respectively, at 14-16 C. 
Three-day TL ni values for larvae were approxi- 
mately 9.4, 11.5, and 18.75^ at 9-11, 14-16, and 
22-24 C, respectively. 

Swimming activity of the pediveligers was found 
to be related to temperature and salinity as shown 
in table 3. Larvae were found swimming 24 hours 
following their release from the adult in salinities 
of 35.0, 18.0, and 14.0%» at the respective tem- 
peratures of 22-24 C, 14-16 C, and 9-11 C. 

Early boring activity of the larvae was reduced 
with decreasing salinity but increased with an in- 
crease in temperature (Table 4). Initial penetra- 
tion of larvae into wood blocks occurred at salini- 
ties of 18.0 and 25.2& at temperatures of 22-24 



and 14—16 C, respectively. Penetration occurred 
only at normal salinity at 9-1 1 C. It should be 
noted, however, that although larvae were able to 
penetrate the surface of the wood, the formation 
of a protective calcareous cap over the opening, 
which occurs under normal conditions, occurred 
only in salinities down to 21.6 and 31.5%c at tem- 
peratures of 22-24 and 14-16 C, respectively. 

The relationship between reduced salinity and 
burrow and pallet lengths of animals raised over 
a five month period at 14-16 and 22-24 C is rep- 
resented in table 5. As the salinity levels de- 
creased, the rate of burrowing and pallet forma- 
tion diminished. At any given salinity, the burrow 
or pallet length was greater at 22-24 than 14-16 C. 

Some adult specimens raised in isolation from 
the pediveliger stage for a five month period con- 
tained larvae at the end of the experiment. These 
data were summarized earlier (Eckelbarger and 
Reish, 19721, but 40, 25, and 11 percent of the 
animals contained normal larvae at 35.0, 31.5, 
and 28.8^ at 14-16 C; whereas, 30 and 20 per- 
cent contained larvae at 35.0 and 31.5#» at 22-24 
C. This was the first report of self-fertilization in 
the family Teredinidae. 

DISCUSSION 

Effects on larvae of Lyrodus pediceUatus: The 
effects of reduced salinity on the survival of the 
larvae of L. pediceUatus were greatly influenced 
by temperature. The mortality was considerably 
higher at all salinity levels as the temperature in- 
creased. While lethal limits for temperature were 
not investigated, the 9-24 C temperature range 



1972 



El'l'ECTS OF T/CMI'IJUTURE AND SAI IM I ) ON I II hl< I I i 






Table 4. The effects of temperature and reduced Table 5. The relationship ol 

salinity on the boring of Lyrodus pedlcellatus larvae length to reduced salinity in I 

at the end of a 10-day period. two tempcratun 



Salinity 






N 


imbor of Boring Lorvai 


+ 




Numbci 


II 




















' nun ) 








MM 




" 


-IPC 


14-16°C 


22-24"C 


S.ilinii y 


..i 
Animals 










I' 


SI, ,,i, 










2 


Q** 


1 9 




35.0 














3 1 .5 









13** 


15** 






II If, < 






28.8 









8 


2** 


35.0 


1 1 


15 25 


20.9 


2.0 (.1 


2.5 


25.2 









4 


4** 


31.5 


8 


13-21 


15.3 


I.K 2.5 


2.1 


21.6 












3* 


28.8 


9 


7-16 


10.5 


1.1 2.6 


1.7 


18.0 












2 


25.2 


5 


5-12 


8.0 


1.0 1.6 


1.3 


14.4 












I) 


21.6 













10.8 



























7.2 



















22-24 


c 






3.6 



























1.8 












o 


35.0 


III 


15-40 


24.6 


2.0^4.5 


2.7 








31.5 


III 


16-30 


23.4 


2.0-3.9 


2.5 


Total 






2 


34 


46 


28.8 
25.2 


8 

4 


8-26 
9-12 


20.0 
10.3 


1.3-2.8 
1.5-1.8 


2.3 
1.6 














* Bused 


ipon 


30 


larvae 


leslcd at each salinity 


level. 














** Calcareous 


caps fornu 


d. 




21.6 


1 


— 


5.0 


— 


1.0 



used in the experiments is close to that encoun- 
tered under local field conditions. 

From the larval salinity experiments, the three- 
day TL m values showed the limiting salinity levels 
to be approximately 9.4, 11.5, and 18.7/ce at 9-11, 
14-16, and 22-24 C, respectively. Death occurred 
within a few hours at 7.2 r ,V and below at all tem- 
perature levels. The three-day TL m values are rep- 
resentative of the normal salinity tolerances of 
larvae since the majority of Lyrodus larvae pene- 
trate wood during this period. Thus, larvae of 
Lyrodus are quite tolerant of low salinities over 
a short period of time, especially during the criti- 
cal first few days after release from the parent. 
Edmonson (1942) reported that larvae of L. pedi- 
cel lotus from Hawaii were able to tolerate 15're 
for 10 days in the laboratory; a result consistent 
with ours. This was the only laboratory investiga- 
tion of the effects of reduced salinity on this spe- 
cies reported until now. 

The effects of temperature and reduced salinity 
on the activity of the larvae demonstrated that 
larvae were able to swim in lower salinities at re- 
duced temperatures. Swimming ceased at expo- 
sure to 10.8 r rr or below at all but the lowest tem- 
perature level. Reduced temperature also ex- 
tended the swimming period from the normal 18 
to 36 hours as reported by Horvath (1951), Isham 
and Tierney (1953), and Lebour (1946). No 
swimming activity was observed at any salinity 
level 24 hours after release from the parent at 22- 
24 C. Some larvae, however, continued swimming 
for five days in the higher salinities at 14-16 C. 



Larvae were also found swimming 13 days after 
release in higher salinities at 9-11 C with a cor- 
responding reduction in swimming period as the 
salinity decreased. A similar shortening of the 
larval swimming period in response to higher tem- 
perature was reported for T. navalis by [mai, 
Hatanaka. and Sato (1950). But Bulatov (1941) 
found 20-30 C the optimum temperature range 
for T. navalis with swimming ceasing at 10-12 C. 
It should be noted that although swimming activ- 
ity in L. pedicellaius was found in this studs to 
increase at lower temperatures, nesvly metamor- 
phosed pediveligers in the crawling stages dis- 
played very little activity compared to those at 
higher temperature levels. 

Temperature and reduced salinity noticeably 
affected the boring of the larvae. Reduced tem- 
perature and salinity restricted boring while higher 
temperatures increased boring and enabled larvae 
to bore even under reduced salinity conditions. 
Boring occurred down to 1S.0'. at 22-24 C but 
did not occur below 35.0',' at 9-11 C. The calci- 
fication of the mound covering the burrow en- 
trance, however, occurred only in salinities down 
to 21.6 and 31.5& at 22-24 and 14—16 C, respec- 
tively. No caps were formed by boring larvae at 
9-11 C. Larvae failing to secrete a protects 
died. It is apparent then that even though it 
penetration of the substrate can be fished 

at 9-1 1 C and in the lower salinity levels a 
16 C and 22-24 C. successful borinp 
more restricted. Lyrodus larvae are ' 



124 



nnii n\ sourm i<\ California academy of sciences 



VOLUME 71 



able to pencil. iic wood and continue growth much 
below 14 C and below a salinity of 21.6',. (Table 

I ). Imai el «/. ( 1950) reported /. navalis capable 
of boring in brackish water at a salinity of W%<> 
bin were able to bore only between 14-26 C. 
Uul.ilov (l l >41) found 10 C or below lethal for 
the larvae of the same species. It would appear 
then that L. pedicellatus is as capable of boring 
under temperature conditions as low as T. navalis 
but is considerably more restricted by salinity 
conditions than the latter species. 

The cold water species, Bankia setacea, is ap- 
parently more hardy than either Teredo or Lyrodas 
for it has been reported to bore in salinities of 9% 
at a temperature of 7 C (Trussell, 1967). Black 
and Elsey (1948) also found this species settling 
in greatest numbers when the surface salinity in 
British Columbia waters ranged from 9-23%?. 

Salinity alone is probably not a significant fac- 
tor for the survival of Lyrodus larvae in southern 
California since the salinity is seldom altered from 
that of normal sea water except directly following 
a heavy rain (Stone and Reish, 1965). These 
salinity changes are of short duration and usually 
restricted to surface waters. Since larval settle- 
ment generally increases with increase in water 
depth (Menzies el ah, 1963), the effects of these 
salinity changes in southern California are prob- 
ably of minor importance. Temperature, however, 
does fluctuate throughout the year and appears to 
place significant limitations on boring activity. In 
this study, for example, the number of larvae bor- 
ing decreased with decrease in temperature with 
a total of 46, 34, and 2 larvae boring at 22-24, 
14-16, and 9-11 C, respectively. Reduced tem- 
perature also delayed the boring process. Boring 
began within 24 hours at 22-24 C but was delayed 
for 48 hours at 14-16 C and for five days at 9- 

I I C. These findings differ from those of Horvath 
(1951) who reported L. pedicellatus unable to 
bore 24 hours after release from the parent. Coe 
(1941) stated that boring in this species can occur 
up to two weeks after release but observations 
in this investigation have never shown boring to 
occur after such an extended period. 

Settlement of Lyrodus larvae appears to be a 
year-round phenomenon in southern California 
waters although seasonal differences can be ob- 
served which probably reflect temperature fluc- 
tuations. Horvath (1951) indicated that Lyrodus 
larvae were released throughout the year in Los 
Angeles-Long Beach Harbors but that survival and 
development did not occur during the cold months 
of December through March. Menzies et al. 



(1963) also indicated that in the same harbor, 
Lyrodus larvae settled during all seasons of the 
year with maximum intensity occurring during the 
warmer months of the year. The authors noted, 
however, that boring larvae failed to mature when 
temperatures reached a low of 12 C. Greenfield 
(1952) also reported L. pedicellatus actively 
breeding and boring throughout the year at 
Miami, Florida; an area approximating hydro- 
graphical conditions in southern California. 

These field studies tend to agree with the find- 
ings of our laboratory investigation with heaviest 
settlement and boring occurring when tempera- 
tures reached 22-24 C. At the lowest tempera- 
tures tested, however, larval mortality was high, 
with few larvae successfully penetrating the sub- 
strate. Utilizing the temperature data of Moore 
and Reish (1969) in Alamitos Bay, California, 
the annual temperature ranged from about 13 to 
24 C, with the warmer peaks occurring during the 
months of March through August. These data 
probably explain the maximum infestation ob- 
served during the period from April to September 
of wood blocks suspended bimonthly in this study, 
from August 1967 to February 1969. 

Effects on adults of Lyrodus pedicellatus: Ex- 
periments on the effects of temperature and re- 
duced salinity on the adults of L. pedicellatus dis- 
closed that the adults were more tolerant to 
environmental stress than larvae. A comparison 
of the seven-day TL m values at 22-24 C (Table 
2) showed that larvae and adults displayed values 
of 28.8 and 2\.6%o, respectively; whereas, these 
values dropped to 18.0 and 9.9?«, respectively, at 
14-16 C. This greater salinity tolerance in adults 
is probably best explained on the basis of both 
physical protection from the environment and de- 
velopmental stage. Animals used in the experi- 
ment had been boring for 14 days and all had 
formed caps and pallets. As is probably true in 
all shipworms, the sealing of their burrow en- 
trances with their pallets provided an effective 
deterrent to osmotic stress. 

White (1929b) discussed the protection afforded 
teredinids by their burrow and pallet mechanism. 
He found Bankia setacea less tolerant of reduced 
salinities when directly exposed to sea water than 
when in intact burrows with death occurring in 
one hour in 1.5%« and within 12 hours in 13.7%« 
at 15.5 to 19.5 C. Roch (1940) reported 25%» 
the minimum salinity required for normal burrow 
activity and 20%» lethal for L. pedicellatus. 

During the course of the reduced salinity ex- 
periments with Lyrodus, disintegration of the pal- 



i<m 



lil'MC'l'S ()!■ TEMPERATURE AND SALINITY on i iu.dk in | 






lets and protective burrow caps occurred in speci- 
mens at low salinities. The formation of thinner 
shells and other calcareous structures in reduced 
salinities was reported in other molluscs hy New- 
combe and Kcsslcr (1936). It is probable that the 
inability of Lyrodus to maintain these protective 
structures accelerated death due to the direct ex- 
posure of the animals to the external medium. 

Temperature and salinity greatly affected 
growth in L. pedicellatus over a live month period. 
Total growth was determined by measuring the 
burrow ami pallet lengths since neither varies with 
the spawning condition of the animal (Quayle, 
1959). The average burrow lengths of experi- 
mental animals decreased with decrease in tem- 
perature and salinity. Reduced salinity presum- 
ably reduced growth by halting or slowing boring 
and feeding. Roch (1940) found 25%a sufficient 
to reduce boring activity in L. pedicellatus. M'- 
Gonigle (1926) reported no boring by T. navalis 
adults at 3.5%« or below, and Blum (1922) ob- 
served that T. navalis did not produce fecal pellets 
at salinities of 4.0 to 5.0#r. Shipworm growth 
then is probably halted because the animal seals 
its burrow and remains relatively inactive until 
more favorable conditions return. Similarly, the 
bivalve Mytilus edulis remained closed and failed 
to produce byssal threads at reduced salinities 
(Reish and Ayres, 1968). It is also possible that 
under reduced salinity conditions the rate and 
efficiency of food metabolism is reduced as Kinne 
(1963) reported for M. edulis. 

The effects of temperature on the growth rate 
of L. pedicellatus were readily apparent from this 
study with the average burrow lengths of animals 
raised at 22-24 C greater than those at the same 
salinity levels raised at 14-16 C. Isham, Smith, 
and Springer (1951), investigating L. pedicellatus, 
M'Gonigle (1926) and Needier and Needier 
(1940), both working with T. navalis, and Quayle 
(1959) studying B. setacca, all reported the maxi- 
mum growth rate in the field corresponding to 
maximum seasonal temperature. 



ITII.KA II RE Mill; 

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Barrows, A. I 1917. An unusual extension ol the 
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Blum, H. F. 1922. On the effect of low salinity on 
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Bulatov, G. A. 1941. Response of the larvae of the 
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ACKNOWLEDGMENTS 

The authors wish to express their gratitude to Ruth 
D. Turner, Museum of Comparative Zoology. Har- 
vard University, for verifying the identification of 
specimens used in this study and for her critical re- 
view of the manuscript and many helpful suggestions. 
Thanks also to John L. Culliney, also of Harvard, 
for his suggestions and use of unpublished data. 



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126 



111 III ll\ SOI'TIIEI<\ CAIIEOHMA ACADEMY OF SCIENCES 



VOLUME 71 



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1972 



EFFECTS OF TEMPERATURE AND SALINI1 Y ON I PI D/< ELLATl 






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loguc oi iIm i. i . Jjnidac I Moll 

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no. 6, 28 pp. 

While. F. D, 1929a. Studies on marine wood borers. 
III. A note on the breeding season of Bankia 
(Xylotrya) setacea in Departure Bay. B. C. Con- 
trib. Canad. Biol. Fish., (n.s.) 4:19-25. 



— . 1929b. Studies on marine wood borers. 
II. Effect of experimental variations in salinity 
and hydrogen ion concentration upon the wood 
borers of the Pacific Coast of Canada. Contrib. 
Canad. Biol. Fish., (n.s.) 4:11-18. 



Accepted for publication May 22. 1972. 



TWO NEW SPECIES OF POLYCHAETOUS ANNELID WORMS FROM BAFFIN BAY 

AND THE DAVIS STRAIT 

James A. Blake 1 



Abstract: Two new species of polychaeta of the families Lumbrineridae and Scalibreg- 
midae from Baffin Bay and the Davis Strait are described. Both species occur in relatively 
deep water and were obtained by qualitative dredge hauls. A table emphasizing some taxo- 
nomic characteristics of some scalibregmid genera is presented. 



During August 1968, the author participated on 
the shakedown cruise of the National Science 
Foundation's new polar research vessel, the R V 
HERO to the waters of the Labrador Sea. Baffin 
Bay, and the Davis Strait. The present paper de- 
scribes two new species of polychaeta in the fami- 
lies Lumbrineridae and Scalibregmidae. The type 
material is deposited in the United States National 



Museum, Washington. D. C. Complete results of 
the cruise are being compiled and will be pub- 
lished in a subsequent paper. 

This is contribution Number 2S from the Pacific 
Marine Station and was supported by N s 



1 Pacific Marine Station. University 
Dillon Beach. California 94929. 



128 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



Grant GA-1405. Figure 2 was prepared by Floy 
E. MacMillan-Zittin. 



Family I.umbrineridac 

Lumbrineris fauchaldi, new species 

Figure 1 

Material examined: Davis Strait (lat. 66° 29' N-long. 
57 26'W), HKRO Station 28D, collected 17 August 
1968. dredged in 580-610 m on a bottom of soft 
mud (7 specimens. TYPE). 

Description: The material consists of 7 specimens, 
only one of which is complete (Holotype). The holo- 
type is coiled, contains approximately 80 setigerous 
segments and measures about 10 mm in length. The 
longest anterior fragments measure 12 mm (44 seti- 
gers) and 15 mm (65 setigers) in length. The speci- 
mens are light tan in color with no body pigmentation. 

The prostomium is conical and slightly longer than 
wide (Fig. la). There are no visible nuchal organs or 
eyes. The two peristomial segments are achaetous 
and similar in size. There are paired anterior pro- 
longations of the first peristomial segment. 

The body is rather slender, tapering at the posterior 
end and terminating in two anal cirri (Fig. lb). One 
specimen had three anal cirri (Fig. lc). 

Setigers throughout are wider than long. The para- 
podia contain short rounded presetal and postsetal 
lobes. Setigers 1-10 contain fascicles of limbate setae. 
Thereafter, the dorsalmost setae in the fascicle exhibit 
blunted ends. These blunt setae grade into hooded 
hooks over succeeding setigers. The fully developed 
hooks do not occur until setigers 25-35. Some of the 
limbate setae of setigers 20-50 contain greatly enlon- 



gated capillary tips (Fig. Id). The hooded hooks 
completely replace the limbate setae in the last % 
of the body. Hooded hooks arc multidentate with 
about 10 small teeth, of which the lowermost tooth 
is only slightly thicker than the rest (Fig. le). The 
acicula are yellow, straight, and may number two 
per parapodium in posterior setigers. 

The pharyngeal apparatus is as follows (Fig. If): 
the maxillary carriers are short and sharply pointed, 
with a weak lateral notch; maxilla I has a weakly 
curved tip with no distinct teeth along the inner mar- 
gin; each maxilla II has five teeth on the inner margin 
and a posteriorly directed lateral prolongation; each 
maxilla III and IV has one tooth; the mandibles were 
not observed. 

Distribution: Davis Strait, depth ranging from 580- 
610 m. 

Remarks: Lumbrineris fauchaldi belongs to 
species group Il.b.l. (Fauchald, 1970), in which 
there are simple hooded hooks, which are multi- 
dentate in posterior setigers; maxilla III has one 
tooth. Approximately 35 species have been as- 
signed to this group by Fauchald ( 1970) . Of these, 
six are known to have limbate setae in median 
setigers with prolonged hair-like tips. Each of 
these species occurs in deep water. These species 
are L. abyssorum (Mcintosh, 1885), L. ehlersii 
tenuisetis (Mcintosh, 1885), L. longensis Hart- 
man, 1960, L. moorei Hartman, 1942, L. neo- 
zealaniae (Mcintosh, 1885), and L. punctata 
(Mcintosh. 1885). 

Lumbrineris abyssorum was described from 
2,225 fathoms off western South America from 
fragmentary specimens in which the hooks had 



Table 1. Some taxonomic characteristics of five genera of the Scalibregmidae having 

anterior acicular setae. 



Genera 


Branchiae 


Parapodial Processes 


Acicular Setae 


Reference 


Asclerocheilus 


Absent 


Absent 


Present in both 


Ashworth, 1901 


Ashworth, 1901 






rami of Setigers 1-3 


Fauvel, 1927 
Day, 1967 


Cryptosclerocheilus 


Present 


Absent 


Limited to both 


This paper 


new genus 


Setigers 
2-5 




rami of Setiger 2 




Parasclerocheilus 


Present 


Ventral cirrus 


Limited to notopodia 


Fauvel, 1928 


Fauvel. 1928 


Setigers 
2-7 


on posterior 
setigers 


of Setigers 1-4 


Day, 1961 


Sclerobregma 


Absent 


Dorsal and ventral 


Limited to both 


Hartman, 1965 


Hartman, 1965 




cirri in posterior 
setigers 


rami of Setiger 1 




Sclerocheilus 


Absent 


Ventral cirrus on 


Present in both rami 


Ashworth, 1901 


Grube, 1863 




posterior setigers 


of Setigers 1-3 


Fauvel, 1927 



1972 



NEW SPECIES OF POLYCHAETOUS ANNELID 







Figure 1. Lumbrineris fauchaldi, new species: a. anterior end in dorsal view: b. posterior end 
in dorsal view; c, posterior end in dorsal view; d. setiger 35 in anterior view: e. hooded hook 
from mid-body setiger: f. pharyngeal apparatus; a-c all to the same scale. 



been lost or broken: L. ehlersii tenuisetis from off 
northeastern North America in 1,340 fathoms has 
five teeth on each side of maxilla II and two teeth 
on maxilla III; L. longensis has prolonged pre- 
and postsetal lobes in posterior setigers, in addi- 
tion to black acicula; L. moorei has hooks from 



approximately setiger 25. but there are no transi- 
tional setae as in L. fauchaldi: L. neozealanic 
apparently described from a group of species and 
type material needs to be reexamined: and L. 
punctata has only two teeth on the left - 
maxilla II and three on the rieht. 



130 



i:illlll\ \<>l lllli;\( \III()R\IA l( .HUM) (II S(//\(7S VOLUME 71 





mmmm 





llf§! 



^- 




pg^gj 




,0.04 mm 



1972 



NliW SPECIES <>l POLYCHAETOl S I ELID 






It is a pleasure to name this species for l)i 
Kristian Fauchald of the Allan Hancock Founda- 
tion, University of Southern California, in recog- 
nition of his monographic work on the super- 
family Eunicea. 

Family Scalibregmidae 
Cryptosclerocheilus, new genus 

Type species'. Cryptosclerocheilus baffinensis, new 

species. 

Diagnosis'. Body fusiform and elongated. I'rosto- 
mium T-shaped, with two long frontal lohes. Without 
eyes. Peristomium achaetous. Proboscis unarmed. 
Branchiae limited to anterior segments. Parapodial 
lohes reduced lo short elevations throughout the body; 
no cirriform processes present in cither anterior or 
posterior regions. Lateral organs not evident. Acicular 
setae delicate and limited to setiger 2 where they oc- 
cur in both the neuropodium and notopodium. Fur- 
cate setae hegin on setiger 3 and continue on succeed- 
ing segments. Pygidium a simple lobed ring. 

Remarks: Cryptosclerocheilus belongs to the 
group of genera having anterior acicular setae and 
a reduction in parapodial lobes and processes. The 
relationship of Cryptosclerocheilus with related 
genera is presented in table 1. 

Cryptosclerocheilus baffinensis, new species 

Figure 2 

Material examined: Southern Baffin Bay (lat. 67°49' 
N— long. 60°46'W to lat. 67°38'N— long. 60°38'W). 
HERO Station 26. collected 16 August 1968, dredged 
in 1.830 m, on a bottom of brown sticky mud (4 
specimens. TYPE). 

Description: Length up to 55 mm, width up to 6 
mm at the inflated portion. Segments number 24 to 
30 in the four specimens available. The body is 
greatly inflated through segments 6 to 10. Thereafter 
it narrows to a slender abdominal region. The entire 
body is areolated and marked off with small rectangu- 
lar raised areas (Fig. 2a). The parapodia are reduced 
throughout the body. There are no parapodial 
processes. The posterior end terminates in a simple 
lobed ring. 

The prostomium is bifid with two prominent lobes 
(Fig. 2a. b). There are no eyes. The base of the 
prostomium is retracted into the achaetous buccal seg- 
ment. The first setiger contains only capillary noto- 



setac :niii ncuroHctai 1 1 
ger 2 arc arranged in two tit 
slender curved ai ii Dial Ctai I I 

furcate setae; the * I pillarici 

The ncurosctae have a similar arrangement, < 

that there are more "i the aciculai and ft 

and fewer capillaries, ["he furcali • i ai omplclcly 

replace the aciculai setae On * 

menl of one bundle ol apillai 

bundle of smaller furcate el uccccd- 

ing setigers to the end ol the bud;.. The rut 

have one tine longei than the other. Each tine u 

spinous along the inner bordei 'I ig. 2d). 

Distribution: Southern Baffin Bay. depth ol 1,830 
m. 



I.I HERATURE (III D 

Ashworth, J. H. 1901. The Anatomy of ScaWj 

inflation Rathke. Quart J. Micro. Sci.. 45: 
309. 

Day, J. H. 1961. The polychactc fauna of South 
Africa. Part 6. Sedentary species dredged off 
Cape coasts with a few new records from the 
shore. J. linn. Soc. Zool.. 44:463-560. 



1967. 



A monograph on the Polychacta 
of Southern Africa. Part 2. Sedentaria. The 
British Museum (Natural History), publication 
656:459-878. 

Fauchald. K. 1970. Polychaetous annelids of the 
families Eunicidae, Lumbrineridae. Imphitimidae. 
Arabellidae. Lysaretidac and Dorvilleidae from 
western Mexico. Allan Hancock Monogr. Mar. 
Biol., (6): 1-335. 

Fauvel. P. 1927. Polychetes Sedentaires. addenda 
aux Errantes. Archiannelides. Myzostomaires. 
Faune de France. 16:1—492. 

. 1928. Annelides polychetes nouvelle de 

llnde. Pt. 2. Bull, du Museum National d'- 
Histoire Naturelle. Paris. 34:159-165. 

Hartman. O. 1942. The identity of some marine 
annelid worms in the United States National Mu- 
seum. Proc. U. S. Nat. Mus.. 92(3142):101-140. 

. 1960. Systematic account of some marine 

invertebrate animals from the deep basin - 



Figure 2. Cryptosclerocheilus baffinensis. new species: a. entire animal in dorsj 
anterior end in ventral view: e. capillary seta from anterior setiger: d. furcate set; 
3; e, acicular seta from setiger 2. 



132 



IWIII ll\ SOI l/lll<\ CAI.IFORMA ACADEMY OF SCIENCES 



VOLUME 71 



southern California. 
pod.. 22:69-216. 



Allan Hancock Pacific Ex- 



— . 1465. Deep water benthic polychaetous 

annelids off New England to Bermuda and other 
North Atlantic areas. Allan Hancock Found. 
Occas. Pap.. (28): 1-378. 



Mcintosh, W. C. 1885. Report on the Annelida 
Polychaeta collected by H.M.S. CHALLENGER 
during the years 1873-76. Challenger Repts. 
Zool., 12:1-554. 



Accepted for publication December 17, 1971. 



FLUCTUATIONS IN POPULATION DENSITY OF THE HISPID COTTON RAT: 
FACTORS INFLUENCING A "CRASH" 

Eugene D. Fleharty, Jerry R. Choate, 1 and Michael A. Mares 2 



Abstract: A population of hispid cotton rats, Sigmodon hispidus, inhabiting a remnant 
grassland in west-central Kansas, near the northern limit of the range of the species, was live- 
trapped from April of 1965 through February of 1969. Until 1968, the population exhibited 
a pronounced annual cycle of abundance, varying from most abundant in autumn to least 
abundant in spring. Annual declines began at the time of normal cessation of breeding with 
the onset of winter, and were augmented by harsh winter weather. Monthly ecological densi- 
ties ranged from 0.0-65.5 rats per hectare in the favored habitat. Population turnover was 
92 percent complete in six months. In 1968, the population continued to exhibit a typical, 
seasonal pattern of fluctuation until autumn, but then underwent a "crash" that completely 
decimated the population. Biotic and environmental factors that might have influenced the 
crash included normal autumnal cessation of breeding coupled with predation, parasitism, and 
severe weather conditions. The balance between adverse winter weather and physiological and 
behavioral adaptations to survive suboptimal weather conditions probably is largely responsible 
for the location of the northern limit of the geographic range of 5. hispidus. 



During the course of ecological and physiological 
studies on cotton rats, Sigmodon hispidus, inhabit- 
ing a remnant grassland in west-central Kansas, 
we had the opportunity to monitor a "crash" in 
population density. Inasmuch as the population 
under study was near the northern limit of dis- 
tribution of the species (Hall and Kelson, 1959: 
673), we anticipated that analysis of factors that 
influenced the crash might reveal natural mecha- 
nisms that regulate northern dispersal of this im- 
migrant Neotropical species. Therefore, the pur- 
poses of this paper are: 1 ) to present data on 
population dynamics of cotton rats at the latitude 
of Kansas: 2) to summarize factors that regulate 
annual fluctuations in population density of cotton 
rats; and 3) to relate those factors to the delicate 
balance between adaptation and environment that 
determines the location of the northern limit of 
distribution of the species. 



METHODS 

The 5.6-hectare remnant prairie in which the 
study was conducted is in the southeastern quarter 
of section 1, T. 14S, R. 19W, Ellis County, Kan- 
sas. The study area consists of six vegetative com- 
munities (Brock, 1968) that have remained rela- 
tively undisturbed in a climax condition for more 
than 60 years (Martin, 1960). Cotton rats were 
live-trapped, marked, and released during the first 
half of all but two months from April of 1965 
through February of 1969. Both the study area 
and the procedures followed in gathering data 
were described in greater detail by Fleharty and 



1 Dept. Biological Sciences and Agriculture, and Mu- 
seum of the High Plains, Fort Hays Kansas State 
College. Hays, Kansas 67601. 

2 Dept. Zoology, University of Texas, Austin, Texas 
78721. 



IV 7 2 



POPULATION DENSITY 01 nil HISPID COTTO R U 






Mares (1972). Determination of population den- 
sities was based on numbers of individual cotton 
rats caught (.luring each trapping period. I he 
paucity of previously uncaptured rats obtained at 
the close of each trapping period indicated thai 
most rats on the grid had been accounted for (see 
Davenport, 1964; of. Van Vleck, 1968). Ecologi- 
cal density (number per preferred habitat — Odum, 
1971 ) was estimated by assuming that the popula- 
tion was distributed throughout the different habi- 
tats in the proportions shown by trap success (see 
Fleharty and Mares, 1972). From October of 
1967 through February of 1969, weights of rats 
were recorded at the time of capture and the "cap- 
ture calendar" method (Andrzcjcwski, 1963, 1967) 
was used to determine age structure of the popu- 
lation. Meteorological data were obtained from 
the Fort Hays Agricultural Experiment Station, 
which is located about two miles from the study 
area. 



RESULTS 




S A 
1965 



W S 



SAWSSAWS 
1966 1967 

SEASONS 



Figure I. line graph showing number of 
caught on grid in spring, summer, autumn, 
of 1965-1968. 



SAW 
1968 



cotton rals 
and winter 



If data pertaining to the last year of study are dis- 
regarded, crude density (number per entire study 
area) ranged from a low of 0.2 per hectare (one 
rat on grid) in May of 1965 to a high of 20.6 per 
hectare (115 rats on grid) in October of 1967. 
Greatest densities occurred in autumn of each 
year, whereas least densities occurred in the springs 
of 1965, 1966, and 1968, and in summer of 1967 
(Fig. 1). Males were significantly more numer- 
ous than females (P < .05); a total of 516 males 
(57 percent) and 390 females (43 percent) were 
captured. 

Ecological densities were greatest in low-lying 
plant communities dominated by Andropogon 
gerardi (Fleharty and Mares, 1972). Monthly 
estimates of density in a draw, where the vegeta- 
tion consisted principally of A. gerardi, Kochia 
scoparia, and Hclianthus animus, reached a peak 
of about 45.5 cotton rats per hectare in three of 
the four autumns, thus suggesting that the carry- 
ing capacity of this habitat was attained almost 
every year. The greatest ecological density re- 
corded during the 47 months of trapping was 65.5 
per hectare in October of 1966. 

Rate of turnover of the population averaged 66 
percent during the month following first capture, 
and turnover was 92 percent complete by the end 
of the sixth month. These figures do not differ 
appreciably from those reported by Odum (1955) 
and Goertz (1964). Average time spent on the 
study area was 52.1 days for males and 54.3 for 



females; however, these times are minimal esti- 
mates because no means of determining precise 
dates of natality and mortality were used. Two 
males were on the study area for at least 336 and 
335 days, respectively, and one female survived 
at least 363 days. Goertz (1964) mentioned a 
male that survived for at least 12 months and a 
female that survived at least nine months. 

Age structure of the population during a 15- 
month period suggests that most reproduction 
occurred during summer and autumn (Fig. 2). 
Juveniles were most numerous in autumn, and 
adults became most numerous in late winter when 
the surviving juveniles attained the weight (60 g) 
at which they were considered mature. Old adults 
were most numerous in summer, when the surviv- 
ing adults attained the weight (110 g) that has 
been used to delimit the old adult age class (see 
Fleharty and Choate. 1972). 

In November of 196S. when the population be- 
gan its annual decline in density, the only notable 
difference in age structure as compared with No- 
vember of 1967 was the fact that fewer than half 
as many old adults w r ere present in the population 
(Fig. 2). By December of 196S. no old adults 
remained and the population contained only about 
half as many rats as it had during December of 
1967. In January of 1969. only !6 cotton rats 
were caught on the study area. .:•; comp 
86 only three months previously, and 1 



134 



BULLETIN SOI lllll<\ CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



100 



so 



i- 60 

< 

DC 



GC 

CD 

Z 



U 



O N D J 
1967 



□Q 



I 






F M 



A M J J A S 
1968 

MONTHS 



O N D J F 
1969 



Figure 2. Age structure of population of cotton rats 
from October of 1967 through February of 1969. 
Diagonal lines represent old adults; plain represents 
adults; and stippled area represents juveniles. 



were juveniles. No cotton rats remained on the 
study area in February of 1969, and subsequent 
snaptrapping in and around the remnant grass- 
land resulted in no captures during March, April, 
or May of 1969. 

DISCUSSION 

Annual cycles of abundance, with greatest densi- 
ties occurring in autumn and least densities in 
spring, were described for cotton rats by Komarek 
(1937), Odum (1955), and Provo (1962). Other 
authors (e.g., Schendel, 1940; Cockrum, 1952; 
Fitch, 1958; Goertz, 1964; Hoffmann and Jones, 
1970) have commented on the marked reduction 
in numbers of cotton rats that takes place between 
autumn and spring and have postulated that vari- 
ous aspects of winter weather might be respon- 
sible for population declines. However, our data 
indicate that annual population declines began in 
November, when there were few days in which 
the maximum temperature failed to exceed 0°C. 
It seems likely, therefore, that severe weather dur- 
ing late winter might act to accelerate an already 
declining population, but that biotic factors such 
as predation (see especially Schnell, 1968) and 



the normal cessation of breeding interact in au- 
tumn to initiate the annual reduction in numbers. 

At the latitude of Kansas, cotton rats reproduce 
mainly in summer and autumn (Bancroft, 1969); 
most breeding on the study area took place during 
the period June through October although, when 
data from all 47 months of the study were pooled, 
a few rats were trapped in reproductive condition 
in every month of the year (Fleharty, unpublished 
data). Coincident with decreased breeding, the 
number of juveniles that entered the population 
declined sharply in late winter and spring (Fig. 2). 
Therefore, animals lost from the population in 
late winter, for whatever reason, were not replaced. 

A substantial percentage of cotton rats lost from 
the population in autumn and early winter likely 
were removed by predators. Schnell (1968) re- 
garded a density of about 37 cotton rats per hec- 
tare as the "predator-limited" capacity for the 
species. With one exception in this study, density 
of cotton rats exceeded 37 per hectare in preferred 
habitats only in autumn and early winter, just be- 
fore the advent of population declines. During 
these periods of high density, when the carrying 
capacity of preferred habitats probably was ex- 
ceeded, rats moved out into less favorable habitats 
with reduced overstory (Fleharty and Mares, 
1972) and undoubtedly were thereby subjected to 
increased predation by raptors. Sealander and 
Walker (1956) reported that cats, coachwhip 
snakes, and red-tailed hawks prey on cotton rats, 
and Schnell (1968) presented evidence in sup- 
port of Odum's (1947) observation that the 
marsh hawk is a primary predator on cotton rats. 
Although we lack actual data on predation, marsh 
hawks were especially numerous over the study 
area in late autumn of each year (Ely, 1971), and 
likely fed largely on cotton rats during their peak 
period of abundance. Also, long-tailed weasels, 
snakes, and other predators were known to in- 
habit the study area, and probably preyed regu- 
larly on cotton rats, especially during their peak 
of abundance. 

Acting together, predation (and other forms of 
mortality) and cessation of reproduction would 
be sufficient to initiate annual declines in abun- 
dance. However, severe winter weather undoubt- 
edly augments those declines. The behavior of 
cotton rats in winter reflects their recent arrival 
from a region where the severity of winter weather 
is less than that which they experience today near 
the northern limit of their range. Cotton rats are 
not known to store food (Schendel, 1940), and 
periods of extreme cold effectively restrict forag- 



1972 



POPULATION DENSITY (>l THE HISPID COTTOI RA1 






ing behavior (Goertz, 1964) and induce "huddling" 
(e.g., Schendel, 1940; Dunaway and Kaye, 1961; 
Wicgcri and Mayenschein, 1966). Inactivity dur- 
ing cokl weather thus would force cotton rats to 
expend their reserves of energy to maintain their 
hotly temperature, and might lead to "cold weather 
starvation" (Howard, 1951) during protracted 
periods of severe weather. 

II cessation of reproductive activity, coupled 
wilh predation and other forms of mortality and 
augmented hy starvation during severe winter 
weather, are responsible for annual declines in 
abundance of cotton rats, then the crux of the 
problem has to do with the factor or factors that 
acted wilh different intensity on the population 
to produce the crash in 1968-1969. Two factors 
that could have been involved are disease and 
parasitism. Stoddard (1931) and Davis (1958) 
both suggested that disease is an important cause 
of periodic population declines in cotton rats. 
Certainly, the high densities that are attained dur- 
ing autumnal peaks in density would be conducive 
to rapid spread of disease, but we have no evi- 
dence that disease was involved. However, we do 
have at least indirect evidence for a high incidence 
of parasitism. In conjunction with other research, 
it was noted that numerous cotton rats, especially 
old adults and adults, collected on and around the 
study area contained large numbers of nematodes 
— in some instances the stomachs were distended. 
Furthermore, if parasitism were involved in the 
population decline this might account for the fact 
that old adults were the first to exhibit a marked 
decrease in numbers (Fig. 2). Whatever the case, 
few cotton rats remained on the study area in 
December of 1968 when the effects of winter 
weather came to bear on the population. 

Goertz (1964) discussed the effects of severe 
winter weather on a population of cotton rats in 
Oklahoma. He concluded (p. 376) that a "long 
period [11 days] of below-freezing weather, with 
snow on the ground and ground frozen . . ." con- 
tributed to a marked decline in population density 
of cotton rats. However, our data do not indicate 
what specific characteristic of winter weather was 
responsible for decimation of the population re- 
ported herein. In December of 1965. 57 cotton 
rats were present on the study area. No decline 
in numbers occurred during January and Febru- 
ary of 1966 even though those months were con- 
siderably colder than normal; the average maxi- 
mum temperature for the last 14 days of January 
and the first three days of February was less than 
-6°C, and 8-10 centimeters of snow covered the 



ground foi 16 days during thit pt riod In i < 
her of 1966, 7f cotton rats t/cn present on the 
study area, bul bj February ol 19', 7 the popula 
lion li.nl declined to only (8 rats i hi 
maximum temperature foi the last nim 

December was aboul 6 ( , ami .1 15 CCntimctCI 
snow cover was present, Some ol thit snow re- 
mained on ihe ground foi the lu. 1 12 di 
January, [n Decembei ol 1967, 53 cotton rata 
were present on the Study area. No decline in 
numbers occurred timing January or I ebni. 

1968 although temperatures averaged slightly be- 
low normal. In December ol 1968, 30 cotton rats 
were present on the area, bul b\ February ol 1969 
none remained. Temperatures during December 
were the lowest recorded for that month since 
1963; the average maximum temperature lor the 
last I I days was -5 ('. and at no time during that 
period did the maximum temperature exceed f . 
Also, eight centimeters of snow blanketed the 
ground for the last 13 days ol December. I lie 
average maximum temperature for the lirst day and 
the last nine days of January of 1969 averaged 
well below freezing, and three centimeters of snow 
covered the ground for the first five days of the 
month. Even so, the weather conditions (at least 
so far as temperature and snow cover are con- 
cerned) were no more severe in the wintei ol 
196S-1969 than they were in 1965-1966. 

These data indicate that decimation of the pop- 
ulation of cotton rats during the winter of 1968- 

1969 probably did not result entirely from weather 
conditions. If a cause-and-effect relationship were 
involved, then the population inhabiting the area 
during the winter of 1965-1966 seemingly would 
have experienced a crash like that which occurred 
three years later unless, for some reason, the rats 
inhabiting the area during the winter of 1965- 
1966 had more energy reserves to draw from than 
did the rats inhabiting the area during the winter 
of 1968-1969. 

In an attempt to estimate the extent and utility 
of energy reserves in ox'erwintering populations of 
cotton rats, we performed the following computa- 
tions on the population that succumbed in the 
"crash" of 196S-1969. Energy contained within 
the population in January of 1969 was distributed 
among only 12 juveniles and four adults (mean 
weights, 56 and 78 grams, respectively). The body 
weight of juveniles consisted of 7.56 percent lipid, 
whereas that of adults consisted of 10.72 percent 
lipid (Fleharty el <;/.. 1972). If we assume tl 
all but three percent of the lipid content was avail- 
able for respiration as energy (Kleiher. 196 P 



136 



HI' I. LET l\ SOlTHEliS CALIEORSIA ACADEMY OE SCIENCES 



VOLUME 71 



that the lipid released 9.093 keal g (Fleharty cl 
id., I')72) when respired, then lipid reserves of 
juveniles and adults amounted to 37.28 and 73.74 
keal. respectively. If the average dry weight of 
full stomachs is 1.04 in juveniles and 1.56 in 
adults (Fleharty, unpublished data) the assimila- 
tion efficiency of stomach contents is 88 percent 
(Davis and Golley, 1963). and the caloric content 
of the food eaten is 4.55 keal g (Fleharty, unpub- 
lished data), then the reserve energy of full stom- 
achs is 4.14 keal in juveniles and S.I I in adults. 

I hese \ .dues added to caloi ic \ allies ol lipids \ ield 
totals of 41.43 and 81.85 keal available to juve- 
niles and adults, respectively, if severe weather 
resulted in a cessation of foraging. These figures 
might be increased somewhat if cotton rats re- 
sorted to feeding on dead nest mates (Schendel, 

1949) or to cannibalism. 

Pearson (1960) and Gorecki (1969) noted the 
importance of "huddling" in reducing the rate of 
metabolism; however, the metabolic rate of huddl- 
ing cotton rats remains unknown. In order to 
bracket the actual metabolic rate of huddling rats 
(which undoubtedly are exposed to microclimatic 
conditions less severe than those above the vege- 
tation), the rates (Gaertner, 1968) at 0°C (3.20 
ccOo g hr) and at 32°C (1.26 cc 2 /g/hr) were 
used to compute the lengths of time that juveniles 
and adults could survive if totally dependent on 
stored energy. Assuming an oxy-caloric coefficient 
of 4.82 calories for both juveniles and adults, juve- 
niles could survive using metabolic reserves for no 
more than 2.5 days at 0°C and only 5.1 days at 
32°C; adults could exist on metabolic reserves for 
no more than 2.8 days at 0°C and only 8.1 days 
at 32°C. The actual length of time huddling rats 
could survive under field conditions doubtlessly 
falls somewhere between these two extremes. It 
should be pointed out, also, that as density de- 
creased during the crash the huddling phenomenon 
to withstand cold would become less effective be- 
cause it would be increasingly difficult to locate 
other rats. 

Our calculations regarding length of survival of 
cotton rats when dependent strictly on energy 
reserves illustrate an important point. Huddling 
rats must make occasional forays in search of 
food or they could not survive even a relatively 
moderate winter near the northern limit of their 
distribution. Such forays, of course, increase the 
probability of predation, but the selective advan- 
tage resulting from replenishment of energy stores 
undoubtedly outweighs the effects of removal by 
predators of a few individuals from the popula- 



tion, at least when the population is not already 
depleted by other factors. In any case, if we 
assume that winter weather was at least indirectly 
responsible for decimation of the population in 
1969, then it is reasonable to suggest that energy 
reserves and huddling simply were not sufficient 
to enable survival of any of the cotton rats that 
remained on the study area during that winter 
period. 

Therefore, it seems likely that predation and /'or 
other forms of mortality, coupled with the normal 
cessation of reproductive activity, was responsible 
for initiation of annual declines in density of cot- 
ton rats, whereas winter weather acted primarily 
to accelerate declines by causing starvation of ani- 
mals that lacked sufficient energy reserves to with- 
stand periods in which foraging was restricted. 
Severe weather undoubtedly was at least indirectly 
responsible for final decimation of the population 
during the crash, but the effects of protracted cold 
might not have been so devastating if old adults 
and more adults had been present or if the remain- 
ing adults had not already been deprived of some 
of their energy reserves, perhaps by parasitic 
infection. 

An alternative explanation for the crash is that 
it merely represented the depression phase of 
cyclic oscillation in population density. The pos- 
sibility that populations of cotton rats exhibit 
periodic cycles of abundance was first suggested 
by Stoddard (1931). Komarek (1937) reviewed 
data on population fluctuations of cotton rats in 
the southern and southeastern United States, and 
concluded that this species undergoes regular 
cycles of abundance with a periodicity of four- 
five years. Observations of several other workers, 
on the other hand, would seem to indicate that 
oscillations in population density of cotton rats 
have a periodicity of about 10 years. Pronounced 
declines in population density of cotton rats were 
recorded in Oklahoma during the winter of 1939— 
1940 (Schendel, 1940); in Kansas during the win- 
ter of 1948-1949 (Cockrum, 1952); in Kansas 
during the winter of 1957-1958 (Martin, 1960); 
and in Oklahoma (Goertz, 1964), Tennessee 
(Dunaway and Kaye, 1961), and Texas (Haines, 
1963) during the winter of 1959-1960. The crash 
of 1968-1969 reported herein occurred approxi- 
mately when it would have been expected if S. 
hispidus undergoes cyclic oscillations in density 
with a periodicity of 9-11 years, but the low 
points shown in figure 1 might also be interpreted 
to indicate a periodicity of four years. We regard 
existing data on periodic cycles of S. hispidus as 



1972 



I'Oi'UI.ATlON DENSITY <>i i ill. iiisriD COTTO 






inconclusive. Imi admit thai regularly-expressed 
intrinsic factors might have prompted ihc crash 
observed in 1968-1969. 

Whether the crash was a manifestation of cyclic 
intrinsic mechanisms or was caused by preclalion, 
disease, parasitism, or one or more of these com- 
bined with "cold weather starvation," the delicate 
balance between weather conditions and energy 
(lipid) reserves undoubtedly is an important factor 
in determining the success of populations of cot- 
ton rats near the northern limit of their distribu- 
tion (sec Fleharty ci <d., 1972), and likely is the 
single most important factor in determining the 
location of the northern limit of the range of the 
species. Northward dispersal of cotton rats in 
Recent time reflects a gradual increase in tem- 
perature that accompanied the last glacial retreat, 
but northward movement during the last century 
(Cockrum, 1948; Genoways and Schlitter, 1966; 
Hoffmann and Jones, 1970) has proceeded at a 
more rapid rate than is explicable on climatic 
grounds alone. It seems likely that this rapid 
northward dispersal has resulted in part from 
adaptations (Fleharty et ah, 1972; Fleharty and 
Choate, 1972) by cotton rats to facilitate survival 
in increasingly severe winter weather through 
stockpiling of energy reserves in the form of lipids 
that can be drawn upon during periods of limited 
activity. 

ACKNOWLEDGMENTS 

Lowell L. Getz (University of Illinois) and Robert L. 
Packard (Texas Tech University) critically reviewed 
the manuscript. Clerical assistance was provided by 
Sherry Quint and Fi Choate. 



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cotton rat in Kansas i rans, t ad. v > 

51 ;306 3 12. 

— . 1952. Mammals ol Kansa i n>- t 



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Davis. D. I!., and F, B. Golley. 1963. Principles in 
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xiii -|- 335 pp. 

Davis, W. B. 1958. Cotton rats plague south 
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Dunaway, P. B.. and S. V. Kayc. 1 96 1. Cotton rat 
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265-268. 

Ely, C. A. 1971. A history and distributional lisi 
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Fitch. H. S. 1958. Home ranges, territories, and 
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Fleharty. E. D.. and J. R. Choate. 1972. Bioener- 
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Fleharty. E. D.. and M. A. Mares. 1972. Habitat 
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Kansas. Southwestern Nat., (in pre 



Bancroft. W. L. 1969. Notes on reproduction of 
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kansas, Fayetteville. 149 pp. 



138 



III III ll\ sol [HERN CALIFORNIA ACADEMY OF SCIENCES 



VOLUME 71 



Genoways, H. H.. and D. A. Schlitter. 1966. North- 
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Goertz, J. W. 1964. The influence of habilat qual- 
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1971. Fundamentals of ecology. W. B. 



Saunders Co.. Philadelphia, xiv + 574 pp. 



Gorecki. A. 1968. Metabolic rate and energy budget 
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Hall. E. R.. and K. R. Kelson. 1959. The mam- 
mals of North America. Ronald Press, New 
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Hoffmann, R. S.. and J. K. Jones, Jr. 1970. Influ- 
ence of late-glacial and post-glacial events on the 
distribution of Recent mammals on the northern 
Great Plains. Pp. 355-394, in Pleistocene and 
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Howard, W. E. 1951. Relation between low tem- 
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Komarek. E. V. 1937. Mammal relationships to 
upland game and other wildlife. Trans. Second 
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152-160. 

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habitat selection, temperature and light in the 
regulation of a Sigmodon population. Unpubl. 
Ph.D. thesis, Univ. Georgia, vi + 73 pp. 

Schendel. R. R. 1940. Life history notes of Sigmo- 
don hispidus texianus with special emphasis on 
populations and nesting habits. Unpubl. M.Sc. 
thesis. Oklahoma State Univ., 40 pp. 

Schnell, J. H. 1968. The limiting effects of natural 
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Sealander, J. A., Jr., and B. Q. Walker. 1955. A 
study of the cotton rat in northwestern Arkansas. 
Proc. Arkansas Acad. Sci., 8:153-162. 

Stoddard, H. L. 1931. The bobwhite quail. Charles 
Scribners Sons, New York, 550 pp. 

Van Vleck. D. B. 1968. Movements of Microtus 
pennsylvanicus in relation to depopulated areas. 
J. Mamm., 49:92-103. 

Wiegert, R. G., and J. C. Mayenschein. 1966. Dis- 
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Mamm.. 47:118-120. 

Accepted for publication July 25, 1972. 



THE CORAL SNAKE MICRURUS NICROCINi /' 
IN HONDURAS (SERPENTES: ELAPIDA1 I 

Larky David Wilson 1 and John R. Meyer 



Abstract: Analysis of seven characters of scutellation and coloration ol matt i ial ol \lf in 
rus nigrocinctus from Honduras indicates thai variation in these characters is more complex 
than has formerly been recognized. The characters studied exhibit discordant patterns o( 

variation but populations of M. nigrocinctus from northern Honduras an ■ mil < ii-.t inc i 

from those from southern Honduras in a number of features. Populations from intermediate 
areas exhibit discordant admixtures of these characters. We suggest that application ol i 
to infraspecific taxa in Honduras tends to obfuscate rather than illuminate variational trends 
in this coral snake. 



Roze (1967, 1970) interpreted the variation in 
color pattern and scutellation exhibited by speci- 
mens of the coral snake Micrurus nigrocinctus 
(Girard) from Honduras as indicating the pres- 
ence of two geographic races within that country. 
One subspecies, M. n. divaricatus (Hallowell), 
was envisioned as inhabiting northern and central 
Honduras, extending as far north as British Hon- 
duras. The other subspecies, M. n. zunilensis 
Schmidt, was considered to range along the Pacific 
slopes of southern Honduras, extending north to 
Chiapas, Mexico. Micrurus n. divaricalus was 
characterized as having light bands present, 1 I to 
26 (usually more than 19) black body bands, and 
the black pigment of the red bands irregularly 
distributed (Schmidt, 1933; Roze, 1970). Micru- 
rus n. zunilensis was characterized as having the 
yellow bands variable in length (0 to 2 scale rows 
long), 12 to 26 (usually less than 19) black body 
bands, and with very little or no black pigment 
on the red bands (Schmidt, 1933; Roze. 1970). 

During examination of material of this species 
of coral snake from Honduras in preparation for 
a report on the snakes of that country, it became 
apparent that color pattern variation was much 
more complex than had formerly been recognized 
and that the taxonomic arrangement discussed 
above could not be accepted in its entirety. It is 
the purpose of this paper to discuss color pattern 
and scutellational variation of Micrurus nigro- 
cinctus in Honduras and to suggest that the recog- 
nition of two subspecies in Honduras obfuscates, 
rather than elucidates the variational picture. 

Eighty-four specimens of Micrurus nigrocinctus 
are available from widely scattered areas in Hon- 
duras. Unfortunately, most of these snakes were 
collected at a few localities, as about SO percent 



of the known specimens are from 4 ol the 1 I 
populations characterized below. 

For the purpose of discussion of variation in 
color pattern and scutellation. we segregated the 
material into II samples (Fig. I I. Samples 
were set up utilizing specimens from geograph- 
ically and altitudinally restricted areas with a more 
or less uniform vegetational cover. These samples, 
termed populations from this point on. may he 
characterized as follows. 

Population A (23 specimens). — From localities in 
the Sula Plain at elevations from about 20 to 100 m 
in mixed scrub forest (Tropical Dry Forest formation 
of Holdridge. 1962). 

Population B (24 specimens). — From localities in 
the western portion of the Nombre de Dios Piedmont 
at elevations from sea level to 30 m in tropical lowland 
rainforest (Tropical Moist Forest formation of Hold- 
ridge, 1962). 

Population C (13 specimens). — From localities in 
the Sierra de Sulaco and nearby ranges at elevations 
from 1200 to 1300 m in disturbed or undisturbed 
broadleaf evergreen forest (Subtropical Wet Forest 
formation of Holdridge. 1962). 

Population D (2 specimens). — From localities in 
the eastern portion of the Nombre de Dios Piedmont 
and the Sierra de Nombre de Dios in tropical lowland 
rainforest (Tropical Moist Forest formation of Hold- 
ridge. 1962). 

Population E (3 specimens). — From localities in 
the Aguan-Negro Plain and the Sierra de Nombre de 
Dios at elevations from sea level to about 200 m i 
tropical lowland rainforest and short-tree savanna 
(Tropical Moist Forest formation of Holdridge. 1< 



1 Miami Dade Jr. College. Div. Interdisciplir 
Studies. South Campus. Miami. Florida 3 

2 Bartram School. 2264 Bartram Road. J 
Florida 32207 



139 



140 



BULLETIN SOUTHERN CALIFORNIA ACADEMY OE SCIENCES VOLUME 71 




Figure I. Distribution in Honduras of population 
samples utilized in this study. See text for description 
of samples A-K. 



Population F (1 specimen). — From one locality in 
the valley of the Rio Tinto at an elevation of about 
430 m in tropical lowland rainforest (Tropical Moist 
Forest formation of Holdridge. 1962). 

Population G (3 specimens). — From localities on 
the Mosquito Coast near sea level in lowland pine 
savanna (Tropical Moist Forest formation of Hold- 
ridge. 1962). 

Population H (4 specimens). — From localities near 
Lake Yojoa at elevations from 700 to 750 m in 
disturbed or undisturbed broadleaf evergreen forest 
(Subtropical Wet Forest formation of Holdridge, 
1962). 

Population I (1 specimen). — From one locality in 
the Valle de Comayagua at an elevation of 580 m 
in open thorn forest (Tropical Arid Forest formation 
of Holdridge, 1962). 

Population J (7 specimens). — From localities in or 
near the Valle de Yeguare at elevations from 800 to 
1000 m in mixed scrub forest (Subtropical Dry Forest 
formation of Holdridge, 1962). 

Population K (1 specimen). — From one locality in 
the Pacific lowlands at an elevation near sea level in 



dense scrub forest (Tropical Dry Forest formation of 

Holdridge, 1962). 



ANALYSIS OF VARIATION 

Snakes of the genus Micntrus are notorious for 
the small number of external distinguishing fea- 
tures. Dorsal scale row number and the condition 
of the head scales and the anal plate arc all 
invariant. Diagnoses of the subspecies of Micrurus 
nigrocinctus have utilized the following characters: 
numbers of ventrals, position of the nuchal band, 
numbers of black body bands, presence or absence 
of yellow bands, and condition of the black tipping 
on the scales of the red bands. It is the variation 
in these characters as well as the variation in the 
number of subcaudals and the size of the black 
head cap in the Honduran specimens with which 
we are here concerned. 

Scutellation. — Coral snakes are highly sexually 
dimorphic in ventral and subcaudal numbers, with 
females having the greater number of ventrals and 
males the greater number of subcaudals. 

In Honduras, there appears to be a tendency for 
both males and females from the southern portion 
of the country to have higher numbers of ventrals, 
although this trend is not well documented (Table 
1 ) . Variation in subcaudal number is limited and 
exhibits no discernible pattern (Table 1). 

Black body band number. — The number of 
black body bands in Micrurus nigrocinctus over 
the entire range of the species varies from 10 to 
29. The range of Honduran specimens is almost 
the same, 1 1 to 29. Variation in black band 
number appears to be of no geographic signifi- 
cance. Both the highest and lowest ring numbers 
occur in the department of Atlantida in northern 



Table 1. Variation in ventrals and subcaudals in Honduran Micrurus nigrocinctus. Population statistics are 

arranged as follows: range (mean) sample size. 



Ventrals 



Subcaudals 



Population 



Mates 



Females 



Males 



Females 



A 

B 

C 

D 

E 

F 

G 

H 

I 

J 

K 



190-203(198.4)9 

191-206(196.8)13 

193-202(198.6)5 

198 
190-201(195.5)2 

203 
200-202(201.0)2 

208 

201 
191-206(200.0)4 

207 



208-220(213.0)13 

206-217(210.4)9 

206-217(212.5)8 

208 

210 

211 
215-218(216.5)2 

218-224(220.6)3 



47-51(49.6)8 
46-52(49.3)8 
45-51(47.0)5 

45 
49-51(50.0)2 

46 
54-56(55.0)2 

50 

46 
42-50(47.5)4 

51 



34-42(37.6)12 

34-39(36.4)9 

32-39(36.8)6 

36 

37 



36-39(37.7)3 
34-43(38.0)3 



19 72 



THE (OHM, SNAKE MK KUKIJS NIOKOf IN' ITJS II HOI 



Table 2. Variation in black body band number in 
Honduran Micrurus nigrocinctus. Population statistics 
are arranged as follows: range (mean) sample size. 



Population 


Mules 


Females 


A 


13-24(17.6)9 


14-23(19.1)13 


H 


11-21(13.8)13 


1 1-20(16.3)1 1 


C 


15 19(17.4)5 


16-21 ( 18.6)8 


D 


27 


29 


E 


14-201 17.0)2 


22 


F 


25 


- 


G 


18-19(18.5)2 


23 


II 


20 


17-221 19.0)3 


1 


22 


- 


J 


IK 19(18.5)4 


18-20(19.0)3 


K 


15 


- 



Honduras, whereas, intermediate numbers are 
found elsewhere (Table 2). A single specimen 
(MCZ 32011) has a saddle pattern as has been 
described for Micrurus fulvius (Neill, 1963). 

Yellow body bands. — Only a few of the speci- 
mens examined have retained the original color 
distinction between the red and yellow bands 
sufficiently intact so that an accurate determina- 
tion of the length of the yellow bands can be made. 
It is apparent, nevertheless, that the yellow bands 
range in length from to 2 scales. Yellow bands 
appear to be absent in all representatives of pop- 
ulations D, E, and F (Roze's, 1970, comments to 
the contrary notwithstanding). Yellow bands are 
present in all representatives of populations C, G, 
H, I, J, and K (Figs. 2A and 3A). Most of the 
specimens comprising population B appear to 
have had yellow bands, although in about 7 speci- 
mens they appear to have been absent. These 
specimens are all old and faded and no fresh 
material is available to corroborate these state- 
ments. Within population A some specimens are 
bicolor (5 of 23 specimens) and some are tricolor 
(at least 13 of 23 specimens still show evidence of 
yellow bands). The change between these two 
conditions is not abrupt, as is demonstrated in a 
series of 1 1 specimens from the vicinity of La 
Lima in the private collection of John Dickson. 
Three specimens are bicolor, showing no evidence 
of yellow bands. Seven specimens are tricolor and 
the yellow bands range from 1 to VA scales in 
length. Three other specimens show various stages 
of intermediacy between the bicolor and tricolor 
conditions. In these specimens yellow bands are 
absent or ill-defined on some amount of the pos- 
terior portion of the body. 

Black tipping on scales of red bands. — In M. 
nigrocinctus throughout its range, black pigment 



y 




Figure 2. Dorsal color pattern of (A i I SI \l/ 24421 
from 4 miles N San Lorenzo. Depto. Vallc. Honduras 
and (B) LSUMZ 22451 from the mountains above 
Trujillo, Depto. Colon. Honduras. 



distribution within the red rings varies from pig- 
ment being absent to being present and evenly 
distributed (approximately one spot per scale) to 
being present and unevenly distributed (some spots 
occupy more than one scale). Subspecies ol W. 
nigrocinctus have been partially diagnosed on the 
basis of the type of pattern of black pigment 
deposition ( Roze. 1970). Micrurus n. zunilensis 
Schmidt. M. n. coibensis Schmidt (Isla de Coiba. 
Panama), and M. n. melanocephalus (Hallowell) 
(Pacific versant of Nicaragua and Costa Rica) 
have very little, if any. black pigment present on 
the scales of the red bands. Micrurus n. nigro- 
cinctus (Girard) (Pacific versant of southern 
Costa Rica. Panama, and adjacent Colombia I. M. 
n. mosquitensis Schmidt (Atlantic versant of east- 
ern and southern Nicaragua to northwestern 
Panama), and M. n. babaspul Roze (Isla del \\.u7. 
Nicaragua) have well-defined black spots present 
on the tips of each of the scales of the red bands. 
Micrurus n. divaricatus has a relatively large 
amount of black pigment irregularly distributed on 
the scales of the red bands. 

The amount of variation in black pigment 
tribution in specimens of M. nigrocincn 
Honduras is so great as to coincide wit 
of variation for the entire species. 1 
up seven pattern categories in order 






Ill III ll\ \<>t ////AW CAUEORXIA ACADI.M) Ol SCIENCES VOLUME 71 

Ota 








Figure 3. Dorsal color pattern of (A) LSUMZ 24422 
from Tancin. Depto. Gracias a Dios, Honduras and 
(B) FMNH 8083 from the Garcia Plantation, Depto. 
Yoro, Honduras. 



A 



Figure 4. Dorsal color pattern of (A) USNM 20282 
and (B) USNM 20283 from Patuca, Depto. Gracias 
a Dios. Honduras. 



the spectrum of variation for facility of discussion. 
The pattern categories may be characterized as 
follows. 

1. Black pigment on the red bands is absent, in- 
distinct, or restricted to a few scattered scales (Fig. 
2A, B). 

2. Black tipping is present and more or less uni- 
form on all scales of the red band (Fig. 3A). 

3. Black pigment is scattered and some is clumped 
into larger spots, but tends to be arranged in the 
middle of the band (Fig. 3B). 

4. Black tipping is present on all scales of the red 
bands but some black tips are fused to form larger 
spots covering one or two dorsal scales (Fig. 4A). 

5. Black pigment is scattered all over the red band, 
but pigment fusion has produced definite paravertebral 
accessory incomplete bands; a black spot or scattered 
black pigment may be present on the venter opposite 
the dorsal spots (Fig. 4B). 

6. Black pigment of red bands is restricted to 
lateral accessory incomplete bands (Fig. 5A). 

7. Accessory incomplete bands fused to form a 
complete band analogous to the primary black bands 
(Fig. 5B). 

Pattern 1 is characteristic of M. n. zunilensis 
and perhaps M. n. coibensis and M. n. melano- 
cephalus. Pattern 2 characterizes M. n. nigro- 
cinctus, M. n. mosquitensis, and M. n. babaspul, 
and patterns 4 and 5 are found on M. n. divari- 



catus. Patterns 3 and 5 have not been previously 
described. The single specimen exhibiting pattern 
7 is discussed below. Distribution of the 7 pattern 
types among the 1 1 populations is shown in Table 
3. 

Several features are apparent in the distribution 
of color patterns (Table 3). First, patterns 1, 2, 
and 4 are relatively widespread geographically. 
They are also represented in the majority of the 
specimens. Patterns 3, 5, 6, and 7 are restricted 



Table 3. Distribution of the types of black pigment 

deposition on the red bands in Honduran Micrurus 

nigrocinctus. 











D attern types 






Population 


l 


2 


3 


4 


5 


6 


7 


A 


8 


10 


_ 


4 


1 


_ 


— 


B 


7 


12 


- 


3 


1 


- 


- 


C 


1 


9 


- 


3 


- 


- 


- 


D 


- 


- 


- 


- 


- 


1 


1 


E 

F 
G 


1 


- 


2 
1 


- 


- 


- 


- 


_ 


i 


_ 


2 


_ 


_ 


H 
I 

J 


3 
1 

4 

1 

26 


- 


- 


1 


- 


- 


- 


3 


- 


- 


- 


- 


- 


Total 


35 


3 


11 


4 


1 


1 



1972 



THE (ORAL SNAKE MK KUKiiS NIGRO( IN< n , / HO 






lo the more northern portions of the country and 
are represented in very lew of i he specimens. I he 
populations wiih the largest number ol members 
exhibit the widest range of pattern types. I hus. 
caution is required in drawing conclusions about 
the significance of these pattern types and their 
distribution because of the biased samples. Never- 
theless, patterns 6 and 7 are known only from 
population I). Also, patterns I, 2, and 4 are found 
to the exclusion of oilier pattern types in the 
southern populations (H through K). 

Position of the nuchal band and the size of the 
black head cap. — The position of the anterior edge 
of the nuchal band in Honduran M. nigrocinctus 
ranges from 2 dorsal scale lengths posterior to the 
parietals to I 'j dorsal scale lengths onto the pos- 
terior edge of the parietals. The black head cap 
may occupy as little as the internasals and pre- 
frontals or as much as the internasals, prefrontals, 
frontal, supraoculars and the anterior one-fourth 
of the parietals. The size of the light head band 
is a reflection of the size of the black head cap 
and Ihc position of the nuchal band, and is shortest 
in populations G, I, J, and K, longest in popula- 
tions B, C, D, E, and F, and of intermediate size 
in populations A and H. 




Figure 5. Dorsal color pattern of (A) I Si MZ 2177 
from the mountains above ( oro/al. Depto. Atlantiila. 
Honduras and (B) TCWC 21181 from 5', miles f 
La Ceiba. Deplo. Atlantida. Honduras. 



STATUS OF THE MAINLAND 

SPECIMEN OF -'MICRURUS 

RUATANUS" 

Roze (1967) reported M. ruatanus, formerly 
known only from the Bay Islands of Honduras, 
from the adjacent mainland of Honduras on the 
basis of TCWC 21181 from 5 Vi miles east of La 
Ceiba, Depto. Atlantida. This record has been 
questioned by Meyer (1969) and Wilson and 
Halm (in press). We have examined this speci- 
men and consider it to represent nigrocinctus and 
offer the following explanation to account for its 
resemblance to Micrurus ruatanus. In size and 
habitus TCWC 21181 resembles nigrocinctus. It 
measures 734 mm in total length, a size approxi- 
mated by numerous nigrocinctus from Honduras, 
whereas the largest ruatanus from the Bay Islands 
known to us measures 620 mm. The head of adult 
nigrocinctus from the northern coast of Honduras 
is most often well distinct from the neck, whereas 
that of ruatanus is scarcely distinct from the neck: 
the La Ceiba specimen resembles nigrocinctus in 
this respect. 

The number of black body bands (29) on 
TCWC 21181 is lower than that recorded for any 
ruatanus from the Bay Islands (the range is 34 to 



45 according to Schmidt. 1933. and a range of 
33 to 45 is given for 8 specimens examined by 
Wilson and Hahn, in press). 

The ventral count for TCWC 21 181 (a female i 
is 208, within the range of 207 to 220 given for 
M. n. divaricatus by Schmidt (1933, 1936), but 
not within that recorded for females of ruatanus 
(Glinther, 1895. recorded a range of 200 to 204 
and Wilson and Hahn. in press, a range of 193 
to 203 ) . 

The resemblance of TCWC 21181 to \/. rua- 
tanus in pattern, particularly in the relatively high 
number of black body bands, is only superficial. 
The specimen appears to have been bicolor in 
life, as is the case with fresh material from this 
general area of Honduras and with M. ruatanus. 
Nevertheless, the black body bands of ruatanus 
are alternately longer and shorter and the shorter 
ones are frequently interrupted laterally, especially 
on the posterior portion of the body (Schmidt. 
1933: Wilson and Hahn. in press). On the other 
hand, the bands of TCWC 21181 are all about 
the same length (Fig. 5B) and none are broken. 
We suggest that this specimen, with its high 
number of black body bands (for nigi 
represents the end stage of a morphocl 
increasing development and conce 
pigment. We envision this ■ 



144 



Hll II. I IS sol nil KS < M.lfOKNIA K IDEM1 01 SCIENCES VOLUME 71 



nate in ;i condition that we have designated as 
pattern 2. where black tipping is uniformly dis- 
tributed on all scales of the red hand. This dine 
continues through pattern 4. where some Mack 
tips are fused together to form larger spots, and 
patterns 5 and 6, where either paravertebral or 
lateral accessory incomplete hands are present. 
The eline ends with pattern 7 (that of TCWC 
21 ISI ). where the accessory hands are complete. 
Complete fusion of accessory bands would almost 
double the number of bands. If the number of 
accessory bands of TCWC 21181 is subtracted 
from the total number of 29, the number of pri- 
mary black bands is 15. a number within the 
range of nigrocinctus, as envisioned by Schmidt 
( 1933). It is worthy of note that patterns 4, 5, 
and 6 are most frequently or solely encountered 
in the populations on the northern coast of Hon- 
duras. We suspect that additional collecting in 
the area around and between La Ceiba and Trujillo 
will demonstrate the presence of specimens with 
patterns intermediate between those designated as 
patterns 5 and 6. 

In conclusion, we feel that TCWC 21181 from 
near La Ceiba, Honduras is a M. nigrocinctus, 
albeit a peculiar one, and that M. ruatanus is not 
known from the mainland of Honduras. 



GEOGRAPHIC TRENDS AND 
SYSTEMATIC CONCLUSIONS 

Although the characters examined in this study 
exhibit patterns that are discordant, the popula- 
tions of the north coast of Honduras (B, D, and 
E) are distinct from the populations of southern 
Honduras (I, J, and K) in a number of features. 
The populations from intermediate areas exhibit 
discordant intermixtures of these characters. 
Specimens from northern Honduras generally are 
bicolor and have a low number of black body 
bands and ventrals and a long pale head band. 
Specimens from southern Honduras generally are 
tricolor with a high number of black body bands 
and ventrals and a short pale head band. 

In Honduras, at least, we see no reason to 
recognize infraspecific taxa. Such an action would 
require an entirely arbitrary decision as to which 
of the characters we have studied is most impor- 
tant (i.e., most representative of the genetic rela- 
tionships) and we have no confidence in making 
such a judgment. In addition, the "zone of inter- 
gradation" is larger than the range of either the 
northern or southern populations. Finally, we 
would like to have the opportunity to examine 



sufficient material from other sites before extend- 
ing our remarks further than we have here. 

MATERIAL EXAMINED 

For easy reference, we have arranged the specimens 
examined under the 1 I populations we recognize. 

Population A.— ATLANTIDA: Berichiche (MCZ 
38773); Guaymas District (FMNH 16116-20); Ulua 
River, near Tela (MCZ 19945). CORTES: Lake 
Ticamaya (FMNH 5311); La Lima (LSUMZ 24425; 
JD-II); W of San Pedro Sula (FMNH 5312); 15.5 
mi S Puerto Cortes (LSUMZ 24426). YORO: Garcia 
Plantation (FMNH 8083). 

Population B.— ATLANTIDA: Lancetilla (AMNH 
46968-71, 70237-38; FMNH 21783; MCZ 29412-13, 
32010-12, 32014, 33342, 38772-73); Colorado Dis- 
trict (FMNH 16115; MCZ 21772-73); Tela (FMNH 
16115. 25215; USNM 82166, 84568-69, 129428). 

Population C. — YORO: Mataderos Mountains 
(MCZ 38770-71); Portillo Grande (FMNH 21893- 
94, 34720-22, 34747, 35559; MCZ 38774-77). 

Population D. — ATLANTIDA: mountains above 
Corozal (LSUMZ 21773 ); 5% mi E La Ceiba (TCWC 
21181). 

Population £.— COLON: Puerto Castilla (MCZ 
22433-34); mountains above Trujillo (LSUMZ 
22451). 

Population F.~ OLANCHO: 2 km SW Dulce 
Nombre de Culmi (LACM 45179). 

Population G.— GRACIAS A DIOS: Patuca 
(USNM 20282-83); Tancin (LSUMZ 24422). 

Population H.— CORTES: Agua Azul (AMNH 
70157, 70216-17); 2 mi N El Jaral (LSUMZ 24424). 

Population 1. — COMAYAGUA: Comayagua 
(AMNH 70155). 

Population J.— FRANCISCO MORAZAN: Las 
Mesas (AMNH 70156); slopes of Cerro Uyuca 
(AMNH 70204); El Zamorano (AMNH 70153; MCZ 
49889-92). 

Population K. — VALLE: 4 mi N San Lorenzo 
(LSUMZ 24423). 

ACKNOWLEDGMENTS 

We wish to thank the following curators who were 
kind enough to loan us specimens (museum abbrevia- 
tion indicated parenthetically): Hymen Marx, Field 
Museum of Natural History (FMNH); Ernest E. 
Williams, Museum of Comparative Zoology (MCZ): 
Charles M. Bogert. formerly of the American Museum 
of Natural History (AMNH); John W. Wright and 
Arden H. Brame. Jr., Natural History Museum of Los 
Angeles County (LACM); James A. Peters, United 
States National Museum (USNM); James R. Dixon, 
Texas A and M University (TCWC); and Douglas A. 
Rossman, Louisiana State University Museum of Zool- 
ogy (LSUMZ). 



1972 



Till. COIiAI. SNAKE MICRURUS NldKOf IN' M . / HO IDl I 






We ;il so extend our gratitude to John Dickson <ii 
La Lima, Honduras for the opportunity to exa s 

specimens in his personal collection anil to Diana Dec 
Dugas for her help in gathering data. 

LITERATURE CITED 

Giinthcr. A. G. L. C. 1895. Biologia Centrali- 

Americana. Keplilia and Halrachia. London, xx 
+ 326 pp. 

Holdridgc, L. R. 1962. Mapa ecologico de Hondu- 
ras. Organizacion de Los Eslados Americanos. 
San Jose, Costa Rica. 

Meyer, J. R. 1969. A biogeographic study of the 
amphibians and reptiles of Honduras. Ph.D. 
Thesis, Univ. Southern California. 589 pp. 

Neill, W. T. 1963. Polychromatism in snakes. 
Quart. J. Florida Acad. Sci.. 26:194-216. 



Ko/e. J, A. 1967. A check list ..I thl 

venomous coral snal i (Elupid 

lions of new forms. Aim' r '.I 

I 60. 
— . 1970. Mi, mm:. Pp. 196 220, In • 



loguc of the Neotropical Squamata ' i ••. P 
and B. Orejas-Miranda, eds.) Pari I Sr 
Smithsonian Institution Press Wa hington D I 

347 pp. 

Schmidt. K. P. 1933. Preliminary account of the 

coral snakes of Central America and MCXICO 
Field Mus. Nat. Hist.. Zool. Sci . 20 29 40 



1936. 



Notes on ( entral American and 



Mexican coral snakes. Field Mus. Nat. Hist.. 
Zool. Ser., 20:205-216. 

Wilson. L. IX and D. E. Hahn. 1972. The herpeto- 
fauna of the Islas dc la Bahia. Honduras. Bull. 
Florida State Mus.. (in press). 

Accepted for publication May 11. 1972. 



A NEW SPECIES OF ROCK DWELLING DENDROCHIROTE 
HOLOTHURIAN FROM CATALINA ISLAND 

Josephine Yudkin Yingst 1 



Abstract: A new species of the genus Cucumaria Blainville ( 1830) is described from Santa 
Catalina Island. It is a filter feeder and occurs abundantly nestled in subtidal rock reefs. It is 
distinct from other members of the genus described from the Pacific coast of North America 
in its coloration, spicule types, and internal anatomy. 



An undescribed species of Cucumaria Blainville 
( 1830) was found to be abundant in subtidal rock 
reefs at Santa Catalina Island (lat. 33° N — long. 
119°W) which lies off the coast of southern 
California. The species is unique in its habit of 
nestling deep into rock crevices. These cucumbers 
differ greatly in size, color, internal anatomy and 
spicule types from other species of Cucumaria 
known from the Pacific coast of North America, 
namely C. cra.x Detchmann. 1 940, C. curata Cow- 
les, 1906. and C. vegae Theel. 1886. A descrip- 
tion of the genus Cucumaria may be found in 
Deichmann (1941). 



CLASS: HOLOTHUROIDEA 
Order: Dendrochirota 

Genus CUCUMARIA Blainville (1830) 
Cucumaria salma. new species 

Material: Holotype (AHF Echinoderni Collection 
No. 61). total length 6 cm (contracted) and 3 para- 
types (AHF Echinoderni Collection) were collected 
at a depth of 14 m on the north face of Bird f 



1 Allan Hancock Foundation. Universi! 
em California. Los Angeles. Califom 



146 



HI III ll\ Mil lllli;\ ( A 1.1 rOHXt A ACADEMY OF SCIENCES VOLUME 71 




0< 






C. QE,F. G. 



Eigure 1. A, B, supporting rods from tentacles; C, perforated plates from tentacles; D, E, F, 
rosettes from tentacles; G, calcareous ring. 



Santa Catalina Island. The animals were concealed 
in rock crevices and removed by use of a geologist's 
pick. 

Description: Ten dendritic tentacles of equal size 
form a single circlet bordering the buccal membrane. 
Retractile tube feet arranged in pairs along and re- 
stricted to the ambulacra. Ventral and dorsal am- 
bulacra not differing markedly from each other. 

Calcareous ring with distinct posterior projections 
and long anteriorly projecting processes (Fig. 1G): 
interradials and radials fused; anterior processes of 



the radials distinctly cleft. Four Polian vesicles pre- 
sent. Three or four short stone canals, 3—4 mm in 
length grouped near the dorsal mesentery or more 
distantly spaced from each other on the ring. Each 
canal terminates in a slightly elongate madreporite. 

Gonad composed of numerous elongated unbranched 
tubules united in one basal tuft attached to the dorsal 
mesentery. Gonoduct opens to exterior directly behind 
tentacles in central dorsal interambulacrum. 

Spicules smooth, without knobs. Body wall spicules 
with two types of perforated buttons (Fig. 2F-H), and 



1972 



A NEW SEA CUCl \4BER FROM < ATALINA I I 














A, B. C. D E. F, G. 

Figure 2. A, perforated three-armed rod from introvert; B. C. supporting rods from tube feet: 
D, E, perforated plates from tube feet; F, G, H. perforated buttons from body wall. 



perforated rods (Fig. 3F). three-armed perforated 
rods (Fig. 3A, E), and perforated plates (Fig. 3B-D). 
Tube feet with a small number of slender supporting 
plates (Fig. 2D, E) and three-armed rods (Fig. 2B. 
C); when perforated usually a small number of open- 
ings. No indication of end plates. Spicules of intro- 
vert distinct from those of tentacles, resembling those 
of body wall, consisting of three-armed perforated 
rods and perforated plates (Fig. 2A). Tentacles with 
numerous large narrow rods with perforated ends 



(Fig. 1A. B). smaller perforated plates (Fig. lO. and 
very small rosette-like plates with irregular marginal 
processes (Fig. 1D-F). 

Color: Primarily salmon with a concentration of 
black pigment toward the anterior end. Introvert, 
buccal membrane, and tentacles motile. 
and black patches of pigment: tentacul 
black. 

Measurements: Length of rebxe, 1 ; : 
10 cm. 



MS 



HI III ll\ SOI llll K.\ ( Al.IFORSIA ACADEMY OF SCIENCES VOLUME 71 





-■ 






0.5mm 


0.5mm 


Oimm T 




. 


.. 



Figure 3. A, E, perforated three-armed rods from body wall; B, C, D. perforated plates from 
body wall; F, perforated rod from body wall. 



Distribution: Known from Santa Cruz Island, Santa 
Barbara County (Given, pers. comm.) to Point Loma, 
San Diego County (Turner, Ebert, and Given, 1968). 
Occurs at depths of 3 to 20 m in rocky reefs. 

Observations: The aborescently branching ten- 
tacles are highly retractile. The papillae on the 
tentacles project above the epithelium and are 
independently retractile. These papillae are ad- 
hesive and used to collect food particles. 

The tube feet nearest the base of the tentacles 
can elongate to a length of 3 mm. Retracted holo- 
thurians utilize their tube feet to pick up particles 
from the substrate and to transfer them into the 
introverted oral end. When the tentacles are ex- 
tended these anteriormost tube feet may also select 



particles from the substrate and transfer them onto 
the tentacular papillae. 

In the field the animals nestle in the rock 
crevices and extend their tentacles to catch sus- 
pended particulate matter. The body wall is ex- 
tremely plastic, which permits the cucumbers to 
wedge themselves into crevices deeper in the rock 
and away from the rock face. 

Remarks: Cucumaria sahna [salma from Latin 
salmo = salmon] is suggested because of the prin- 
cipal color of the cucumbers. 

Panning (1949, 1951, 1962) revised the genus 
Cucumaria and incorporated into other genera 
several of the species referred to by Deichmann 
(1941) from the Panamic region and by Clark 



1972 A NEW SEA CUCUM HlCli I- ROM CA1 \L1NA 1SLA 

Table I. Comparison of wcsi coasl species in the Menus Cucumarla 






C. curata 



C. crux 



C. vegai 



Up to 2.5 cm; 
ventrum flat 

Unequal 



Few; double row in 
dorsal ambulacrum 



Well-developed 



1/1 /none 

Perf. buttons, 4, 6 
holes, undulated 
margins; plates few, 
20+ holes 

No end plate; 
few supp. rods 



Numerous supp. rods 



Small, 1.5 cm 
(contracted ) 

Equal, bushy 

Large; retractile: 
only in ambulacra 



SIZE 
Small, 1.8 cm; 

ends tapered 

TENTA( l l s 
Unequal 

TUBE FEE l 

Double row; in 
intcrambulacra 



Medium 10 



I qual, bushy 

Retractile; double row; 
only in ambulacra 



Posl prolongations; 

ant processes w/clcft 



CALCAREOUS KING 

Simple; post. Slender; no post, 

prolongations prolongations 

STONE CANAL/POLIAN VESICLES/GKNITAI. PA I'll I \l 

1/2/? 1/1/5 3-4/4 /none 

SPICULES-BODY 
Buttons, 4-8 marginal Few, smooth rods, 1-3 Smooth perf. rods 
holes, not all entirely holes in each end and buttons; plates 

closed 



SPICULES-TUBE FEET 
Rudimentary end Fragment of end plate, 

plate; three-armed or none; no supp. 

supp. rods rods; smooth plates 

w/undulating margins, 
varying no. of holes 

SPICULES-TENTACLES AND INTROVERT 
Tent.; buttons intro.; 
numerous rods w/ 
perf. ends 



Dorsum and tent, 
purple brown; vent, 
and oral disc yellow 

Intertidal, rocks, 
mussel beds, central 
California 



Mottled brown body: 
ambulacra lighter; 
tent, black 



COLOR 

Vent, yellow, gray: 
body ends brown 



DISTRIBUTION 
Sand, west coast Baja Intertidal rock 



California-South 
America 



crevices, north 
Pacific coast 



No end plate: few 
supp. plates: three- 
armed rods w/few 
holes 



Tent.: large rods w/ 
perf. ends; perf. 
plates; minute 
rosettes intro.: three- 
armed rods 

Body salmon: intro. 
and tent, mottled: 
tent, podia black 

Rock crevices. 3-15 m 
deep, southern California 



(1901a, 1901b) from the Pacific coast of North 
America as Cucumaria. Panning (1949) changed 
the species C. dubiosa Semper, 186S. to Pseu- 
docnus dubiosus and in 1964 he changed C. cali- 
f arnica Semper, 1868, to Pseudocmts calif '-amicus. 
Stimpson (1964) changed Cucumaria populifera 
to Pentamera populifera Panning ( 1949). Pavvson 
(pers. comm.) suggests that Cucumaria miniata 
Brandt. 1835, C. hibrica Clark, 1901, and C. 



piperata Stimpson. 1S64. do not belong in the 
genus Cucumaria and probably should be placed 
in the genus Pseudocnus. 

Cucumaria salma is easily distinguished from 
the remaining three species C. cra.x. C. curata. an< 
C. vegae; all of which are found on the 
coast of North America. Cucumaria 
about 10 cm in length: has three or four stone 
canals: four Polian vesicles: is salnv sd: and 



150 



in in n\ soi mi i<\ ( u.iroiisiA academy or sc n:\crs volume 7/ 



has distinct spicules. The killer three species range 
in size from 1.5—4 cm; have one stone canal: one 
or two Polian vesicles; and are gray or brown in 
color i i able I ). 

ACKNOWLEDGMENTS 

I would like to thank D. L. Pawson. Smithsonian 
Institute for his verification of my findings, D. 
Straughan and K. Fauchald, Allan Hancock Foun- 
dation for reviewing the manuscript, and R. Given 
and Douglas Yingst. Allan Hancock Foundation, for 
their assistance in collecting specimens. 

LITERATURE CITED 

Clark. H. L. 1901a. The holothurians of the Pacific 
coast of North America. Zool. Anz.. 24:162-171. 



Part 1. Dendroehirota. 
Exped., 8(31:61-196. 



Allan Hancock Pacific 



Panning. A. 1949. Versuch einer neordnung der 
familie Cucumariidae. Zool. Jb. Abt. Systematik, 
78(4):404-470. 



1951. 



Ober Psciidocnus leoninus (Semper) 
und verwandte Arten. Zool. Anz., 146:73-80. 

— . 1962. Bermerkungen liber die holothurien- 
familie Cucumariidae (ordnung Dendroehirota). 
3 Teil. Die gattung Pseuclocniis Panning. Mitt. 
Hamburg Zool. Mus. Inst.. 60:57-80. 

. 1964. Bermerkungen iiber die holothurien- 

familie Cucumariidae (ordnung Dendroehirota). 
3 Teil. Die gattungen Stereoderma, Stawotiiyone, 
and Trachythyone. Mitt. Hamburg Zool. Mus. 
Inst.. Kosswig-Festschrift., 61:159-174. 



. 1901b. Synopses of North American inver- 
tebrates. XV. The Holothuroidea. Amer. Nat., 
35:479-496. 

Deichmann, E. 1941. The Holothuroidea collected 
by the Velero III during the years 1932 to 1938 



Turner. D. H., E. Ebert, and R. Given. 1968. The 
Marine environment off shore from Point Loma, 
San Diego county. California Dept. Fish and 
Game. Fish. Bull., 140:76. 

Accepted for publication May 3, 1972. 



A NEW SPECIES OF SEA OTTER FROM THE LATE 
PLEISTOCENE OF NORTHWESTERN CALIFORNIA 

Frank H. Kilmer 1 



Abstract: A new species of sea otter is described from a Late Pleistocene marine deposit, 
Humboldt County, California. The species is closely related to the living sea otter, Enhydra 
lutris, but differs significantly in possessing larger P ;! , P 4 , Mi, and presumably Ms, shorter 
ascending ramus with respect to total length of mandible, and a more simply formed angle. 
Proportions of the mandible suggest that the skull of the new species may have been slightly 
longer and lower than that of E. lutris. No special affinities with particular living populations 
of E. lutris are recognized. 



The prior fossil records of sea otter in the north- 
eastern Pacific has, with but one exception, in- 
cluded only postcranial elements and all fossil 
discoveries have occurred in two widely separated 
areas, Oregon and southern California. The Ore- 
gon record consists only of a right femur of 
Pleistocene age from the Elk River Formation 



(Leffler, 1964). The southern California records 
(Mitchell, 1966) include a lower, deciduous tooth 
of Early Pleistocene age from the Timm's Point 
Silt, a right humerus of Late Pleistocene age from 



1 Dept. Geology and Earth Science, California State 
University, Humboldt, Areata, California. 



1972 



NEW FOSSIL SEA OTTER FROM CALIFOl 






i he Palos Verdes Sand, and various limbs and 
vertebral elements from Late Pleistocene deposits 
of Santa Rosa Island, Leffler referred the Oregon 
femur u> Enhydra sp. while Mitchell concluded 
thai all the southern California specimens probafelj 
were referable u> Enhydra lutris (Linnaeus). 

In May 1967, a well preserved mandible of a 
fossil sea oiler, lacking only a lew teeth, was 
recovered from a Late Pleistocene marine deposit, 
located on the Humboldl County coast, north- 
western California. The mandible is evidently the 
first known of a fossil sea otter from ihe north- 
eastern Pacific and il provides the first major 
indication of (he cranial morphology of fossil sea 
otters. The Humboldt County mandible is inter- 
preted to be that of a new species of sea otter 
referable to the genus Enhydra. 

The left and right rami of the mandible were 
discovered at the same locality by different persons 
at different times. The close morphologic agree- 
ment of the two rami and near perfect fit of the 
symphysis make it virtually certain that they are 
from the same individual. 

GENUS ENHYDRA FLEMING, 1822 

Enhydra macrodonta, new species 

Figures 1 and 2 

Holotype: California State University, Humboldt. 
Department of Geology and Earth Science, specimen 
No. 745-1, a mandible lacking left P», M,, and M 2 ; 
right incisors and M-. The medial projection of both 
condyles has been broken off. 

Type locality: California State University, Hum- 
boldt, Department of Geology and Earth Science, 
locality No. 745. A moderately steep, north-facing, 
excavated slope located approximately 1 1 miles north 
of Areata. Humboldt County, California, at the junc- 
tion of U.S. 101 and Crannell Road. The exposure 
consists of three, nearly flat-lying, sedimentary units 
including (from bottom to top): 1) a basal 12 feet 
of gray, medium-grained, pebbly, fossiliferous. semi- 
consolidated sand: 2) 50 feet of gray, clayey, fine- 
grained, fossiliferous sand: and 3) 70 feet of light- 
brown, medium-grained, semi-consolidated, pebbly 
sand. 

The rami were found, as float, lying on the exposed 
surface of the gray clayey sand unit. The left ramus 
was found in a narrow gulley at the foot of the ex- 
cavated slope about 230 feet east of the centerline 
of Little River Road (the frontage road for U. S. 101). 
The right ramus was found directly upslope from the 
left ramus a distance of approximately 25 feet. The 
gray, clayey sand matrix adhering to both rami is 
identical with the lithology of the gray clayey sand 
unit and it is virtually certain that the two were eroded 
from this stratigraphic unit. 










n 






' f 


s 


H 










M 2 


/Ml 


p 4 


P 3 P 2 c 


-H 2 







H 

Figure I. Key to mandibular measurements utilized 
in this study and presented in Table I. Dimensions 
A, B, and D are measured from the posterior margin 
of the condylar process along the axis of the ascend- 
ing ramus. AH measurements are in millimeters. 



Formation: The otter-bearing unit has not been 
formally described or named and it can be definitch 
traced only about % mile south of the Crannell Road 
locality. Its relationship with the well known Ceno- 
zoic sedimentary sequence (Ogle. 1953) exposed in 
the Eel River Valley 40 miles to the south is not 
known. However, reconnaissance mapping and in- 
direct paleontologic evidence suggest that the otter- 
bearing unit may be older than the Hookton Forma- 
tion (Late Pleistocene) and either younger or partially 
equivalent to the Carlotta Formation (Plio-PIeisto- 
cene): these formations are in depositional contact 
with each other in the Eel River Valley sequence. 

Age: Late Pleistocene (see discussion below). 

Diagnosis: Relative to Enhydra lutris. the only 
other species included in the genus: horizontal rami 
thick, occlusal outline of P.-. P,. and NT approximately 
1 .-i larger and plump: Ms presumably slightly larger 
based on slightly greater length and width of the 
alveoli: and ascending ramus short with respect to 
length of the mandible: angle sharply rounded, right- 
angled, with ascending and horizontal margins narrow 
and rounded. 

Description and comparison: The close corre- 
spondence between the fossil mandible and the 
of the compared specimens of Enltydi 
(Table 1) with respect to size, shape, and der 
morphology indicates that E. macr 
related to E. lutris and is certair 



152 



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NEW FOSSIL SEA OTTER FROM < u iiaKNlA 











Figure 2. A, Enhydra macrodonta, new species: joined left and right mandibular rami, occlusal view (CSUH. 
Dept. of Geology and Earth Science, No. 745-1). B, Enhydra lutris (Linnaeus); mandible, occlusal view (U.S. 
Geological Survey, No. 6801, adult male, Asilomar State Beach. Monterey Co.. California, captured 14 April 
1966). C, Enhydra macrodonta, new species; right mandibular ramus, lateral view (CSUH. Dept. of Geology 
and Earth Science, No. 745-1 ). D. right mandibular ramus, medial view (CSUH. Dept. of Geology and Earth 
Science, No. 745-1). Scale lX- 



the genus Enhydra. The presence of permanent 
dentition coupled with the lack of extensive wear 
suggests death at an early adult stage of life. 

Although the gross morphology of the E. mac- 
rodonta closely resembles that of E. lutris it differs 
significantly from specimens of comparable or 
even greater jaw length in that the horizontal 
ramus is more robust and strongly built, to accom- 
modate the larger teeth, and the toothrow is pro- 
portionately longer. The medial surface of the 
jaws tends to be rather flat and featureless as in 
E. lutris but the lateral surface differs in being less 
concave below P H -P| an d substantially more con- 
vex below M, and M 2 . There are two large, oval- 
shaped mental foramina on each ramus below P 3 
and P 4 on the lateral surface. These foramina 
are larger than the mental foramina seen in E. 
lutris but their location is essentially the same. 

The ascending ramus is more lishtlv built, more 



slender, and shorter than those of E. lutris of 
comparable jaw length. The lateral and medial 
surfaces of the ascending ramus are essentially 
parallel and without the surficial irregularities 
typical of E. lutris. The anterodorsal margin of 
the coronoid crest is narrow and slightly curled 
medially whereas, in E. lutris the margin is more 
strongly curled and thicker. When the toothrow 
is placed in a horizontal position, the anterior 
edge of the ascending ramus rises at a steep angle 
and the posterior margin is nearly vertical. In 
this position the coronoid crest does not project 
posterior to the condylar process although it over- 
hangs it slightly. 

The masseteric fossa is a deeply excaval 
depression which shallows anteriorly, dimir 
to an arrowhead-shaped point. Tr. 
directed anteriorly, not downward as is common 
in E. lutris and terminates jus- b 



/.w 



HI III ll\ sol THERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 



half of M|. In / . lulris the anterior extremity of 
the masseteric lossa almost invariabl) lies below 
M_. and occasional!) lies below the posterior mar- 
gin oi M,. The condyles are about the same size 
ami diameter as the largest specimens of E. lutris 
examined and the) are connected to the posterior 
margin of the ascending ramus by strongly built 
gussets of bone. The angle consists of a narrow, 
sharply rounded corner, the posterior and ventral 
sides ol which form an angle of about 90°. The 
backward-pointing, plate-like structure, which 
commonly forms the base of the angle in E. lutris, 
is absent in the Humboldt form. 

The horizontal angle formed by the medial 
surfaces of the mandible, when the jaws are placed 
in the position of "best fit" at the symphysis, 
measures approximately 63°. Although the mag- 
nitude of this angle must be considered only an 
estimate, since the jaws were not found articulated, 
it is close to a number of adult specimens of E. 
lutris in which the same angle ranged from 56° 
to 60° (Fig. 2A, B). 

Although the toothrow in E. macrodonta is 
significantly longer than in those specimens of 
E. lutris having comparable jaw length (Table 1), 
the teeth in the fossil are spaced and individually 
oriented in the jaws almost identically to those 
of the living species. The line of teeth form a 
fairly well-pronounced arc which bows outward 
in occlusal view. Whether this curvature is a 
reflection of growth stage or is a constant charac- 
ter is not certain; in adult E. lutris curvature of 
the toothrow also occurs but it seems to be much 
less pronounced (Fig. 2 A. B). 

The Mo is lost in both left and right jaws of 
E. macrodonta but the alveoli are well-preserved 
and their size (Table 1) suggests that M 2 was 
slightly larger than that found in E, lutris. The 
alveolus of M 2 lies in a large, rounded bulge which 
swells medially from the base of the anterior edge 
of the ascending ramus. The alveolus is positioned 
considerably higher than that for the M 1 . Its 
shape and orientation suggest that it was occupied 
by a single-rooted, ovate tooth, whose long axis 
was oriented transversely to the jaw and whose 
occlusal surface tilted forward at a moderate 
angle. The general orientation and position of the 
alveolus in E. macrodonta closely resembles that 
found in specimens of E. lutris in which the growth 
stage is considered to be late juvenile/ early adult. 

The Mj is a bunodont tooth with the cusps 
arranged in a pattern almost identical to that of 
E. lutris. The tooth width and the distance be- 
tween the protoconid and metaconid are, however, 
proportionately greater, and the occlusal area is 



about ':i larger. The alveolus for M ( in the left 
jaw is figure-eight shaped in occlusal outline. Near 
its center there are distinct inward projections of 
bone, one of which connects to a centrally located 
transverse ridge. This suggests that the alveolus 
was divided into anterior and posterior sockets as 
is standard in E. lutris. 

The P.j of E. macrodonta is approximately % 
larger in occlusal area than that of E. lutris. The 
constriction of the root, evident below the enamel 
base and above the alveolar margin, suggests that 
the tooth possesses two roots as in E. lutris. The 
occlusal morphology of the P., closely resembles 
that of E. lutris except that in E. lutris the ridge 
radiating posteriorly along the lateral side of the 
tooth is larger and developed into a more distinct 
cusp-like structure. 

The P s in E. macrodonta has nearly twice the 
occlusal area of that in E. lutris. It is about % the 
size of P 4 but is similar in occlusal