Woe We yeti fy) Sf) a ee Dan Oo ae q F i t ; a? , -_ . 7 — 4 ca _- i , : . os ey i x Te 7 } 7 j i i ; a ni , : fa y Ty Migs : ! B F i ; I i i in v i a a - ' - ei gua ; = ': i ie ir ‘ 7. Pa: ; SOUTHERN CALIFORNIA ACADEMY OF SCIENCES BOLLETIN Volume 70 Number 1 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. Annualviembersmaese eran Students Membersigemis see cine ieee ILS IMINO Cosasuuccsdocsesbenee 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 AuGustT 27, 1971 NUMBER | 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 ............. A new species of Simopelta from Costa Rica (Hymenoptera: Formicidae). By Roy RAM STLCLULFI Omen Mts ca ra ts ee nach s Pan eters mOlacnt oh notes Sdede ia aise a cadena vosedevial shee say Sintas Cee A revision of the genus Loandalia Monro with description of a new genus and species of pilgargiid polychaete. By Raymond R. Emerson and Kristian REDO Wis etrats riche CHENG Otc: OO cso Ge DOG PCR OR CRN Pet acre UES Taxonomy and distribution of the Arctic species of Lucicutia (Copepoda: Cal- fanoida) By Julio Vidal: Mehta eke c. chaaerae oe otetel cere enciaee lake chegetel leaoeene A review of the genus Boccardia Carazzi (Polychaeta: Spionidae) with descriptions of two new species. By James A. Blake and Keith H. Woodwick ......... Parasitic mites of Surinam. VIII. A new genus and species of chigger, Fauranius atecmartus, and additional records of species (Acarina: Trombiculidae). By amesVEBLeEnnaiy and Hale uKOSCAUS mea itachi cine icici nearer: Variation and geographic distribution in some Argentine and Chilean Osmylidae, with a new species of Kempynus (Neuroptera). By Phillip A. Adams ...... RESEARCH NOTES Distribution of some southern California Kangaroo rats. By Blair A. Csuti ..... An evaluation of two methods of measuring metacarpal length in Artibeus lituratus (Olfers) (Chiroptera: Phyllostomatidae). By Donald R. Patten ........... A new synonymy in the genus Chilicola (Hymenoptera: Colletidae). By Roy R. ISHICLLIN QUE er re era ak MUA ater ct tories sia si ecosb stro ene Graccesorshat ct sorekee eites pene eval eiieree Cuvier’s beaked whale, Ziphias cavirostris, from Barbados. By David K. Caldwell, Warrenvl. Rathjen, and Melba‘G: Galdwell). =.=... 40. - 44-25-2556": Additional specimens of the colubrid snake Amastridium veliferum Cope from Costa Rica, with comments on a pseudohermaphrodite. By Larry David Wilson:and’Douclasi CV Robinson aaa. ee eee er one sie ee INEWSTANDBINOTES wees eto yaa cay arco ers ea cee as Gace oe a ose Sabi oy cue ieee 23 31 42 45 50 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! 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, 196S5a: 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 (196Sa, Fig. 8.1). He then assigned a numerical weight of | 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” (196Sa, Table 8.1). He also presented these weights in two large tables (1959a, Tables I and II) with 189 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 1Department of Biology, California State College at Long Beach, California 90801 11971 shave a practical advantage over long term studies in that both the earlier and the later surveys can be ‘made by the same observer, jpopulations before and after, 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. Boney (1968) investigated the effects of various detergents on Ascophyllum, Cladophora, Bryopsis, Prasino- cladus, Porphyra, Acrochaetium, and Polysiphonia. ‘Okamura, ef al. (1926) correlated heavy mortality jof intertidal Porphyra in Tokyo Bay with episodes lof heavy fog. They demonstrated experimentally \the lethal effect of soot particles from the fog con- jtaining 70% to <0.7% adsorbed sulfur dioxide and jalso of air containing more than 0.01% free SO. | 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- jment of new techniques that the person who comes jto make the second part of the study should be dissatisfied with the first. In this context, it is perhaps jironic that the paper by Okamura, er al. (1926) ishould 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 a larger set would be affected by the RESON ‘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 jalong 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 Conclusions derived from the observations of SOUTHERN CALIFORNIA ALGAE =f work, were used by the author to check the identi- fications made. These specimens were deposited in the herbarium of the California State College at Long Beach. Taxonomic problems, — Many intertidal algae 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. Ulva 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 (196Sa, 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 a) @ Reoccupied 1968-70 © Best 1956-9 * Others BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 Los x Angeles Verdes 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 particular 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 of 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 in 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 stations 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 assignec to high interfidal, mid-intertidal, low intertidal, o1 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 1971 60 50 OF SPP. eo > ° ° NO. 20 19 18 2 MALIBU SeMONICA 7 33 PALOS VERDES SOUTHERN CALIFORNIA ALGAE 5 16 LONG BEACH 9 21 ORANGE CO. STATIONS 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 (1961). 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 during the summer 6 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 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- VOLUME 70 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. Corallina vancouveriensis 400 Gigartina canaliculata 400 Bossiella WwW c fe} oO n J 10000 _MPN/cm?2 coliform bacteria in bottom sediments No maximum month MONTH SOUTHERN CALIFORNIA ALGAE /] 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). obtained at certain of 106 sampling stations. Off the Orange County outfall (Rittenberg er al., 1958, Fig. 1) 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 1.06 +0.9 South Laguna 0.6 0.56 =O Laguna Beach 1.0 2.05 +1 Orange County 55.0 119¢ +64 White’s Point (3) 277.0 3609 +83 Hyperion (effluent) 275.0 340d +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). dHertel, pers. comm. 12 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 TABLE II. Description of intertidal sampling stations. 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 =) —12% 9 11 11 4 3 2 10 Big Rock Beach as) — 15% 10 9 10 3 2 2 2 Topanga Canyon ti $ — 44% 5 10 9 2} 3 2 1 Sunset Blvd. =7/ — 48% 2 1 8 I 4 2 7 Flat Rock Pt. 5 — 40% | 6 5 5 I 2 25 Lunada Bay a5} 3290 6 1 4 6 3 3 33 Portuguese Pt. +8 +17% 15 15 2 1] 1 2 17 White Pt. 0 + 16% 11 3 1 10 3 2 16 Pt. Fermin (Cabrillo Beach) 414 —23% 4 4 3 9 2 2 9 (Little) Corona del Mar — I? — 43% 3 5 6 12 2 2 42 Crystal Cove +8 —3% 13 13 1 13 1 2 21 Laguna Beach +6 — 16% 8 2 15 14 2 2 22 Dana Point North s (Salt Creek) =? +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 1956-9 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, Gastroclonium, and Gracilaria/Gracilari- 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 L.A. Airport San Pedro SOUTHERN CALIFORNIA ALGAL 13 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 crinale/coulteri, and Pterosiphonia 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 crinale/coulteri 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 vancouveriensis/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 vancouveriensis/gra- cilis has a similar effect on the March group, while both Pelvetia and Rhodoglossum affine have @ 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 OF SCIENCES HYPERION 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, 1960, 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 VOLUME 70 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 (196Sa: — 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 luxuriance during the winter and spring and an increase during the 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 1). 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 ja 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 _ damaged plants could be showing the effects of | pollution and when they are simply the effects of | drying winds. » It would also be advisable to use species which ‘appear to have decreased in response to pollution, SOUTHERN CALIFORNIA ALGAE 15 rather than those which have increased, since there are so few of the latter. Gastroclonium coulteri and Chaetomorpha aerea are two species which are easily identified, have little seasonal variation, and have shown a consistent decline both before 1956 and after. Their decline could perhaps be used as a general measure of ‘pollution’. Pelvetia Rhodoglossum affine, Bossiella spp., fastigiata, Gigartina armatalspinosa, and Gelidium pulchrum|purpurescens 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 313 pp. Allan Hancock Foundation. 1965. Chapter 1, Introduc- tion /n 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.) 16 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 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: 1170-1173. Boney, A. D. 1968. Experiments with some detergents and certain intertidal algae. /n J. D. Carthy and Don R. Arthur, Ed., The Biological Effects of Oil Pollution on Littoral Communities. Field Stud. Council 2 (Suppl.) pp. 55-72. California Air Resources Board. 1969a. California Air Quality Data for September, October, November 1968. Vol. | 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. J. Appl. Ecology 6: 133-142. Dawson, E. Yale. 1959a. A primary report on the benthic marine flora of southern California /n 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. /bid. pp. 219-264. . 1961. A guide to the literature and distributions of Pacific benthic algae from Alaska to the Gala- pagos Islands. Pacific Sci. 15: 370-461. VOLUME 70 . 196Sa. [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. Table IX. /bid. Appendix Data pp. 351-438. Emery, K. O. 1960. 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 | 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. 231 pp. Stevenson, R. E. 1959. The marine climate of southern California Jn 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! 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 S. 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 1Natural History Museum of Los Angeles County, Los Angeles. California 90007. observations on the behavior of S. oculata Gotwald and Brown made by Brown in Costa Rica, These observations reaffirmed the army antlike behayior of the foraging workers. The following new species 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 | 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 S. 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 S. williamsi, those of the occiput and pronotum, especially, are decidedly transverse; in | S. 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; ~CI71-81. Color, basically blackish brown, mandibles, fla- ' gellum, legs brownish. Pronotum with brownish _ areas of irregular extent. Gastric apex light brownish. Holotype and 21 paratype workers, 4 mi south of San Vito de Java, Puntarenas Province, Costa 5 1971 NEW SPECIES OF ANTS 17 SO 1 mm Figure 1. 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 S. 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, S. 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 LOANDALIA MONRO WITH DESCRIPTION OF A NEW GENUS AND SPECIES OF PILARGIID POLYCHAETE! RAYMOND R. EMERSON? AND KRISTIAN FAUCHALD2 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 Muller (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 OF'SPINES oS ae te Qe erento 2. Notopodia without emergent hooks OFiSpimes) 5 \e.3)5.A0 ees ec ae eer eRe ote 1. 2. Notopodia with recurved emergentshooks'2). \) seiner 3h 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) 05.4554. eee eee Cabira — Dorsal and ventral cirri well developed, parapodia well developed, body dorso-ventrally flattened ................ By | 5. Antennae shorter than palps; integument papillated .......... Ancistrosyllis Antennae longer than palps; integument:smooth’ |. 2) ae Sigambra 6. Prostomial antennae and peristomial cirri present, parapodia sharply set off | fromitheibody* 3:4 52). see eee Synelmis Prostomial antennae and peristomial cirri absent, parapodia distinct, but not set off from the body ....53.Saa000 Parandalia 7. Prostomial antennae and peristomial CIPEI absent. iii .;2. soe ee ee eee Loandalia Prostomial antennae and peristomial Nadya sadaddawoloocgesoseccuede 8. 8. Prostomium with two antennae, biarticulate palps present ............ Pilargis Prostomium with three antennae, palps without palpostyles ............ Otopsis 1Contribution from the Allan Hancock Foundation. 2Allan Hancock Foundation, University of Southern California, Los Angeles, California 90007 18 1971 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, biarticulate palps with button- shaped palpostyles 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 eversed and may terminate with several sensory papillae. The proxi- ‘jmal surface of the proboscis is smooth. The first parapodia are uniramous, further back all parapodia are biramous. A thick, stout, straight, crystal-clear spine emerges from the median and posterior notopodia. All notopodia are poorly developed and have at most one or two capillary setae in addition to the spine. Neuropodia where fully developed, are slightly tapering with blunt, jevenly rounded tips. Dorsal cirri are absent; ventral jcirri are present as small, papillar lobes. Branchiae The neuropodia have several geniculate, pointed, simple setae. Each seta has a cylindrical stalk and a 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 aberrans Monro (1936) showed that notopodial spines are absent and that branchiae are well developed in the posterior part of the body. In addition, the two first neuropodia have thick, black Jacicula; a feature not found in any other described pilargiid. The species presently assigned to Paran- dalia include: P. americana (Hartman, 1947); P. gracilis (Hartman-Schréder, 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. oculafis, new species. Key to Species of Parandalia IPAllparapodia biramous ................ 2: Anteriormost parapodia uniramous, ekothersibinamOuse . sashes cece see kaa 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 | NEW SPECIES OF POLYCHAELTE 19 3. Eyes present; six-seven neuropodial setae in each parapodium ...... P. ocularis Eyes absent; twelve-fifteen neuropodial setae in each parapodium ..... ; 4, 4. Emergent notopodial spine from SCLISCI EZ enn te natalie cen tea tte P. americana Emergent notopodial spine from SOUIDENPorcriny sieresaogs eect sete exe ars eke P. fauveli Parandalia ocularis, new species Material examined: AHF VELERO 12856, April 3, 1969, 34°23’12"N, 119°38’18" W, 24 fms silty- clay, gear: Campbell grab, paratype (1). AHF VELERO 12860, April 3, 1969, 34°22'58”"N, 119°38’05"W, 23 fms, silty-clay, gear: Campbell grab, HOLOTYPE. AHF VELERO 12864, April 4, 1969, 34°21’15” N, 119°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 dorsolaterally 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 biarticulate 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 OF SCIENCES Figures 1-3. Parandalia ocularis, new species. Figure 1. Anterior end in dorsal view x50. Figure 2. Para- podium 14 from left side in anterior view x100. 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 | 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). VOLUME 70 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 neurosetae 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 flat 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 POLYCHALTE 2/1 Figures 8, 9. Loandalia aberrans, Monro. Figure 8. Distal end of notoseta x950. Figure 9. Posterior end in dorsal view x50. near San José Light, Guatemala in 12-13 fms. P. indica Thomas (1963) is known from the West Coast of India from muddy substratum in 7-10 fms. P. gracilis Hartman-Schréder (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°13’00" 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 bars on each side of the notopodia in median and posterior setigers, The prostomium is short and wide; prostomial palps are large compared to the size of the pros- tomium. A single peristomial segment is present; peristomial appendages are absent (Fig. 7). The 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 | 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 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 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. VOLUME 70 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., 10: 383-445. Ditlevsen, H. 1917. Annelids, I. The Danish Ingolf Expedition, 4: 1-71. Fauvel, P. 1923. Polychétes 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-Schréder, 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: 499-511. Monro, C. C. A. 1936. Polychaete worms. Discovery Reports, 12: 59-198. Miller, F. 1858. Einiges tber die Annelidenfauna der Insel Santa Catharina an der brasilianischen kuste. Arch. Naturg., 24: 211-220. 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. Annélides polychétes 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(1 ): 23-30, 1971 TAXONOMY AND DISTRIBUTION OF THE ARTIC SPECIES OF LUCICUTTIA (COPEPODA: CALANOIDA) JULIO VIDAL! AbsTRACT: The taxonomy and distribution of the Arctic species of Lucicutia were studied. Analysis of approximately 400 plankton hauls from the Arctic Ocean revealed the presence of Lucicutia polaris Brodsky, L. pseudopolaris Heptner, and L. anomala Brodsky in samples collected in deep waters. The two former species are closely related but 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 Artic 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 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, 2000-3000m, 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. i) uy 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 (2 only); Hulse- mann, 1966 (d 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 Ist exopodal segment reaches the distal third of the outer spine on 1Department of Biological Sciences, University of Southern California, Los Angeles, California 90007 Present address: Department of Oceanography, University of Washington, Seattle, Washington 98105 24 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 90° Lucicutia pseudopolaris L: pseudopolaris & L. polaris L. anomala 90° 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 11 to 15 times longer than wide. The antennules, 24-segmented, extend beyond the furca by the last 2-3 segments. Segments | 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 maxillule has 14 spines on the Ist inner lobe; 3 setae each on inner lobe 2 and 3; 3 setae on the basipodal segment 2; 9 setae on the endopodite; 11 setae on the exopodite and 5 setae on outer lobe |. The first pair of legs (Fig. 2f) has 3-segmented exopodites and 2-segmented endopodites. The outer spine on exopodal segment | 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 | 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 | 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 | to 4 of the metasome are as those of the female. The basipodal segment | of the right 5th leg (Fig. 21) is longer than the corresponding segment hoomm, — 9,b,),k ose mM, 6.d,0.9 ogomm, Ad vd-d.+) | Figure 2. Lucicutia 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 ) views 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 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 | outer subapical ‘spine. The left side of the fifth pair of legs has a 3-segmented exopodite and endopodite. Exopodal segment 3 has | outer, apical, and inner spine. Copepodite Stage V. Body shape (Figs. 4), 1) is as the adult. The cephalosome and metasomal segment | 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. ARCTIC COPEPODS 25 The fifth pair of legs has 2-segmented endopodites and exopodites in both sexes (Figs. 4k, m). The distal segment corresponds to fused segments 2 and 3. In the male the right side of leg 5 is shorter than the left. Copepodite Stage IV. The specimens of this stage have a 3-segmented abdomen (Figs. 4e, f). Exopodites and endopodites of legs 1-4 have 3 segments (Fig. 4g). The fifth pair of legs has 1- segmented endopodites and exopodites. Distribution. L. pseudopolaris has a wide geo- graphical distribution in the Arctic Ocean. It has been recorded by Heptner (1969) from an area between 76°38’ N and 81°12’ N and 173°44’ Eand 189°35’ E, and at 88°10’ N 82°22’ W. In our collections it was found between 72°42’ N and 83°27’ N and 9°15’ W and 157° 12’ W. Lamm, abijmng ose mm ccdeg o2zemm, fhkopr 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 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Lucicutia polaris Brodsky, 1950 Figures 3a-r, 4a-d. Lucicutia polaris Brodsky, 1950 (2 only); John- son, 1963; Heptner, 1969. Not Lucicutia polaris Brodsky, 1950; mann, 1966; Grice and Hulsemann; 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 | 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 Hulse- VOLUME 70 the adults. In L. 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 L. pseudopolaris (Table 1). Distribution. L. polaris has been recorded from the central part of the Aretic 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 173°36’ W. L. polaris has also been reported from the Indian Ocean and southeastern Pacific Ocean (Htlsemann, 1966; Grice and Htilsemann, 1967). These records probably correspond to species from the southern Pacific and Indian Ocean rather than to the species L. polaris described by Brodsky from the Arctic Ocean. Lucicutia anomala Brodsky, 1950 Figures 4n-s. Lucicutia anomala Brodsky, 1950; Grice and Hulsemann, 1965; Huilsemann, 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, 0) 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 I-2, 19-21, and 22-23 are fused. The right antennule reaches the end of the furca. The Ist 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 | 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 11 small teeth (Fig. 4s); exopodal segment 3 bears | apical and | inner spine. Distribution. This species has been recorded from the Arctic Ocean (Brodsky, 1950) and north- eastern Atlantic Ocean (Grice and Hulsemann, 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 _ 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 | 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 slightly 1971 ARCTIC COPEPODS 27 TABLE I, Measurements and length ratios of the Arctic species of Lucicutia PROSOME-UROSOME ABDOMEN-FURCA TOTAL LENGTH (MM)* LENGTH RATIO LENGTH RATIO N Mean Range St. Dev. Mean Range St. Dev. N Mean Range | Lucicutia pseudopolaris | Adult female Si) 4.277 3.81-4.75 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 hay sere i 1.56-2.00 0.114 18 0.98: 1 0.88-1.10 ‘Copepodite ‘Stage V female ON Sh2i7) 3.10-3.56 0.148 2.11: 1 1.90-2.27 0.127 9 1.04: | 0.91-1.20 | Copepodite ‘Stage V male USAT 3.13-3.41 0.089 2.07: 1 1.86-2.3 1 0.135 7 1205: 1 0.94-1.18 | Copepodite Stage IV 3) 2255 2.46-2.64 0.064 2.32: 1 2.26-2.41 0.055 5 1.08: | 1.04-1.11 | 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 DP DES 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 50:85271 0.81-1.04 | Lucicutia anomala Adult female Die SOM | 2.432258 10043" 1G75:a 1.62-1.85 0.058 9 0.74:1 0.66-1.06 Adult male Ue 2e 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 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 CALIFORNIA ACADEMY OF SCIENCES TABLE II. Rate (%) of incidence of suctorians on the species of Lucicutia L. polaris L. pseudopolaris L. anomala Female 65 11 . 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 (1963) 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 VOLUME 70 organic matter in the sea (Parsons and Strickland, | 1962; Riley, 1963; Menzel and Goering, 1966; Jgrgensen, 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 10u in L. pseudopolaris and L. polaris — and slightly smaller than 5u in L. anomala. This fine mesh and the structure of the oral appendages / 0.50 mm__ 1.0 wwomm. a.befil wo mm. p,s 20mm, c.d,g.h,j- m.q.r —=——S SS aA ws Figure 4 — Lucicutia polaris Brodsky. Stage 1V 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 60 50 40 30 20 Number of specimens Percentages of relative abundance 1000 1000- 1500 1500- 1500 2000 Depth | 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 In to several mm, and on small organisms of the nannoplankton. The absence of morphological differences in the structure of the oral appendages (especially between L. pseudo- qpelaris and L. polaris) suggests that the above species are feeding on the same or ona very similar 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 spp; Cyclothone spp; Stomias spp;) and gave exam- dles of vertical exclusion. Other examples of sharp in ARCTIC COPEPODS 29 Lucicutla pseudopolaris } Lucicutia polaris Lucicutia anomola 2000 2000- 3000 3000 below 3000 meters Figure 5 — Percentages of relative abundance and number of specimens of the arctic species of 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 SCIENCES 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: 116-126. Banse, K. 1964. On the vertical distribution of zoo- plankton in the sea. Progress in Oceanography, 2255-125: Brodsky, K. A. 1950. Veslonogie rachki Calanoida dal’nevestochnykh morei SSSR i polyarnogo bas- seina. Opred. Po Faune SSSR, Akad. Nauk SSSR, 35: 1-441. Coachman, L. K. 1963. Water masses of the Arctic. /n, 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. /n, Proc. Arctic Basin Symp., October 1962, Arctic Inst. N. Amer., p. 184-196. Grice, G. D. and K. Htilsemann. 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. Mus., 122: 1-67. VOLUME 70 Heptner, M. V. 1969. Systematic status of Lucicutia 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. Jgrgensen, 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. /n, 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. |The present paper is fourth in a series on Boccardia jtaxonomy. The first three (Woodwick, 1963a; /1963b; Blake, 1966) dealt with B. tricuspa, B. Jeolumbiana, B. proboscidea and B. hamata. Adults were described and observations on ecology record- fare described. Boccardia berkeleyorum comes from jcentral California while B. chilensis comes from | the coast of Chile. In addition, B. ligerica, a characteristics of Known species of the genus are summarized in ‘Table I. Analysis of the critical characteristics | ceveals that several well-defined groups of species “are present. Discussion of these relationships follows | he species descriptions. | Specimens of Boccardia berkeleyorum were ‘collected by us during the years 1961-63. The B. | chilensis material was included among the collec- ions of the Lund University Chile Expedition 1948-49, the sedentary polychaetes of which are ow at the Allan Hancock Foundation. We are grateful to Dr. Kristian Fauchald for the loan of | his material. Specimens of B. ligerica were kindly sent by the Zoological Museum of Amsterdam _ZMA) and Dr. Francois Rullier of the Universite de 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 “iis 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 and will be published in a subsequent paper. } SYSTEMATICS Genus Boccardia Carazzi, 1893 Type species: Boccardia polybranchia (Haswell) 1885. Synonyms: ?Perialla Kinberg, 1866. BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(1 ): 31-42, 1971 A REVIEW OF THE GENUS BOCCARDIA CARAZZ]I (POLYCHAETA: SPIONIDAE) WITH DESCRIPTIONS OF TWO NEW SPECIES JAMES A, BLAKE! AND KEITH H. Woopwick? ABSTRACT: Taxonomic characteristics of 14 species of Boccardia are compiled in the 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 B. chilensis are described. Diagnosis: Prostomium rounded or bifid 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, 1938) . B. ligerica Ferronniere, 1898 (* Polydora redeki Horst, 1920) . B. natrix (Sdderstrom), 1920 . B. perata (Khlebovitsch), 1959 . B. polybranchia (Haswell), 1885 . B. proboscidea Hartman, 1940 (?Spio californica Fewkes, 1889) (?Polydora californica Treadwell, 1914 HOMONYM) 11. B. pseudonatirx Day, 1961 12. B. tricuspa (Hartman), 1939 13. B. truncata Hartman, 1936 14. B. sp. fo) nA BW th owoanan \Pacific Marine Station, University of the Pacific, Dillon Beach, California 94929. 2Department of Biology, Fresno State College, Fresno, California 93710. 32 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES (Polydora redeki sensu Okuda, 1937, not Horst, 1920) Boccardia ligerica Ferronniere Emended Figure | 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 11, 1932, ZMA 1156 (30 specimens); France, Benouville, April 25, 1953, coll. Frangois 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 | 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. 1i). 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 | 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. 1h). 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. 1b) alternating with lanceolate companion setae (Fig. |c-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 VOLUME 70 ventral to it is a small tuft of pointed geniculate setae (Fig. If). 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 (1920). His figures clearly show branchiae on setigers 2, 3, -, -, -, 7. Fauvel (1927) repeated Horst’: 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. Rullier (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 figure of the pygidium are like Ferronniere’s figure 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: proved that specimens from the two localities wer the same. Only the material from Holland contain: complete specimens. The anterior ends of thi specimens from the two localities were identical Branchiae were limited to setigers 2, 3, -, -, -, 7 197] NEW SPECIES OF BOCCARDIA Figure 1. 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. 33 34 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 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 Horst (1920), Ferronniere (1898) and Rullier (1960) 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. Muller) 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 VOLUME 70 Edith and Cyril Berkeley who have made significant 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 1; 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 . . NEW SPECIES OF BOCCARDIA FF EEZE- ess az gr 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. shy 36 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 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 siniple falcate structure, but if viewed froma 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 Polydora. The species, P. quadrilobata, P. caulleryi, and P. armata 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 (11 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- TYPE and 2 PARATYPES); Sta. 131, July 1, 1949, Iquique, southern part of town, exposed rocky VOLUME 70 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° 1019" W (2); Bahia de Concepcion, central part, SE of Isla Quiriquina 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 NEW SPECIES OF BOCCARDIA 37 l Figure 3. 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: i h, hooded hook from a far posterior setiger; i, posterior end in dorsal view. 38 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES TABLE I. Some Taxonomic Characteristics of the Known Species of Boccardia Species af Boccardia basilaria berkeleyorum chilensis columbiana hamata ligerica natrix Prostomial n a b Cc. Crt? Ge » T # » » Cae uchal ridge . prostomium . caruncle eyes incised to end of setiger 3 4 rounded to end of setiger 4 .0 incised into setiger 2 0 - rounded to end of setiger 3 4 incised to end of setiger 3 .4 weakly incised to end of setiger 2 .4 incised . to end of setiger 2 .4 VOLUME 70 Spines of Branchial Setiger 5 distribution Notosetae Posterior a. number of a. anterior on Notopodial types b. posterior Seliger | Spines b. description of types Ap2,7345-10 a. two b. absent b. 1. falcate posterior absent absent 2. bristle- one-half topped with constricted neck a. 2,3,4,-,6 a. two b. absent b. 1. falcate posterior absent acicular 2. bristle- one-fourth topped with accessary tooth a. 2,3, 4,-,6 a. two b. present present absent b. 1. falcate 2. distal concavity a.2,3,4,-,6 long a. two b. absent last capillaries b. 1. falcate few setigers presentin absent 2. bristle- spreading topped fascicle a. 2, 3, -,-, 6 long a. one b. absent falcate b. simple posterior absent recurved falcate one-fourth hooks EERE chen a. one = 7 long b. simple b. absent absent falcate falcate posterior recurved two-thirds hooks a. 2, 3,4, -,6 a. two b. present present absent b. 1. falcate 2. bristle top with 2 heavy smooth bosses Pygidium Other Features a. pouched glands semi- in neuropodia circular of setigers disc, with 10-17 2 ventral b. hooded hooks lappets unidentate in posterior setigers bores in four small calcareous lobes materials simple a. nuchal tentacle weakly present lobed b. up to 16 collar, hooded hooks ventrally per incised neuropodium four equal = lobes small ring dorso-median with 2 channel ventral blade- between like lappets posterior each with a notopodia terminal process dorso-median flattened channel between plate with 2 posterior terminal cirri notopodia; inhabits brackish waters four lobes —— Reference Hartman (1961; 1969) Berkeley (1927) Woodwic (1963a) Blake (1966) This pape Soderstré (1920) Hartman (1948) | boscidea NEW SPECIES OF BOCCARDIA TABLE I (CONTINUED). Some Taxonomic Characteristics of the Known Species of Boccardia Prostomial nuchal ridge a b c, . prostomium a. b. . caruncle eyes . incised . into setiger 5 3-6 . incised . to end of setiger 3 6-8 . rounded . to end of setiger 3 .4 . incised . to end of setiger 2 5 - rounded . to end of setiger 3 .4 . incised . to end of setiger 2 .4 . incised . to end of setiger 3 4 Branchial distribution anterior posterior SOREN CHE) . absent posterior one-third . 2, 3, 4, -, 6 . absent posterior one-half 42,954) = 6 . absent last few setigers . 2, 3, 4, -, 6 . absent posterior one-third Leroi =10) . absent posterior two-thirds . 2,3, 4,-, 6 . absent posterior one-fifth ei DorSeiaain3 0. b. absent posterior one-half Notosetae — Posterior on Notopodial Setiger | Spines present absent absent absent present absent (short) present absent absent absent absent absent absent ? Spines of Setiger 5 a. number of types b. description of types Pygidium Other Features IY References a, two b. 1. falcate 2. bristle top and club-shaped a. two b. 1. falcate 2. inverted, bristle-topped cone with oblique base a. two b. 1. simple falcate 2. bristle- topped a. two b. 1. simple falcate nr . central cone with raised margins a. two b. 1. falcate 2. tricuspid a. one b. simple falcate a. one b. simple falcate simple collar with several weak lobes disc-like 4 lobes dorsal pair smaller than ventral two flattened glandular cushions 4 small lobes disc-like witha dorsal gap large forwardly- directed “lateral fans” on ventral side of setigers 1-4 ventral oral groove to anterior of setiger 3 pigment along prostomium; diverse habitats of shell, sand mud... palps barred bores in calcareous materials nuchal tentacle present Khlebovitsch (1959) Carazzi (1893) Mesnil (1896) Fauvel (1927) Hartman (1940; 1941) Woodwick (1963a) Day (1961; 1967) Woodwick (1963b) Hartman (1936; 1969) Okuda (1937) 40 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES rior end. Anteriorly they are 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. 31). 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 polybranchia 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 typensimplesstalcatevam jer sae he scene ei 2 1b Heavy spines of setiger 5 of two types, first type simple, falcate; second type highly modified ............ 5 2ZasNuchaletentacle presenti iscsi B. sp. 2biNuchalatentaclevabsent*manc rics ecieaecr 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 ...B. ligerica Sa 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 ..8B. 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 penNeuropodiumyse me. eet ee B. chilensis 7b Nuchal tentacle absent; hooded hooks number only 6 per neuropodium .......... B. pseudonatrix VOLUME 70 8a) Prostomiumirounded 4st sneer 9 8b Prostomium incised 9a Posterior acicular spines present; bristle-topped spines of setiger 5 with an accessory tooth; notosetae absent from Setigenjl tearacciicrser ener B. berkeleyorum 9b Posterior acicular spines absent; no tooth on bristle-topped spines of setiger 5; notosetae present on Setiger 'l: ¢. AM, smaller leg index because of shorter leg II, smaller tarsalae I and I, 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 260u. 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 18, 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 3lu, 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, Moy- 1971 NEW SPECIES OF CHIGGER 44 °. fj / fy < a EZ joi —— N= _ > ier oY \ \ - ts Reet Kine Figure 1. 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 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 35y and 34y ; length of other dorsals 20 to 294 and 25 to 3lu; eyes distinct 2/2 and eyes not discernible; lengths of tarsalae I and II 18u, 17u and l4u. l2u. in Pseudoschoengastia in their “Aeci Crotiscus desdentatus tissoti Fauran, 1960 Six specimens off Proechimys guyannensis, Lelydorp, 18 December 1969. Euschoengastia (s. 1.) desmodus (Brennan and Dalmat, 1960) Two larvae from ventral wing membrane of Glossophaga soricina, Brownsweg, 9 February 1970; 1 off Carollia perspicillata, 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 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Eutrombicula alfreddugesi (OQudemans, 1910) Five 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; | off D. marsupialis, Brokobaka, 21 February 1970; 2 off Philander opossum, Coronie, 10 January 1970; 6 off Rattus rattus frugivorus, Brokobaka, 21 February 1970; 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; | 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 (1946), Jenkins (1949), and Fuller (1952). Eutrombicula goeldii (OQudemans, 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, 31 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; | off Philander opossum, Coronie, 28 November 1969; 2 off P. opossum, Coronie, 12 February 1970; | 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 1969 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. VOLUME 70 1971 Tecomatlana (Hooperella) vesperuginis (Brennan and Jones, 1960) Thirteen larvae off 2 Carollia perspicillata, Zandery, 7 January and | 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 Glossophaga soricina, Brokopondo, 2 February 1970. The following chiggers of Surinam, not listed here, were recorded recently by Brennan (1970). Arisocerus amapensis Brennan, 1970 Boshkerria punctata (Boshell and Kerr, 1942) Colicus icomi Brennan, 1970 Colicus oblonga (Fauran, 1959) Eutrombicula alfreddugesi (Oudemans, tropica form of Ewing, 1925 Pseudoschoengastia tricosa (Brennan and Jones, 1961) Tecomatlana (Hooperella) saccopteryx (Brennan and Jones, 1960) Trombicula (s. 1.) palmigera Fauran, 1960 1910), LITERATURE CITED ‘Brennan, J. M. 1970. A small collection of chiggers NEW SPECIES OF CHIGGER from Surinam (Acarina Calif. Acad, Sci. 69 Trombiculidae), Bull 32-37 Fauran, P. 1960. Description de quatre especes et d'une nouvelle sous-espece de trombi nouvelle culides de Guyane frangaise. Arch. Inst. Pasteur Guyane frang. et Inini, No. 459, 20 p Fuller, H. S. 1952. The mite larvae of the family Trombiculidae in the Oudemans collection: Tax onomy and medical importance. Zoologisch Verhandelingen, No. 18. E. J. Brill, Leiden. 261 p Geest, J. C. and R. B. Loomis. 1968. Chiggers of the genus Pseudoschoengastia (Acarina: Trombicu lidae) from Costa Rica. Los Angeles Co. Mus. Nat Hist., Contrib. Sci. No. 150; 1-49. Jenkins, D. W. 1949. Trombiculid mites affecting man IV. Revision of Eutrombicula in the American hemisphere. Ann. Ent. Soc. Amer. 42: 289-318. Michener, C. D. 1946. Observations on the habits and life history of a chigger mite, Eutrombicula batatas (Acarina: Trombiculinae). Ann. Ent. Soc. 39: 101-118. Amer. 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! 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 Chiloé and Aysén, 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 confiuent 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 Fullerton, 1Department of Biology, California State College, California 92631 46 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF 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, 2, 28.0, 28.5, 29.0; 3 26.0. Antenna length, 2, 7.0, 8.0, 8.3; oF Tas bk Holotype. Chile, Prov. Nuble, Cordilleras Chil- lan, Las Trancas, 12 km E. Recinto, 1-10 Dec. 1964, 2, leg. L. E. Pena, Museum of Comparative Zoology, Harvard [ca. 800 mt., 71°37’ W, 36°51’ SIF Allotype. Argentina, Prov. Neuquén, Pucara, Parque Nacional Lanin, Jan. 1951, 3, leg. S. S. Schajovskoy, Museum of Comparative Zoology, Harvard (ex coll. E.G. MacLeod). Paratypes. | 9, same data as holotype, E. G. MacLeod collection. Argentina, Prov. Neuquén, Pucara, 30 Nov. 1959, 2, 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 1971 NEW SPECIES OF LACEWING 47 Figure 2. Reproductive structures. A, C, E-H, Kempynus crenatus. B, D, K. falcatus. 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. Figure 3. Striped wing pattern: A, Phymatosmylus caprorum; B, Kempynus falcatus. 48 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Kempynus the apical projections of the female eighth sternite are more slender and more separated basally. Kempynus falcatus Navas Figures 2B, D; 3B Kempynus falcatus Navas 1912, 1928, 1930, 1936, Kimmins, 1949. Type: 2, 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 66° 34° ® K. FALCATUS 9 K. CRENATUS + P. CAPRORUM Figure 4. Known distribution of South American Kempyninae and Stenosmylinae. VOLUME 70 | otherwise noted, are in my collection and were collected by L. E. Pena. Chile: Curic6, Buchén, — Mar. 1956, 2, (“farm in preandean foothills’); Curic6, La Jaula (Los Quenes), Andes, 14-18 Feb. — 1965, 6; Linares, Fundo Malcho (“in the high mountains”) Dec. 1952, 2; Malleco Prov., Termas — de Tolguaca, 30 km NE Curacautin, 20 Jan. 1959, — 6; Cautin 30 km N Villarica, 1-30 Jan. 1965, 6 3 4 2; Chiloé, Dalcahue, Feb. 1954, 3; Aysen, Pto. Cisnes, Feb. 1961, 2; Aysén, Rio Maniguales [vic. — Puerto Aisén] 27 Jan. 1961, 18, 12. Argentina, Neuquén, Pucara, Parque Nacional Lanin: 31 Nov. 1959, 2, purch. ex. F. H. Walz; Feb. 1951, 2, Feb. 1952, 3, leg. S. S. Schajovskoy, E.G. MacLeod col- lection. | Color variation. In my collection are single specimens of Phymatosmylus caprorum Adams (Fig. 3A @, from Las Trancas, Cord. Chillan, Chile, 1-10 Jan. 1964, L. E. Pena leg.) and Kempynus falcatus Navas (Fig. 3B, 2, 30 km N. Villarica, Prov. Cautin, Chile, 1-30 Jan. 1965, L. E. Pena 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 occur in related Osmylidae (Kimmins, 1940). 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 Chiloé and Aysén, 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 /sostenosmylus, 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 OF LACEWING 49 ABBREVIATIONS Bed — bursal gland duct, egr — colleterial gland reservoir, cop b—copulatory bursa, CuA — cubitus anterior, CuP — cubitus posterior, fe — fertilization canal, gcx — gonocoxite, go — geni- tal opening, gs — gonarcus, hyi —hypandrium 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 planipennian genitalic homologies. Postilla, 141: 1-11. ‘Kimmins, D. E. 1940. A revision of the osmylid subfamilies Stenosmylinae and Kalosmylinae (Neu- roptera). Novitates Zoologicae, 42: 165-201, pl 1-8, Kriiger, L. 1913, 1914, Beitrage zu einer Monographie der Neuropteren-Familie der Osmyliden. Srettiner entomologische Zeitung, 74: 3-294; 75: 4-140 Navas, L. 1912. Insectos neurépteros nuevos 6 poco conocidos. Memorias de la Real Academia de Ciencias y Artes de Barcelona, 10: 135-202 . 1928. Insectos neotrépicos. 4a serie. Revista Chilena Historia Natural, 32: 106-128 . 1930. Algunos insectos de Chile, 4a serie Revista Chilena Historia Natural, 34: 350-366 1936. Insectos neurépteros de Chile, poco conocidos. Revista Chilena Historia Natural, 40: 179-181. Accepted for publication September 4, 1970. 50 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 RESEARCH NOTES Bull. So. Calif. Acad. Sci. 70(1): 50-51, 1971 DISTRIBUTION OF SOME SOUTHERN CALIFORNIA KANGAROO RATS The ranges illustrated by Grinnell (1922) and Hall and Kelson (1959) for southern California kangaroo rats were drawn by encircling known records of occurrence in the latter case 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 those previously known from_ the Tehachapi 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 and this locality (as illustrated by Hall and Kelson, 1959) — 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 California. Unpublished M.S. thesis, San Fernando Valley State College, Northridge, California, 75 pp. Grinnell, J. 1922. A geographical study of the kangaroo rats of California. Univ. Calif. Publ. Zool., 24: \- 124. . 1933. Review of the Recent mammal fauna of California. Univ. Calif. Publ. Zool., 40: 71-234. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. Ronald Press Co., New York, 1083 pp. Munz, P. A., and D. D. Keck. 1959. A California Flora. Univ. California Press, Berkeley, 1681 p. 5 Vaughan, T. A. 1954. Mammals of the San Gabriel Mountains of California. Univ. Kansas Publ., Mus. Nat. Hist., 7: 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 ARTIBEUS LITURATUS (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 the scapholunar. 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 RESEARCH NOTES / length of the second, third, fourth and fifth meta carpals. The series was obtained during a seven day period, 10 to 17 May 1963, from adjacent sides of the Rio Coco [=Rio Segovia, Wanks River] which separates Honduras and Nicaragua, at a locality 78 mi. ENEE Danli, Honduras. All speci mens are deposited in the Texas Cooperative Wild life Collection, Texas A & M University A. statistical Yomparison of both methods is shown in Table |. The variance-ratio (F = 52;/s%s) was used to test the hypothesis that one method consistently yielded observations with smaller vari- ance and was therefore the more desirable. None of the F-values was significant at the 5 per cent level The method including the carpals consistently yielded smaller coefficient of variation values but 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 topography 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, | 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 MEAN MC RANGE s2* cv INCLUDING CARPALS 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 S318) 3:52 67.9 5.28 3.38 0.98 $9.0-71.5 62.3-75.0 4 64.0 5.97 3.82 66.2 5.15 3.43 1.16 §7.6-69.9 60.0-72.0 5 65.2 5.82 3.69 68.5 5.87 3.54 0.99 §9.6-71.6 62.6-75.6 *s2 and CV, variance and coefficient of variation. respectively. **The F-value is obtained from the variance-ration. s2,/s%9. Sample size is 50 throughout: measure- ments in mm. 52 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES ACKNOWLEDGMENTS Appreciation is extended to Gail Glick for clerical assistance, and to Dilford C. Carter, William B. Davis and Andrew Starrett for 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., 51: 105-122. Starrett, A., and L. de la Torre. 1964. Notes on a collection of bats from Central America, with the third record for Cyttarops 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 I 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. Ent. 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, Z/PHIUS CAVIROSTRIS, 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 (1961) 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 H 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 VOLUME 70 | 1971 to the site before the carcass was buried. This 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 C, 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 condylobasal 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 macrorhyncha, 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. Mammial., 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. Bull. U.S. Natl. Mus., 246: 1-259. Norris, K. S. (Ed.). 1961. Standardized methods for measuring and recording data on the smaller cetaceans. J. Mammaal., 42(4): 47 1-476. 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 RESEARCH NOTES of Florida, Gainesville, Florida 32601: Warren 4 Rathjen, Caribbean Fisheries Development Project Food and Agriculture Organization of the United Nations, Barbados (present address Ma rine Fisheries Service, Woods Hole, Massachusetts) and Melba C. Caldwell, Communication Sciences Laboratory, of Florida, Florida. National University Gainesville Accepted for publication September 26, 1970 Bull. So. Calif. Acad. Sci. 70(1): 53-54, 197] ADDITIONAL SPECIMENS OF THE COLUBRID SNAKE AMASTRIDIUM VELIFERUM COPE FROM COSTA RICA, 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 versant. The locality, San José, given for the holotype of Fleischmannia obscura Boettger, 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 level, the second approximately 1200 m. The only other known Pacific slope occurrence for the species is in Chiapas, México. 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 2819 is as follows: 17 243-498 15 (111). Furthermore. MNCR } Ses 2+3 (106) : E 2819 has a single anal plate (there is only at race 54 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES of a groove on the anterior portion of the scale), all other specimens have a divided anal plate. The number of subcaudals in this specimen is 70, 2 less than the lowest count for males from throughout the range (the subcaudal range is 72 to 86), given by Wilson and Meyer (1969), but the tail length- total length ratio (0.326) is greater than that previously reported for 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 (l1+2, the normal condition, on the right), a condition seen in only one other specimen (from Panama). As is the case for specimens of 4A. 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 2819 should show so many external morphological peculiarities is, at first analysis, striking. The species Amastridium 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- VOLUME 70 ities in the reproductive system of what, on the basis of external examination, appeared to be a male specimen. This specimen has one everted 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 pseudohermaphroditic 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. hemipenis, 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. Sci., 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. ith this issue, \ y the Southern Cal- ifornia Academy of NEW FORMAT Sciences begins a new era of publications. The ‘Bulletin has been redesigned and will appear three ‘times yearly. Hopefully, the Bulletin will now serve its community better by providing more information ‘to all members. During the period of ‘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 31, and December 31. 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. COVER ILLUSTRATION new section has been 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 NEWS SECTION 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 O n April 19, 1971 AND Alec R. Stra- ENVIRONMENT chan, formerly with the California Department of Fish and Game, Gn NEWS AND NOTES 350 Golden Shore, Long Beach, accepted a new industrial position with Southern California Edison Ron will be investigating the environmental impact of thermal discharges, antifouling equipment, and impregnation studies to prevent fouling. Ron is a fulltime employee of Southern California Edison as biologist-diver. OIL (ee of the two POLLUTION volume Univer- REPORT sity of Southern Cali- fornia Oil Pollution Report are now available. Copies may be purchased through Mrs. Dorothy Halmos, Librarian, Allan Hancock University of Southern California, Los Angeles. California 90007. Foundation, ICHTHYOLOGICAL dward Raney and ASSOCIATES Ed Bates, both of Ichthyological Associates, are consultant contractors to Southern California Edison to study the fishes in and about the generating plants in southern California. They 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 Oo’ May 21, 1971, ENDANGERED 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 95814. 56 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES In the early morning LLOYD W. BARKER APRIL 25, 1949 hours of Tuesday, June 29, 1971, a vessel under tow by 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 accident 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 VOLUME 70 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. MacMillan A new species of /phitima (Polychaeta) from Cancer antennarius (Crustacea: Decapoda). By John Pilger INSTRUCTIONS FOR AUTHORS Contributions to the BULLETIN may be in any of the fields of science, 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 BULLETIN style will be 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 Journals is 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% x 11 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 84% x 11 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 1971-1972 1971-1974 Jack W. Anderson George F. Fisler Phillip A. Adams Roberta S. Greenwood Robert M. Chew William J. Morris Joseph Arditti Daniel Gutherie Kristian Fauchald Nancy Nicholson 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 16 1972 NEW YORK BOTANICAL GARDEN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES BULLETIN Number 2 RWRMAO ARASA\ §-* 41n? /141A74\ ATTIC ?TrCOT,T ta77 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. PATITE UATE TIN CLS perch estes ete e sees a eirat aoa ah ener Rees Chie eS ah ct $ 8.00 StudentoMembers tise rciaschoke es eihaisls Bie sichoneie GeO Glace tees eee 5.00 MF OWMIVICIMND RSH kts hb estes pausase Ste carqeuene ge Wa coe eea ee ea oot eerie caunilee cet od Set a 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 NUMBER 2 CONTENTS The larvae of Pagurus samuelis (Decapoda: Anomura) reared in the laboratory. BYERLOV REM MVIACIMILLAMm Mranyina Menai ar are secct ti esiei «4 ele) al asuie ied e chs Salona 6 58 Descriptions and notes concerning some oriental Aphelocheirus (Hemiptera: pPHelocheinidac) sy KaveauRLVers aire rsiyare eilwis's «a «s/t icles «le ulely 69 New species of Polydora (Polychaeta: Spionidae) from the coast of California. By dames A. Blakerand Kerth H. Woodwick 7... 16.5... oer dene iz A new species of Hunterotrema (Digenea: Campulidae) from the Amazon River Dolphin (Inia geoffrensis). By Murray D. Dailey ........0..000 000000. 79 A new species of Funnel-eared bat (Natalus stramineus) from western Venezuela. BVAOMAIIS NTEINQHES Fics: csey ere (e. eiekt oslo eyes is Tose, Oeaiton ce Wyola e109 pone fohel ote fos le 81 A new species of [phitime (Polychaeta) from Cancer antennarius (Crustacea: Decapoda) sy ByiWOhM Pil ee ric gecwceapey tis toteie, sisuome eicslsyel sue) eee) Spare) of ere eta 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 Arlo W. Fast ............-0..020.4- 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. We bb MU rea reat CdS te eA RE cured Matera AUoHi Mee cati cise ce odsyie) by elisieire: ek aye; ss) ox 102 nH N 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! ABSTRACT: Pagurus samuelis (Stimpson) larvae from northern California were studied. Larvae were raised at 15 and 17°C. Illustrations and descrip- tions of the four zoeal stages and glaucoth6e 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 (1964), Greenwood (1966), 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 glaucoth6e. 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-19344). 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. 58 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 Artemia 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°%po9 to 35%oo. Water was changed every other day and fresh Artemia 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. 1Pacific Marine Station, University of the Pacific, Dillon Beach, California 94929. 1971 THE LARVAE OF PAGURUS SAMUELIS $9 Figure 1, Pagurus samuelis (Stimpson). Dorsal views of zoeal stages I-IV. Scale 0.5 mm. FIRST ZOEA (Figure | I, 2 1) Size: CL, 1.1 mm; TL 2.4 mm Duration: 6-7 days at 15 and 17°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 | 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 1) 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 eyes 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 chromatophores posterior to each antennule and 8 60 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Figure 2. Pagurus samuelis (Stimpson). Lateral views of zoeal stages I-IV. Scale 0.5 mm. VOLUME 70 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 | 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 | 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. 7) PPP PPP PI??? Figure 3. Pagurus samuelis (Stimpson). Telson of zoeal stages I-IV. Scale 0.2 mm. 1971 THE LARVAE OF PAGURUS SAMUELIS —— ———— OO” aN SX ZV Sere ss = = ib | / Figure 4. Pagurus samuelis (Stimpson). Top, antennules of zoeal stages I-IV. Scale 0.1 mm. Borfom, antennae o zoeal stages I-IV. Scale 0.2 mm. 6] 62 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Scaphognathite of the maxilla (Fig. 5 1) has 5 plumose setae along the margin. The endopodite bears 3 plumose setae terminally and 2 setae sub- terminally on the medial margin. The distal and proximal lobes of the basal endite each have | subterminal and 3 terminal plumose setae. The distal lobe of the coxal endite has 2 terminal setae and | subterminal seta. The proximal lobe has 4 terminal setae. Basipodite of the maxilliped | (Fig. 6 1) has a pair of plumose setae at the anteromedial corner and 6-7 hairs on the medial margin. The exopodite is composed of 2 indistinct segments and terminates in 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 setulés 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 | 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 | sub- terminally. The other segments each have | heavy plumose seta and | 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 | II, 2 II) Size: CL 1.4 mm; TL 2.6 mm Duration: 6-8 days at 15 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 Il) 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. VOLUME 70 Antennae (Fig. 411) have elongated but are identical to that of zoeal stage | 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- bearing 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 | III, 2 IID) 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. 4III) are segmented and biramous. The distal segment terminally bears | large and 2 small aesthetes, | plumose seta and 2 hairs. Subterminally, there is an additional pair of aesthetes. The segment representing the ventral 1971 THE LARVAE OF PAGURUS SAMUELIS 64 = Je SS AE Ly t Z \ ; | Figure 5. Pagurus samucelis (Stimpson). Top, mandibles of zoeal stages I-IV. Center, maxillules of zoeal stages I-IV. Bottom, maxillae of zoeal stages I-IV. Scales 0.1 mm. flagellum 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 8 natatory setae. Maxilliped 3 (Fig. 8 III) has an endopodite terminating in | small plumose seta. The exopodite has 7 natatory setae. 64 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 4A* 4A EEA AL Figure 6. Pagurus samucelis (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 ZOEA (Figure | TV, 2 1V) Size: CL 2.2 mm; TL 3.4 mm Duration: 7-8 days at 17°C General Characteristics and Coloration Pleopod buds are present on abdominal somites 2-5. Telson (Fig. 3 1V) is almost 3 times as long as wide. Protopodites of the uropods are articulated with the exopodites but not with the endopodites, There is an additional yellow chromatophore on the rudimentary cheliped and a red chromatophore just above the basipodite of maxilliped 3. Appendages The process representing the dorsal flagellum of the antennule (Fig. 41V) bears 3 aesthetes, | plumose seta, and 2 hairs. In addition, there is a longitudinal row of 4-5 subterminal aesthetes. The ventral flagellum has lost its terminal seta but has retained the plumose seta at its base. The endopodite of the antenna (Fig. 4 1V) 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 | (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 | terminal plumose seta and | 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. 10a) 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 | 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 LARVAE OF PAGURUS SAMUELIS ae lee apes Wi vey LY= Figure 8. Pagurus samuelis (Stimpson). Third maxil lipeds of zoeal stages I-IV. Scale 0.5 mm Red chromatophores are prominent on the eye- stalks, antennules, antennal basipodites, and on the anterolateral region of the carapace. There are a great number of pale yellow chromatophores scattered over the anterior dorsal region of the carapace, the eyestalks, and the fifth periopod Appendages The ventral flagellum of the antennule (Fig. 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 are 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 | 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 long plumose setae. The endopodite consists of 4 segments, each with a few short setae. Exopodite of maxilliped 3 (Fig. 10 K) has tn > 66 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 Figure 9. Pagurus samuelis (Stimpson). Glaucothée. Lateral and dorsal views. Scale 1.0 mm. 1971 THE LARVAE OF PAGURUS SAMUELIS 67 Figure 10. Pagurus samuelis (Stimpson). Glaucothée. 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-I) 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 only 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 OF SCIENCES VOLUME 70 TABLE |. Morphological differences between Japanese and California zoea larvae of Pagurus samuelis. Characteristic Japanese Species Rostral length extends to base of antennal scale spine Antenna endopodite scale forked broad, concave outer . margin with six plumose California Species extends beyond tip of antennal scale spine not forked narrow, straight outer margin with five plumose setae tooth at base articulated setae not articulated Fifth abdominal prominent lateral spines short lateral spines segment Telson wide — convex outer narrow — straight outer margin margin Coloration chromatophores all on ventral margin scattered on lateral on Carapace chromatophores present at base of rostrum and on antennal protopods 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. surface absent 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, AF 2NT-239% 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, ACADEMY OF SCIENCES 70(2); 69-72, 1971 DESCRIPTIONS AND NOTES CONCERNING SOME ORIENTAL APHELOCHEIRUS (HEMIPTERA: APHELOCHEIRIDAE) IRA LA Rivers! ABSTRACT: Two new species of Aphelocheirus are described from India and A. pallens Horvath is noted from additional localities in New Guinea. 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, 1800 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 | 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 69 Figure 1. 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%): and (2) median length to greatest width 20: 60 (33° Scutellum: Large, blackish, shiny; ratio of three 1Biological Society of Nevada, Verdi, Nevada 89439. 70 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Hemelytra: Translucent brownish; claval sutures distinct; membranes very large, comprising almost half 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 I with a prominent, whitish, narrowly elongate static sense organ (sfatexta) 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 |.1 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 VOLUME 70 D. Q. Cavagnaro, 110 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 anteriorly, eyes essentially flush with general surface. Labrum prominent, broad and weakly pointed, ratio of length-to-width 16:22 (73%), Head ratios are: (1) Total median length-to-width (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) anterior 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 to greatest width 27: 87 (31 %);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, ORIENTAL HEMIPTERA A 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. Con- 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. Trochanter prominent. Femur weakly incrassate. flattened, well-furred along tibial closure edge: ratio of length-to-width 65: 24 (37%), length 2mm Tibia long, narrow, tubular, bearing a_ three- segmented tarsus whose first segment is minute, the remaining two elongate and terminating in two prominent claws. angularly-globular 72 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Mesoleg — Coxa and trochanter similar to proleg except larger. Femur weakly incrassate, 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 of reddish spines, particularly along outer edge and a loose, trans- verse, terminal row; ratio of length-to-width 48: 8 (17%), 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%), 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; | winged paratype from Coimba- tore, Nathan, 1959 February. All specimens in the VOLUME 70 collection of the Biological Society of Nevada, Verdi, Nevada. | 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 east, 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) 1803 group to the west, A. nathani is quite close to the Phillippine species described by Usinger, A. philippinensis 1938 and A. uichancoi 1938, particularly the latter, which is about the same size. From A. uichancoi, A. nathani differs in the lighter color, somewhat smaller size and much different head proportions, uichancoi 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 uichancoi. LITERATURE CITED Horvath, G., 1899. Monographia generis Aphelochei- rus. Termeszetrajzi Fiizetek, 22: 256-267. Usinger, R. L. 1938. The Naucoridae of the Philippine Islands (Hemiptera). Philippine J. Sci., 64: 299- 309. Accepted for publication October 23, 1970. BULLETIN SO. CALIF. ACADEMY OF SCIENCES 70(2): 72-79, 1971 NEW SPECIES OF POLYDORA (POLYCHAETA: SPIONIDAE) FROM THE COAST OF CALIFORNIA JAMES A. BLAKE! AND KEITH H. WooDWICk2 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 13 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. 1Pacific Marine Station, University of the Pacific, Dillon Beach, California 94929. 2Department of Biology, Fresno State College, Fresno, California 93710. 1971 Genus Polydora Bosc, 1802 Polydora convexa, new species Figure | Material examined: California: Santa Barbara, August 27, 1961 (1), from shells of hermit crabs; Avila, March 3, 1962 (8, TYPE), from shell of Pododesmus macroschisma; Morro Bay, October 24, 1961 (4), May 18, 1963 (13), from hermit crab shells and Pododesmus; Cayucos, June 28, 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 Macro- cystis, and Dodecaceria colonies; Bodega Harbor, March 3, 1970 (3), from Pododesmus; 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 1. 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. NEW SPECIES OF POLY DORA 13 The prostomium is bifid on its anterior margin (Fig. la). The caruncle extends to the posterior border of setiger 4. There is no nuchal tentacle Eyes may be absent, but when present the four eyes are arranged in a trapezoidal manner with the posterior pair closer together and smaller than the anterior pair. The palps extend posteriorly only to setiger 10 in preserved specimens; in life they are longer and prehensile. Setiger | has capillary setae in both noto- and 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 of short curved capillary setae, a second row of longer, less curved capillary setae, and a third row of long stout capillaries. The number of these setae per notopo- dium 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 by an unusual setal structure which is difficult to interpret (Fig. Ig-h). These structures do not protrude through the cuticle and resemble tight bundles of setal fibers. They are undoubtedly similar to the “organes en filieres” of Claparéde (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. le) 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 | 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 lying anterior to a semi- circular row of alternating heavy spines (Fig. 1b-c) and smaller pennoned companion setae (Fig. 1d). The heavy spines are falcate and have a broad collar on the convex side (Fig. 1b-c). Ventral 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 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES A structure similar in appearance to the “gizzard- like” structure previously described for P. socialis (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 P. 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 incertae 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 (Mésnil 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, VOLUME 70 March 30, 1970, intertidal from a hermit crab shell (1); Morro Bay, December 31, 1963, intertidal from shells of Tivela 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 31 mm in length, is | 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 | 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 hooks begin on setigers 10, 11, or 12, but usually on 11. 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 structure throughout the length of the body (Fig. 2b). Setiger 5 is modified and larger than other 1971 NEW SPECIES OF POLY DORA j | 20mm 905mm 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 pennoned 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 150u 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 of Olivella biplicata occupied by the hermit crab, Pagurus granosimanus At Morro Bay it occurred in shells of Tivela stultorum and at Malibu Beach in shells with hermit crabs. Polydora bioccipitalis, new species Figures 3 and 4 Material examined: California; Malibu Beach (22 TYPE), August 20, 1964 from hermit crabs col lected by James McHenry; Santa Barbara (1), July 30, 1965 from a shell of Olivella biplicata occupied by a hermit crab. Description of the adult: Since both adults and post-larval juveniles are 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 120 segments. There is no body pigmentation. 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- . Polydora bioccipitalis, new species: a, anterior Figure 3 end in dorsal view: b. hooded hook: c. fascicle of heavy spines and companion setae from setiger 5; d. posterio end in dorsal view. 76 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES dora 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 | 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). acémm Figure 4. Polydora bioccipitalis, new species, juvenile: a, anterior end in dorsal view; b, group of setae from setiger 5; c, posterior end in dorsal view. VOLUME 70 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 are 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 fin 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 sharply hooked seta. The remaining heavy spines are 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 are 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- setae 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. NEW SPECIES OF POLY DORA Polydora pygidialis, new specie Figure 5 Polydora ciliata Berkeley and Berkeley 1952: 19-20 (not Johnston, 1838) Material examined: California; Santa Barbara August 27, 1961 (4), from inner harbor piling material; Avila, March 3, 1962 (3), from hermit crab shells; Morro Bay, October 24, 1961 (23) May 18, 1963 (6); Cayucos, June 28, 1961 (5) August 28, 1961 (23, TYPE), December 19, 1961 (19), July 3, 1962 (1), from hermit crab shells keyhole limpet shell, and piling material; Bodega Bay, September 28, 1970 (10), from bryozoa dredged from 18 m. British Columbia; Departure Bay, circa 1936 (4), from hermit crab shells (Berke- ley and Berkeley, 1936). Description: The individuals of this species are very slender. A specimen 9 mm long and having 100 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 widely- spaced and cup-shaped; the posterior pair are close 1936: 4 Dapp P cdiaaqnqy see 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 - , i © setae from setiger 5; g, neurosetae from se posterior end in dorsal view: i, posterior end in lateral view. 78 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES together and irregularly shaped. In some specimens only the anterior pair are present. Setiger | lacks notosetae but has a small conical notopodial lobe (Fig. Sa). 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. 5f), a curved row of large modified spines and alternating pennoned companion setae (Fig. S5c-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. Sa). 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. Sh-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 py gidialis 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). VOLUME 70 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 funebralis|/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. \n 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. Zool. 75: 1-32. Carazzi, D. 1893. Revisione del genero Polydora Bosc, e cenni su due specie che vivono sulle ostriche. Mitteilungen aus der Zoologischen Station zu Neapel, 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. Polychétes 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-Schréder, G. 1959. Zur Okologie der Poly- chaeten des Mangrove-Estero-Gebietes von El Salvador. Beitrage zur Neotropischen Fauna, 1: 69-183. 1971 NEW SPECIES OF Johnston, G. 1838. Miscellanea Zoologica. Ariciadae. Mag. Zool, Bot,, 2: 63-73. MeIntosh, W. C. 1915. A Monograph of the British marine annelids, Polychaeta, Opheliidae to: Am- phictenidae. Ray Soc., London, 3: 1-368. Mesnil, F. 1896. Etude de morphologic externe chez les Annélides. Les Spionidiens des cétes de la Marche. Bulletin Scientifique de la France et de la Belgique, 29: 110-287. Rioja, E. 1925. Anelidos poliquetos de San Vicente de la Barquera (Cantabrico). Trabajas del Museo Nacional de Ciencias Naturales. Serie Zoologica, 53: 1-62. POLY DORA y . 1939, Etudios anelidologicos 1. Observacions acerca de varias formas larvarias y postlarvaria pelagicas de Spionidae, procedentes de Acapul con descripcion de una expecies nueva del gener Polydora, Annales del Instituto de Biologia, 10 297-311. . 1943. Estudios anclidologicos VIIL Dato acerca de las especies del genero Polydora Bo de las costas Mexicanas del Pacifico. Annal« Instituto de Biologia, 14; 229-241 Woodwick, K. H. 1961. species of spionid polychaete from Lower Cali fornia. Pacific Sci., 15: 78-81. olydora rickettsi, a new Accepted for publication February 18, 1971 BULLETIN SO. CALIF, ACADEMY OF SCIENCES 70(2): 79-80, 1971 A NEW SPECIES OF HUNTEROTREMA (DIGENEA: CAMPULIDAE) FROM THE AMAZON RIVER DOLPHIN (INIA GEOFFRENSIS) Murray D. DaAILey! ABSTRACT: A new species of Hunterotrema (Digenea: Campulidae) is described from the lungs of the Amazon river dolphin (/nia geoffrensis). 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 (/nia 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 were stained in Semichon’s carmine or celestine 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. Aver- age measurements are presented with ranges in parentheses. Hunterotrema macrosoma, new species Figures 1-5 Description based on measurements from 15 specimens. Diagnosis: Body slender, elongate, 31 (24-36) long, with distinct forebody 3.7 (3.1-4.7), mid-body 19.6 (12.2-25.4) and hindbody 7.8 (6.0-9.8). Maxi- mum body width 1.1 (0.74-1.46) at acetabular level. hindbody 1.0 (0.53-1.5) in region anterior testis 1Department of Biology, California State College. Long California 90801. SO BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Figures 1-5. Hunterotrema macrosoma, new species. Fig. 1. 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 — uterus; 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, VOLUME 70 extending from posterior end of intestinal cecae to anterior constriction of hindbody. Uterus coiled immediately anterior to ovary, then straightens to run as a sinuous tube to genital pore. Eggs oval 1084 (105-112) by 624 (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 1930-1960. 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. CALIF, ACADEMY OF SCIENCE 70(2): 81-84, 197] A NEW SUBSPECIES OF FUNNEL-EARED BAT (NATALUS STRAMINEUS) FROM WESTERN VENEZUELA OMAR J. LINARES! ABSTRACT: A new subspecies of the funnel-cared bat Natalus stramineu is described from a cave in the Guasare river, Zulia, Venezuela. On the basis of palatal structure, the 3 recognized subspecies of N. major are assigned to N stramineus, The species status of N. tumidirostris and its possible synonymy with N. stramineus are also discussed. Additional records of N. . continentis 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, alc. with skull, Biology Museum, Central University of Venezuela, MBUCV 1-1578, collected by J. A. Tronchoni (original number L-281) from Gavilanes cave (Zu-1, Soc. Venezolana Espel., 1968: 113-118), Guasare river, Zulia, Venezuela, 183 m, 20 October 1967. Paratypes: Three adult females, alc. 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- 81 Figure 1. Palatal emargination and dentition of upper right jaw of: a, Natalus stramineus tronchonii (MBUCV 1-1578. ¢ —holotype), and b, Natalus stramineus mexicanus (TCWC 8507, 2). rostris. Upper molar with the hypocone widest 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). 1Laboratorio de Vertebrados, Instituto de Zoologia T Universidad Central de Venezuela, Caracas, Apt. 59058, zuela (Present address: 3, Rue des Ecoffes, Paris-IV°-France 82 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 TaBLe |. Measurements (mm) of Natalus stramineus tronchonii from Venezuela compared with selected measurements of Natalus stramineus mexicanus from Mexico, Nicaragua and Guatemala. N. s. tronchonii N. 8. mexicanus 1577 1578! 1579 1580 n Range X + ISD 2 3 g g Total length (body + tail) 97.0 97.0 92.0 100.0 — — == Length of tail 52.0 51.0 46.0 53.0 — — — Hind foot 8.0 8.5 8.0 8.5 —_ — ad Ear from meatus 14.0 14.5 13.0 14.0 — —— — Forearm 38.3 39.7 37.1 38.3 10 (36.9-39.8) 38.25+0.85 Third finger, 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 Perl 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 1385: 13.1 13.4 10 (13.3-13.9) 13.54+0.20 palatal length deal 7.0 7.8 ell 10 (8.7-9.6) 9.02+0.29 zygomatic width Tal. 8.1 el. = 10 (7.9-8.3) 8.11+0.19 width of braincase 7.4 Wed Ted 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 — el 9 (7.3-7.5) 7.41+0.09 interorbital width 3.3 3.2 3.2 SRD. 10 (2.8-3.1) 3.00+0.10 width across molars 5.0 5.2 Sal S22 9 (5.1-5.3) 5.26+0.06 width across canine — 3.5 3.4 325) 9 = (3.5-3.8) 3.61-0.10 upper toothrow, c-m? 6.5 6.8 6.5 6.7 9 (6.8-7.0) 6.86 + 0.08 lower toothrow, c-m3 7.0 al 6.8 Wal 9 (7.2-7.5) 7.33+£0.08 length of mandible ley 12.0 11.8 11.9 8 (11.4-12.5) 11.79+0.32 '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- tinentis Thomas. On the basis of an analysis of the form of the palate, only 2 groups can be admitted: stramineus- 1971 ts —— Cea] et <——— _—— aa fondylevacat length SaaS J _=— La) Wea Narcarsstiaieet i et J \ —— ia) een ——— — a cepa width acters eingute aaninee 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 /aeviventris 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 /aeviventris 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 VOLUME 70 of Anolis laeviventris. Anolis laeviventris at present appears to be restricted to intermediate elevations of both versants from southern México to western Panama. Anolis sminthus Dunn and Emlen 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. sminthus is related to A. altae and A. con- color. In our opinion this was an erroneous associa- tion, and A. sminthus 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, sminthus by Dunn and Emlen (1932) from San Juancito, 2100 m, Francisco Morazan, Honduras, A. haguei by Stuart (1942) from 2 km S Finca Chichén, Alta Verapaz, Guatemala, and A. ani- solepis by Smith, et al., (1968) from 17 km SE San Cristobal de Las Casas, Chiapas México. In 1968 we secured a series of what we consider to be Anolis crassulus, in sympatry with A. sminthus 3 km NE La Esperanza. Departamento de Intibuca, and there is no problem in separating this series of crassulus from all known specimens of sminthus. Anolis sminthus 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 (3 and 2), a narrow white mid-dorsal stripe (2 only), or a series of faint paravertebral inverted V’s (3 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 (2) and 41 to 46 (¢) 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 (¢ and @); tail laterally compressed in cross-section (strongly so ind), the scales of the dorsal caudal row strongly keeled and ridge-like; mid-ventral scales strongly keeled, 29 to 30 (2) and 30 to 36 (2) between axilla and groin. The above characters consistently separate smin- thus 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 sminthus in Honduras, specimens of the crassulus group from Guatemala and México 1971 have a bewildering array of admixtures of the dis- tinctive characters observed in Honduras, except that nowhere outside of Honduras and El Salvador (see below) are smooth or weakly-keeled mid- ventrals encountered. The inter-relationships of the populations that have received the names sminthus, crassulus, haguei, and anisolepis are exceedingly complex, and after examining specimens of all 4 supposed forms from México 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 (UMMZ 117647, 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 are 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 males of 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 AMPHIBIANS AND REPTILES OF HONDURAS 109 extending onto shoulders (color and pattern variable in crassulus group, but never approaching that of bouvierti), Anolis bouvieril 1s clearly not related to the crassulus group, nor in our opinion does it have any close relationship to cohanensis or any other anole from northern Middle America. Stuart (1955) based the supposed relationship of A, cobanensis and A. heteropholidotus (= sminthus) on the presence of smooth to weakly keeled ventrals in the two species. Examination of more specimens of sminthus than were available to Stuart indicates that there is a tendency toward strong keeling in all but the mid-ventral scales, so that approaches having the strongly keeled ventrals characteristic of all other species of the crassulus group. In addition to the difference in ventrals between cobanensis and the crassulus group, there are no enlarged postanal scales in cobanensis (pres- ent in all crassul/us group males), the dorsals are not abruptly enlarged from the laterals in cobanensis (abruptly enlarged in crassulus group), the ventrals range from 53 to 61 between the axilla and groin in both sexes of cobanensis (no more than SO 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). sminthus Sphaerodactylus dunni Schmidt This species was previously known only from the type, described by Schmidt (1963) from Rio Naco, near Cofradia, Departamento de Cortés, Honduras. In July 1968, we collected 3 specimens (LACM 47302-04) of this species, 12 km S La Ceiba, 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 Rio 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 alonga 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 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES a reddish-brown nasal blotch and a dark reddish to orange-brown parietal blotch, outlined by a Y- shaped white to pinkish-white marking, the arms of which are confluent with the light ground color of the head; lateral head color orange-brown, with pinkish-white upper 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- leyi, 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 VOLUME 70 juvenile from the mountains above Zanatepec, Oaxaca, México, resembling the pattern depicted by Stuart (1941) for dorsalis, and on the fact that adults from the same area are stripeless and are allocated to melanolomus. The distribution of the two species as pictured by Stuart (1941) is suggestive of a subspecific 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 Cortés, and the same year Ernest A. Liner collected a juvenile melanolomus (EAL 2200), 6 km N Agua Azul, Departamento de Cortés. 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, pl. 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 species distinct from melanolomus, especially in view of the somewhat confused situation presently existing with material from the Tehuantepec region of México. Lynch and Smith (1966) have discussed this situation and regard the subspecific status of the circumisthmal populations of melanolomus as in doubt, and their allocation of the Zanatepec juvenile to dorsalis is in need of confirmation. Lampropeltis triangulum Lacepede A specimen (LSUMZ 23824) from 4.5 km SE Catacamas, Departamento de Olancho, is unusual in having a color pattern composed only of red and black rings. In life the dorsum was dark red with 41 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 triangulum micro- pholis from near the Canal Zone in Panama. Our specimen was taken at night in a partially flooded cornfield. Ninia atrata (Hallowell) 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 112622) 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 112621 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 AMPHIBIANS AND REPTILES OF HONDURAS //] close to, the ranges given for atrata by Burger and Werler (1954), The Honduran specimens, all males, have 142 to 146 ventrals and 59 to 60 subcaudals In view of the fact that only the lack of a nuchal collar distinguishes the Honduran specimens from typical afrata, and since some Venezuelan speci- mens also lack collars, we regard the Hondu- ran specimens as representatives of Ninia atrata. Whether the Honduran population is character- istically collarless cannot be answered until more material becomes available. We accept the use of the name afrata instead of lansbergi for this species of Ninia, in accordance with the decision presented in Opinion 644 of the International Commission on Zoological Nomen- clature (Bull. Zool. Nomencl., 20: 26) to supress the combination Coluber atratus Gmelin, 1768 and to validate the combination Coluber atratus Hallowell, 1845. Rhadinaea kinkelini Boettger This snake has previously been recorded only from Nicaragua by Boettger (1898) 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 | 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 23829) 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, 1960). The range of S. anthracops is poorly known, and only 3 of the 112 BULLETIN SOUTHERN CALIFORNIA specimens cited by Peters (1960) and one cited by 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, Sth, and 6th entering on the right; no preoculars; loreal single, elongate, entering the orbit; postoculars 2-3; temporals | + 2/1 + 2; maxil- lary teeth 15; total length 664 mm, tail length 153 mm. 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! 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 this coloration extending laterally almost to lip, posterolaterally 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 1967, about 3 m ina 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, El 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 ACADEMY OF SCIENCES VOLUME 70 parietal and one secondary temporal in both; total length 379 mm, 421 mm; tail length 61 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 cyrtopsis (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 cyrtopsis 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 Field work in 1967 was made possible by funds from the Biomedical Sciences Support Grant, FR 07012-01, from the National Institutes of Health (senior author) and a Field Research Fellowship from Louisiana State University (unior author), and in 1968 by a Graduate School Travel Fellowship from the University of South- ern California (senior author) and a Sigma Xi Grant- in-Aid of Research (senior author). “es LITERATURE CITED Bocourt, M. F. 1873 (1872-82). Recherches zoologiques. Etudes sur les reptiles. /n Mission Scientifique au Mexique et dans l’Amerique Centrale. Livr 3: 28-1000. Boettger, O. 1898. Katalog der Reptilien-Sammlung im Museum der Senckenbergischen Naturfor- schenden Gesellschaft. Tiel II (Schlangen). Frank- fort; Gebriider 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. Dunn, E. R. 1930. Notes on Central American Anolis. Proc. New England Zool. Club., 12: 15-24. 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 José, Costa Rica. Lynch, J. D. 1965. A review of the rugu/osus group of Eleutherodactylus in northern Central America. Herpetologica, 21: 102-113. 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. AMPHIBIANS AND REPTILES OF HONDURAS 1/3 Mertens, R. 19524, Neues liber die Reptilienfauna von El Salvador, Zoologische Anzciver, 148: 87-93 . 1952b. Die Amphibien und Reptilien von El Salvador, auf Grund der Reisen von R. Mertens und A. Zilch. Abhandlungen der Senckenhergis chen Naturforschenden Gesellschaft, 487: 1-120 Peters, J. A. 1960. The snakes of the subfamily Dipsadi- nae. Misc. Publ. Mus. Zool,., Univ. Michigan, 114: 1-224. Peters, W. 1863. Eine mittheilung tiber neue Arten der Saurier-Gattung Anolis. Monatshericht Akademie Wissenschaftlichen Berlin, 1862 (1863): 135-149. Rand, A. S. 1957. Notes on amphibians and reptiles from El Salvador. Fieldiana: Zool., 34: 505-534. Roze, J. A. 1966. La taxonomia y zoogeographia de los ofidios de Venezuela. Servicio de Distribuciones de Pulicaciones, Biblioteca, Universidad Central de Venezuela, Caracas, 362 pp. Schmidt, K. P. 1933. New reptiles and amphibians from Honduras. Field Mus. Nat. Hist., Zool. Ser., 20: 15-22. 1936. New amphibians and reptiles from Honduras in the Museum of Comparative Zool- ogy. Proc. Biol. Soc. Washington, 49: 43-S0. 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 (Reptilia: Lacertilia) from Mexico. J. Herp., 2: 147-151. 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. Mus., 199: 1- 258. 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 new 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., Univ. Michigan, 91: 1-31. 114 . 1963. A checklist of the herpetofauna of Guate- mala. Misc. Publ. Mus. Zool., 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. . 19S4a. A review of the Lizards of Costa Rica. Univ. Kansas Sci. Bull., 38: 3-322. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 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 I1, 1971. BULLETIN SO. CALIF. ACADEMY OF SCIENCE 70(3): 114-124, 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! 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. The number of species found in each of the four areas studied along the coast (Santa Barbara, Pt. 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 (lat. 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 . Naples Reef, 13-15 m, shale reef and rubble; 21 July 1970. 2. Ellwood pier, 1-11 m, concrete, wooden, and steel pier pilings; 21 July 1970. - 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. Ww 1Pacific Bio-Marine Supply Company, P.O. Box 536, Venice, California 90291. 1971 ASCIDIANS OF SOUTHERN CALIFORNIA 115 Order Enterogona S. sp. (barnharti) S. gibbsi 44,5 Aplidium californicum ‘ A, solidum A. arenatum | S. plicata | | Euherdmania claviformis 2,3,4 Moleula verrucifera 2,4 3 3 Pyura haustor 2,3.4,5 Boltenia villosa p Polyclinum planum Synoteum par-fustis Didemnum carnulentin 1,2,3,4 Number of species found: 32 Trididemnum opacum 13 Pre DURE nen Unidentified didemnid | (ee Diplosoma macdonaldi 3 Figure 3 Clavelina huntsmani 4 Gystodytes lobatus 2 1. Paradise Cove, bait receiver, 1-2 m, nylon mesh Distaplia occidentalis 14 fish net; 16 June 1970. Eudistoma psammion 13 2. Little Dume Pt., 3-5 m, shale reef; 16 June 1970. EL ritteri 3 3. Leo Carrillo, 5 m, granitic rocks; 19 June 1970. E. diaphanes 3 4. Paradise Cove, No. 8 reef, 10 m, shale reef; 26 June Sigillinaria aequali-siphonis I 1970. S. pulchra ‘ | 5. Malibu Pt., 3 m, basaltic rocks; 28 June 1970. Wvcidiaiceratodes 1.2.3.5 6. Pt. Dume rocks, 3-10 m, basaltic rocks; 2 July 1970. Chelyosoma productum 4 7. Little Dume Cove, 3 m, shale reef; 2 July 1970. Gionaintestinalis 5 8. Latigo Cove, 10 m, basaltic rocks; 26 August 1970. Perophora annectens 23 9. West end of Zuma Beach, 6 m, granitic rocks: | September 1970. Order Pleurogona Botryllus sp. S85) Order Enterogona Botrylloides sp. 4,5 Styela montereyensis 1,2,3,4,5 Aplidium californicum 2,3,4,5 S. truncata 253,55: A. solidum 3.7 SANTA BARBARA Pt Conception PT. DUME AREA 2 Venice Leo Carrillo Beach co Carrilfo Be B Marina de/ Rey PALOS VERDES-LOS ANGELES AREA @-~X Lf Newport Bay MAINLAND COAST OF SOUTHERN CALIFORNIA 116 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 A. arenatum 2,6 REDONDO, PALOS VERDES, A. propinquum 2 Los ANGELES HARBOR AREA Euherdmania claviformis 2,5 Figure 4 Polyclinum sp. 8 ean ae 9 , ‘ a KH Lets ; i |. Palos Verdes, Long Pt. Marineland reef, 12 m, San tdemintion carnidenttan = . . re Breccia; 18 June 1970. Trididemnum opacum SPE y/ Onetre Breccia HEU TE TAY at Unidentihedtdidennid 4 2. Marineland pier, 3-S m, wooden pilings; 15 June nidentined dide C j : . 1970. AH Cult be 2 3. King Harbor (Redondo Beach) boat slips, NE of Bee ; ne a x harbor office, floats; 29 June 1970, 10 August 1970. Fearne aa li - 4. East end of King Harbor breakwater, 7 m, breccia istaptta occidentatts 4 9 rocks; 29 June 1970. Eudi i 2,4,5 - : : ey psammion ae 7 5. Rocky Pt., Palos Verdes, 7 m, San Onofre Breccia s. ritteri aye = rocks; 29 June 1970. Dyr76 . 2? te 5 eis UNE Shae ; 7 6. Alamitos Bay, I-3 m, floats and wooden pilings; 3 7 . , “Sip Peete ro Ei a oe i ad nate July 1970, 12 July 1970, 30 August 1970. ae 2 Sua cr ° 15.7 7. Los Angeles Harbor breakwater, 4-10 m, granitic lyoso roduc By 5 x 8 le ie ma pre om um 3 and breccia rocks and debris; 3 July 1970, 12 July erophora annectens > xz 1970, 30 August 1970. Jiplos : " 5 r Diplosomaimacdonaldt S 8. Watchorn Basin, San Pedro, 0-2 m, floats and wood- Order Pleurogona en piling; 15 August 1970. 9. Pt. Vicente, 7 m, breccia rocks; 15 August 1970. Metandrocarpa taylori 4 10. Redondo, 45-50 m, sandy mud; 11 September 1970. Mi dura 27 11. Abalone Cove, 6 m, breccia rock; 21 September Styela montereyensis 17235 1970, 26 September 1970. ; Suiruncata 1,3,4,5.7 12. Resort Point, 5-10 m, breccia rock; 28 September Pyura haustor 3 1970. 13. Abalone Cove, 3 m, breccia rock; 28 September Number of species found: 27 1970. N KEY MAP _ WO SCALE Oreet Sonta Barbara / Oil Pt. Coes Goleta Pt. apne Pt Castillo Santa Barbara Pt ONCu Cuda 7z SANTA BARBARA AREA FIGURE 2 WTA Order Enterogona Aplidium californicum A. solidium A. propinquum Euherdmania claviformis Synoicum par-fustis Didemnum carnulentum Clavelina huntsmani Distaplia occidentalis Cystodytes lobatus Eudistoma psammion Ascidia ceratodes Ciona intestinalis Diplosoma macdonaldi Order Pleurogona Metandrocarpa taylori Botrylloides sp. Botryllus sp. Unidentified styelid Styela truncata S. sp. (barnharti) S. gibbsii S. plicata Halocynthia igaboja Pyura haustor Eugyra arenosa Number of species found: 24 TR 2EME pane Yf PT. DUME AREA ce ASCIDIANS OF Dedeiie lide 13 Se lileilZ 36,10 3 3 3,6,7 3,6 6 Dale l 2 3,6 7,8, 11 SOUTHERN CALIFORNIA i) Aplidium californicum A. solidum Euherdmania claviformis Polyclinum planum . Entrance channel 117 SAN DIEGO AREA Figure § . Dana marina, Mission Bay, 0-1 m, floats and wood en pilings: 5 June 1970. . Pt. Loma, 27 m, shale rock, transect opposite intake line for naval station (Fig. 5); 11 June 1970 . 23 m, shale rock; 11 June 1970 . 20 m, shale rock; 11 June 1970. . 14m, shale rock; 11 June 1970. . 7m, shale rock; 11 June 1970. cast jetty inside Mission Bay, 2-3 m, breccia rock; 11 June 1970. . Ventura Bridge, 3 m, wooden pilings; 14 June 1970. . Ventura Bridge, 0-5 m, rock rip rap; 18 June 1970, 12 September 1970. . New Hope Rock Transect line Pt. Loma, 25-30 m, shale rock; 21 October 1970. . 1S m, shale rock; 21 October 1970. . 10 m, shale rock; 21 October 1970. . 3m, shale rock; 21 October 1970. . Bird Rock, La cobble bed; 28 November 1970. Jolla, intertidal, Order Enterogona NmwnNnNM— KEY MAP WO SCALE Malibu Pt Latigo Pr. 118 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES King Redondo Beach Harbor ~ 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 et 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 densus (Cushman), Pseudohastigerina micra (Cole), S. patagonica, and Truncorotaloides spinuloinflatus (Bandy) brackets deposition between the base of the Middle Eocene and medial Middle Eocene according to the work of Bandy (1949), Bolli (1957, 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 Globigerapsis 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. suteriand Truncorotaloides collacteus (Finlay) suggests equivalence with the Orbulinoides beckmanni Zone of the standard zonation according to the work of Bolli (1957, 1966), Pessagno (1961), 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 (1971). 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 (Bolivina pisciformis and Anomalina regina minor). The relative abundances of major foraminiferid groups and dominant ben- thonic species are listed in Table |. and the complete 128 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 TABLE 2. Faunal reference list ARDATH SHALE BENTHONIC 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) = Cibicidoides coalingensis minor Smith, 1957 : Cibicides praecursorius (Schwager) = Discorbina praecursoria Schwager, 1883 Cibicides sandiegensis Cushman and Hanna, 1927 Cyclammina clarki (Hanna) = Nonionia? clarki Hanna, 1923 Cyclammina pacifica Beck, 1943 Dentalina communis (d’ Orbigny) = Nodosaria communis d@ Orbigny, 1826 Elphidium sp. Eponides dorfi Toulmin, 1941 Eponides minima Cushman, 1933 Fissurina sp. Globulina sp. Gyroidina orbicularis planata Cushman, 1935 Hormosina sp. Karreriella elongata Mallory, 1959 Lenticulina antipoda (Stache)= Robulina cultrata antipodum Stache, 1865 Lenticulina inornata (a Orbigny) = Robulina 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 Textularia 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, 1851 Anomalina coalingensis Cushman and Hanna, 1927 Bulimina capitata Yokoyama, 1890 Bulimina schwageri Yokoyama, 1890 Buliminella sp. Cibicides sassesi Cole, 1927 Dentalina capitata (Bouvigner) = Nodosaria capitata Bouvigner, 1852 Dentalina colei Cushman and Dusenbury, 1934 Dentalina consobrina @ Obrigny, 1846 Dentalina obliquistriata 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 Globulina landesi (Hanna and Hanna)= Polymorphina 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 Lenticulina articulata texana (Cushman and Applin)= Cristellaria articulata texana Cushman and Applin, 1926 Lenticulina gyroscalpra (Stache) = Cristellaria gyroscalprum Stache, 1865 Lenticulina rotula (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 (1971) determined the upper depth limit of Cyclam- mina at 500 m. Size parameters of 5O 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 Anomalina, Gyroidina, Bulimina, and Cibicides is strikingly similar to the abyssal biofacies (1500 m and deeper) of Bandy and 1971 LOCENE FORAMINIFERA 129 Arnal (1960). All factors seem to 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 and species of Stadium Conglomerate foraminifers are listed in Table |. 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, 1960), Olsson (1960, 1963), and Nogan (1964). No milio- 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 shelf 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. LITERATURE CITED Bandy, O. L. 1949. Eocene and Oligocene Foraminifera from Little Stave Creek, Clarke County, Alabama. Bull. Amer. Paleo., 32: 1-222. 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 Sedimentological Trends in the Tertiary Section of Tecolote Tunnel, California. Micropaleo., 9: 117- 170. Bellemin, G, J., and R. H. Merriam, 1958. Petrology and Origin of the Poway Conglomerate, Bull Geol, Soc, Amer,, 69; 199-220. Berggren, W. A, 1968. Phylogenetic and Taxonomic Problems of Some Tertiary Planktonic Lineages. Tulane Studies in Geol, 6; 1-22. Bolli, H. 1957. Planktonic Foraminifera from the Eocene Navet and San Fernando Formations of Trinidad, B. W. I. Bull. U.S. Nat, Mus., 215: 155- 12% . 1966. Zonation of the Cretaceous to the Plio- cene Marine Sediments Based on Planktonic Foraminifera. Boletin Informativo Assoc. Vene- zolana Geologia, Mineria y Petroleo, 9: 3-32. Bukry, D., and M. P. Kennedy. 1969. Cretaceous and Eocene Coccoliths at San Diego, California. Cali- fornia Div. Mines and Geol. Spec. Rept., \00: 33-43, Clarke, B. L., and H. E. Vokes. 1936. Summary of Marine Eocene Sequence of Western North Amer- ica. Bull. Geol. Soc. Amer., 47: 851-878. Cushman, J. A., and A. N. Dusenbury. 1934. Eocene Foraminifera of the Poway Conglomerate of Cali- fornia. Contr. Cushman Lab. Foram. Res., \0: 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. Hanna, M.A. 1926. Geology of the La Jolla Quadrangle. Univ. California Publ. Geol. Sci., 16: 187-246. Hertlein, L. G., and U.S. Grant, IV. 1944. The geology and paleontology of the marine Pliocene of San Diego, California. San Diego Soc. Nat. Hist., 27 2ipp: 1954. Geology of the Oceanside-San Diego Coastal Area, Southern California. Bu/ll., Cali- fornia Div. Mines, 170: 53-64. Jenkins, D. G. 196Sa. 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 Globorotalia Col- lactea Finlay, 1939. New Zealand J. Geol. Geo- physics, 8: 843-848. Laiming, B. 1943. Eocene Foraminiferal Correlations in California. Bull. California Div. Mines, 118: 193-198. Kennedy, M. P., and G. W. Moore. 1971. Stratigraphic Relations of Upper Cretaceous and Eocene Forma- 130 tions, San Diego Coastal Area, California. Bull. Amer. Assoc. Pet. Geol., 55: 709-722. Mallory, V. S. 1959. Lower Tertiary Biostratigraphy of the California Coast Ranges. Amer. Assoc. Pet. Geol., Tulsa, 1-416. Milow, E. D., and D. B. Ennis. 1961. Guide to Geologic Field Trip of Southwestern San Diego County. Geol. Soc. Amer., Cordilleran Section; Guidebook; 24-44. Moore, G. W., and M. P. Kennedy. 1970. Coastal Geology of the California-Baja California Border Area. Amer. Assoc. Pet. Geol., Pacific Section, Guidebook: 4-9. Natland, M. L. 1933. The Temperature and Depth Dis- tribution of Some Recent and Fossil Foraminifera in the Southern California Region. Bull. Scripps Inst. Ocean., 3: 225-230. Nogan, D. S. 1964. Foraminifera, Stratigraphy, and Paleoecology of the Aquia Formation of Maryland and Virginia. Cushman Lab. Foram. Res., Special Pub., 7: 1-50. Olsson, R. K. 1960. 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. Pet. Geol., 47: 643-665. Pessagno, E. A. 1961. The Micropaleontology and Bio- stratigraphy of the Middle Eocene Jacaquas Group, Puerto Rico. Micropaleo., 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. Phleger, F. B., and F. L. Parker. 1951. Ecology of Fora- BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 minifera Northwest Gulf of Mexico. Mem. Amer. Geol. Soc., 46: 1-64. Saito, T. 1962. Eocene Planktonic Foraminifera from Hillsborough Island. Trans. Paleo. Soc. Japan, 45: 209-225. Samuel, O., and J. Salaj. 1968. Microbiostratigraphy and Foraminifera of the Slovak Carpathian Paleocene. Geologicky Ustayv Dionyza Stura, Bratislavia: 1- 260. 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. 1971. Age and Corre- lation of the Eocene Ulatisian and Narizian Stages, California. Bull. Geol. Soc. 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. Pet. Geol., 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: 309-313. 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. Accepted for publication September 24, 1971. BULLETIN SO, CALIF. ACADEMY OF SCIENCE 70(3); 131-135, 1971 A SECOND RECENT SPECIES OF NOETIA, SENSU STRICTO (MOLLUSCA: BIVALVIA) IN THE TROPICAL EASTERN PACIFIC OCEAN BARRY Rotu! ABSTRACT: Recent specimens of an arcoid bivalve differing in some partic ulars from Noetia reversa (Sowerby) are reported from Central American localities, and tentatively referred to Noetia magna MacNeil, described from the (?)Late 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 | 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 131 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 Arca (Noetia) reversa Gray, Grzybowski, 1899: 634-635, pl. 17, figs. 1, la [Not Arca reversa (Gray MS) Sowerby, 1833: 20]. (2) Arca (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 1Department of Geology, California Academy of Sciences, San Francisco, California 94118. (Present address: 1217 Waller St., San Francisco, California 94117). ~ i) disk, although this might be a matter of individual variation. It compares with N. reversa in having strongly opisthogyrate, low, and very posterior beaks; these being at or even behind the posterior end of the cardinal area. The two species are extreme in this respect. Length 80 mm, height 72 mm, convexity 37 mm. The margins are eroded, so that only approximate measurements can be 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 | out of every 10 valves had between 42 and 47 radiating ribs; the remainder gave rib- counts of 31-37. 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. 5. valves in the California Academy of Sciences, Natural History Museum of Los Angeles County, and Stanford University collections showed most falling between 31 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 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. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 70 the California Academy of Sciences collection, fine- ribbed Recent specimens, resembling the holotype of Noetia magna MacNeil (1938), range from Corinto, Nicaragua (Loc. 27226, CAS) to near Mazatlan, Sinaloa, Mexico (Loc. 27583, CAS). The latter lot of 3 living specimens was trawled in 10-17 fathoms by the Templeton Crocker 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 fine-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 Noetia magna 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 Grzybowski’s 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 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 FOSSIL PELECY PODS 133 Figures 11-15. Noetia reserva (Sowerby). “Possible syntypes” of Arca 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 Noetia 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 nota synonym of Arca 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 Arca reversa Sowerby was located at the same institution, but photographs of a “possible syntype™ supplied by Dr. Taylor (Figs. 11-15) 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. Arca hemicardium Koch in Philippi (1843) was 134 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES described as having 20-22 ribs on the “posterior” (actually anterior) portion of the shell and 11-12 on the (posterior) slope. The accompanying figure shows a shell with about 38 ribs. Philippi (1844) himself synonymized Arca hemicardium and Noetia reversa, and many later authors have repeated the synonymy. Synonymies for N. reversa were given by 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 alta for high-beaked shells from Costa Rica and Noetia macdonaldi truncata 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. 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. VOLUME 70 Dall, W. H. 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. 1946. A study of two arcid pelecypod species from western South America. J. Palco., 20: 38-51. Gray, J. E. 1840. Mollusca. Pp. 146-152 Jn Synopsis of the contents of the British Museum. 42nd Edition. British Museum, London. . 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. Neues Jahrbuch fiir Mineralogie, Geologie und Palaeonto- logic, 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. Surv., Prof. Paper 189-A: 1-49. Newell, N. D. 1969. Family Noetiidae Stewart, 1930. Pp. N261-N264 Jn R. C. Moore, et al., eds. Treatise on invertebrate paleontology, part N, 1 (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. Palco., 9: 341-482 (169-310). . 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 half of the Panamic- Pacific faunal province (Panama to Peru). Panamic- Pacific Pelecypoda. Paleo. Res. Inst., Ithaca, 574 pp. Philippi, R. A. (1842)-1845. 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 . 1843, 1844, see Philippi, [1842]-1845, Pilsbry, H. A., and A. A, Olsson. 1941. A Pliocene fauna from western Ecuador. Proc. Acad. Nat. Sci. Philadelphia, 93: 1-79. Reinhart, P. W. 1935, Classification of the pelecypod family Arcidae, Bulletin du Musée royal d'Histoire naturelle de Belgique, V1: 1-68. FOSSIL PELECY PODS 135 Say, T. 1822, An account of some of the marine shells of the United States. J. Acad. Nat, Sci. Philadel- phia, 2: 221-248, 257-276. Sowerby, G. B. 1833, Characters of new species of Mollusca and Conchifera. Proc. 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Finally, as shown by the history of the BUL- LETIN, a journal belongs not to its editors but rather to its readers and contributors. High pro- fessional standards and careful attention to the Instructions to Authors (inside back cover) on the part of contributors, and a demand for schol- arly worth and clear communication on the part of readers will assure continued growth in use- fulness and stature of the SCAS BULLETIN. Patrick H. Wells, Technical Editor James Dale Smith, Managing Editor 1 Opinions expressed in editorials are those of the authors, and are not official points of view adopted by the membership or Board of Directors of the Southern California Academy of Sciences. GENERIC PARTITIONING OF THE SOUTH AMERICAN LEPTODACTYLID FROG GENUS EUPSOPHUS FITZINGER, 1843 (SENSU LATO) JOHN ABSTRACT: D. LYNcH! 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 Niceforonia). 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 16 species (E. coppingeri, E. frog genus Eupsophus contains bolitoglossus, E. columbianus, E. illotus, E. lutzi, E. vanus, E. petropolitanus, E. miliaris, E. nodosus, E. per- 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 (1965) and Gallardo (1965) who recommended removing three Brasilian species (/utzi, miliaris, and petro- politanus) 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 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 with Craspedoglossa_ sanctae- 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 'Museum of Natural History, University of Kan- sas, Lawrence, Kansas 66044 (Present address: Dept. Zoology, University of Nebraska, Lincoln, Nebraska 68508). 1972 the hyoid, thigh muscles, and jaw muscles), ex- ternal morphology, and larval traits (including egg size, deposition sites, and larval biology). Data for some traits are not available for all species but 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 (KU), Naturhistorisches Museum zu Wien, Vienna (NMW), University of Illinois Museum of Natural History (UIMNH), University of Michigan Museum of Zoology (UMMZ), United States Na- tional Museum (USNM), and Universitetes Zoo- logiskke 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. lutzi (KU), E. miliaris (KU, UIMNH, UMMZ), E. 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. lutzi, 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) . SOUTH AMERICAN LEPTODACTYLID FROGS j CHARACTERISTICS OF GROUPS Cei (1960, 1962) and Grandison (1961) defined three species groups of Chilean Eupsophus on the basis of position of the prevomerine teeth relative to the choanae, size of the omosternum, notching of the sternum, and exposure of the tympanum Cei (1960, 1962) frogs sometimes included in Eupsophus and termed also commented on the other these the peruanus-werttsteini group (central Peru) Gallardo (1965) pointed out osteological bases for separat- and Thoropa (southeastern Brasil). ing Thoropa from Eupsophus (shape of the ter- minal phalanges and of atlantar cotyles ). Before listing the characteristics of the species arrangement groups of frogs included in Eupsophus by various authors, it is germane to comment on the use of certain characteristics of the pectoral girdle in de- All of the species included in Eupsophus have arciferal pec- fining species groups of Eupsophus. toral girdles, cartilaginous sterna, and an omo- sternum. Several authors have used the presence or absence of a posterior notch in the xiphisternum (= 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 (sniliaris) has a notched sternum, whereas the smaller species have no notch in the sternum (/utzi and petropolitanus). I have not used this characteristic because it may simply be size-dependent and thus agree with a classification based on average adult size. Three species (/utzi, miliaris, and petropolitanus) 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 /utzi, 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- 4 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Plantar views of feet of Thoropa miliaris (left, KU 92852) and /schnocnema quixensis (right, KU 99010). Figure 1. ent; nasal bones separated; frontoparietal fonta- nelle present; occipital condyles widely separated; terminal phalanges T-shaped. Three species (i//otus, 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- VOLUME 71 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 pads, first finger 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 SOUTH AMERICAN LEPTODAC TYLID FROGS Figure 2. Skulls of Ischnocnema quixensis (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. Prevomerine 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 wertsteini does not; the occip- ital condyles are narrowly separated in juninensis and widely separated in wefrtsteini; 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 lacking. For (left, UIMNH 59643) and Eupsophus roseus purposes of discussion this group is termed the peruanus group. DISCUSSION The six groups defined above are each homoge- neous with the exception the group (juninensis, peruanus, sylvestris, and wettsteini). The major difficulty with that group is the paucity of specimens and information. of last 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. 6 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES The taeniatus group (taeniatus) is distinctive in tadpole morphology and osteology (no quadrato- jugal bone, 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 been 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 taeniatus; Lynch, 1971). The widely separated occipital condyles in Batrachyla and Eupsophus taeniatus 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 taeniatus 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_ taeniata (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 (antartandica, 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 VOLUME 71 of the quixensis group in Eupsophus rests on the simple digital tips (7.e., no pads) and ignores the other osteological data. The oldest generic name for the quixensis group is Ischnocnema Reinhardt and Litken, 1862 (type-species Leiuperus verru- cosus Reinhardt and Litken). Leiuperus verru- cosus Reinhardt and Litken 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 but concealed beneath the skin in illotus and nodosus, and present and visible externally in roseus and vertebralis). 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- basalticus, and reverberii), as well as the three nodosus group species (illotus, monticola, and nodosus) in the genus Alsodes. 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 1s 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 hauthali, and jelskii), (3) the enlarged forearms in reproductive males of the nodosus group are no more distinctive than the condition seen in some Leptodactylus, and (4) the proper generic name for Gallardo’s genus is Mammatodactylus, not Alsodes, if Fitzinger’s (1843) 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- teological 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 fontanelle, 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 dubium. 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 faeniatus group. SOUTH AMERICAN LEPTODACTYLID FROGS 7 Content.—Three species, 2B 1967, B. leptopus Bell, 1843, and 2. taeniata (Girard) 1854. 1843, is earlier name for Batrachyla taeniata (see Cei, 1962) Remarks.—Lynch (1971) pointed out that Bou lenger’s (1882) and Myers’ (1962) combination antartandica Barrio Litoria glandulosa Bell, probably ar of Batrachyla with Eleutherodactylus was in error Eupsophus Fitzinger, 1843 Eupsophus Fitzinger, 1843:31 [Type-species by orig inal designation, Cystignathus roseus Duméril and Bibron, 1841]. Hammatodactylus Fitzinger, 1843:31 [Type-species by original designation, Cystignathus nodosus Duméril and Bibron, 1841]. Bell, 1843:34 [Type-species by sequent designation (Lynch, 1971), Borborocoetes grayi Bell, 1843. Preoccupied by Borborocoetes Schoenherr, 1842 (Insecta: Coleoptera) }. Alsodes Bell, 1843:41 [Type-species by Alsodes monticola Bell, 1843}. Cacotus Giinther, 1868:482 [Type-species by mono- typy, Cacotus maculatus Ginther, 1868}. 2Phrynopus Peters, 1873:416 [Type-species 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: bones widely separated; frontoparietal fontanelle present: occipital condyles narrowly separated (convex atlas): terminal phalanges knobbed. Content.—Eight species, E. (Gallardo). 1970, new combination, E. illotus (Barbour), 1922. E. juninensis (Shreve), 1932, E. montanus (Lataste), 1902, new combination, E. monticola (Bell), 1843, E. nodosus (Duméril and Bibron), 1841, EF. roseus (Duméril and Bibron), 1841, and E. Grandison, 1961. Two other taxa may belong here but incomplete data preclude Cystiganthus sylvestris Tschudi, 1845, and Phrynopus Borborocoetes sub- monotypy, nasal gargolus vertebralis generic assignment: peruanus Peters, 1873. Remarks—Gallardo’s (1970) Alsodes has not been followed here for the reasons cited in the discussion. Before the nodosus group Tecognition of 8 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 Figure 3. is placed in a separate genus the relationships of the Patagonian Telmatobius must be more fully documented. Ischnocnema Reinhardt and Litken, 1862 Ischnocnema Reinhardt and Liitken, 1862:239 [Type- species by monotypy, Leuiperus verrucosus Rein- hardt and Liitken, 1862]. Oreobates Jiménez de la Espada, 1872:87 [Type- species by monotypy, Oreobates quixensis Jiménez de la Espada, 1872]. Definition.—see definition of quixensis group. Dorsum, hand, and side of head of the holotype of Leuiperus verrucosus (ZMK 1180). (=Jschnocnema verrucosa). Content.—TIwo species, I. quixensis and I. 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 Ischnocnema 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 1972 SOUTH AMERICAN terminal phalanges of some fingers knobbed, At present [ am unwilling to place the two species of Ischnocnema in the genus Lleutherodactylus, al- though the possibility cannot be eliminated that the two species represent a single morphological divergence from the binotatus group of Lleuthero- dactylus. a 2s Reinhardt and Liitken (1862) named a single specimen of a frog from Brasil as Leuiperus ver- rucosus in allusion to its warty skin (Pig. 3). In an appendix to their paper, the authors proposed the generic name Ischnocnema. They stated 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- den, and Gorham, failed to notice this generic name. Apparently, the next author to examine the holotype 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 by Nieden (1923). Parker (1927), without exam- ining the unique holotype, 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 Liit- 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 LEPTODACTYIID FROGS ) name to be valid and correctly ass ociated gen ically, proposed a replacement name, Eupsophu Versus, SINCE Verrucosus 18 preoccupied in the VCNUS Eupsophus by Borhorocoetes verrucosus 1902 ( Duméril and Bibron, 1841). Since it now appears that the Philippi, Eupsophus nodosus Brasilian species is valid, a redescription of the type is presented below. Apparently only two specimens have been collected, cach being the holotype of specific epithets of verrucosus. Ischnocnema verrucosa (Reinhardt and Liitken), 1862 Leuiperus verrucosus Reinhardt and Liitken, 1862:171 [Holotype: Zoologiske Museum Kjgbenhayns (Co- penhagen) 1180, Mina Gerais, Brasil]. Ischnocnema verrucosa: Reinhardt and Liitken, 1862: 239. from near Juiz de Féra, Paludicola verrucosa: Boulenger, 1882:171; Neiden, 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 de Janeiro), from Rio Mutum, Coluna, Espirito Santo, Brasil, not examined]. Eleutherodactylus verrucosus [verrucosus of Miranda- Ribeiro, 1937, not Reinhardt and Liitken, 1862] (non Hylodes verrucosus Jiménez de Ja Espada, 1875): Bokermann, 1966:40. Eupsophus versus Gorham, 1966:120 [Replacement name for Eupsophus verrucosus Miranda-Ribeiro, 1937; preoccupied in Eupsophus by Borborocoetes verrucosus Philippi, 1902 (= Eupsophus nodosus Dumeéril 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% 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: fold absent: fingers long and slender (Fig. 3) with prominent subarticular discoidal 10 BULLETIN SOUTHERN CALIFORNIA 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 tips 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 Jiménez de la Espada, 1875, and H. verrucosus Jiménez 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, 1841]. ACADEMY OF SCIENCES VOLUME 71 Definition.—see definition of miliaris group. Content.—Three species, 7. lutzi Cochran, 1938, T. miliaris (Spix), 1824, 7. petropolitana (Wandol- leck), 1907. Remarks.—Excellent accounts of the species of this genus are available in the works of Boker- mann (1965) and Cochran (1955). Ololygon Fitz- inger is frequently cited as a synonym, whereas in fact the generic name Ololygon was proposed in 1843 for Hyla strigilata Spix and is therefore a synonym of Hyla. The species included in Eupsophus by Gorham (1966) but not allocated to Batrachyla, Eupsophus, Ischnocnema, or Thoropa here are listed below with their taxonomic disposition. Borborocoetes bolitoglossus Werner, 1897, is presently considered a synonym of Craspedoglossa sanctaecatherinae (Bokermann, 1966). Borborocoetes columbianus Werner, 1899, is presently included in the genus WNiceforonia (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. Insuetophrynus acarpicus, un nuevo leptodactilido firmisternio sudamericano (Am- phibia, Anura). Physis, 30:331-341. Bell, T. 1843. Reptiles. /n C. Darwin [ed.] The Zoology of the Voyage of H.M.S. Beagle under the command of Captain Fitzroy, London. pt. 5, S1 pp. 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 sdbre as espécies de Thoropa Fitzinger (Amphibia, Leptodactyli- dae). Anais da Academia Brasileira de Ciéncias, 37/25 25—5317. — 1966. Lista anotada das localidades tipo de anfibios brasileiros. Servico documentacao. Sao Paulo, RUSP, 181 pp. 1972 SOUTH AMERICAN Boulenger, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collections of the British Museum, 2nd ed., London, 503 pp. Cei, J. M. 1960. A survey on the leptodactylid frogs, genus Expsophus in Chile. Breviora, 118: 1-13. ——. 1962. Batracios de Chile. Santiago, 128 ++ cviii pp. Cochran, D.M. 1955. Frogs of southeastern Brazil. Bull. U.S. Nat. Mus., (222), 423 pp. Cope, E. D. 1865. Sketch of the primary groups of Batrachia Salientia. Nat. Hist. Rev., 5:97— 120. Fitzinger, L. 1843. Systema Reptilium. Vindobonae. 106 pp. Gallardo, J. M. 1962. Los géneros Eupsophus y Batrachyla (Anura, Leptodactylidae) en la Ar- gentina, y la verdadera identidad de Paludicola illota Barbour. Rev. Mus. Argentino Cs. Nats. Cienc. Zool., 8(10):113—122. 1965. #*#mpecnoo 20 30 40 70 80 50 60 90 WEIGHT OF PORPOISE (kilograms) Figure 1. 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” @. 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. Stenella graffmani. Weight of heart relative to total body weight for 29 males and 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% ). HEART 200 z 150 100 8 50 O 10 20 30 40 ORGAN WEIGHTS OF PORPOISES IMMATURE MALE IMMATURE FEMALE SEXUALLY MATURE MALE SEXUALLY MATURE FEMALE # PREGNANT FEMALE gweedo 70 50 60 80 90 WEIGHT OF PORPOISE (kilograms) Figure 2. and 10 females. TABLE 3. Log-log regression analyses of relationships between organ weights and total body weight in S. graffmani and S. cf. S. longirostris. Sample at bt rh size S. graffmani Heart -1.25 0.769 0.948 47 Lungs —0.92 0.880 0.990 35 Liver 2.18 1.114 0.922 5 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. S$. 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 1Values of a, b, and correlation coefficient r in linear regression equation Log Y=a-+b Log X, where Y = organ weight and X = total body weight, in gms. Stenella cf. S. longirostris. Weight of heart relative to total body weight for 4 males 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 weighed from 191-272 gm and from 0.37-0.52% (average: 0.46%) of total body weight. The data are insufficient to determine whether relative weight of the heart decreases during development in this species. weight) Lungs S. graffmani (Fig. 3). 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%) of total weight. S. longirostris (Fig. 3). The lungs of 4 individ- uals, all calves and subadults (from 118-149 cm total length and from 18.1—33.6 kg total weight weighed from 599-1112 gm and from 3.09-—3.39 (average: 3.28%) of total weight. WEIGHT OF LUNGS (grams) BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 2600 “ aol: LUNGS S. graffmani o IMMATURE MALE 2200 0 IMMATURE FEMALE e SEXUALLY MATURE MALE ° = SEXUALLY MATURE FEMALE 2 ° MALE FETUS 2000 4 FEMALE FETUS S. cf. S. longirostris 4S IMMATURE MALE ® IMMATURE FEMALE 1800 1600 1400 1200 1000 800 600 400 200 O 10 20 30 40 50 60 70 80 WEIGHT OF PORPOISE (kilograms) Figure 3. Stenella graffmani. Weight of lungs relative to total body weight for 22 males and 15 females. Stenella cf. S. longirostris. Weight of lungs relative to total body weight for 2 males and 2 females. 90 1972 ORGAN WEIGHTS OF PORPOISES ASG LIVER o IMMATURE MALE 1800 0 IMMATURE FEMALE e SEXUALLY MATURE MALE = SEXUALLY MATURE FEMALE 4# PREGNANT FEMALE 1600 4 MALE FETUS + FEMALE FETUS Oo iS S 1400 SH tu > 1200 =I) ve oO 1000 —_ ae © 800 uJ = 600 400 200 O O 10 20 30 WEIGHT OF Figure 4. Stenella graffmani. Weigh Liver S. graffmani (Fig. 4). The livers of 16 adults (from 175-218 cm total length and from 55.4— 84.0 kg total weight) weighed from 1281-2195 gm and from 2.06-3.17% (average: 2.53%) of total weight. Relative weight of the liver appears 40 50 60 70 80 PORPOISE (kilograms) t of liver in 36 males and 27 females. to increase early in development (avera fetuses and infants was 1.84‘ 2.22% ) and remain nearly constant ther S. longirostris (Fig. 5). The liver (168-177 cm total length and 43.5 weight) weighed from 832-997 gm an 2.10% (average: 1.90%) of total we 90 26 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES LIVER — 1600 wn = < 2S 1400 oO > 5 1200 LL oe) 1000 Kk ac 2 lJ ~800 = 600 400 200 fo) fe) To) 20 30 40 VOLUME 71 © IMMATURE MALE © IMMATURE FEMALE @ SEXUALLY MATURE MALE ® SEXUALLY MATURE FEMALE 4 PREGNANT FEMALE 90 80 50 60 70 WEIGHT OF PORPOISE (kilograms) Figure 5. Stenella cf. S. longirostris. males and 10 females. of this species have smaller livers than subadults of S. graffmani; the livers of 21 calves and sub- total weight weighed from 974-1457 gm and from 2.08-3.00% (average: 2.42%) of total weight. Weight of liver relative to total body weight for 4 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 WEIGHTS OF PORPOISES 1000 a © 900 KIDNEYS ATURE MALE =" o IMMATURE EB 800 0 IMMATURE FEMALE ‘ © SEXUALLY MATURE MALE =™ SEXUALLY MATURE FEMALE B 4 PREGNANT FEMALE 700 + MALE FETUS > 4+ FEMALE FETUS ~ 600 500 400 300 WEIGHT OF KIDNEYS (grams) 200 fo) 10, 20> 30° «40% « 50) 60" 70" 80 90 WEIGHT OF PORPOISE (kilograms) Figure 6. Stenella graffmani. Combined weight of kidneys relative to total body weight for 35 males and 27 females. neonatals and near-term fetuses (74-86 cm total from 129-178 gm and from 0.67—1.11% (average: length and 3.4—6.8 kg total weight) ranged from 0.84%) of total weight, and those of 15 adults 49-70 gm (average: 59 gm) and from 0.91— (175-218 cm total length and 55.4—84.0 kg total 1.44% (average: 1.09%) of total weight. The weight) ranged from 426-968 gm and from 0.78- kidneys of 31 calves and subadults (129-178 cm 1.20% (average: 0.98%) of total weight. The total length and 23.6-53.2 kg total weight) ranged animal is asymmetrical with respect to weight of LEFT KIONEY LARGER RIGHT KIONEY LARGER (14) (43) w a a =) ° = fa) = 15 10 5 (okie) 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: 0.04-0.15% (average: 31 gm) and from 0.11%) of total weight, 28 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 and 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- 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. parable size of S. graffmani. 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), (1.055). That of S. longirostris (0.820) falls between those for D. 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 and Phocoena phocoena delphis and S. coeruleoalba. 1972 ORGAN WEIGHTS OF PORPOISES © IMMATURE MALE 0 IMMATURE FEMA 900 KIDNEYS e SEXUALLY MATURE MA @ SEXUALLY MATURE FEMALE # PREGNANT FEMALE 800 700 600 500 400 300 WEIGHT OF KIDNEYS (grams) 200 O 10 20 30 40 50 60 70 BO 90 WEIGHT OF PORPOISE (kilograms) Figure 8. Stenella 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 Pilleri’s (1969) value of b for Phocoena phocoena both species dealt with here (0.965 and 0.940) (0.685) is very much lower than those for the 4 are lower than those obtained by Pilleri for §. delphinids (sensu stricto). coeruleoalba (0.977) and D. delphis (1.089). The odontocete spleen has a relatively large 30 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 110 100 SPLEEN o IMMATURE MALE 90 0 IMMATURE FEMALE e SEXUALLY MATURE MALE = SEXUALLY MATURE FEMALE # PREGNANT FEMALE 80 ~ MALE FETUS 4+FEMALE FETUS WEIGHT OF SPLEEN (grams) 20 WEIGHT OF PORPOISE (kilograms) Stenella graffmani. Weight of spleen relative to total weight for 37 males and 30 Figure 9. 28 females. 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 ie) 40 VOLUME 71 50 60 70 80 90 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. 1972 ORGAN WEIGHTS OF PORPOISES 110 © IMMATURE MALE O IMMATURE FEMALE 100 SPLEEN © SEXUALLY MATURE MA @® SEXUALLY MATURE FEMA # PREGNANT FEMALE WEIGHT OF SPLEEN (grams) O 10 20 30 40 50 60 70 80 WEIGHT OF PORPOISE (kilograms) Figure 10. Stenella cf. S. longirostris. Weight of spleen relative to total weight for 4 males and 9 females. ACKNOWLEDGMENTS Commission. 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. LITERATURE CITED Support was provided by the National Marine Fish- Cowan, D. F. 1966. Observations on the 90 pilc eries Service and the Inter-American Tropical Tuna +f t ww te whale Globicephala melaena: organ weight and growth. Anat. Rec., 155:623-628. Gihr, M., and G. Pilleri. 1969. On the anatomy and biometry of Srenella styx Gray and Delphinus delphis L. (Cetacea, Delphinidae) of the Western Mediterranean. /n Pilleri, G. [ed.], Investigations on Cetacea. Vol. I, Bentelli A. G., Berne, 219 pp. Jubb, K. V.. and P. C. Kennedy. 1963. Pathology of domestic animals. Academic Press, New York, 2:613 pp. Huxley, J. S. 1932. Problems of relative growth. Dial Press, New York, 276 pp. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 Perrin, W. 1969. Using porpoise to catch tuna. World Fishing, 18(6):42—45. Pilleri, G., and M. Gihr. 1969. Zur Anatomie und Pathologie von J/nia geoffrensis De Blainville 1817 (Cetacea, Susuidae) aus dem Beni, Bolivien. In Pilleri, G. [ed.], Investigations on Cetacea. Vol. I, Bentelli A. G., Berne, 219 pp. Slijper, E. J. 1959. problems in porpoises and seals. Zool. 13 (1st suppl.) :97—113. Organ weights and symmetry Arch. Neéer. 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! 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 centrations of a few forms (glycine and proline, for example) and low concentrations of a large number of amino acids, and 3) the pool is sig- nificantly small in’ terrestrial and fresh water species. Awapara (1962) postulated that the free amino acid pool of marine invertebrates 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 Limmnoria were subjected to sublethal levels of reduced dissolved oxygen concentrations (Anderson and FREE AMINO ACID POOL OF NEANTHES ARENACEODENTATA Reish, 1967). Egg production was suppre hemoglobin compensation occurred in Neanthi arenaceodentata when it was subjected to lethal concentrations of dissolved oxygen (D 1969; Raps and Reish, 1971). The laboratory induced changes in the organism discussed above serve to indicate that metabolic changes may be occurring within the organism and that perhaps changes at the organismic level Therefore, it is the purpose of this study to measure the ef and molecular level can be correlated. fects of lowered dissolved oxygen concentrations and lowered salinities on the free amino acid pool of an inbred strain of Neanthes arenaceo- dentata and to correlate these results with those previously gathered with 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 this species on the be employed as a monitoring measure of marine pollution. METHODS The Reish strain of Neanthes arenaceodentata (Moore) was chosen as the experimental animal because of its uniform genetic background. The stock colony was fed powdered alfalfa and the green alga Enteromorpha 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 Enteromorpha crinita). The flasks were sealed with a rubber stopper and in the case of the dissolved oxygen experiments each flask was flushed with Ne gas according to the method of Reish and Richards (1966). Reduced chlorinity conditions were made by diluting 100% sea water (19.2 %« chlorinity) to 90% (16.4 %c-) and 70% (12.8 %-). All flasks were kept in a cold bath at 18° C for seven days. At seven days, in the case with the dissolved oxygen experiments, the rubber stopper was removed and the dissolved oxygen measured with an electrode. flasks placed into separate vials containing 80% ethanol. The worms were removed from the and Extraction of amino acids was accomplished by leaving the worms in the 80% ethanol for 48 to from the 50 hours. The worms were removed 34 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 Means, standard deviations,’ and Student /-test® of the 14 most abundant amino nitrogen compounds at varying concentrations of dissolved oxygen. Control (5.8 mg/l) 2.7 mg/1 1.46 mg/1 0.83 mgz/1 Amino Acid Spot Means, S.D. Means, S.D. t-test Means, S.D. f-test Means, S.D. t-test Aspartic Acid DrA0 ENT 2.08 + 0.61 1.25 2 US =10:5 2 ede 0.70 + 0.33 4.47%" Glutamic Acid 2.77 + 0.96 273210574) (ONL 3572/0164 287*" 2.870 N250129 L-amino Adipic Acid 0.47 + 0.30 0.23 + 0.20 0.00 0.52 + 0.20 0.00 0.28 + 0.31 0.00 Serine 4.04 + 1.11 3.81 +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)== A Ole LS L-threonine Sypyl ey tal) 1121063 6:28** W777 126) Si48h"* 306 eEsOSm ONS. Glutamine 2.65 + 0.96 Sahel els 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 MS 707) 4 SS 2199 a 40. 0'30 2.38 + 0.80 2.23* Valine, norvaline, L-methionine 3:35/-= 2611 3 21eE pO 28" 4242 i- 12 5 O85 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 = eS 0358) 3.21+0.90 0.24 2.08 + 0.95 2.73* Proline 14.35 + 4.65 16.64 + 3.84 1.26 0.98 2445+ 4.66 5.10* 15.99 + 3.05 1 Means and standard deviations recorded as per cent values 2 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 HCl. 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 mp wavelength for the purple and pink spots and at 300 my, 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 t-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 t-tests. 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/l, six and possibly eight compounds 1972 FREE AMINO ACID POOL OF TABLE 2. Means, standard deviations! and Student /-test’ NEANTHES ARENACEODENTATA ‘of the 13 most abundant amino nitrogen cor under varying salinities, Control (normal salinity) Amino Acid Spot Means, 8.D. Aspartic Acid DAO ras0 27) Glutamic Acid 2.77 + 0.96 Serine 4.04 - 1.11 Glycine 29.03 + 6.00 L-threonine top hy feu bel US) Glutamine 2.65 + 0.96 Citrulline OPS See 10122 L-alanine 2.92 + 0.96 Tyrosine 3.14 + 0.89 Valine, norvaline, L-methionine 53 '572e 2761 Phenylalanine, leucine, isoleucine 11.04 + 4,29 Asparagine Shl2.==10'93 Proline 14.35 + 4.65 1 Means and standard deviations recorded as per cent values 2 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, tryrosine, at spot 32 which contained valine, norvaline, and methionine. At least three compounds differed significantly at mean dissolved oxygen levels of 1.46 mg/I; namely, glutamic acid, threonine, and spot 35 which may contain phenylalanine, leucine, and isoleucine. At the mean dissolved oxygen level of 0.83 mg/l, 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, I-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 90% Salinity 10 linit Means, S.D, f-tent Means, 5.D t 1.62 +1.50 1.34 1.60 0,462 1.76 Zep un OFS) 1869 1.60 + 0.69 0.29 4.27+1.25 0.46 3.79 4+- 0.57 0.67 4218: 7.92 4,39"™ 43,58 + 10.73 3.86** Vivace 12: 93.76"" 3.16 2.13 0.48 SIS) am UI Ppa 1,27 0.75. 3.837" 0.80 + 0.46 1.78 0.85 + 0.92 1.15 2.49 + 1.40 0.86 LS8ek P27) 2912" 1.74+1.30 3.04** 154+ 1.14 3.80** 2:8 = 1641) 2.85" 3.96+2.49 1.28 6:25 3 736n 2920 (SiS 38.04 3.62 + 0.84 1.38 3.25 + 0.84 0.36 1.28 13.38 + 3.30 0.56 16.61 + 3.58 additional difference was noted in the totals of the optical densities of all ninhydrin-positive 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 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 is com- taurine in this study; therefore, no estimation of its presence could be made. Other Camien et al. (1951), Kittredge et al. (1962). and Clark (1968b) have found similar results in all of the major marine invertebrate phyla. workers, The free amino acid pool of N. arenaceodentata also showed a large amount of variability one animal to another. Most of the vai found among the amino acids present in the 36 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Clark (1968b), and Mearns and other concentrations. Reish (1969) polychaete species. studied were from natural field populations. The observed similar results in However, the organisms they specimens of Neanthes arenaceodentata used in this study 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- mentals 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 amounts of amino acids at each level of lowered relative VOLUME 71 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 N. 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 POOL OF Thus far, all explanations advanced for changes in the free amino acid pool are based on the as- sumption that under low dissolved oxygen condi- tions, the organism must compensate in some manner for loss of the terminal electron acceptor, oxygen. If 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, pers. 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,, 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 NEANTHES ARENACEODENTATA increase in the relative amount of glycine some of the decreases were not signitn least eight and possibly 12 (counting the multipl of one spot) amino acids decreased in relative proportions at 90% salinity and at least ten possibly 15 at 70%. However, the glycine cor centration remained the same from 90% to 70% salinity. All of these results indicate that the total free amino acid pool is osmotically diluted by decreased salinity and that glycine is maintained at a constant minimum level while the other free amino acids decrease. No explanation is advanced herein to explain this observed glycine stability. From the results of both the dissolved oxygen and salinity experiments, it is apparent that the free amino acid pool is affected, either directly or indirectly by changes in this organism's en- vironment. Alterations in dissolved salinity concentrations accompany some forms of water pollution. Schafer (1961, 1963) has pre- oxygen and viously noted distinct changes in the free amino acid pool of organisms living in a polluted en- vironment. Therefore, monitoring the effects of these variables on the free amino acid pool may 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 cuneata (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. 1962. Free amino acids in inverte- brates: a comparative study of this distribution and metabolism. Pp. 158-175. Jn Holden, J. T.. [ed.], Amino acid pools. Distribution, formation Awapara, J. and function of free amino acids. Elsivier. Amsterdam. Ball, G. H., and E. W. Clark. 1953. Species dif- : : Zool., 2:138—141. Camien, M. H., H. Sarlet, G. Duchateau, and M Florkin. 1951. Non-protein amino muscle and blood of marine and er crustaceans. J. Biol. Chem., 193:881—885 Clark, M. E. 1968a. A survey of the effect of 38 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES osmotic dilution on free amino acids of various polychaetes. Biol. Bull., 134:252-260. —. 1968b. 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 heteronereis. Biol. Rev., 36:199-236. Davis, D. 1969. The effects of lowered dissolved oxygen levels on the reproduction of the poly- chaetous annelid Neanthes 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, Mus musculus. 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. Jn 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. Mahler, H. R., and E. H. Cordes. 1968. Basic Biological Chemistry. Harper and Row, New York, 509 pp. 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. 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. VOLUME 71 Micks, D. W., and F. J. Gibson. 1957. The char- acterization of insects and ticks 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. Raps, M. E., and D. J. Reish. 1971. The effects of dissolved oxygen on the hemoglobin levels of the polychaetous annelid, Neanthes arenaceodentata. Mar. Biol., 11:363—368. Reish, D. J. 1967. varying dissolved oxygen concentrations. Water Poll. Res., 3:199-266. Relationship of polychaetes to Adv. 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- ternat. J. 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 Neanthes arenaceodentata. Mar. Biol., 3:252-255. Roberts, E., and P. Borges. 1955. Patterns of free amino acids in growing and regressing tumors. Canadian Res., 15:697-699. 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. 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 1972 FREE AMINO ACID POOL OF 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, J. W., K. Allen, and J. Awapara. 1959. Free amino acids in some aquatic invertebrates. Biol. Bull., 117:371-381. 1963. Stephens, G. C. Uptake of Organic material NEANTHES AREN {CLODENTATA by aquatic invertebrates », Accum amino acids by the bamboo worm, Clymene torquata, Comp. Biochem, Physiol., 10:19] Stephens, G. C., and R. A. Virkar. 1966. Uptake « organic material by aquatic invertebrates. 4, The influence of salinity on the brittlestar, Ophiacti arenosa. Biol. Bull., 131:172—185 Accepted for publication February 8, 1972 AN ANNOTATED KEY TO THE CERCARIAE THAT DEVELOP IN THE SNAIL CERITHIDEA CALIFORNICA W. E. Martin! ABSTRACT: An annotated key is provided to the 18 species of trematodes 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 found. Perhaps the cercaria of Ascocotyle 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 Ascocotyle 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. more will be KEY TO CERCARIAE INFECTING CERITHIDEA CALIFORNICA 1. a. Tail forked 2 b. Tail nonforked _ 4 1 Biology Depi., University of Southern California. Los Angeles, California 90007. 40 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 \ \ a O *\ [( \\ 6 \ (e | jus OG" 0 “aX \w ] »\| | | \ / ~ \ A ra | \ | } A r I Li | i I} WS ){/ &| Se 4 WTA \ ££ a Ss a LN OE ae =a \ 6 SSS 7), AS mm_, SSS mee eee Figures 1-4. 1. Schistosomid. 2, 3. Strigeids. 4. 4 \| y Notocotylid. | | \| Vi 9g 10., Un. 2-72; With-eyespots,_ Avian blood ifluke Develops: 7 ras) 51. #5) 10s (EchinostomidsMmOmeletecoohyid! ini sporocysts) (Hig) sj Uhis mayabe the! same 7 ee) IMicrophallidsmo mie Philoputhalaide as Austrobilharzia penneri Short and Holli- Rag Pa ates man, 1961 and Holliman (1961) which uses Cerithidea scalariformis as intermediate host 4 4 Tail withfins...... 14 and chicks, parakeets, and pigeons as experi- bee iaill without tins) rrr 5 mental adult hosts. However, we have ex- ? posed chicks and pigeons to large numbers 5. a. With QYCS IOUS accesses 6 of the cercaria from C. californica with nega- b. Without eyespots 8 tive results. Therefore this may be a different 6. a. With a pouch at each _posterior-lateral species of Austrobilharzia. Further work is in margin of body—Catatropis johnstoni Martin, progress. 1956 (Fig. 4). Develops in rediae, encysts braawithouteeyespots, eee 3 on almost any object, adults experimentally 3. a. Flame cells in groups of twos—Strigeid ial ea OP GUase IG cyl Cesemled Ly cercaria (Fig. 2). Develops in sporocysts, Martin (1956). i cercariae encyst in muscles of Fundulus b. Without a pouch at each posterior-lateral parvipinnis. Life cycle unknown. Smaller margin of body -_________-____-__- 7 than 3b. 7. a. Encysts in dish—Himasthla rhigedana Dietz, b. Flame cells in groups of threes—Meso- 1909 (Fig. 5). Develops in rediae, encysts stephanus appendiculatus (Ciurea, 1916) on almost any object, adults in willet and ex- Lutz, 1935 (Fig. 3). Develops in sporocysts, perimentally in ileum of chick. Life cycle encysts in muscles of Fundulus parvipinnis, described by Adams and Martin (1963). adults experimentally in intestine of chicks, Cercaria has median pigment spot nearly in naturally in shore birds. Life cycle described line with eyespots. by Martin (1961). b. Does not encyst in dish—Pygidiopsoides 1972 Figures 12-18. icolids. a my ws a b. Without a stylet KEY TO THE CERCARIAE OF CALIFORNIA HORNSHELL SNAILS os 12,17. Echinostomids. 13, 14. Ren- 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). With stylet (small “spear”) near mouth— Microphallids 1 and 2. 1. Develops in sporocysts (Fig. 7). than 2. 2. Develops in sporocysts (Fig. 8). Smaller than 1. Sarkisian (1959) described Maritrema wuca 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. Larger 9. a. With pouch opening at end of tail 10 b. Without pouch opening at end of tail 11 10, a. With spined collar—VParorchis (Nicoll, 1906) (Fig. 11). Develops in rediae encysts on almost any object, adults in cloaca of shore birds. Life Stunkard and Cable (1932) as P. avitus which was declared a synonym of P. acanthus by Cable and Martin (1935). I have established infections in chicks by injecting them with metacercariae per cloaca but the percentage of success is low. This cercaria is larger than 10b. Without spined ganensis McIntosh, 1938 (Fig. 9). Develops 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. acanthu cycle described by collar—Cloacitrema michi- With spined collar 12 Without spined collar 13 Encysts in radular muscle—A canthoparyphium spinulosum Johnston, 1917 (Fig. 10). De- velops in rediae, encysts in Cerithidea cali- fornica especially in radular muscles, adults experimentally in intestine of chicks. Life cycle described by Martin and Adams (1960, 1961). Cercaria larger than 12b. 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. Usually forms clusters by adhesions of proximal portions of tails, Y-shaped ex- cretory bladder—Renicola buchanani (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. Does not form clusters, Y-shaped excre- tory bladder—Renicola cerithidicola Martin, 1971 (Fig. 14). Develops in sporocysts, encysts in gills of Fundulus parvipinnis, young adults experimentally in kidneys of Larus californicus, Martin (1971). With dorso-ventral tail fin only Se A bs} With dorso-ventral and lateral tail fins __ 16 With eyespots—Phocitremoides ovale Martin. 1950 (Fig. 15). Develops in rediae, encysts under scales of Fundulus parvipinnis and Atherinopsis californica (Girard), adults ex- perimentally in chicks and cats. Life cycle described by Martin (1950c). Without eyespots—Echinoparyphiumi sp. (Fis. 17). Develops in rediae, sometimes forms weak-walled cysts in dish, life cycle unknown. 42 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Flame cells in groups of twos—Euhaplorchis Martin, 1950 (Fig. 16). De- velops in rediae, encysts on brain of Fundulus parvipinnis, adults experimentally in small intestines of chicks, Larus californicus, cats (Bamberger and Martin, 1951), monkeys, etc. Life cycle described by Martin (1950a). Gear californiensis b. Flame cells in groups of threes—Stictodora (Parastictodora) (Martin, 1950) (Fig. 18). Develops in rediae, encysts in various tissues of Fundulus, parvipinnis and Gillichthys mirabilis Cooper, adults experi- mentally intestines of chicks and Larus californicus. Life cycle described by Martin (1950b). hancocki in small LITERATURE CITED Adams, J. E., and W. E. Martin. 1960. Life history of Himasthla sp. an echinostome trematode. J. Parasit., 46:(supple.):15. 1963. Life cycle of Himasthla rhigedana Dietz, 1909 (Trematoda: Echinostomatidae). Trans. Amer. Micros. Soc., 82: 1-6. 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 californiensis 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., III, 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 californiensis 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. VOLUME 71 1950c. Phocitremoides ovale 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: Notocotylidae). 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. 1964. Life cycle of Pygidiopsoides spin- dalis Martin, 1951 (Heterophyidae: Trematoda). Trans. Amer. Micros. Soc., 83:270-272. 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 uca, 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 Short, R. B., and R. B. Holliman, 1961. Austro- bilharzia penneri, a new schistosome from marine snails. J. Parasit., 47:447—452. Stunkard, H. W., and R. M. Cable. 1932, ‘The life KEY TO THE CERCARIAE OF CALIFORNIA HORNSHELL SNAILS history of Parorchis avitus (Linton) a ty from the cloaca of the Biol, Bi 328-338. gull Accepted for publication September 25, 1970 NEW SPECIES OF ROBBER FLIES OF THE GENERA WILCOXIA AND METAPOGON (DIPTERA: ASILIDAE) J. WiLcox! 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. 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 cinerea. were made with an_ ocular micrometer; the face and wings at 40x, and the antennae at 75x. Wilcoxia martinorum, 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 (.54) width of one eye. Antennae black, grayish pollinose; hairs white; bristles white, two weak bristles below on segment | 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. lateral spot on each side and a central posterior spot. and 7 with a small central posterior spot. Hairs white. the antennae 15/28 2-6 with an anterior sparse, long on sides of 1-3: five lateral bristle-l hairs on 1. Sternites gray pollinose. hairs erect wi becoming shorter apically. Genitalia white. brown. 121171 Mohler Place. Anaheim, California 92806 44 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Legs black, fore and middle femora dorsally and laterally, hind femora dorsally and sides of apical third, gray 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. 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., 86, 72, Organ Pipe Cactus National Monument, 2 mi N Headquarters, 11 No- vember 1966 (R. J. Hamton) EF; Yuma Co., 82,12. 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., 6, Death Valley National Monument, 1.5 mi SW Wildrose Station, 3100 ft, 6 November 1968 (P. H. Arnaud, Jr.) at flowers of Chrysothamnus peniculatus (Gray) Hall, CAS; River- side Co., 2, Coyote Creek, 6 November 1963 (E. I. Schlinger) UCR; 56, 22, 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; 44, 72, Joshua Tree Na- tional Monument (JINM); Pleasant Valley, Fried Liver Wash, 22 October 1967, 29 October 1965 (E. L. Sleeper) part blacklight, CSCLB; 64, 109, 5 and 10 mi S Oasis, 17 November 1947, 10 December 1962 (A. F. Howland, I. J. Wilcox, and J. Wilcox) JW; 24, Palm Desert, 10 October 1964 (E. Fisher) EF; 448, 32, Palm Springs, 12 November 1944, 19 No- vember 1943 (A. L. Melander) USNM; 774, 739, Palm Springs, 23 December 1950, 29 December 1953 to 8 January 1954, 21 January 1953 (P. H. Arnaud, Jr.) JW; 54, 22, Palm Springs, 21 January 1953, 16 December 1964 (M. W. Stone and J. Wilcox) JW: 94, 1392, Palm Springs, Chino Canyon, 22 to 26 Female. VOLUME 71 December 1950 (P. H. Arnaud, Jr.) JW; 5@, 1792, Palm Springs, Tahquitz Canyon, 24 November 1944 (A. L. Melander) USNM; 464, 1359, White Water, 23 November to 27 December 1947, 6 January 1948 (A. F. Howland, C. H. Martin, D. Martin, I. J. Wilcox, and J. Wilcox) CHM, CIS, JW, UCD; 154, 129, White Water, 11 November 1966, 13 November, 16 December 1962 (EF. Fisher and R. J. Hamton) EF. NEVADA. Nye Co., &, Rhyolite, 2 October 1936 (A. J. Basinger) CAS. MEXICO. Sonora, 36, 22, 13 mi SW Sonoyta, 12 November 1966 (E. Fisher) EF. Wilcoxia 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 11-11-40-12 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., 266,42, Mammoth, 9 August 1957 (I. J. Wilcox and J. Wilcox) EF, JW; Nevada Co., 9, 3 mi N 1972 Boca, 23 July 1961 (Ff. D. Parker) UCD; 9, | mi S Hobart Mills, | September 1957 (EB. G. Linsley) Chrysothamnus viscidiflorus var. typicus, CIS; 4, Prosser Dam, 15 July 1966 (D. R. Miller) UCD; 4,29, Truckee, 17 August 1955 (EB. G. Linsley) Com- positae, CIS; Tulare Co., 9, Mineral King, 8 August 1959 (W, E. Simonds) CDA; 4, 2, 0.5 mi E Smith Meadow, Nine Mile Canyon, 7850 ft, 5 August 1961 (C, W. O’Brien) CIS. NEVADA. Douglas Co., 4, Topaz Lake, 17 August 1960 (A. S. Menke) UCD. Wileoxia 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 (1 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. Female. 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, NEW SPECIES OF ROBBER FLIES always collected rare and interesting specie many trips. Other specimens examined; NEW MEXICO. C; tron Co., 3, 29, Datil, Continental Divide July 1930 (T. F. Winburn and R. H. Painter) RHP, JW. ARIZONA Apache Co., Chis Lee (Chinle), 26 July 1935 (Brues) USNM. UTAH Kane Co., 9, 16 mi W Glen Canyon, 23 September 1969 (P. H. Timberlake) Sutienzia sp., UCR; Millard Co., 6, 23 mi W Delta, 4900 ft, 4 September 1965 (R. H. Painter and E. M. Painter) RHP; Uintah Co., 9, 16 mi SW Vernal, 5000 ft, 7 September 1965 (R. H. Painter and E. M. Painter) RHP; county un determined, 9, Showell, 20 August 1932 (G. F. Knowlton) JW, [anterior fascia on abdomen of this specimen much broader and faint fascia on tergite 6). Wilecoxia pollinosa, new species Description: Male. 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 (.74) width of one eye. Antennae black: short sparse hairs white: one weak white bristle below on segment | 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, 1 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 36/57 (.63) length of discal cell: third vein branched s beyond end of discal cell: posterior cells broadly open anal cell narrowly open. Female. Length 10 mm. Weak bristle below 2. Mesonotal pollen with a yel- on antennal segment 2. lowish cast: two posterior and four anteri central bristles: two scutellar bristles. ments 1-6 and 7 terminalia and spines reddish-brown, hairs yellowish basally, grayish ose, 8 46 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Hairs and bristles of legs white to yellowish. Anterior crossvein at 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. Timberlake) UCR. Allotype: female, New Mexico, Eddy Co., 0.5 mi N State Line, Hwy. 180 and 62, 2 October 1962 (C. S. Papp) UCR. Other specimens examined: NEW°MEXICO. Eddy Co., 2, 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- notum reddish; 3 scutellar bristles]; Torrance Co., 2, Gran Quivira, 11 August 1931 (R. H. Painter) RHP [third antennal segment missing and scutellum crushed]. TEXAS. Hudspeth Co., 2, 30 August 1940 (D. J. Knull and J. N. Knull) OSU. KEY TO THE SPECIES OF WILCOXIA 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 tErcites eas ee SERS Se ae 2 Scutellum all pollinose; wings hyaline; abdomen with at least some pollen on the dorsum of the tergites i) 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 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 Ww 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—S, gray pollinose; length 5-6 mm (Arizona, New Mexico, and Utah) Femora except apical fourth and tibiae except Se eS MEE A painteri Wilcox VOLUME 71 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 to antennae 1-2. Bristles white, 2—3 presutural, | 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. carinatus Wilcox, the anterior hairs on the central stripe are usually black, but a few M. carinatus have white hairs; in M. gibber there are no anterior bristles 1972 on the central stripe whereas, M. carinafus has about 30 black bristles. Other specimens examined; NEVADA. Nye Co., 24,2 miS Beatty, 24 December 1964 (D. L. Coates) UI, JW. ARIZONA, Maricopa Co., 9, Tempey 21 November 1967 (D. Plantz) ASU. CALIFORNIA, Riverside Co., 9, Blythe, 28 October 1940 (K. S, Snyder) Tamarix, CIS; San Bernardino Co., &, 9°, Needles, 20 March 1967 (R. M. Bohart and D. S. Horning, Jr.) UCD; 646, 39, Twentynine Palms, 23 October, 6 November 1967 (FB. 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 I, 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, | 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. NEW SPECIES OF ROBBER FLIES Female, Length 6 mm, Median line of mesonot golden. Abdominal segments 1-7 grayish polling 8 shining black; apical spines black, short hairs white Holotype: male, California, San Luis Obispo Co Baywood Park, 24 October 1970 (J. Wilcox) CA‘ Allotype: female, same data as holotype, 25 October 1970. In the key to the species (Wilcox, 1964) this specie runs to couplet 10; with its long pulvilli it would be close to M. tarsalus Wilcox from which it differs by having the abdomen all pollinose and is without bare spots on the mesonotum. Other specimens examined: CALIFORNIA. San Luis Obispo Co., 414, 289%, Baywood Park, 23 to 25 October 1970 (J. Wilcox) CAS, 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]. {collected on the ACKNOWLEDGMENTS I 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. (Dipteria-Asilidae), a New Genus. Ent. Soc., 30:1-5. Accepted for publication October 21, 19 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 (Nototeredo) norvagica (Lebour, 1938, 1946; Nair, 1962), Teredo (Psiloteredo) megotara (Sigerfoos, 1908; Nair, 1962), Bankia setacea (Coe, 1941; Quayle, 1953), B. indica (Nair, 1956a, b), dunlopei (Smith, 1963), B. australis 1971), and perhaps B. gouldi (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. Nausitora (Turner, 48 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 «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 %, 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 (Nototeredo) norvagica although it had been earlier postulated by Sigerfoos (1908) in Bankia gouldi. Coe (1933-41) 1972 Taste 1. The effects of temperature and reduced chlorinity on survival and reproduction of Lyrodus pedicellatus. No. of Sur- No. of Sur- viving Test No, With — viving Test No. With Animals Veligers Animals Velipgers Chlorinity ————__——_- —_—_—_—— (%o) 14-16C 22-24 C 19.3 10 4 10 3} 16.0 8 2 10 2 13.0 ) I 8 0 11.0 4 0 4 0 9.0 0 0 1 0 found Teredo navalis, B. 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 methods of Intersci. Barnes, H. 1959. Apparatus and oceanography, Part one: Chemical. Publ. Inc., New York, 341 pp. Becker, G. 1959. Biological investigations on marine RESEARCH NOTES 49 borers in Berlin-Dahlem. /n: D. L. Ra Ex Marine and fouling Washington Press, Seattle, pp. 62-76. horing organism Ur Clapp, W. F. 1951. Observations on living Tere dinidaec. Fourth Progress Report (Rept. no 7550), William F. Clapp Laboratories, In Duxbury, Massachusetts, 9 pp. Coe, W. R. 1933. Sexual phases in Teredo. Biol Bull., 65:293—303. 1934. Sexual rhythm in the pelecypod mollusk Teredo. Science, 80:192. 1936, in Teredo. Sequence of functional sexual phases Biol. Bull., 71:122-133. 1941. Biol. Sexual phases in wood-boring mol- lusks. Bull., 81: 168-176. Eckelbarger, K. J., 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 Occas. Pap. Bishop Mus., 17:97—150. in Hawaii. Grave, B. H. 1928. Natural history of the ship- worm, Teredo navalis, 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) pedicellata 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. Jn: 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. 1938. Notes on the breedi some lamellibranchs from Plymouth larvae. J. Mar. Biol. Assoc. U.K., 23: 1946. The species of Teredo from Ply 50 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES outh waters. J. Mar. Biol. Assoc. U.K., 26: 381-389. Menzies, R. J., J. Mohr, and C. M. Wakeman. 1963. The seasonal settlkement of wood-borers in Los Angeles-Long Beach Harbors. Wasmann J. Biol., 21:97-120. Moore, D. R., and D. J. Reish. 1969. Studies on the Mytilus edulis community in Alamitos Bay, California. TV. 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. of marine fouling and of western Norway. 1962. Ecology wood-boring organisms 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. (In press.) Yonge, C. M. 1926. Protandry in Teredo norvegica. Quart. J. Micros. Sci., 70:391-394. VOLUME71 KEVIN J. ECKELBARGER, Marine Science Institute, Northeastern University, Nahant, Massashusetts 01908, and DoNALD J. RE1sH, Dept. Biology, Cali- fornia State College, Long Beach, California, 90801. Accepted for publication December 1, 1971. RHADINAEA GODMANI, AN ADDITION TO THE SNAKE FAUNA OF HONDURAS Rhadinaea 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 Pinus pseudostrobus 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 | 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 on supralabials; supralabials mostly ivory, supralabials | through 4 with dark brown upper and lower edges, dark blotch on upper portion of supralabial 6 ex- 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 pale diagonal stripe from eye to lip); infralabials ivory with dark brown mottling on anterior ones; chin ivory; iris dark brown. The other specimen (LSUMZ 24398) agrees in essential aspects with the above description. Keels are present on the dorsal scales above the vent in LACM 72070 but not in LSUMZ 24398. Mertens (1952a) described Rhadinaea zilchi on the basis of a single specimen from El! Salvador, distin- guishing this taxon from Rhadinaeca godmani on the basis of differences in dorsal color pattern and the 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 (1970) along with the erroneous conception that pattern differences exist between the two. Inasmuch as the name zilc/i 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 (Ginther) 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 a/tamontana, keels not present in godmani). We have not examined the holotype of a/tamontana but we discount the impor- RESEARCH NOTES tance of supra-anal keeling and point ou description of the color pattern of the hol very similar (but not identical, according to Tay description) to the color pattern of godmani suggest that Rhadinaeca altamontana Taylor is pre ably a synonym of R. godmani and thus support tt implication of Scott (1969) that alfamontana is not recognizable. to occur known (1450-2660 Presently, Rhadinaea godmani is at moderate and intermediate elevations m) of both versants from southwestern Guatemala to has thus far not been recorded from Costa Rica; it Nicaragua. LITERATURE CITED Carr, A. F., Jr. 1950. Outline for a classification of animal habitats in Honduras. Bull. Amer. Mus Nat. Hist., 94:567-594. Holdridge, L. R. 1962. Mapa ecologico de Hon- duras. Organizaci6n de los Estados Americanos, San José, Costa Rica. Mertens, R. 1952a. Neues tiber die Reptilienfauna von El Salvador. Zoologische Anzeiger, Leipzig, 148:87-93. 1952b. Die Amphibien und Reptilien von El Salvador. Abhandlungen 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. the snakes of Costa Rica. Southern California, 390 pp. A zoogeographic analysis of Ph.D. Thesis. Univ. Slevin, J. R. 1939. and amphibians from Guatemala. I. Snakes. Proc. California Acad. Sci., ser. 4, 23:393-414. Notes on a collection of reptiles Stuart, L.C. 1951. The herpetofauna of the Guate- malan Plateau, with special distribution on the southwestern highlands. Con- trib. Lab. Vert. Biol., Univ. Michigan. 45:1-71. reference to its 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 nev species of the genus Rhadinaea from Guatemala Occas. Pap. Mus. Zool., Univ. Michigan. 442 1-11. 52 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 1951. A brief review of the snakes Univ. Kansas Sci. Bull., 34:3-188. Taylor, E. H. of Costa Rica. 1954. Further studies on the serpents of Costa Rica. Univ. Kansas Sci. Bull., 36:673—800. Uzzell, T. M.. IJr., and P. Starrett. 1958. Snakes from El Salvador. Copeia, 1958:339-342. Larry Davip Witson, Horizon School, 9025 Sunset Drive, Miami, Florida 33143, and JOHN R. MEYER, The Ranch, Elgin, Arizona 85611. Research Accepted for publication October 21, 1971. A NEW SPECIES OF BAETODES FROM ARIZONA (EPHEMEROPTERA: BAETIDAE) The first record of Baetodes 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 Baetodes, 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. 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 VOLUME 71 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). The nymph of Baetodes sigillatus, new Figures 1-2. species: Fig. 1, abdominal terga, lateral view; Fig. 2, abdominal sterna, ventral view. RICHARD K. ALLEN and EsTHER S. M. CHaAo, Dept. Biology, California State College, Los Angeles, California 90032. Accepted for publication October 21, 1971. 1972 AN INTRODUCED POPULATION OF SOCIAL WASPS, POLISTES APACHUS, THAT HAS PERSISTED FOR 10 YEARS (HYMENOPTERA: VESPIDAE) The large social wasp Polistes apachus Saussure often builds its nests under the eaves of houses, This species has a typical life cycle for wasps of the genus. Over- wintering queens, usually in groups of two to four, establish nests during 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 < 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- RESEARCH NOTES This site nests of any species had been found in the «¢ fornia, was chosen because no Pol overhangs of any of the buildings on camp because nests on the roof of the science building not be disturbed. The nest boxes were fastened row to the top of a safety wall on the perimeter of the roofs. Each box was about 30 cm from any othe box. The new nest site was 4 miles east and 1.3 mil south of the original location in Chatsworth After the they served through the summer and fall. All ten nests increased in 10 nests were relocated, were of size, produced sexual forms, and ter minated in the fall. (1963), P. apachus queens were observed flying about During the following spring in the vicinity of the science building, and a number of new nests were started under an eave-like 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 State 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 Bell for encouraging the research and for obtain proval from the college for placement of boxes. Davip L. Gino, Erindale College. Toronto, Clarkson, Ontario. Accepted for publication February 8, 1972 54 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES THE PHYLLOSTOMATID BAT, VAMPY RESSA BROCKI, IN COLOMBIA The 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, 11.0, 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 (MI-—M1), 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, Shs SII 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- VOLUME 71 BA XR XR BK RK XK RX x2 10 U AK BA BB AK Figure 1. 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. nymphaca 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. Ropert 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, 1960; 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 CLOSING SPATHE OPEN-——->st-e SPATHE CLOSED+> +) (STAMENS » STYLES J/EMERGE a (eo) “DROSOPHILA a oa SPADIX BASE T. a °o PP LEAF T. At ~T\ hathas TEMPERATURE °C 1) a 20 15 7 12 8 24 6 2 18 24 6 12 18 24 JULY 10 JULY 11 JULY 12 DAY | DAY 2 DAY 3 Figure I. Spadix, leaf, and ambient temperatures of a single flower (17) of Philodendron selloum. 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; spadiv base, within spadix 2” below beginning of female flowers; air temperature, within spathe but 1” away from and 2” above bottom of spadix; and /eaf, 3” inside leaf base and 3” below spadix. Morning temperatures were taken between 0830-0930, afternoon: 1300-1700, night: 2000— 2200 hrs. 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 Drosophila. 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 Drosoph- 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- RESEARCH NOTES + —OPATHE OPEN 45 : 4oTAMEN oO 40 BPADIX BASE T o << rd 35 t 4 {5 30 44 a. rf E | » ——1 AIR ?) | ine =aed 3-4 | 5-12 DAY | DAY 2 Figure 2. Summary of spadix and ambient temper- atures of 28 Philodendron selloum flowering cycles. Vertical lines indicate range and dots indicate means. ment Co., for calling my attention to this problem. Portions of this work were supported by the Na- tional Science Foundation (GB-2307, Postdoctoral Fellowship, 56017). and a Senior LITERATURE CITED Dormer, K. J. in Arum. 1960. The Truth about Pollination New Phytol., 59:298-301. Fischer, H. standen. 1960. Atmung von Bliiten und Bliiten- Encyc. Plant Physiol., 12:521—535. Forward, D. F. Respiration. 1960. Effect of Temperature on 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. 3 } Amer., Thomas, M. 1960. History of Plant Respiration Encyc. Plant Physiol., 12:1—46. BayarD H. BRATTSTROM, Dept. State University, Fullerton, California 92631 Accepted for publication April 5. 19 56 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES RANGE EXTENSIONS OF PORCELAIN AND HERMIT CRABS‘ IN THE GULF OF CALIFORNIA During 1970-1972 the senior author made numerous collections of intertidal 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, México, 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 México: Guaymas, Sonora, México (Haig, Hopkins, and Scanland, Trans. San Diego Soc. Nat. Hist., 16(2):13-32, 1970). VOLUME 71 New record: Cabo San Lucas, Baja California Sur, México; November 28, 1968; Alex Kerstitch. Petrolisthes hirtispinosus Lockington Former northernmost record: Bahia de Tepoca, Sonora, México (Haig, Allan Hancock Pac. Exp. 24: 1-440, 1960). New record: Puerto Penasco, Sonora, México; 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, México; 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 Hai, Curator of Crustacea, Allan Hancock Foundation, Los Angeles, California 90007. Accepted for publication February 29, 1972. INSTRUCTIONS FOR AUTHORS The BULLETIN is published three times cach year (April, August, and November) and includes articles in English in any field of science. Non-members will be assessed a page charge of $40.00 per page. Manuscript submitted for publication should contain results of original research, embrace sound principles of scientif investigation, and present data in a clear and concise manner. The current AIBS Style Manual for Biological ~ Journals is recommended as a guide for contribufors, Consult also recent issues of the BULLETIN. Author should strive for directness and lucidity, achieved by use of the active voice. 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Evaluation of a paper submitted to the BULLETIN begins with a critical reading by the technical editor; several referees also check the paper for scientific con- tent, originality, and clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent to the author at which time he may be requested to rework portions of the paper considering these recommendations. The paper is then re-submitted and may be re-evaluated before final acceptance after which it is sent to the managing editor. Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He should promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations legends, and bibliographical references. He marks corrections on the galley (copy editing and proof proce- dures in Style Manual) and promptly returns both galley and manuscript to the managing editor, James Dale Smith, Department of Biology, California State University, Fullerton, California 92631. Manuscripts and origi- nal illustrations will not be returned unless requested at this time. Printing charges accruing from excessive addi- tions to, or changes in, the proofs must be assumed by the author. Reprint orders are placed with the printer. not the Managing Editor. CONTENTS Ree | ' 1? Rditorial 11418). od BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES SUBFAMILY MACROCEPHALINAE Genus Macrocephalus SWEDERUS, 1787 Macrocephalus similis, new species Figure | Description: Female. Elongate ovate; head, 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; Il almost globose, III evenly enlarged toward tip, TV 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. Scutellum 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, middle and hind with smaller granules tibiae densely laterally; granulate. Color VOLUME 71 4 _— ~ 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 and inflated portion of disc, and connexiva I, ochraceous or slightly brownish; anterolateral portions of scutellum (roughly punctured) brown; ventral side of body 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 of pronotum 3.00 mm; width of abdomen 4.00 mm. Male. Smaller and 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 III, 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 I 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 IIT); 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, NEW AMBUSH BUG FROM ARIZONA and probably was confused with the latter Vay be separated from it by without pronotum and connexivum generally finer larger size, pronotal knob or ridge anteriorly, granulatior and not relatively longer, and abdomen slightly longer than its ma width, Holotype: Female. Arizona, Cochise Co., 5 mi W Portal, 25 June 1959 (L. A. Stange) LACM. Allotyp: Male. Arizona, Santa Cruz Co., Pena Blanca Lake 22 May 1970 (T. Halstead) LACM. Paratypes: 5 2 dense, head relatively shorter, pronotum imum and 7 2, some were collected with the allotype, and others in the same locality on 15 November 1969, T. Natural History Angeles and in the collections of Department of Entomology University of Arizona, Tucson, and of the author. Halstead coll. Deposited in the Museum of Los County 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 L. Hogue, Senior Curator of Entomology. I am indebted to T. Halstead, Phoenix, Arizona, for the gift of Phymatidae collected by him in Arizona. also Accepted for publication November 16, 1971. THE ASCIDIANS STYELA BARNHARTI, S. PLICATA, S. CLAVA, AND S. MONTEREYENSIS IN CALIFORNIAN WATERS DONALD P. ABBoTT! AND JEFFREY V. JOHNSON” ABSTRACT: Reexamination of the holotype of Styela barnharti Ritter and Forsyth, 1917, 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, 1968). 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 S. barnharti by Ritter and Forsyth (1917) gave ‘a very poor idea of the size and usual character of the species.” 2 1 Hopkins Marine Station of Stanford Universit Pacific Grove, California 93950 2Dept. Zoology. University of Nebras! Nebraska 68508. 96 BULLETIN SOUTHERN ( barnharti Holotype of Styela (aky plicata): positions of apertures marked in right hand Figure 1. view. vided a new description. Subsequent investigators 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 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). have resembles THE HOLOTYPE OF STYELA BARNHARTI RITTER AND FORSYTH, 1917, AND COMPARISON WITH S. PLICATA (LESUEUR, 1823) The holotype of S. 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, pl. 38. 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 {LIFORNIA ACADEMY OF SCIENCES VOLUME 71 7 LY gn 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, pl. 42), though no testes exhibit such attenuated branches as those 1972 A, Styela clava, San Diego, California. B, Styela montereyensis, Monterey, California. Figure 3. shown for an old and large individual of §. plicata 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, pl. 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 S. 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 CALIFORNIAN STYELID ASCIDIANS within the known range of species; Tokioka (1953:268) rey | right and 1—3 on the left in Japan: and Tucker (1942) recorded 2—1] and 1-3 on the left in individuals fr Carolina, The first correct published record of currence of §S. plicata, under that name, on west coast of North America is that of Reis} (1963), based on material collected in the sprir of 1960 at Alamitos Bay, Long Beach, Californi: An earlier Richmond, Sar Francisco Bay (Abbott, 1954) is erroneous: ré they are § clava. New records of S. plicata in southern Cali and determined by one of us (Abbott) report of S. plicata from Point examination of the specimens shows fornia are given by Fay and Johnson (1971). It i now clear that S. plicata was present at San Diego 1915, Ritter Forsyth’s material was collected. Its present known at least as early as July when and distribution on this coast extends only from San Diego to Santa Barbara, California, at depths of 0-3 m. Atlantic, Pacific, and Indian Oceans, and has been Elsewhere it is widely distributed in the the subject of numerous field and laboratory in- vestigations. Specimens examined: Point Loma, San Diego, California, 24 November 1971, coll. G. Lingle: Mission Bay, San Diego, California, 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 R. F. Hutton; Beaufort, North Carolina, 13 December 1915, coll. unknown; and Wakanoura, Kii, Japan, 1955, coll. 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 _ especis large series from Venice, California was examine in connection with studies of growth (J. V. Johnson, 1971). taken intertidally or in the shallow subt on floats, pilings, or rocks: only the specin the Hyperion sewer outfall comes depths (50 ft). Van Name’s description of S os BULLETIN SOUTHERN ( Styelid {LIFORNIA ACADEMY OF SCIENCES VOLUME 71 Figure 4. ascidians found growing together, Santa Barbara harbor, California. 1-6, Styela montereyensis (specimen 4 bearing an S. plicata). 7-11, Styela clava. where it deals with the placement of the stomach Dorsal tubercle—The aperture often shows and intestine, is generally accurate, though it does not always convey the range of variation to be The following account augments or modifies only those portions of the description expected. which require this. External features —Our 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; largest 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 and not al- ternating areas of thick and thin tunic. The oral relatively thin and leathery tunic, 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. 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 S. clava. ANA yes Outlines of testes from holotype of Styela barnharti (= S. plicata), showing nature of branch- ing. Figure 5. 1972 CALIFORNIAN STYELID ASCIDIAN 120 w 250 P 100 r uu ai U A= < 80 2° 200 Ww 4 ¢ ps = z Z < iti | 1150) Ps O O Z. 6 40 e = 100 2 4 6 LENGTH (cm) Vp) (a) a 2 5 0 u : : ©) Z O 7) O Za 2 4 6 LENGTH (cm) LENGTH (cm) A HOLOTYPE OF S. barnharti Gd BOCA CIEGA BAY, FLORIDA @ SAN DIEGO, CALIF. ZL, CEDAR KEY, LEVY CO., FLORIDA mM NEWPORT BAY, CALIF. O BEAUFORT, NORTH CAROLINA q LONG BEACH, CALIF. O WAKANOURA /KIl , JAPAN Figure 6. Styela plicata: relation of total length (body and stalk) to numbers of ¢ 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 to 76 (in a 90 mm individual). Total number of internal longitudin Pharynx.—The pharyngeal folds are low but — sides) ranged from 137 in a: 100 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 350 300 & im 250 ep) WY Ww > < Z 200 Q =) = O Zz OM iso < STYELA clava Z er ~ Ww = 100 m VENICE, CALIF. E, O SANTA BARBARA , CALIF. VL STYELA montereyensis 50 A MONTEREY, CALIF. @ SANTA BARBARA , CALIF. 5 10 15 20 25 LENGTH (cm) Figure 7. Styela clava and S. montereyensis; relation between total length (body and stalk) and number of internal longitudinal vessels on pharynx. 1972 TABLE 1. CALIFORNIAN STYELID Feature S. clava Tunic Usually with conspicuous tubercles, at least anteriorly (Figs. 3A, 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). Siphons Both siphons straight, directed anteriorly (Fig. 4). Pharynx Esophageal aperture located halfway down pharynx; branchial folds with numerous internal longitudinal vessels (Fig. 7). Gut Stomach in descending position (Fig. 8A); Atrial tentacles Ovaries Testes West coast dis- tribution and habitat 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—5S 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. Distinguishing features of Styela clava and Styela montereyens ASCIDIANS 10 S, montereyensi Tubercles lacking, or few and inconspicuou larger animals with a few loose warts of tunic material, easily detached, adhering near the siphons (Figs, 3B, 4) Longitudinal ridges and grooves on body and stalk extending anteriorly nearly to siphon and representing respectively thicker and thinner regions of the test (Figs. 3B, 4) Oral siphon curved, directed ventrally or posteriorly (Figs. 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 (rarely 1 or 3), seldom branched posteriorly (Fig. 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 25 mm -Left 4(16)3(12)3(14)3(11)3 -Right 6(35)4(31)3(31)5(21)4 DL 100 mm -Left 6(30)4(29)7(26)7(17)4 -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. 8 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. of the specimens examined by us was the placement of esophagus and stomach and the course of the intestinal loop at all similar to situation portrayed by Van Name (1945:310, fig. Gonads (Fig. 8 A).—Wan Name (1945) found We In none the 205). 5-9 ovaries on the right and 3—5 on the left. find 4—9 (average 6) on the right and 2—5 (aver- age 4) on the left in a series of 26 individuals 25-160 mm in total length. Most specimens I = z ncterinrl at least one ovary which is branched posite 102 BULLETIN SOUTHERN CALIFORNIA Figure 8. ACADEMY OF SCIENCES VOLUME 71 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 1972 (Kobe, Honshu, Japan; USNM 11703); 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 Californian individuals, but this is a character in which both American and Asiatic populations show enormous and widely overlapping variation. A slight 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 harnharti agrees with the Japanese specimens and with recent de- scriptions of Styela clava (Millar 1960; 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. Feinberg placed at our disposal the earliest Californian collection of S$. clava in the American Museum of Natural History, New York (AMNH 931, taken in Newport Bay on 15 No- vember 1933 by G. E. MacGinitie, and noted on the label to “occur abundantly on piles, etc.’’). The specimens, mostly dissected already, had been identified by Van Name as 8S. barnharti. All are typical §. clava as regards the gut and other features. A search through the other collec- tions of S. 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 §. 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 S. clava is regularly abun- dant in sheltered areas from San Diego to Santa CALIFORNIAN STYELID ASCIDIANS Barbara (Fay and Johnson, 197) f Point Conception its occurrence is more sp It is recorded once from Morro Bay, and | most years from Monterey Bay. It can be found in most years in San Francisco Bay, though pop ulations in some localities disappear after period of heavy rains (W. E. Charles Spowart pers. comm.). The species has never been noted Berg, in either Tomales Bay or Bodega Bay to the north of San (WanB: and C. H Hand, pers. comm.), despite the seeming Francisco Gladfelter Suita bility of the habitat and extensive collecting by field biologists. Specimens examined from: Point Loma, San Diego, California, 24 November 1971, coll. G. Lingle; 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. E. Mac- Ginitie; Hyperion outfall, Verdes 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. R. C. 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. sewer Palos AND S. IN- COMPARISON OF S. CLAVA MONTEREYENSIS, AND THE TRODUCTION OF S. 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 reasonable guess as to the date S. clava was inti Californian waters. Ritter collected ascid tensively over a period of two decades 104 1917, especially in southern California, and it seems unlikely he could have missed this species had it been common in his time. It is also un- likely that Ritter mistook it for S. montereyensis. He mentions taking the latter “from many points on the coast from San Diego to Mendocino” (Ritter 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 1917. No species resembling S. clava is reported by Johnson and Snook (1927), though these authors, too, collected extensively in areas where S. clava is now abundant, 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 S. 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. and 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. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 LITERATURE CITED Abbott, D. P. 1954. Key to the littoral ascidians of the central California coast, p. 301-310. Jn Light, S. F., R. I. Smith, F. A. Pitelka, D. P. 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. ural history of marine animals. Graw-Hill, New York, 523 pp. 1968. Nat- 2nd ed. Mc- 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. connues des cétes de la Manche. Biologie Marine, 11:145—152. Sur quatre Ascidies rares ou mal Cahiers de Reish, D. L. 1963. ganisms attributed to the “red tide” in southern Mass mortality of marine or- 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 Ricketts, B. F., and J. Calvin. 1939, Between Pacific tides. Stanford Univ. Press, 320. pp. 1968. Between Pacific tides. 4th ed. (J. W. Hedgpeth, rev.), Stanford Univ. Press, 614 pp. Ritter, W. E. 1909. Halocynthia johnsoni n. sp., a comprehensive inquiry as to the extent of law and order that prevails in a single animal species. Univ. California Publ. Zool., 6:65—-114. Ascidians Univ. Ritter, W. B., and R. A. Forsyth. 1917. of the littoral zone of southern California. California Publ. Zool., 16:439-512. Tokioka, T. 1953. Ascidians of Sagami Bay col- lected by His Majesty the Emperor of Japan. Iwanami Shoten, Tokyo, 315 pp., 79 pl. 1955a. Record of Styela clava Herdman from the European waters. Zool. Mag. Tokyo, 64:200. CALIFORNIAN STYELID ASCIDIANS 1955b, Contributions to Jay fauna XI. Sporadic memoranda (2) Mar. Biol. Lab., 4:205-218 ——. 1959. Contributions to Japanese ascidias fauna XV. Sporadic memorandum (6). Styela clava var. symmetrica nov., a new pedunculats styelid from the Inland Sea. Publ. Seto Mars Biol. Lab., 7:457—463. 1967. National 1-247. Pacific Tunicata of the United State Museum. Bull. U.S. Nat. Mus., 25] Tucker, G. H. 1942. The histology of the gonad and development of the egg envelopes of an ascidian (Styela_plicata Morph. 70:81-113. Lesueur). J. Van Name, W. G. American ascidians. 84:1-476, 31 pl. 1945. The North and South Bull. Amer. Mus. Nat. Hist., Accepted for publication June 10, 1972. RESEARCH NOTES A NEW SPECIES OF PUGNUS FROM COCOS-KEELING 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(4):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 Marginellidae, 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. Pugnus maesae, new species Figure | Pupnus sp. Maes, 1967, p. 141, pl. 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 106 Sciences, Philadelphia, No. 288324. Eight paratypes, ANSP No. 32474, from same locality as holotype. 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; Ostheimer, Orr, Ross, collectors, 11 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 Figure 1. Pugnus maesae, new species. Ventral view of holotype. Copy, Maes, 1967, pl. 14, fig. D. 1972 for the species by Hedley, These specimens also had perforate apices. Familial placement of the genus Pugnus is) un- certain. It was described by Hedley (1896) as a member of Ringiculidae, and Zilch (Handbuch der Paliiozoologic, 6:1—-835, 1959) placed it in Cephalas- pidea. Maes (1967) transferred it to Marginellidae, remarking on its resemblance to the [?marginellid] genus Marginellopsis Bavay, 1911. The character of the columellar folds and the lip denticulation appear typically marginellid. The wavy, incised spiral sculp- ture recalls that of some cephalaspidean genera, such as Acfeon Montfort, 1810. Knowledge of its true relationships awaits an anatomical study. Barry Roru, /2/7 Waller San California 94117. Street, Francisco, Accepted for publication May 24, 1972. A NOTOPHYCID POLYCHAETE FROM CALIFORNIA 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 errantiate 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, Notoplhycus 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. RESEARCH NOTES 10) Phyllodocella bodegae, | Figures | and 2 Material examined: Mason's Marina, Bodega Has bor, California, July 20, 1971, from a gelatinou sac attached to a colony of Obelia sp.; 15 cm deptt on a floating boat dock; one specimen, Holotyys deposited in the collections of the Allan Hancock Foundation. Description: The holotype is a complete, sexually mature female with 24 segments that is & mm long and 2 mm wide without setae. It is white with pigment alcohol preservation. reddish spots over the anterior end in The pygidium is a small, rounded cushion with out anal cirri; the anus is dorsal. The prostomium (Fig. 1) two pairs of long, slender antennae near the anterior margin. A pair of small, distinct frontal lobes are present on the anteroventral margin. pairs of eyes are present at the middle and mid-posterior part is pentagonal and has Two of the prostomium; the anterior pair is lensed; the posterior pair is semi-lunular in shape. The peristomial segment is a complete ring, forming 1. Phyllodocella bodegae. new species view, right dorsal ten Figure anterior cirrus broken, S70! dorsal M7505 B: end, anterior en 108 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES | A Figure 2. Phyllodocella bodegae, new species. A. seta from parapodium 7, x 950. B. parapodium 7, anterior view, x 52. 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. 1B) project strongly ventrally with the setae pointing anteriorly and ventrally; the second neuropodia are slightly more lateral in position. Noto- and neuropodia project dorsolaterally and ventrolaterally 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- VOLUME 71 phycus minuta Knox and Cameron (1970) 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 P. 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 Notophycidae are quite small and are 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. A KEY TO THE FISHES OF THE FAMILY GOBIDAE (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). 1972 RESEARCH NOTES FAMILY GOBIILDAE row of teeth tricuspi } dilated, (Fig. 2b) Dorsal fins separate, the pelvic fins goby—Tridentiver — trivonoce) G united to form a flaring cone-shaped sucking disc, free from the body (Fig. 1). Figure 4. 2. a. Eyes normal; no specialized folds and flay in front and below the eyes 4 b. Eyes very small, vestigial or nearly so in the adults; cheeks much enlarged, tumid; skin of head with tactile organs well developed (Fig. 4) 3 3. a. Dorsal spines VI Halfblind goby—Lethops connectens Hubbs b. Dorsal spines U1 Blind goby —Typhlogobius californiensis Steindachner. 4. a. Dorsal spines VII or less Broure tl b. Dorsal spines VIII ae Yellowfin goby—Acantho eobius flavimanus (Temminck and Schlegel). 1. a. Without five dark bands on leading edge of both the first and second dorsal fins; teeth in jaws simple, neither clubbed nor broad and rounded (Fig. 5a) incised at their extremities (Fig. 2a) 2, tn a. Caudal fin normal, not greatly elongate: 6 Figure 2. b. Five dark bands on leading edge of both the first and second dorsal fins (Fig. 3); outer b. Caudal fin greatly Sb) Figure 3. lus longicaudus (Jen} 110 8. a. a. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES x \ Figure Inner margin of shoulder girdle without dermal flaps ‘ et iT Inner margin of shoulder girdle with 1 to 3 dermal flaps (Fig. 6) ee lil Maxillary not extending beyond the posterior margin of orbit (Fig. 7a) ————____- 9 Maxillary extending beyond the posterior margin of orbit (Fig. 7b) —. Figure 7. Dorsal V to VI-I, 8 to 13; anal fin elements 9 to 14 oe =a __.. Long- jaw mudsucker—Gillichthys mirabilis Cooper. Dorsal IV to V-O to I, 14 to 17; anal fin Anal fin elements 13 or less ~ es AMY) Anal fin elements 14 or more Blue- banded goby—Lythrypnus dalli (Gilbert). Anal I, 11(10 to 12): head and sides with- out crossbars encircling posterior part of VOLUME 71 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). body Il. a. Anal fin elements 13 or more 12 b. Anal fin elements 12 or less Tidewater goby—Eucyclogobius newberryi (Giratd). Maxillary not prolonged beyond the posterior margin of orbit peer 5 2 ls) 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— /lypnus gilberti (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 affinis. Habitat: rocky intertidal. Range: Cerros Island, Baja California to Point Vincente, California. 1972 Acanthogobius flavimanus.—Color variable, Four- teen soft rays in dorsal fin with eight dusky spots, each somewhat larger than the eye-diameter, ar- ranged in a nearly evenly-spaced series down each side, the first three being concealed beneath | the pectoral fin; the last forming a more prominent spot at the base of the caudal fin (Brittan, Albrecht, and Hopkirk, 1963). Habitat: lagoons and estuaries ascending rivers to fresh water. Range: Elkhorn Slough to Tomales Bay, California (L. Dempster, pers. comm.). 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 Californian gobies at least on the basis of the cephalic-lateralis system. Habitat: la- goons and estuaries. Range: San Diego Bay, Cali- fornia to Guayaquil, Equador. 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 U pogebia 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 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 Morro 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 in crevices RESEARCH NOTES (CSCLE range extension for the University, Long Beach a northern Pale margin on the first dorsal fin A fleshy crest on top of | Coryphopterus nicholsi black Large black eyes. yellow or olive } a distinctive tends posteriorly from behind eyes to origi f first dorsal fin, Habitat: 5 and 50 m, resting on sand bottoms close to rock greater commonly found betweer and interfaces; (it has been than 700° m. further than a few Range: San Martin and Turner, 1962) British Columbia. captured at depth This gobi seldom venturing meters from protective Baja California to the Queen Charlotte cover (Ebert Islands, Island, Lythrypnus zebra.—Cherry red with 15 blue bars oriented dorsoventrally. Bars alternating with a thin blue line along the length of the body. Habitat: caves and under rocks or well within deep crevices: commonly found to depths of 30 m. Range: Island, Baja California to fornia. Cedros Ventura County, Cali- Eucyclogobius newberryi.—Maxillary 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. reaching to Quietula y-cauda.—The length of the maxillary increases with age and may not reach to posterior margin of orbit in specimens 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 “vy”. Enters burrows of invertebrates during low tide or when threatened. Habitat: channels of tidal less mudflats. Range: Lower California to Morro Bay. California; northern Gulf of California, Baja Cali- fornia. Ilypnus_ gilberti—Maxillary extending to below middle of the eye. Conspicuous metallic blue-black spot on opercle. Habitat: shallow bays and channels of tidal mudflats. Range: northern Gulf of Cali- fornia: Bahia Magdalena, Baja California to Tomales Bay, California (L. Dempster. pers. comm.). Lepidogobius lepidus.——Distinctive black on vents surface of head and tips of all fins. Pal l green, with rust colored blotches on si Muddy bottoms to 100 m depth. Rang California to Vancouver Island, 1 112 ACKNOWLEDGMENTS I wish to thank Camm Swift 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 Lilian Dempster, California Academy of Sciences, San Francisco distributional information and specimens of Tridentiger trigonocephalus. for LITERATURE CITED Barlow, G. W. 1961. Gobies of the Genus gillichthys, 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. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 Clothier, C. R. 1950. A key to some Southern California fishes based on vertebral characters. California Dept. Fish and Game, Fish Bull., (79) :1-83. 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. 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. Macponatp, Dept. Biology, California State University, Long Beach, California 90801. Accepted for publication February 16, 1972. | INSTRUCTIONS FOR AUTHORS _ The BULLETIN is published three times each year (April, August, and November) and includes articles ty | English in any field of science. Non-members will be assessed a page charge of $40.00 per page. Manuscript submitted for publication should contain results of original research, embrace sound principles of «cient! investigation, and present data in a clear and concise manner. The current AIBS Style Manual for Biolo Journals is recommended as a guide for contributors. Consult also recent issues of the BULLETIN. Author should strive for directness and lucidity, achieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous reference of pronouns, and logically placed modifiers. 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Special attention should be given to the description of new taxa, designation of holo- type, etc. The literature cited section should include six or more references; entries for books and articles should take these forms. een McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii 4- 326 pp. Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54: 452-458. Brattstrom, B. H. 1969. The condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), Univ. California Press, xii -+ 635 pp. | If fewer than six references are cited they should be inserted in text as follows: (McWilliams, Insect mimicry, p. 216, 1970); (Holmes and Speak, J. Mamm., 54: 452-458, 1971): (Bratt- strom, The Condor in California, Pp. 369-382, in Vertebrates of California, 1969). Tables and figures (line drawings, graphs, or black and white photographs) should not repeat data con- tained in the text. 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Evaluation of a paper submitted to the BULLETIN begins with a critical reading by the technical editor; several referees also check the paper for scientific con- tent, originality, and clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent to the author at which time he may be requested to rework portions of the paper considering these recommendations. The paper is then re-submitted and may be re-evaluated before final acceptance after which it is sent to the managing editor. F Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He _ should promptly and carefully read the proof sheets for errors and omissions in text. tables. illustrations legends, and bibliographical references. He marks corrections on the galley (copy editing and proof pro. _ dures in Style Manual) and promptly returns both galley and manuscript to the managing editor, Jame _ Smith, Department of Biology, California State University, Fullerton, California 92634. Manuscripts an nal illustrations will not be returned unless requested at this time. Printing charges accruing from excessiv: _ tions to, or changes in, the proofs must be assumed by the author. Reprint orders are placed with the not the Managing Editor. es a 5 CONTENTS John Adams Comstock, 1883-1970. By Lloyd M. Martin 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 A review of Eucyllus Horn (Coleoptera: Curculionidae, Brachyrhininae, Peritelini). By Frank W. Pelsue and Elbert L. Sleeper Two new species of North American flat bugs. By Nicholas A. Kormileyv A new spécies of Ambush bug from Arizona. By Nicholas A. Kormilev The ascidians Styela barnharti, S. plicata, S. clava, and S. montereyensis in Californian waters. By Donald P. Abbott and Jeffrey V. Johnson RESEARCH NOTES A new species of Pugnus from Cocos-Keeling Barry Roth A notophycid polychaete from California. By Kristian Fauchald and Bruce W. Belman A key to the fishes of the family Gobiidae (Teleostomi) of California. By Craig K. Macdonald .. cover: Larva of Adelocephala heiligbrodti hubbardi. Reproduced from a painting by Dr. John Adams Comstock. POL HERN CALIFORNIA ACADEMY OF SCIENCES BULLETIN Number 3 BCAS-A71(3) 113-164 (1972) November 1972 Southern California Academy of Sciences Founded 6 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 H. Wells, Technical Editor James Dale Smith, Managing Editor Roberta S. Greenwood, Index Editor BOARD OF DIRECTORS 1970-1972 1971-1973 1972-1974 Donald Bright John J. Baird Wayland D. Hand Jules Crane, Jr. Takashi Hoshizaki Donald R. Patten John E. Fitch Elbert Sleeper Donald J. Reish William J. Morris David L. Walkington Andrew Starrett Charles R. Weston Stuart L. Warter 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. AninualipMembersvscr- ccs: vox) o ote eave ne is er Tee POT ota See $ 8.00 StudentiMem bersissinrs cis tierepeecleesorssest okt ecco apoe el ake veoh vars Pslosic else oe ee ene aCe 5.00 Beife Mem bers! ai nero cree ges ie Secale ro Say aus cae ro Mtanenesanes mnehene in rel iene oie Rune ee eI ee 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. OF THE ACADEMY BULLETIN VOLUME 71 ORNITHOPHILY AND EXTRAFLORAL SOW T EE ARIN = OF DECEMBER 30, ALIFORNI SCIENCES 1972 NUMBER COLOR PATTERNS IN COLUMNEA FLORIDA MORTON (GESNERIACEAE ) C. EUGENE JONES, JR.! AND PAT VICKERS RICH® ABSTRACT: Hummingbirds, such as the Litthke Hermit (Phiaethornis longuemareus) and the Green-Crowned Brilliant (//eliodoxa jacula) apparently are attracted to the hidden flowers of Columnea florida 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 Columnea (Gesneriaceae) possess a group of characteristics which tends to delineate ornithophilous plants (Morley, 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 Columnea 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 Columnea. 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 Columnea, 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 tlowers 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 Columnea_ florida pattern, Morton, an understory, epi- phyte, is conspicuous in the flora of the Premon- tane Wet Forest (Holdridge, 1967) of southeast- ern Costa Rica. The leaves of this species are strongly anisophyllous; the larger leaf may be 40 cm long and 11 cm wide, whereas the smaller is less than 1 cm long. The larger leaf completely conceals the relatively small (about 142-2 cm long), yellowish-orange axillary 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) 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 | February 1972 at Finca Las Cruces, near San Vito. P nas, Costa Rica. Visitations by pollinators were 1 Dept. Biology, California State University. 1 ton, California 92634. > Dept. of Vertebrate Paleontology. Amer seum of Natural History, New Yo Ne 10024. 113 114 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES © | Figure 1. VOLUME 71 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 (Heliodoxa 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 location of the food source. Consequently, these birds spent several seconds inspecting various por- tions of the plant before finding the flowers. It 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. 1b). 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. Ic). 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 away ORNITHOPHILY AND EXTRAFLORAL COLOR PATTERNS j] from the superior ovary and into contact with the nectar which collects in a slight depression in the Since the flower are horizontal to the ground, nectar collects in the lower portion of the corolla. depression in the corolla by gravity. An examina tion of 25 flowers from 10 separate plants, re vealed no damage to the ovaries from the probing bills of the Litthe Hermit hummingbirds. Behavioral responses of the Litthe Hermits were similar in 1972 to those noted in 1970, except six birds flew directly to the flowers of C. 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 florida, thus insuring successful pollination. AI- flowers of C. though the attraction of hummingbirds to flowers by extrafloral mechanisms is well documented in the case of Castilleja and others, (Faegri and van der Pijl, 1966; Grant and Grant. 1968). the mech- le he leaf does anism for C. florida is unique in that t not simulate a portion of the flower, but ner simply acts as a visual signal denoting the pres- ence of nectar in the immediate Vicinity 116 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 B. H. Bratt- strom, T. L. Hanes, and C. B. Heiser 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. and Their Flowers. York, 115 pp. 1968. Hummingbirds Columbia Univ. Press, New Grant, V. 1950. The Protection of the ovules of flowering plants. Evolution, 4:179-201. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 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. 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! AND DONALD J. REISH? 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%. 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%. 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 and about 11 C. Reproduction occurred above 28.8%. in the temperature range of 14 to 24 C. 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 more prevalent (Menzies, Mohr, and Wakeman, 1963). Little data are available on the tolerances of L. pedicellatus from the Pacific Coast. Barrows (1917) found this species able to temporarily withstand salinities as low as 10% in San Fran- cisco Bay. Menzies ef al. (1963) 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%. 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 17%. Watson (1957) reported L. pedicellatus in Queens- land, Australia only in salinities above 25%. 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 EFFECTS OF TEMPERATURE AND SALINITY ON L. PEDICELLATU: |] pipetted from the container and the wood bloc} were returned to the cold bath. The largest num ber of larvae were obtained by using the wood blocks the day they were removed from the field but they could be used as a further source of larvae in the future with decreasing success in obtaining large numbers. Larval Salinity Experiments: An experiment was conducted to determine the effects of tem- perature and reduced salinity on the swimming, survival of pediveliger Eleven concentrations of sea water were used: 35:0) 31-5, 28:8, 25:2; 21.6; 18:0) 14:4) 10:85 722, 3.6, and 1.8% salinity. boring, and the stage. Solutions were prepared by dilution with appropriate volumes of distilled water and the salinity determined (Barnes, 1959). A single 350 ml finger bowl containing a 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 hours and larval swimming, crawling, and boring activity including the formation of the calcareous cap was noted. 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, SSLONSIES 28:85 25-25 21-05 18:05 4410.85) cee, 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- pre- every 6 The experiment was conducted 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 9-11 C temperature bath was unavailable for this e periment. Sea water in each flask was replacec by a fresh solution every five days. The ment was terminated at five months at \ VOLUME 71 RN CALIFORNIA ACADEMY OF SCIENCES BULLETIN SOUTHE 118 007 TS6I ‘AQTINH ynoqe yeod pue ydjey pur[eaz MON SUIpse1q = = = = = syo.isny piyuvg "CIE JUSU9T}IJ9S (vJvuuadiq =) TL6I ‘Jeinouey ySBOD AlOAT — = = —= —= umMuwIxeur sisuappispg piyuvg (Apmis qey) PUI[OIeD 6961 ‘AeutIND YON 0 0E-S'LT am =z aa ar =v 0 07-91 PS6l “HINIL Avg ye sulsaq °AOE—6 U! pue ewa}[ayos ayradesayD Ajqeqo.id — uOUIUIOS = = = Ip[nos piyuvg (Apnys qey) °*vT "hb OP-91 9961 RIquInjOD AOTaq JUST UI JadI[aA “0 Ja Ka|SUMO ysnuig = = — -dojaaap ou — 0} pastel Od “moqiey O OT MOTAaq 6S61 ‘AIAENO yuusApe Ty pue °% 87-47 = —- = = = °DET—6 He Sr6l “Aas|a RIquinjo) jUaWaT}as pue yor[g ysnug = = = = = ysoysiy Ceol “JATIN pure uosuyor punos josng OTI-L = — == sa a °*9L MOlEq Aeg oosio “IY $7 ul AVATIOR 9761 “APTN BE HALES a Teyia] ’7¢°9 peonpol = a == RIquIn[OD Ja}UIM UL 46761 “UUM ysnig "2/8 ST eaoge Sat — iam a = (Apmis qe]) “IY ZITT eiquinjoy) UP yeqse] P6cé6l ANU Ysniig sy "AL EI-S'L a aa — = Avg oosio 1761 “PIOJOM SUB I UES = = °AOT PAOGR = = = Daovjas vIyUDg 203n0S AyyRIOT surumedsg TBATAINS AVATIOWV TPBATAINS sullog AVWATIOW satoadg 1oj unwndo oy Joy wnwndo snoiediag WnUTUTyAy Wn WUT yy wn wiUTyy te SPrulpelaL INpVy SpluIpolay [PAIVT *spluIpalay, UO s}Oayja AjUI[RS pue simjeIdduia} UO sINJVIO] JO MIIADY ‘[ ATAV /t MOT Aojaq Sulsoqg ou epeury 4OL-1T LT] “uouy mnuEpy = D gp wnoge pur 5 g] a — —_ _ (vg OasIo SAUP p Ul > OT6T “JOqTNIN “uv ues DAOQE 10 2256 [BYII] UP —_— — — _ x ~OP juswdojaaap = (Apnis qe]) VQ] MOlAG aaoge JO CL Joy jeundo 5 OTe] “AHBwOH FY —- BNOIS BAON — = suliog Ou MOTAG 'O OL — O 8°t7 a (Apnis qey) = {eg oosi9 uni 3u0] ul Sj TZ6l “wNig -uevi4 US = Teyiay 77,9 aA0ge JO 77,6 oo 7 aa = Aeg oosio 2° IT6I “PlOJON -uelj URS _ 2A~OI-S°L _ °%¢61-Ol —_— —_— SIDADU Opada ~ > 2amog Ayyeso 7 SuruMeds TeATAINS AWANOY TRATAINS sullog AWAY sa19adg > 10} wnwndoO oj Oj wnwndo snoiediAiey s wnuwiruryAy WInWTUryAy WIN UITUT Ay wa y-110Y4g = sprurpasaL NPV SprurpasaL [eae y ” 3 e961 “WS Be NsNy = [QC6] “UOSIB AA ‘puejsusend) 2.Q] MOTAq = 2A%Q] MOTAG — — — 1ado]UNp DAOJISNDN 3 "41 E-6T Ye (n918pA10U < (Apnis qeq) peonpal opasajojoN =) S OF6l “YOON oneUpy O 0£-0¢ °%8T Sul10q =a a os SNjNIIAIN OpadsaL »~ z x (Apmis qe]) = puejsuq DIsYVALOU (Opasat = TT6T “UOW;ULIeH “yynowlsyd ed aa = "*OL are — -OJON) Opada = mS (Apnjs ge]) (9jpU1ADI =) SS OF6T “YSO" oneHpy O ¥T—-0E a a aay re — DUIUIU DIYUDT 2 196] ‘weue —-eIpUy “IOQIeH ~yE-1T Ul Q -ysnygesen, woujedoyyestA = — uowWWO0S = — = piojjaupdup) niyung Q & 30mog AyyeI0T suramedg yeATAINS AVATIOW JeATAING sull0g AVATIOW saisadg mE Io} wnumdo 1oj Bio} s wnundQ snorediAag = uN UTUT AL TaN TOYA wanUTUryAy sprurpelaL NPV Sprurperal [BAe T = (panuyuoDg) “T alavh XK oN 1 VOLUME 71 MY OF SCIENCES ACADE SOUTHERN CALIFORNIA TIN BULLE 120 £961 “PASPTOXYIN pure aoyrysqraAy £961 ‘SIAR pur jyoursooT Bag Yor (Apnys qey) “UUOD “PIF 0 0c-9T Paseayar avAIey] faaoge 10D pT TPYI9] O Of TOG “YRUIAISeg (Apnis qr]) aaoqe 10 -PAOUIPNY, UOIUL) JAIAOS —_ — — O OT MoTaqg OS6I uedey ‘Avg aaoqge “]p ja Tew] BMESPUC pue od gl _— — = (Apnys qey) O Se aA0ge Tr6l “Aojeing Bag YORI = = = SO. ZT Mojaq (Apnis qe]) OF6l Yoo" oneupVy O rc-IT Fas a ici MOOG OF6I1 ‘121P9AN QDUdIMET pure 9 0] 0} pue 12/paaN IS FIND = as UMOP Y]MOIS — DZI 40 876] ‘AARID 2]OH S,poom IT aa0qge — —— — (Apnjs qry) L761 “JPTIN Avg oosio uni Suo] pur plojoy -UPI URS = Ul ?%¢ BAOR 10 °%,6 ae 301n0S AWyeo0'7T surumedsg TRAIAINS AVWANOW TRATAINS Joy umund9g 10j WIN UWITUTTAY wnuUuryAy SpluIpsjay ynpy (panuyuod) ‘| aATavL 2 97 aaoqe JOD +1 MOT2g 2701 sullog Joy wn WwIUuryAy OD LI-+7Z sins90 yudUTeT}}08 wunwixeur quswido]aaap IO} °%~T] VAOGR *O LT-81 OTI-OL ye poyyey SUTUTWIMS *O 0-07 a SIDADU Opa.Ja T, AVATIOW wunwnyd9Q SPlulpaiay, [BAe] saloadg snoredt Ale] wuday-)10Ys ATU PEDICELL AND SALINITY ON L. 7 RATURE 7 4, MPI TS OF TE 7 5) EVVE 1972 SOT “UOSIEAL (pnys Wasaig TPE] ‘UOsuOWPA RIpEsNy ‘purysuaend (Apnis qe]) HIRO “yovag su07] (Apnis qey) HEME +9 7-41 2239°ST age 4ST MOI2Q punoj jou O TT noge °%467-T7 Jwau 2%4L] ul skep ZI paatains O FcFI £2467 Baoqu "%ST MOTE °L0T £O OT oe o%S] ur Shep QT PeataAins AQT WOgR ‘D pI-ZI avou »%ST AA0Qe ‘D ye-Pl (Apnis qey) poonpal OF6l “GOON oneUpy O6I-01 °%07 Surg mS: 4 iam Aeg oosio LI6] ‘sMoreg -ueijt ues = 2A7Q] noe = = = = (Apmis qe) °% LT Ul Shep ome T ye Shep TF6l “‘UOsuOWpy TRAE = TI PeATAins = OT PeAtAins = = ML PAOGR €96] ‘weue (Apnis gry) SUIUIWWIMS -ysnygesen RIpuUy — —_— —_ 24,9 — jews0u Bleysny 2A, MOTAG LO6T WOSIe A, ‘puejsusen() — uouwwo0S — — — — 2a0mno0s AVEIOT surumeds TeATAINS AVATIOW JBATAINS sull0og AYWATIIOV oy mnundoO oj Joy mnwiydO perranersasug ig WIN UWITUTA UINUWITUTAL sprurpaieL Wnpy sprurpelal [BAe y (panuyuod) “| AlavL (snppppaaipad SNpOLK] =) NSIA) OPIdI.L snypjjaaipad snpolnT 1498}ADg Opada L (v4afiainf{ =) snyppplainf opataL dafynaod opasa [, satsads snoiediaieTy wiaj-du0T 122 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Taste 2. A comparison of the survival of Lyrodus pedicellatus larvae and adults under reduced salinity conditions at the end of seven days at two temperature levels. = a 14-16°C 22-24°C Salinity Larval Adult Larval Adult (%o) Survival Survival** Survival* Survival* * 35.0 90 100 70 100 31.5 65 100 70 100 28.8 70 100 50 100 25.2 60 100 10 90 21.6 50 100 8 50 18.0 50 100 3 40 14.4 35 100 0 40 10.8 0 60 0 10 Hee, 0 20 0 0 3.6 0 0 0 0 1.8 0 0 0 0 * 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,,) 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,, values for larvae were approxi- mately 9.4, 11.5, and 18.7%: 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—24C, 14-16C, and 9-11C. 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 VOLUME 71 TaBLe 3. The effects of temperature and reduced salinity on the larval swimming activity of Lyrodus pedicellatus 24 hours after release. Number of Swimming Larvae* Salinity 9-11°C 14-16°C 22-24°C 35.0 6 6 3 Shile) 9 3 0 28.8 21 9 0 25.2 18 3 0 21.6 15 i) 0 18.0 6 3} 0 14.4 3 0 0 10.8 0 0 0 7.2 0 0 0 3.6 0 0 10) 1.8 0 0 0 * Based upon 30 larvae tested at each salinity level. and 14-16 C, respectively. Penetration occurred only at normal salinity at 9-11 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%. 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, 1972), 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 pedicellatus: The effects of reduced salinity on the survival of the larvae of L. pedicellatus 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 EFFECTS OF TEMPERATURE AND SALINITY ON L. PEDICELLATU: TABLE 4. The effects of temperature and reduced Taste 5. The relationship of burroy f salinity on the boring of Lyrodus pedicellatus larvae length to reduced salinity in Lyrodus pedicetlan at the end of a 10-day period, two temperature level Number of Boring Lorvac® a Burrow Length Pallet J Salinity — —_ Number (mm) ( (%o) 9-11°C 14-16°C 22-24°C Salinity of (ho) Animals Range Mean Kar “he 35.0 2 gen 7 31.5 0 13%" 14—16°C Bee 0 35.0 7 15-25 209 20-31 2 Bore A ; 31.5 8 (3-20 415.31" aig=as) eas é ( 28.8 9 7-16 10.5 1.12.6 137 18.0 9 ) 25.2 5 5-12, 8.0) L.0=1h6 13 14.4 0 0 0 216 0 E 10.8 0 0 0 ; UP 0 0 0 22-24°C 3.6 0 0 0 ¢ 35.0 10 15—40 24.6 2.04.5 2.7 1.8 0 0 0 £1 (5) 10 16-30 23.4 2.0-3.9 2.5 Total 2 34 46 28.8 8 8-26 20.0 1:3=2'8, 23 25.2 9-12 8 ) “ Based upon 30 larvae tested at each salinity level. 4 AZ 10.3 1.5-1.8 1.6 *“*" Calcareous caps formed. 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,, values showed the limiting salinity levels to be approximately 9.4, 11.5, and 18.7% at 9-11, 14-16, and 22—24 C, respectively. Death occurred within a few hours at 7.2%: and below at all tem- perature levels. The three-day TL,, 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- cellatus from Hawaii were able to tolerate 15%c 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% 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 JT. navalis by Imai, 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. pedicellatus was found in this study to increase at lower temperatures, newly 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 18.0% 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 9-11 C. Larvae failing to secrete a protective caf died. It is apparent then that even though penetration of the substrate can be accomplishe at 9-11 C and in the lower salinity levels 16 C and 22-24 C, successful boring : more restricted. Lyrodus larvae are probably un- 124 able to penetrate wood and continue growth much below 14 C and below a salinity of 21.6%. (Table 1). Imai et al. (1950) reported 7. navalis capable of boring in brackish water at a salinity of 10% but were able to bore only between 14-26 C. Bulatoy (1941) 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 Lyrodus 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%c. 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 et al., 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— 11 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 ef al. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 (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,, values at 22-24 C (Table 2) showed that larvae and adults displayed values of 28.8 and 21.6%, respectively; whereas, these values dropped to 18.0 and 9.9%c, 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 7.5% 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- 1972 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 by New- combe and Kessler (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 five month period. Total growth was determined by measuring the burrow and 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% sufficient to reduce boring activity in L. pedicellatus. M’- Gonigle (1926) reported no boring by 7. navalis adults at 3.5%- or below, and Blum (1922) ob- served that 7. navalis did not produce fecal pellets at salinities of 4.0 to 5.0%. 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 Needler and Needler (1940), both working with T. navalis, and Quayle (1959) studying B. setacea, all reported the maxi- mum growth rate in the field corresponding to maximum seasonal temperature. 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. EFFECTS OF TEMPERATURE AND SALINITY ON L PEDICELLAT LITERATURE CITED Anonymous, 1927, Destruction of marine pilin borers. Canada, National Res Report of the President (1925 Council, Canada 26):42-43 Barnes, H. 1959, ography. Part one: Chemica. Interscience Publ Inc., New York, 341 pp. Apparatus and methods of ocean Barrows, A. L. 1917. An unusual extension of the distribution of the shipworm in San Francisco Bay, California. Univ. Publ. Zool 18:27-43. California Black, E. C., and C. R. Elsey. 1948. wood-borers in British Columbia waters. Fish. Res. Bd. Canada, 80, 20 pp. Incidence of Bull. Blum, H. FP. T. navalis. 368. 1922. On the effect of low salinity on Univ. California Publ. Zool., 22:349 Bulatov, G. A. 1941. Response of the larvae of the Black Sea 7. navalis to different water tempera- tures. Compt. rend. (Doklayd) de l’Acad. des Sci. SSSR, (n.s.) 32:291-292. Coes WwW. IR: molluscs. 1941. Sexual phases in wood-boring Biol. Bull., 81:168—176. Culliney, J. L. 1969. Larval biology and recruit- ment of the shipworms Teredo and Bankia gouldi, in the Newport Estuary, North Carolina. Unpublished Dissertation, Duke Univ. navalis 174 pp. Eckelbarger, K. J., and D. J. Reish. 1972. A first report of self-fertilization in the wood-boring Family Teredinidae (Mollusca, Bivalvia). Bull. So. California Acad. Sci., 71:48—50. Edmonson, C. H. 1942. Teredinidae of Hawaii. Oc- cas. Pap. Bernice P. Bishop Mus., 17:97—150. Grave, B. H. 1928. worm Teredo navalis Biol. Bull., 55:260—282. Natural history of the ship- at Wood’s Hole. Mass. Greenfield, L. J. 1952. The distribution of marine borers in the Miami area in relation to ecological conditions. Bull. Mar. Sci. Gulf and Caribbean. 2:448—464. Harington, C. R. 1922. Report of work done Marine Biological Station, Plymouth Sept. 18, 1920. Rep. Comm. (Deter. Struc Inst. Civil Engin., London, 2:35—42 126 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Horvath, C. 1951. A study of the teredinidae of Los Angeles-Long Beach Harbors. Unpublished Master’s thesis, Univ. So. California, Los Angeles, 145 pp. Imai, T., M. Hatanaka, and R. Sato. 1950. Breed- ing of marine timber borer, Teredo navalis L., in tanks and its use for anti-boring tests. Tohoku J. Agric. Res., 1:199-208. Isham, L. B., F. G. W. Smith, and V. Springer. 1951. Marine borer attack in relation to conditions of illumination. Bull. Mar. Sci. Gulf and Caribbean, 1:46-63. Isham, L. B., and J. Q. Tierney. 1953. Some as- pects of larval development and metamorphosis of Teredo (Lyrodus) pedicellata de Quatrefages. Bull. Mar. Sci. Gulf and Caribbean, 2:574—589. Johnson, M. W., and R. C. Miller. 1935. The sea- sonal settlement of shipworms, barnacles, and other wharf-pile organisms at Friday Harbor, Washington. Univ. Washington Publ. Oceanogr., 2:1-18. Kinne, O. 1963. The effects of temperature and salinity on marine and brackish water animals. 1. Temperature. Oceanogr. Mar. Biol. Ann. Rev., 1:301-340. Kofoid, C. A. 1921. Factors controlling the dis- tribution and ranges of the marine borers. Proc. Amer. Wood-Preserv. Assoc., 17:327—339. Kofoid, C. A., and R. C. Miller. 1927. Biological Section. Pp. 188-343 in Marine borers and their relation to marine construction on the Pacific coast. (C. L. Hill, and C. A. Kofoid, eds.), San Francisco Bay Piling Comm., San Francisco, ix + 357 pp. Kudinova-Pasternak, R. K. 1962. Effect of sea water of a decreased salinity and of various tem- peratures upon the larvae of Teredo navalis. Zool. Zhur. Akad. Nauk. SSSR, 41:49—-57. Lebour, M. V. 1946. The species of Teredo from Plymouth waters. J. Mar. Biol. Assoc. U.K., 26: 381-389. Loosanoff, V. L., and H. C. Davis. 1963. Rearing of bivalve mollusks. Advances in Mar. Biol., 1: 1-136. Menzies, R. J., J. Mohr, and C. M. Wakeman. 1963. The seasonal settlement of wood-borers in Los VOLUME 71 Angeles-Long Beach Harbors. Wasmann J. Biol., 21:97-120. M’Gonigle, R. H. 1926. A further consideration of the relation between distribution of Teredo navalis (Linne), and the temperature and salinity of its environment. Nat. Res. Council, Canada Rept. no. 20, 31 pp. Miller, R. C. 1926. Ecological relations of marine- boring organisms in San Francisco Bay. Ecology, 7:247-254. Moore, D. R., and D. J. Reish. 1969. Studies on the Mytilus edulis community in Alamitos Bay, California. IV. Seasonal variation in gametes from different regions of the bay. The Veliger, 11:250-255. Nagabhushanam, R. 1961. Ecological factors re- lated to the distribution of Bankia campanellata in Visakhopatnam Harbour. Amer. Zool., 1:243. 1963. of the shipworm, Teredo furcillatus. Exp. Biol., 1:232. Effect of low salinity on the veligers Indian J. Needler, A. W. H., and A. B. Needler. 1940. Growth of young shipworms (Teredo navalis) in Mal- peque Bay. J. Fish. Res. Bd. Canada, 5:8-10. Newcombe, R. C., and R. F. Kessler. 1936. The effects of salinity on shell formation in molluscs. Ecology, 17:429-443. Quayle, D. B. 1959. The early development of Bankia setacea Tryon. Pp. 157-174 in Marine Boring and Fouling Organisms. (D. L. Ray, ed.), Univ. Washington Press, xii + 536 pp. Ralph, P. M., and D. E. Hurley. 1952. The settling and growth of wharf-pile fauna in Port Nichol- son, Wellington, New Zealand. Zool. Publ. Vic- toria Univ. College, 19:1—22. Rancurel, P. 1971. Les Teredinidea (Mollusques Lamellibranches) dans les lagunes de Cote d’Ivo. Mem. Orstom n°47, Paris, 235 pp. Reish, D. J., and J. L. Ayres, Jr. 1968. Studies on the Mytilus edulis community in Alamitos Bay, California. III. The effects of reduced dissolved oxygen and chlorinity concentrations on survival and byssus thread formation. The Veliger, 10: 384-388. Roch, F. 1940. Die Terediniden de Mittelmeeres. Thalassia, 4:1—147. 1972 Ryabchikoy, P. 1, and G. G. Nikalaeva, 1963, Set- tlement on wood of the larvae of Teredo navalis L. (Teredinidae, Mollusca) in the Black Sea. Trudy Inst. Okeanol., Akad. Nauk. SSSR, 70: 179-185. Scheltema, R. S., and R. V. Truitt. 1954. Ecologi- cal factors related to the distribution of Bankia gouldi Bartsch in Chesapeake Bay. Chesapeake Biol. Lab., Publ. 100, 31 pp. Smith, M. L. 1963. The teredinidae of the Queens- land coast from Cairns to Brisbane. Unpublished MSc. thesis, Zoology, Univ. Queensland, Bris- bane, 206 pp. Stone, A. N., and D. J. Reish. 1965. ‘The effect of fresh water runoff on a population of estuarine polychaetous annelids. Bull. So. California Acad. Sci., 64: 111-119. 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. Trussell, P. C. 1967. Teredine borers. Sea Fron- tiers, Internat. Oceanogr. Found., 13:235—243. EFFECTS OF TEMPERATURE AND SALINITY ON L. PEDICELLATUS Turner, R. D. 1966. A survey and illustrated logue of the teredinidae (Mollusca: Bivalvia) Harvard Univ. Cambridge, Massachusetts, 26° pp. Watson, C. J. J. 1936. Marine borers destroying timber in the port of Brisbane. Destruction of timber by marine organisms in the port of Brisbane. (C. J. J. Watson ef al.) Queensland Forest Serv. Bull., 12:x 4- 107 pp Pp. 10-30 in 1957. ing timber: The effect of stream flow on and destruction of Turpentine piling by marine borers in the Brisbane River. Queensland Forest Serv., Forest Prod. Res. Notes, Project O.P. &, no. 6, 28 pp. Studies in marine organisms attack salinity White, 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? 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 28 from the Pacific Marine Station and was supported by N.S.F 1 Pacific Marine Station. University of the Pacific Dillon Beach, California 94929. 128 Grant GA-1405. Figure 2 was prepared by Floy E. MacMillan-Zittin. Family Lumbrineridae Lumbrineris fauchaldi, new species Figure | Material examined: Davis Strait (lat. 66° 29’ N-long. 57° 26’ W), HERO 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. 1b). One specimen had three anal cirri (Fig. 1c). 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- BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 gated capillary tips (Fig. ld). The hooded hooks completely replace the limbate setae in the last 1% of the body. Hooded hooks are 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 II.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 Asclerocheilus Absent Absent Ashworth, 1901 Cryptosclerocheilus Present Absent new genus Setigers 2-5 Parasclerocheilus Present Ventral cirrus Fauvel, 1928 Setigers on posterior 2-7 setigers Sclerobregma Absent Dorsal and ventral Hartman, 1965 cirri in posterior setigers Sclerocheilus Absent Ventral cirrus on Grube, 1863 posterior setigers Acicular Setae Reference Present in both rami of Setigers 1-3 Ashworth, 1901 Fauvel, 1927 Day, 1967 Limited to both rami of Setiger 2 This paper Limited to notopodia of Setigers 1-4 Fauvel, 1928 Day, 1961 Limited to both rami of Setiger 1 Hartman, 1965 Ashworth, 1901 Fauvel, 1927 Present in both rami of Setigers 1-3 \ 0.02mm \ Figure 1. 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; 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 NEW SPECIES OF POLYCHAETOUS O.2mr Lumbrineris fauchaldi, new species: ANNELIDS L a, anterior end in dorsal view: b, posterior end a-c all to the same scale. approximately setiger 25, but there are no transi- tional setae as in L. fauchaldi; L. neozealaniae was apparently described from a group of species an type material needs to be reexamined; and I punctata has only two teeth on the left side of maxilla IJ and three on the ri 19 130 BULLETIN SOUTHERN CALIFORNIA ACADEMY SAS LS Eine te i}, aie a pA £7 Beh OF SCIENCES VOLUME 71 1972 NEW SPECIES OF It is a pleasure to name this species for Dr. 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. Prosto- mium T-shaped, with two long frontal lobes. Without eyes. Peristomium achaetous. Proboscis unarmed. Branchiae limited to anterior segments. Parapodial lobes reduced to short elevations throughout the body; no cirriform processes present in either 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 begin 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- POLYCHALTOUS ANNELID setae and neurosetae (Fig. 2c). The s the first (Fig le) and fr ger 2 are arranged in two tier slender curved acicular setae furcate setae; the second group are long cay The neurosetae have a similar arrangement, « pt that there are more of the acicular and furcate setac and fewer capillaries. The furcate setae complete! replace the acicular setae on setiger 3. This arrange ment of one bundle of capillary setae and bundle of smaller furcate setae continues on succeed ing setigers to the end of the body. The furcate setac have one tine longer than the other. Each tine i spinous along the inner border (Fig. 2d) Distribution: Southern Baffin Bay, depth of 1,830 m. LITERATURE CITED Ashworth, J. H. 1901. The Anatomy of Scaligregma inflatum Rathke. Quart. J. Micro. Sci., 45:237 309. Day, J. H. 1961. The polychaete 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 Polychaeta 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, Lysaretidae and Dorvilleidae from western Mexico. Allan Hancock Monogr. Mar. Biol., (6):1—335. Fauvel, P. 1927. Polychétes Sédentaires, addenda aux Errantes. Archiannelides, Myzostomaires. Faune de France, 16: 1-492. 1928. Annélides polychétes nouvelle de lInde. 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. invertebrate Systematic account of some from animals the deep ba Figure 2. 3; e, acicular seta from setiger 2. Cryptosclerocheilus baffinensis, new species: a, anterior end in ventral view: c, capillary seta from anterior setiger: d. fur entire animal in dorsal vie ate seta from se southern California. Allan Hancock Pacific Ex- ped., 22:69-216. 1965. Deep water benthic polychaetous annelids off New England to Bermuda and other North Atlantic areas. Allan Hancock Found. Occas. Pap., (28): 1-378. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 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,! AND MICHAEL A. MARES? Apstract: SS) N y Ni % oo XS Re bs > x or Cc) 0) Tan ? Q : mle 0) > ~ & oo M ®D NS : TA LOS é r ws ws 405 lice) \ we Oe ea Pe le Figure |. 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 ACADEMY OF SCIENCES VOLUME 71 dense scrub forest (Tropical Dry Forest formation of Holdridge, 1962). ANALYSIS OF VARIATION Snakes of the genus Micrurus 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 are 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, 11 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 Males Females Males Females A 190-203(198.4)9 208—220(213.0) 13 47-51(49.6)8 34-42(37.6)12 B 191—206(196.8) 13 206-217(210.4)9 46-52(49.3)8 34-39(36.4)9 Cc 193-202(198.6)5 206-—217(212.5)8 45—51(47.0)5 32-39(36.8)6 D 198 208 45 36 E 190-201(195.5)2 210 49-51(50.0)2 37 F 203 - 46 - G 200—202(201.0)2 211 54-56(55.0)2 - H 208 215-218(216.5)2 50 36-39 (37.7)3 I 201 - 46 - J 191—206(200.0)4 218-224(220.6)3 42-50(47.5)4 34-43 (38.0)3 K 207 = 51 - 1972 TABLE 2. Variation in black body band number in Honduran Micrurus nigrocinctus., Population statistics are arranged as follows: range (mean) sample size. Population Males Females A 13—24(17.6)9 14—23(19.1)13 B 11—21(13.8) 13 11—-20(16.3) 11 (©; 15—19(17.4)5 16—21(18.6)8 D 27 29 E 14—20(17.0)2 22 F 25) a G 18—19(18.5)2 23 H 20 17—22(19.0)3 I 22 - J 18—19(18.5)4 18—20(19.0)3 kK 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 0 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 11 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 1% 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 THE CORAL SNAKE MICRURUS NIGROCINCTUS // HONDUI { } * 5. 0's e050 Figure 2. Dorsal color pattern of (A) LSUMZ 24423 from 4 miles N San Lorenzo, Depto. Valle, Honduras and (B) LSUMZ 22451 Trujillo, Depto. Colon, Honduras. from the mountains above 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 of M. 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. cinctus (Girard) (Pacific southern Costa Rica, Panama, and adjacent Colombia). !]. n. mosquitensis Schmidt (Atlantic versant of east- ern and southern Nicaragua to northwestern Panama), and M. n. babaspul Roze (Isla del Maiz, Nicaragua) have well-defined black spots present nigro- versant of on the tips of each of the scales of the red bands. relatively Micrurus n. divaricatus has a y large amount of black pigment irregularly distributed on the scales of the red bands. The amount of variation in black pigment dis- tribution in specimens of Af. nigro Honduras is so great as to coincide with th nge of variation for the entire species. up seven pattern categorie 142 BULLETIN 5 a arab Ae Oedl epececetien a esesesesetes 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. 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. SA). 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. melano- cephalus. Pattern 2 characterizes M. n. cinctus, M. n. mosquitensis, and M. n. babaspul, coibensis and M. n. nigro- and patterns 4 and 5 are found on M. n. divari- SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 re %O re Gai yy 9 > » lias are) See Sn ORNS Figure 4. Dorsal color pattern of (A) USNM 20282 and (B) USNM 20283 from Patuca, Depto. Gracias a Dios, Honduras. 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 11 populations is shown in Table 35 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. Pattern types Population 1 2 3 4 5 6 7 A 8 10 — 4 1 - - B 7 12 3 1 C 1 — | 1g ean ig se 5 = D ~ ot > 25 1 1 E a er F = ss he i G = 1 = $e ee H Bh i Sus 1 = I 1 J 4 3 K 1 - Total 2.65 3 11 4 1 1 1972 THI CORAL SNAKE to the more northern portions of the country and are represented in very few of the specimens. The populations with the largest number of members exhibit the widest range of pattern types. Thus, 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 D, Also, patterns 1, 2, and 4 are found to the exclusion of other pattern types in_ the southern populations (H through K). Position of the nuchal band and the size of the black head cap.—Vhe position of the anterior edge of the nuchal band in Honduran M. nigrocinctus ranges from 2 dorsal scale lengths posterior to the parietals to 14 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 the 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. 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% miles east of La Ceiba, Depto. Atlantida. This record has questioned by Meyer (1969) and Wilson and Hahn (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 been 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 MICRURUS NIGROCINCTUS / HIONDI ” > i WK 470; 0-0:0.0.40 4M Se 1676-06-66. 6, 6: LL Figure 5. Dorsal color pattern of (A) LSUMZ 21773 from the mountains above Corozal, Depto. Atlantida Honduras and (B) TCWC 21181 from 5% miles I La Ceiba, Depto. Atlantida, Honduras. 45 according to Schmidt, 1933, and a range of 33 to 45 is given for 8 specimens examined Wilson and Hahn, in press). The ventral count for TCWC 21181 (a female) 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 (Giinther, 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 M tanus in pattern, particularly in the relatively high number of black body bands, is only superficial. The specimen appears to have been bicolor in ruatanus rua- life, as is the case with fresh material from this general area of Honduras and with M. ruatanus. Nevertheless, the black body are alternately longer and shorter and the ones are frequently interrupted laterally. especiall on the posterior portion of the body (Schm 1933: Wilson and Hahn, in press). On the other hand, the bands of TCWC the same length (Fig. 5B) and none are broken that this bands of ruat 21181 are all about We suggest specimen. with its hig number of black body bands (for represents the end sta increasing development and c pigment. We envision 144 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES nate in a condition that we have designated as pattern 2, where black tipping is uniformly dis- tributed on all scales of the red band. This cline continues through pattern 4, where some black tips are fused together to form larger spots, and patterns 5 and 6, where either paravertebral or lateral accessory incomplete bands are present. The cline ends with pattern 7 (that of TCWC 21181), where the accessory bands 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 (J, 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 VOLUME 71 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 11 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-11); W of San Pedro Sula (FMNH 5312); 15.5 miS Puerto Cortés (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 E.—COLON: Puerto Castilla (MCZ 2243334); mountains above Trujillo (LSUMZ 22451). Population F.—OLANCHO: 2 km SW _ Dulce Nombre de Culmi (LACM 45179). Population G.—GRACIAS A_ DIOS: (USNM 20282-83); Tancin (LSUMZ 24422). Population H.—CORTES: Agua Azul (AMNH 70157, 70216-17); 2 mi N El Jaral (LSUMZ 24424). Patuca Population I1.—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: (LSUMZ 24423). 4 mi N San Lorenzo 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 THE CORAL SNAKE MICRURUS We also extend our gratitude to John Dickson of La Lima, Honduras for the opportunity to examine specimens in his personal collection and to Diana Dee Dugas for her help in gathering data. LITERATURE CITED Ginther, A. G. L. C. 1895. Biologia Centrali- Americana. Reptilia and Batrachia. London, xx +- 326 pp. Holdridge, L. R. ras. 1962. Organizacion de Los Estados Americanos, San José, Costa Rica. Mapa ecologico de Hondu- Meyer, J. R. 1969. amphibians and reptiles of Honduras. Thesis, Univ. Southern California. 589 pp. A biogeographic study of the Ph.D. Neill, W. T. 1963. Polychromatism in_ snakes. Quart. J. Florida Acad. Sci., 26:194—216. NIGROCINCTUS IN HONDURA Roze, J. A. 1967. A check list of th venomous coral snakes (Elapidae), with de tions of new forms. Amer. Mus. Novit 1-60. — 1970, Micrurus. Pp. 196-220, in Cata logue of the Neotropical Squamata. (J. A. Peter and B. Orejas-Miranda, eds.) Part I. Snake Smithsonian Institution Press, Washington, D. ¢ 347 pp. Schmidt, K. P. 1933. Preliminary account of the coral snakes of Central America and Mexico Field Mus. Nat. Hist., Zool. Ser., 20:29—40, 1936. Notes on Mexican coral snakes. Zool. Ser., 20:205-216. and Hist., American Mus. Nat. Central Field Wilson, L. D., and D. E. Hahn. 1972. The herpeto- fauna of the Islas de 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! 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. crax Deichmann, 1940, 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 Echinoderm Collectic No. 61). total length 6 cm (contracted) and 3 par types (AHF Echinoderm Collection) were collec at a depth of 14 m on the north face of Bird R 1 Allan Hancock Foundation, University S ern California, Los Angeles. Californ 146 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Figure 1. 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 VOLUME 71 0.025mm 0.5mm 0.3mm A, B, supporting rods from tentacles; C, perforated plates from tentacles; D, E, F, 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 CUCUMBER FROM CATALINA ISLAND 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. 1C). 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 mottled with white and black patches of pigment: tentacular papill black. Measurements: Length of relaxed specimens about 10 cm. 148 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 Figure 3. 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 0.5mm ] | 05mm 0.5mm A, E, perforated three-armed rods from body wall; B, C, D, perforated plates from 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 salma [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 CUCUMBER FROM CATALINA ISLAND C. salma Medium, 10 cm Equal, bushy Retractile; double row: TABLE 1. Comparison of west coast species in the genus Cucumaria C. curata C. crax C. vegae SIZE Up to 2.5 cm; Small, 1.5 cm Small, 3.8 cm; ventrum flat (contracted ) ends tapered TENTACLES Unequal Equal, bushy Unequal TUBE FEET Few; double row in Large; retractile; Double row; in dorsal ambulacrum only in ambulacra interambulacra 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 Dorsum and tent. purple brown; vent. and oral disc yellow Intertidal, rocks, mussel beds, central California CALCAREOUS RING Simple; post. Slender; no post. prolongations prolongations 1/2/? 1/1/5 SPICULES-BODY Buttons, 4—8 marginal Few, smooth rods, 1—3 holes, not all entirely holes in each end 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 COLOR Mottled brown body; Vent. yellow, gray: ambulacra lighter: body ends brown tent. black DISTRIBUTION Sand, west coast Baja Intertidal rock California—South crevices, north America Pacific coast only in ambulacra Post. prolongations; ant. processes w/cleft STONE CANAL/POLIAN VESICLES/GENITAL PAPILLAE 3—4/4/none Smooth perf. rods and buttons; plates 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, 1868, to Pseu- docnus dubiosus and in 1964 he changed C. cali- fornica Semper, 1868, to Pseudocnus californicus. Stimpson (1964) changed Cucumaria populifera the remaining three species C. crax. C. curata. and C. vegae; all of which are found on the Pacific piperata Stimpson, 1864, do not belong in the genus Cucumaria and probably should be placed in the genus Pseudocnus. Cucumaria salma is easily distinguished from to Pentamera populifera Panning (1949). Pawson coast of North America. Cucumaria soln (pers. comm.) suggests that Cucumaria miniata Brandt, 1835, C. lubrica Clark, 1901, and C. about 10 cm in length: has three or four stone canals; four Polian vesicles: is salmon colored: anc 150 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES has distinct spicules. The latter 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 (Table 1). ACKNOWLEDGMENTS I would like to thank D. L. Pawson, Smithsonian Institute for his of my findings, D. Straughan and K. Fauchald, Allan Hancock Foun- verification 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. 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 VOLUME 71 Part I, Dendrochirota. Allan Hancock Pacific Exped., 8(3):61-196. Panning, A. 1949. Versuch einer neordnung der familie Cucumariidae. Zool. Jb. Abt. Systematik, 78(4) :404—470. 1951. Uber Pseudocnus leoninus (Semper) und verwandte Arten. Zool. Anz., 146:73-80. 1962. Bermerkungen tiber die holothurien- familie Cucumariidae (ordnung Dendrochirota). 3 Teil. Die gattung Pseudocnus Panning. Mitt. Hamburg Zool. Mus. Inst., 60:57—80. 1964. Bermerkungen uber die holothurien- familie Cucumariidae (ordnung Dendrochirota). 3 Teil. Die gattungen Stereoderma, Staurothyone, and Trachythyone. Mitt. Hamburg Zool. Mus. Inst., Kosswig-Festschrift., 61:159-174. 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! 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 Ps, Pi, M:, and presumably Mz, 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. Jutris. 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 ' Dept. Geology and Earth Science, California State University, Humboldt, Arcata, California. 1972 the Palos Verdes Sand, and various limbs and vertebral elements from Late Pleistocene deposits of Santa Rosa Island. Leffler referred the Oregon femur to Enhydra sp. while Mitchell concluded that all the southern California specimens probably were referable to Enhydra lutris (Linnaeus). In May 1967, a well preserved mandible of a fossil sea otter, lacking only a few teeth, was recovered from a Late Pleistocene marine deposit, located on the Humboldt County coast, north- western California. The mandible is evidently the first Known of a fossil sea otter from the north- eastern Pacific and it provides the first major indication of the 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 Ps, M:, and M2; right incisors and Ms. 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 11 miles north of Arcata, 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. NEW FOSSIL SEA OTTER FROM CALIFORNIA Figure J. Key to mandibular measurements utilized in this study and presented in Table 1. A, B, and D are measured from the posterior margin of the condylar process along the axis of the ascend- ing ramus. All measurements are in millimeters. Dimensions Formation: The otter-bearing unit has not been formally described or named and it can be definitely traced only about *4 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—Pleisto- 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 Ps, P., and M: approximately 1, larger and plump; Mz 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 those of the compared specimens of Enjiyd (Table 1) with respect to size, shape morphology indicates that E. macr ita is clo related to E. lutris and is certainl VOLUME 71 ‘uonnuap yIIW pajdniauQ yx ‘uonnuep NINA * FeO ‘Aaaysog “[OOZ “WaA WnasnyAl “HED Jo “ATU, (P) ‘yyeO ‘yIeg oUap ‘1ajua_D yseOD s1jloeg ‘ABAING [BdIBOTOIH “§ “Q (9) SCIENCES MY OF BULLETIN SOUTHERN CALIFORNIA ACADE 2 1 “UIPIM MA “ABOTOOZ JO ‘Jdaq “yproquiny “Atuy aeIg “JED (q) ‘ysueT 7 ‘g0udIdg Ye pue Adojoay Jo "‘}daq ‘Ipjoquiny “ATUA, a1eIg “FRO (ke) SS Gh HAC ae OFS) Domes 059 ames Cal 6el Scs OFT T8l ere Pre BIb SPL dA “Ang y i 160-ILW (4) Sis) ys [ASL FEO yb S35 LULL 9€S OOT €8I SBC H8E PEP HSL & ynpe sux te ut 060-TLW (4) 6% VL CHGE AAS Gt3s SR €Cl OPS EST OBI T8T PHBE Ver O6L WW “AUL s i 680-ILW (4) OS hal 6) me o/s Se Sy aoe S VCI 9S 69I PBI S6C S6E LSh BSL é “ANE “ Mm 880-TLW (4) £9 9 VEL GSiL0 Lf. 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OBE SIS PEs DIS W OPV i FRO 1089 (9) OL s9 SOS GAL Otsll G/L OS 7 LOI] S9S T6l OF PLE GSLr £98 606 W yOpVy 4 BYSeTV 7069 (9) SO) /EX) (GO SANE} REGIE Ssdsit 76 OHS TIT 09 LLI 977 SEE TL BPS 8:06 é yopyv SLAIN] u O80-TLW (4) 68 €8 61l vel 8st Sst LS 89 TST PIO E81 96 O'8T GIF TOS 68 4 npe sux — ypyuoposoviu “gy FTO I-Spl (e) Mw T Mw I M I M I H 5) Fs § a da 2) a Vv xag_ aHEIS YIMOID satoadg “90'T “ON ‘oadg &d td 'W (snjoaare) “W “SIOJOUNT][IW UT oe S]UDWWAINseoU! [TY “[ aindi4 Ul UdAIS SI Y-VY S]UOWAINsvaW 0] Ady ‘“sl4iN] “q pue saidads MoU ‘DJUOpOsAIVIU DApPAYUG JO SJUdWAINSvIW Iv[NqIpurlu pa}ssfog “| ATV], 1972 ae ¢ - - Ko ee NEW FOSSIL SEA OTTER FROM CALIFORNIA 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 1x. 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,;—P, and substantially more con- vex below M, and Ms. There are two large, oval- shaped mental foramina on each ramus below Ps and P, 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 lightly built, more slender, and shorter than those of EF. /utris 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. /utris 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 depression which shallows anteriorly, diminishing to an arrowhead-shaped directed anteriorly. not downward as is comm in E. lutris and terminates just belo fossa is a deeply excavated point. The point is 154 BULLETIN SOUTHERN CALIFORNIA half of M,. In E&. lutris the anterior extremity of the masseteric fossa almost invariably lies below Ms and occasionally lies below the posterior mar- gin of M,. The condyles are about the same size and diameter as the largest specimens of E. lutris examined and they 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 of 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 FE. Jutris curvature of the toothrow also occurs but it seems to be much less pronounced (Fig. 2A, B). The Mz is lost in both left and right jaws of E. macrodonta but the alveoli are well-preserved and their size (Table 1) suggests that Ms was slightly larger than that found in E. lutris. The alveolus of Mz 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,. 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 M, 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 ACADEMY OF SCIENCES VOLUME 71 about '% larger. The alveolus for My, 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. /utris. The P, 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. in E. macrodonta has nearly twice the occlusal area of that in FE. /utris. It is about ¥% the size of P, but is similar in occlusal outline, the major exception being a more sharply rounded anterior corner. The low ridge on the medial side of the tooth is proportionately larger in E. lutris and it tends to be more inflated and cusp-like. The tooth is oriented as in E. lutris but does not appear to lean outward and overhang the jaw as is frequently the case in E. lutris. The P, is about % the size of Ps, and is only slightly larger than P., in E. lutris. The long axis of the tooth crosses the jaw axis at an angle some- what greater than in Ps. The tooth does not appear to be directed laterally nor to overhang the jaw to the degree as in E. lutris. A single, nearly round alveolus in the left jaw indicates that the tooth possessed one root, as in E. lutris. The canine is almost identical in size and form to the canine in E. lutris. The first lower incisor of E. macrodonta is a long, slender tooth of which nearly half is exposed root. The cutting edge of the enameled surface is bluntly wedge-shaped, with the long axis oriented transverse to the jaw axis. The second lower incisor tightly fills a narrow space between J, and the canine. The wedge- shaped crown is about twice the length as that of I,. Taken as a group, the close alignment of the canines and incisors suggest that the bite of the animal was effective across the entire anterior margin of the mandible. The incisors show very close correspondence in size to those in E. lutris. DISCUSSION AND CONCLUSIONS Taxonomic assignment: Enhydra macrodonta is assigned to the genus Enhydra because of its close 1972 resemblance to /. /utris in both mandibular and dental morphology. It consistently and signifi- cantly differs from known or examined mandibles of E. lutris in the following features: 1) greater size of Py, Py, M,, and possibly My; 2) longer toothrow; 3) a sharply-rounded angle formed of ascending and ventral margins which are narrow and rounded; and 4) shorter ascending ramus with respect to total length of mandible. These differences are interpreted as justifying the assign- ment of this fossil mandible to a new species. Other morphologic characters which are, to some extent, duplicated in specimens of FE. /utris include: 1) the masseteric fossa extends anteriorly to a point beneath the posterior half of M,, and is anteriorly directed; 2) Ms. occupies a large, medially directed swelling, partially hidden from lateral view by the ascending ramus; 3) the medial and lateral surfaces of the ascending ramus are essentially smooth and without bulges or depres- sions; and 4) the margin of the coronoid crest is narrow and curled only slightly medially. These four characters may represent a transitional stage in growth or perhaps relate to sex and are, there- fore, considered to be of questionable diagnostic value at this time. Morphologically E. macrodonta is sufficiently close to E. lutris to indicate that the two species are closely related. However, the larger teeth and thicker jaw of E. macrodonta seem to suggest that E. lutris was, perhaps, not derived directly from E. macrodonta, but rather that the two species represent separate lineages which presumably had a common origin prior to the Late Pleistocene. One cannot rule out the possibility, however, that selection toward a “small-toothed” E. lutris might have occurred from a “large-toothed” E. macro- donta. If this selection did occur then the general tendencies during the evolution of E. macrodonta to E. lutris would have included: 1) reduction in size of Ps, Py, M,, and possibly Ms, with propor- tionately greater reduction in tooth width; 2) reduction in the thickness of the jaw below Ps, P,, and M,; 3) reduction in the length of the tooth- row; 4) increase in the overall size and complexity of the ascending ramus: 5) increase in height of the ascending ramus with respect to total man- dibular length: 6) development of stronger muscle attachments at the angle and ventral margin of the ramus; 7) maintenance of the same general size of incisors, canines, and Ps: and 8) main- tenance of about the same orientation of the long axes of premolars and molars in occlusal view. The suggestion by Fisher (1941), that the mandible in E. lutris might reflect an evolutionary NEW FOSSIL SEA OTTER FROM CALIFORNIA process tending to progressively shorten the m dible, is generally supported by the present stud If, as Fisher suggested, the teeth have been pro gressively crowded together during the evolution of sea otters, so that their long axe now ume variable angles as they do in £. macrodonta, then that process probably began before the Lat Pleistocene, Enhydra macrodonta does not appear to retain characters which might provide clues into the pre-Pleistocene evolution of sea otters, nor does it seem to offer a test of the hypothesis suggested by McLaren (1960) that phocids were derived from a lutrine should be ranked as a very late development in sea otter evolution one close affinity to the living species. The multicusped, left, deciduous P, from the Timm’s Point Silt (Early Mitchell (1966, p. 1909, figs. 4a—c) referred to Enhydra cf. E. lutris, allows little basis for direct comparison with the permanent dentition of E. macrodonta. Mitchell referred the Timm’s Point fossil to the living species on the basis of obvious resemblance, but recognized that it differed from the compared specimens of FE. /utris in being generally larger and in various details of occlusal morphology. The Timm’s Point tooth was found to be slightly larger than two lower deciduous fourth premolars of E. /utris which were available in the present study (Table 1) and two others examined at the U.S. Geological Survey (Repen- ning, pers. comm. 1972). The fact that the deciduous tooth from Timm’s Point Silt and the permanent premolars and M, of E. macrodonta are larger than their counterparts in E. /utris may be significant and open up the possibility that the Timm’s Point fossil may be more closely related to E. macrodonta than to E. lutris. In addition, if the Late Pleistocene post-cranial elements de- scribed by Mitchell are indeed those of EF. /utris. then it would appear probable that E. macrodonta and E. lutris coexisted in the northeastern Pacific during some part of the Pleistocene. perhaps as geographically separated populations. The right M, of E. macrodonta shows a crude resemblance to the right M, of Lurra reevei New- ton from the Norwich Crag (England) of Middle Pleistocene age, in having the same number and Q It differs principally in having: 1) about twice the occlus ancestry. The species evidently and which shows very Pleistocene), which gross arrangement of cusps and ridges. area; 2) a much squarer occlusal outlit blunter anterior margin: 3) a paraconi and anterior to the protoconid: and + proportionately smaller 156 and metaconid positioned in a relatively more an- terior position. These differences, taken in com- bination, would seem to indicate that E. macro- donta and Lutra reevei are not particularly closely related and that the M, of L. reevei was not a suitable morphologic structure for the derivation of M, in FE. macrodonta in the interval from Middle to Late Pleistocene. Thus the author is in accord with Mitchell (1966) and Repenning (1967) that L. reevei is not ancestral to Enhydra. Lutra reevei, on the contrary, seems more closely allied to the Recent Aonyx capensis Schinz and the Pliocene Enhydriodon africanus Stormer, both from South Africa. Adaptations: In comparison with E. lutris specimens of the same growth stage, the coronoid process in E. macrodonta is consistently shorter with respect to jaw length. This relationship suggests that the skull was not higher than E. lutris and may have been slightly lower. Preserved insertional structures on the mandible of E. macrodonta, including those for the mas- seter, internal pterygoid, and temporal muscles, appear on the whole to be much less strongly formed than their counterparts in E. lutris. This circumstance suggests that the jaw musculature of E. macrodonta was generally less developed than that of the living species. If this line of reasoning is correct then it remains to be explained why E. macrodonta was endowed with substantially larger and heavier posterior teeth than is FE. lutris. In theory this condition would seem to require that the jaw muscles of E. macrodonta would have to be more powerful than those for E. /utris, not less, as a comparison of the muscle insertion parts seem to suggest. No satisfactory explanation has been found for this apparent paradox. It is possible that the musculature of E. macro- donta was, indeed, proportionately stronger than that of E. lutris but that: 1) the insertional struc- tures do not reflect this condition, or 2) the inser- tional structures were not yet fully developed in the young adult of E. macrodonta and would be larger and more strongly formed in the fully adult state. Environment of deposition: Enhydra macro- donta was recovered from a gray, clayey, fine- grained sand containing the following molluscan species: Yoldia sp.: Mytilus californianus; Clino- cardium sp.: Macoma nasuta;: Solen sicarius; Cryp- tomya_ californica: (?) sp.: Epitonium Thais lamellosa; Thais lima; Nassarius fossatus; Nassarius rhinetes; and Olivella biplicata. The configuration of topo- graphically higher exposures of Franciscan Forma- Teredo indianorum: Polinices lewisii: BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 tion rocks (Late Jurassic—Late Cretaceous), north and east of the Crannell Road deposit, indicate that the deposit probably accumulated in a semi- protected marine embayment along an open coast. The fossil assemblage and associated sediment indicate that the site of deposition was a muddy bottom which probably did not exceed five fathoms in depth. Some rocky outcroppings were located nearby. Age: The recency of the otter-bearing deposit at Crannell Road is suggested by the following lines of evidence: 1) general lack of consolidation of the sediments and near horizontal attitude of bedding; 2) except for Clinocardium sp., which may be an extinct species, all the other mollusks in the otter-bearing deposit are presently living along the Humboldt County coast; and 3) fish otoliths found in the gray, pebbly sand, imme- diately below the otter-bearing unit (see descrip- tion of the type locality above) are the same as those from the Palos Verdes Sand (Late Pleisto- cene) of San Pedro (Fitch, 1970). Mammalian remains from the otter-bearing unit and from a marine deposit at Moonstone Beach, located 1% miles north of Crannell Road, suggest somewhat greater precision in dating Enhydra macrodonta. From the pebbly, fossiliferous sand unit, which stratigraphically underlies the otter- bearing unit, C. A. Repenning (pers. comm. to R. Kohl, 1972) has identified an upper premolar apparently inseparable from the sea lion, Eume- topias jubata. Repenning suggests that this tooth, based on limited material from other areas, is no older in age than Rancholabrean (Late Pleisto- cene). The Moonstone Beach deposit, which appears to contain a marine invertebrate fauna slightly older than that of the otter-bearing unit, has yielded the left scaphoid of a mammoth, Mam- muthus sp. Repenning (pers. comm. 1972) con- siders this element to be no older in age than Irvingtonian (Middle Pleistocene), although it may be slightly younger. Thus the available evidence points to a Ran- cholabrean or Late Pleistocene age for Enhydra macrodonta. ACKNOWLEDGMENTS The author extends sincere appreciation to Elizabeth Rose, Eureka, California, and Roy Kohl, Dows Prai- rie, for making the otter remains available for study. Charles A. Repenning provided valuable critical dis- | cussion of the manuscript and loaned specimens from the collection of the U.S. Geological Survey. Warren J. Houck and Robert Jones contributed importantly 1972 NEW FOSSIL SEA OTTER FROM CALIFORNIA to the discussion and loaned specimens for comparison cence) of California, Los Angeles Co. Mu from collections in the Dept. of Zoology, California Hist., Contrib. Sci., no. 199:1-41 State University, Humboldt, and the Museum of Vertebrate Zoology, University of California, Berke- [effler, §. R. 1964, Fossil Mammals from the Plb ley, respectively, Valuable information concerning River Formation, Cape Blanco, Oregon. Jc living sea otters was furnished by Aryan I. Roest Mamm., 45:53-61, (California State University, San Luis Obispo), Karl Kenyon (U.S. Fish and Wildlife Service) and Karl McLaren, I. A. 1960. Are the Pinnipedia biph B. Schneider (Dept. of Fish and Game, Alaska). letic? Syst. Zool., 9:18-28. Warren O. Addicott, U.S. Geological Survey, kindly furnished information on the ecology of marine Mitchell, BE. D. 1966. Northeastern Pacific Pleisto invertebrates. Kenji Sakamoto, U.S. Geological Sur- cene Sea Otters. Jour. Fish. Res. Bd. Canada vey, Menlo Park, provided the excellent photographs 23(12):1897-1911. of the mandible, and Shirley M. Kilmer typed and edited the manuscript. Repenning, C. A. 1967. Palearctic-Nearctic Mam malian Dispersal in the Late Cenozoic. Pp. 288 311, in The Bering Land Bridge. (D. M. Hopkins, LITERATURE CITED ed.), Stanford Univ. Press, Stanford. Fisher, E. M. 1941. Notes on the teeth of the sea Ogle, B. A. 1953. Geology of the Eel River Valley otter. Jour. Mamm., 22:428-434. area, Humboldt County, California. California Div. Mines, Bull. 164, 128 pp. Fitch, J. E. 1970. Fish remains, mostly otoliths and teeth from the Palos Verdes Sand (Late Pleisto- Accepted for publication October 26, 1972. RESEARCH NOTES FIRST RECORD OF THE GIANT SEADEVIL Ceratias holbolli FROM CALIFORNIA, BASED UPON A SPECIMEN FROM THE STOMACH OF A SPERM WHALE A giant seadevil Ceratias holbolli Kr@yer, 1845, the largest of the deep sea anglerfishes, was recovered from the stomach of a 13.0 m male sperm whale Physeter catodon that was killed on 13 November 1970, at lat. 37°SO’N, long. 123°48’W (115 km west of San Francisco). This position is over the outer edge of the continental slope where the water is about 3,500m deep. The whale was taken by the catcher boat ALLEN CODY, Captain Douglas Campbell, of the Del Monte Fishing Company’s whaling station in Richmond, California. Bernard Lenheim, then with the National Marine Fisheries Service (NMFS), examined the stomach of the whale (Field No. 1970-53). Elbert H. Ahlstrom of the NMFS, John E. Fitch of the California Department of Fish and Game, and Robert J. Lavenberg of the Museum of Natural History of Los Angeles County identified the anglerfish, which is deposited in the collection of the latter institute (LACM 31841-1). Although the giant seadevil is virtually cosmo- politan, it has not previously been collected nearer to California than Cocos Island, the Galapagos Islands, the Hawaiian Islands, and Japan (Clarke, 1950). The few metamorphosed specimens that have been collected were caught at depths of 120 to 4390m, but most catches shallower than 600m were made in Arctic waters (Bertelsen, 1951; Clarke, 1950; Grey, 1956). The species was originally described by Krgyer (1845) on the basis of a specimen collected off Greenland. This seadevil was the only one that I and my assistants found in the stomachs of 552 sperm whales taken off central California that we examined from 1959 through 1970. Sperm whales are deep divers (they are known to descend as deep as 1,134m) and feed mostly on large squids, but they also eat many large demersal and mesopelagic fishes. Ceratias holbolli has previously been reported from the stomachs of sperm whales taken in the Antarctic at lat. 61°S, long. 103°E (Clarke, 1950), off Western Australia (Bannister, 1970), near the Falkland Islands (Korabel’nikov, 1959), off South Africa (Penrith, 1967), and around the Azores (Clarke, 1956). Although the specimen is partially digested, it is essentially entire and much of its skin is intact: the illicitum is present, but the esca is missing. The total length of the fish is about 63cm excluding the caudal rays, which are missing. When whole, it was probably about 75 or 80cm long. The maximum length reported for this species is 120cm (Bertelsen, 1951). is doubtless a female; the dwarf males of this species are parasitic on the females. The specimen The vernacular name “giant seadevil” was sug- gested by Carl L. Hubbs, Scripps Institution of Oceanography, and will be used in a forthcoming check list of California fishes. In the spelling of the specific name and the date of its original publication, I have followed the advice of W. I. Follett, California Academy of Sciences. LITERATURE CITED Bannister, J. L. 1970. The biology and status of the sperm whale off Western Australia—an ex- tended summary of results of recent work. Unpublished report presented to International Whaling Commission, Scientific Committee, Special Meeting on Sperm Whale Biology and Stock Assessments, Honolulu, Hawaii, 13-24 March 1970: 10 pp., 22 figs., 29 tables. Bertelsen, E. 1951. The Ceratioid fishes. Dana Rept., 39:1-276. Clarke, R. 1950. The bathypelagic angler fish Ceratias holbélli Kréyer. Discovery Rept., 26: 1-32. 1956. Sperm whales of the Azores. Dis- covery Rept., 28:237-298. Grey, M. 1956. The distribution of fishes found below a depth of 2000 meters. Fieldiana: Zool. Ser., 36(2):73—337. Korabel’nikov, L. V. 1959. O pitanii kashalotov v Antarkticheskikh moryakh. Priroda, 48(3): 103-104. Krgyer, H. 1945. Ichthyologiske Bidrag. Naturhist. Tidsskr., n.s., 1:639-649. Penrith, M. J. 1967. South Africa. Ceratioid angler-fishes from J. Nat. Hist., 1967(1):185—-188. DaLe W. Rice, National Marine Fisheries Service, Southwest Fisheries Center, P.O. Box 271, La Jolla, California 92037. Accepted for publication May 24, 1972. 158 1972 PLIOCERCUS EURYZONUS COPE: AN ADDITION TO THE SNAKE FAUNA OF HONDURAS Studies in progress on the snake fauna of Honduras revealed the presence of a specimen of Pliocercis euryzonus (MCZ 22911) from Tela, Depto. Atlantida, misidentified as Tropidodipsas sartorii. This specimen represents the first record of the species from Hon- duras, even though it is known to occur north and south of this country. The species is now known to inhabit low, moderate, and intermediate elevations along the Atlantic versant from central Guatemala to Brazil (Peters and Orejas-Miranda, Bull. U.S. Nat. Mus., 297:1-347, 1970). It seems likely that the nominal species Pliocercus annellatus Taylor, P. aru- bricus Taylor, and P. dimidiatus Cope are based on specimens of Pliocercus euryzonus (fide Scott, Ph.D. dissertation, Univ. Southern California, 1969, by in- ference). The Honduran specimen, a female, has 25 black bands on the body separated by light (red in life ?) interspaces one and one-half to two scales in length. The head is black to the middle of the parietals except for a light spot on the first and second supralabials. This black cap is followed by a light nuchal band, which extends to the posterior tip of the parietals. There are 134 ventrals, a divided anal plate, and 1 + 1 temporals, in which features P. euryzonus is easily distinguishable from TJ. sartorii, which has over 165 ventrals, a single anal plate, and usually 1 + 2 tem- porals. The tail of MCZ 22911 is incomplete. The locality from which this specimen came lies within the Tropical Moist Forest formation or tropical lowland rainforest. The specimen corresponds to the diagnosis for P. e. aequalis; a subspecies which is stated by Stuart (Misc. Publ., Mus. Zool., Univ. Michigan, 122:1—150, 1963) and Peters and Orejas-Miranda (1970) to occur only in central Guatemala. However, it probably ranges south at least to Costa Rica; the specimens upon which the above-mentioned probable synonyms of P. euryzonus are based came from this latter country. Larry Davip WiLson, Miami Dade Jr. College, Div. Interdisciplinary Studies, South Campus, Miami, Flor- ida 33156, and Diana D. Duaas, 505 S. Kibbe, Erath, Louisiana 70533. Accepted for publication September 1, 1972. A NEW SPECIES OF MICROTROMBICULA (ACARINA: TROMBICULIDAE) FROM HIDALGO, MEXICO Studies of the New World Microtrombicula Ewing by Vercammen-Grandjean (1965), Webb and Loomis RESEARCH NOTES (1970, 1971a, b, and c), and Loomis anc 1 Webb (19 have resulted in the recognition of three subgener 26 species. This new species from Hidalgo, México tentatively placed in the subgenus Scapuscutala, i special interest because of its extremely small size and the absence of certain nude leg setae characterist of other species of Microtrombicula Microtrombicula jacalae, new species Figure | Types.—Larvae: km NE Jacala, Holotype and 21 paratypes from 13 1585 m, Hidalgo, México, from ear of five Peromyscus boylii levipes Merriam, original numbers EMF 70-329 (holotype and 4 paratypes), EMF 70-324 (3), EMF 70-325 (1), EMF 70-327 (2), lacala jacalae, new species. A Ventral Figure 1. Microtrombicula Scutum. B. Dorsal aspect of gnathosoma. C. aspect of palpal tibia and tarsus. D. Setae: S seta; H, humeral; PD, posterior dorsal. E showing nude setae with measurements an F. Leg I. as above branched setae. above. 160 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES and EMF 70-328 (10) taken 30 June 1970, and B69989 (1) from Peromyscus sp. taken 2 September 1965, all collected by Eric M. Fisher. The holotype and one paratype will be deposited in the collection of the Rocky Mountain Laboratory, Hamilton, Mon- tana. Other paratypes now in the chigger research collection at California State University, Long Beach will be sent to appropriate institutions. Diagnosis: Genualae I and II, tibiala IIT and pre- tarsala II absent; eyes absent; | genuala I; coxa III with 5-7 branched setae; coxa II with 2 setae, 1 branched and 1 nearly nude; three pairs of sternal setae; palpotibial claw bifurcate; tarsus HI with 13 branched setae (mastitarsala absent); galeala nude. Description of holotype (measurements in microns, with mean and range of 19 types in parentheses, unless otherwise noted).—Body engorged, 175 by 115; eyes absent. Dorsal setae 2 (humerals)-6-6-6-4-4, total 28; mea- surements of humeral seta 17, anterior dorsal seta 19, posterior dorsal seta 22. Ventral setae 2-2-2 (sternals)-4-2-2-4-4-6-2-2, total 32; measurements of anterior sternal seta 16, anterior ventral seta 15, posterior ventral seta 19. Scutum: Subpentagonal; sensilla extremely short with 8—10 long setules along shaft. Scutal measurements: AW 23 (24.3, 22-26; 18), PW 35 (35.5, 32-38), SB 11 (11.2, 10-12), ASB 20 (20.6, 19-23; 18), PSB 13 (13.9, 12-16), AP 25 (24.4, 22-26), AM 16 (15.6, 14-17; 16), AL 14 (14.8, 14-19; 18), PL 22 (20.1, 19-22; 16), S 12 (12.7, 10-15; 7). Gnathosoma: Cheliceral blade small with small tricuspid cap; cheliceral base and capitular sternum lightly punctate. Galeala nude. Palpal setae B/B/ BBB; palpal tarsus with one nude and five branched setae, tarsala 4; palpotibial claw bifurcate with dorsal prong slightly shorter than the recurved ventral prong. Legs: Segments 7-7-7, extremely short and com- pressed, each leg ending in 2 claws and clawlike empodium without onychotriches. Leg I with genuala 7 and tarsala 9 (8.2, 7-9; 18), pretarsala, subterminala and parasubterminala; leg II with coxa bisetose, 2 tibialae and tarsala 10 (8.9, 8-10), pretarsala absent; leg III with coxa multisetose (S—7 branched setae) and lacking nude setae. Leg measurements (including, in parentheses, the mean and range of five types): I 36 (125, 116-136); II 114 (107, 92-114); III 131 (130, 117-131); total 381 (358, 325-381). Taxonomic remarks.—This species is tentatively placed into the subgenus Scapuscutala, because of the general similarities to the other species. Also the absence of certain nude leg setae may be correlated with the extremely short leg segments. The two setae on coxa II also have been recorded for two other species of Microtrombicula from West Pakistan (Traub and Nadchatram, 1966) and a larva of M. trisetica from Durango, México (Webb and Loomis, 1971c). VOLUME 71 Ecological notes—Of 95 traps set along paths in an oak woodland among or adjacent to limestone outcroppings, five of six trapped Peromyscus boylii had M. jacalae and a few trombiculid larvae of the genera Pseudoschoengastia and Leptotrombidium. ACKNOWLEDGMENTS We extend our sincere appreciation to Sr. Dr. Rudolfo Hernandez Corzo, Director General, Direccion de Caza, Departamento de Conservacion de la Fauna Silvestre, Secretaria de Agricultura y Ganaderia for permission to collect specimens in México. In addi- tion, we thank Eric M. Fisher for collecting this new chigger and its hosts, and Janice F. Fisher for the fine illustrations. Studies upon which this report is based were supported by a research grant AI-03407 from the National Institute of Allergy and Infectious Diseases. LITERATURE CITED Loomis, R. B., and J. P. Webb, Jr., 1972. A new intranasal chigger of the subgenus Crypticula, genus Microtrombicula (Acarina: Trombiculi- dae) from Texas. Bull. So. California Acad. Sci., 70(2):102-103. Traub, R., and M. Nadchatram. 1966. Notes on chiggers of the genus Microtrombicula Ewing, 1950 (Acarina: Trombiculidae) from Pakistan and India, with descriptions of five new species. J. Med. Ent., 3(3—4) :305-321. Vercammen-Grandjean, P. H. 1965. Revision of the genera Eltonella Audy, 1956 and Microtrom- bicula Ewing, 1950, with descriptions of fifty new species and transferal of subgenus Chiroptella to genus Leptotrombidium (Acarina: Trombiculi- dae). Acarologia, 7(fasc. suppl.) :34—257. Webb, J. P., Jr., and R. B. Loomis. 1970. A new subgenus of intranasal chiggers of the genus Microtrombicula from North America and Korea. J. Med. Ent. 7(6) :655-663. 197 1a. Four species of Microtrombicula (Acarina: Trombiculidae) from Mexico and Nicaragua. Bull. So. California Acad. Sci., 69 (3-4): 133-144. —. 1971b. Trombiculid mites of the genus Microtrombicula (Acarina) from Costa Rica. Los Angeles Co. Mus. Contrib. Sci., (207):1-15. 1971c. The subgenus Scapuscutata of the genus Microtrombicula (Acarina: Trombiculidae) 1972 from North America. J. Med. Ent., 8(3):319 329, Ricuarp B. Loomis, Dept. Biology, California State University, Long Beach, California 90840, and JAMES RESEARCH NOTES P. Webs, Jn, The Los Whittier 1000 Ave., South El Monte, California 9173 Angeles County Nature t Narrows Nature Center Accepted for publication October 15, 19° EDITORIAL: CRITICAL REVIEW OF MANUSCRIPTS As indicated in the Instructions for Authors (in- side back cover), all .manuscripts submitted to SCAS BULLETIN are reviewed by referees who critically read for scientific content, originality, and clarity of presentation, and who assist the editors in making judgments as to acceptability for publication. The reviewers also assist authors by offering constructive suggestions which may lead to improvement of the text or illustrations of manuscripts. By contribution of their time and professional expertise the referees measurably en- hance the quality of SCAS BULLETIN. In behalf of our readers, authors, and ourselves we thank the following reviewers for valuable editorial assistance in recent months: Philip A. ANNOUNCEMENT: The 1973 annual meetings of the Southern California Academy of Sciences will be held on May 4 and 5, 1973 at California State University, Long Beach. Technical papers are solicited from the fields of natu- ral and social sciences. Abstracts of no more than 150 words must be typed on 3 X 5 cards and must be received no later than March 26, 1973. The first line of the abstract is to include author(s), student or professional, pre- ferred section (see below), and type of projection equipment, if needed. Send abstracts to: Donald R. of Los Angeles California Patten, Natural History Museum County, Exposition Park, Los Angeles, 90007. Adams, Eric D. Austin, James A. Blake, Bayard H. Brattstrom, Jules Crane, Jr., David Cummings, Murray D. Dailey, Kristian Fauchald, George F. Fisler, John E. Fitch, William Hand, John William Hardy, Charles Lambert, Robert J. Lavenberg, Richard B. Loomis, John W. Mcmenamin, William J. Morris, Martin L. Morton, I. M. Newell, Don- ald R. Patten, Donald J. Reish, Elbert Sleeper, Andrew Starrett, John S. Stephens, Stuart L. War- ter, Joel Weintraub, Harrington Wells, Dieter H. Wilken, Adrian M. Wenner. Patrick H. WELLS, Technical Editor JAMES DALE SMITH, Managing Editor 1973 ANNUAL MEETINGS Technical sessions—Anthropology, Archaeology. Botany, Earth Science, Entomology, Experimental Biology, Folklore, History, Invertebrate Zoology, Ma- rine Science, and Vertebrate Zoology (other sections will be opened to accommodate demand). Student awards will be presented in the Natural Science and Social Science Divisions. First award in each division will be $150.00: second award in each division $75.00. A.A.A.S. Grant in Aid of Researeh—S$150.00 will be awarded to a high school, undergraduate. or uate student submitting the most outstandi proposal in the sciences. Application fo from Takashi Hoshizaki, Depart U.C.L.A., Los Angeles, California INDEX TO VOLUME 70 Adams, P. A.: Variation and geographic distribution in some Argentine and Chilean Osmylidae, with a new species of Kempynus (Neuroptera), 45-49 Algae, Intertidal species, southern California, 2—16 Amastridium veliferum, 53-54 ; Amazon basin, New Hunterotrema from dolphin, 79-80 Anolis humilis, 107; Anolis Anolis sminthus, 108-109 Aphelocheirus pallens, 69 Arctic, Taxonomy and distribution of copepods, 23-30 Argentina, Distribution and variation in Osmylidae, 107-108; laeviventris, 45-49 Ascidians, Distribution in southern California, 114— 124 Bats, Methods of measurement, 51-52; New sub- species of Natalus stramineus, 81-84 Bee, New synonymy in Chilicola, 52 Behavior, Aggregation in milkweed bug, 87—90 Blake, J. A., and K. H. Woodwick: New species of Polydora (Polychaeta: Spionidae) from the coast of California, 72—79 Blake, J. A., and K. H. Woodwick: A review of the genus Boccardia Carazzi (Polychaeta: Spionidae ) with descriptions of two new species, 31—42 Boccardia ligerica, 31-39 Bopyrids, From New Guinea, 99-102 Bossiella spp., 8 Bothrops bicolor, 110 Brennan, J. M., and F. Lukoschus: Parasitic mites of Surinam. VIII. A new genus and species of chigger, Fauranius atecmartus, and additional records of species (Acarina: Trombiculidae), 42-45 Caldwell, D. K., W. F. Rathjen, and M. C. Caldwell: Cuvier’s Beaked Whale, Ziphius cavirostris, from Barbados, 52-53 Caldwell, M. C., see Caldwell, D. K. California, New species of Polydora, 72-79 Central America, Recent arcoid bivalve, 131-135 Central California, Boccardia berkeleyorum, new spe- cies, 31 Chigger, New genus and species, 42-45; New species on mouse, 102-103 Chile, Boccardia chilensis, new species, 31; Variation and distribution in Osmylidae, 45—49 Clam, Asiatic species in San Diego County, 91—98 Copepods, Taxonomy and distribution of Arctic spe- cies, 23-30 Corallina officinalis var. chilensis/pinnatifolia, 8; Corallina vancouverensis/gracilis, 8 Corbicula manilensis, 91-98 Costa Rica, Colubrid snake, 53-54; Simopelta pae- minosa, 16-17 Crab, New Iphitime, 84-87 Crotiscus desdentatus, 43 Csuti, B. A.: Distribution of some southern Cali- fornia kangaroo rats, 50-51 Dailey, M. D.: A new species of Hunterotrema (Digenea: Campulidae) from the Amazon River Dolphin (/nia geoffrensis), 79-80 Danforth, C. G.: Two bopyrids (Isopoda) from New Guinea, 99-102 Dolphin, lung trematodes, 79-80 Dryadophis dorsalis, 110-111 Egregia laevigata, 8 Eleutherodactylus merendonensis, 106 Emerson, R. R., and K. Fauchald: A revision of the genus Loandalia Monro with description of a new genus and species of pilargiid polychaete, 18-22 Enterogona, 114-124 Eocene, Benthonic Foraminifera in San Diego, 125— 130 Eutrombicula alfreddugesi, 44; Eutrombicula batatas, 44; Eutrombicula goeldii, 44; Eutrombicula tin- ami, 44 Fast, A. W.: The invasion and distribution of the Asiatic Clam (Corbicula manilensis) in a south- ern California reservoir, 91—98 Fauchald, K., see Emerson, R. R. Fauranius myoproctae, 43 Fay, R. C., and J. V. Johnson: Observations on the distribution and ecology of the littoral ascidians of the mainland coast of southern California, 114-124 Foraminifera, Benthonic assemblage in San Diego, 125-130 Formicidae, Simopelta paeminosa, new species, 16-17 Gastroclonium coulteri, 8 Gibson, J. M.: Benthonic Foraminifera of the Ardath Shale and Stadium Conglomerate (Eocene), San Diego Basin, California, 125-130 Gigartina armata/spinosa, 8; Gigartina canaliculata, 8; Gigartina leptorhynchos, 8 Honduras, Taxonomy of amphibians and reptiles, 106-114 India, New species of Aphelocheirus, 69-72 Johnson, J. V., see Fay, R. C. Kangaroo rats, Distribution of Dipodomys in southern California, 50-51 Kempynus falcatus, 45, 48 162 1972 Key to: species of Boccardia, 40; Genera of Pilargi- idae, 18; species of Parandalia, 19 Kutcher, S. R.: Two types of aggregation grouping in the Large Milkweed Bug, Oncopeltus fasciatus (Hemiptera: Lygacidae), 87-90 ais Lacewing, Variation, distribution, and new species, 45-49 Lampropeltis triangulum, 111 La Rivers, I.: Descriptions and notes concerning some Oriental Aphelocheirus (Hemiptera: Aphelochei- ridae), 69-72 Larvae, Pagurus samuelis reared in laboratory, 58-68 Linares, O. J.: A new subspecies of Funnel-eared Bat (Natalus stramineus) from Western Venezuela, 81-84 Loandalia aberrans, 21-22 Loomis, R. B., and J. P. Webb, Jr.: A new intranasal chigger of the subgenus Crypticula, genus Mi- crotrombicula (Acarina: Trombiculidae) from Texas, 102—103 Lucicutia anomala, 23-29; Lucicutia polaris, 23-29; Lucicutia pseudopolaris, 23-29 Lukoschus, F., see Brennan, J. M. MacMillan, F. E.: The larvae of Pagurus samuelis (Decapoda: Anomura) reared in the laboratory, 58-68 Meyer, J. R., and L. D. Wilson: Taxonomic studies and notes on some Honduran amphibians and reptiles, 106—114 Microtrombicula fragibarba, 44; Microtrombicula nr. tragulata, 44 Mouse, New chigger from Texas, 102—103 Nasicola annereauxi, 44 New Guinea, Bopyrids, 99-102; New localities of Aphelocheirus, 69 New taxa: Aphelocheirus nathani, 70-72; A phelochei- rus pygmaeus, 69-70; Boccardia berkeleyorum, 31-41; Boccardia chilensis, 31-41; atecmartus, 42-43; Hunterotrema macrosoma, 79-80; Iphitime holobranchiata, 86-87; Kempy- nus crenatus, 45—48; Microtrombicula welbourni, 102-103; Natalus stramineus tronchonii, 81-83; Parandalia, 18—22; Parandalia americana, 18-19; Parandalia fauveli, 18-19; Parandalia gracilis, 18-19; Parandalia indica, 18-19; Parandalia ocu- laris, 18-21; Polydora bioccipitalis, 75-77; Poly- dora convexa, 73-74; Polydora elegantissima, 74-75; Polydora_ pygidialis, 77-78; Pseudione novaeguineensis, 99-101; Simopelta paeminosa, 16-17 Ninia atrata, 111 Noetia magna, 131-135; Noetia reversa, 131-135 Fauranius Odontacarus tubercularis, 44 Oncopeltus fasciatus, 87-90 INDEX 163 Pachydictyon coriaceum, & Pagurus samuclis, 58-68% Parasecia aitkeni, 44; Parasecia manueli, 44; Parasecia valida, 44 Parathelees weberi, 99 Patten, D. R.: An measuring metacarpal length in Artibeus lituratus (Olfers) (Chiroptera: Phyllostomatidae), 51-52 Pelvetia fastigiata, 8 Perissopalla barticonycteris, 44; Perissopalla ipeani, 44 Phyllospadix spp., 8 Pilger, J.: A new species of /phitime (Polychaeta) from Cancer antennarius (Crustacea: Decapoda ), 84-87 Pleurogona, 114—124 Pliocene, Noctia from Peru, 131-135 Polychaeta, New genus from Santa Barbara, 18-22; New species of /phitime, 84-87; New species of Polydora, 72-79; Review of genus Boccardia, 31-42; Revision of genus Loandalia, 18-22 Pterocladia pyramidale, 8 evaluation of two methods of Quadreseta flochi, 44 Ralfsia spp., 8 Rathjen, W. F., see Caldwell, D. K. Rhadinaea kinkelini, 111 Rhodoglossum affine, 8 Robinson, D. C., see Wilson, L. D. Roth, B.: A second Recent species of Noetia, sensu stricto (Mollusca: Bivalvia) in the tropical east- ern Pacific Ocean, 131-135 San Diego, Benthonic Foraminifera, 125-130; In- vasion of Asiatic clam, 91-98; New genus of polychaete, 18—22 Sibon anthracops, 111-112; Sibon carri, 112 Snake, Colubrid specimens from Costa Rica, 53-54 Snelling, R. R.: A new species of Simopelta from Costa Rica (Hymenoptera: Formicidae), 16-17; A new synonymy in the genus Chilicola (Hy- menoptera: Colletidae), 52 Southern California, Distribution of kangaroo rats, 50-51; Distribution of littoral ascidians, 114-124: Intertidal algae, 2-16 Speleocola secunda, 44 Spliaerodactylus dunni, 109-110 Surinam, New genus and species of chigger, 42-45 Tecomatlana vesperuginis, 45 Texas, New chigger on mouse, 102-103 Thamnophis cyrtopsis, 112 Ulva, sp., 8 Venezuela, New subspecies of bat, 81-84 Vidal, J.: Taxonomy and distribution of the Arctic 164 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 71 species of Lucicutia (Copepoda: Calanoida), 23-30 Webb, J. P.. Jr., see Loomis, R. B. Whale, Ziphius cavirostris from Barbados, 52—53 Widdowson, T. B.: Changes in the intertidal algal flora of the Los Angeles area since the survey by E. Yale Dawson in 1956-1959, 2-16 Wilson, L. D., and D. C. Robinson: Additional speci- mens of the colubrid snake Ammastridium veli- ferum Cope from Costa Rica, with comments on a pseudohermaphrodite, 53—54 Wilson, L. D., see Meyer, J. R. Woodwick, K. H., see Blake, J. A., 31-42; 72-79 Ziphius cavirostris, 52-53 INSTRUCTIONS FOR AUTHORS The BULLETIN is published three times cach year (April, August, and November) and includes articles in English in any field of science. Non-members will be assessed a page charge of $40.00 per page. Manuscripts submitted for publication should contain results of original research, embrace sound principles of scientific investigation, and present data in a clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide for contributors. Consult also recent issues of the BULLETIN. 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Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He should promptly and carefully read the proof sheets for errors and omissions in text, tables. illus S legends, and bibliographical references. He marks corrections on the galley (copy editing and proof dures in Style Manual) and promptly returns both galley and manuscript to the managing edito Smith, Department of Biology, California State University, Fullerton, California 92634. Manuscripts nal illustrations will not be returned unless requested at this time. Printing charges accruing from exc tions to, or changes in, the proofs must be assumed by the author. Reprint orders are placed wit not the Managing Editor. CONTENTS Ornithophily and extrafloral color patterns in Columnea florida Morton (Gesneriaceae). By C. BugenenJOoness Ure and sea Vickers RICH fry cscieicrise aisle: statelote eet clench elelel selects Cie ae eae 113 Effects of varying temperatures and salinities on settlement, growth, and reproduction of the wood-boring pelecypod, Lyrodus pedicellatus. By Kevin J. Eckelbarger and Donald J. Reish 116 Two new species of polychaetous annelid worms from Baffin Bay and the Davis Strait. By ames PAs (BIAKC. i jk. whecs cree eke ol eiecewe etehe late ch se ah be lelenoyay he chester halve eee ee 127 Fluctuations in population density of the Hispid Cotton Rat: factors influencing a “crash.” By Eugene D. Fleharty, Jerry R. Choate, and Michael A. Mares ............2-ceueesuees 132 The coral snake Micrurus nigrocinctus in Honduras (Serpentes: Elapidae). By Larry David Wilson ‘and JonntReMeyer oision ose dec 5 oe Gos tee oo eer ohe le eel alee ere 139 A new species of rock dwelling dendrochirote holothurian from Catalina Island. By Josephine YUCK LAN BS ti Gee are: SRE ol ark eodia a lsavs anoriale tee a sale tenn las niet Galaud jerdlenin Yh le eo COL 145 A new species of sea otter from the late Pleistocene of northwestern California. By Frank H. TO, a ec ERE Toa ers eet Cit ANN MERE OIM G Gido pc oD oo cco u00 150 RESEARCH NOTES First record of the giant seadevil Ceratias holbolli from California, based upon a specimen from the stomach’ ofa-Sperm Whales By DalesW. (Rice ).00) Yous deine sec eee ee 158 Pliocercus euryzonus Cope: An addition to the snake fauna of Honduras. By Larry David Wil- Sonand DidnasD Dugas i soi esse yale oo sn OEE HO ee ONE ESATO RO Oe 159 A new species of Microtrombicula (Acarina: Trombiculidae) from Hidalgo, México. By Rich- ard ’B. Eoomisiand Jamessb., Webby Ii iene ce eke: eee ee eka Ae eee 159 Editorial critical review. ‘of manuscripts! jan acee enters ee cries oe eee 161 Announcement: 1973annualsmectings jars aclsins cysts enero eee oe Ee 161 Indextor Volumen7 OW west eur siesta fcistostetvateorsralaioustel ateteicestne sla cuehreiceine StU oe ee eee 162 cover: A Little Hermit (Phaethornis longuemareus) hovering above two red spots on the upper surface near the apex of the leaves of Columnea florida (Gesneriaceae). This drawing depicts a proposed new type of hummingbird pollination system involving an extrafloral attraction mechanism in which the attracting unit, in this case the leaf, does not simply simulate a portion of the flower. Illustration by Susan E. Payne, California State University, Fullerton. INDEX TO VOLUME 7] | Abati, J. L., and D. J. Reish: The effects of lowered dissolved oxygen concentrations and salinity on the free amino acid pool of the polychactous annelid Neanthes arenaceodentata, 32-39 Abbott, D. P., and J. VY. Johnson: The ascidians Styela barnharti, S. plicata, S. clava, and S. montereyensis in Californian waters, 95-105 Acarina, New Microtrombicula, 159-161 Allen, R. K., and E. S. M. Chao: A new species of Baetodes from Arizona (Ephemeroptera: Baeti- dae), 52 Anomura, Range extensions, 56 Aphanopus carbo, 12-13: Arizona, New ambush bug, 93-95; New Bactodes, 52 Ascidians, Reexamination of Californian Styela, 95— 105 Baffin Bay, New Polychaeta, 127—132 Baker, R. J., H. H. Genoways, and A. Cadena: The phyllostomatid bat, Vampyressa brocki, in Colom- bia, 54 Bat, Vampyressa brocki, 54 Behavior, Effects of temperatures and salinities on pelecypod, 116-127; Hummingbird pollination, 113-116; Population density in rat, 132-138; Reproduction in Teredinidae, 48-50; Social wasps, 53 Belman, B. W., see Fauchald, K. Birnie, D., and J. D. Smith: Bibliography of John Adams Comstock, 1883-1970, 60-68 Blake, J. A.: Two new species of polychaetous anne- lid worms from Baffin Bay and the Davis Strait, 127-132 Brattstrom, B. H.: Temperature changes in heat pro- ducing plants, 54-55 Brusca, R. C., and J. Haig: Range extensions of Porcelain and Hermit Crabs in the Gulf of California, 56 Cadena, A., see Baker, R. J. California, Ascidians, 95-105; First record of Giant Seadevil, 158; Key to Gobiidae, 108-112; New fossil otter, 150-157; Notophycid polychaete, 107-108; New holothurian, 145-150 Ceratias holbolli, 158 Cercariae, In Corithidia californica, 39-43 Chao, E. S. M., see Allen, R. K. Choate, J. R., see Fleharty, E. D. Coleoptera, Curcolionidae, Brachyrhininae, Peritelini, 80-91; Scarabaeidae, 69-80 Colombia, Phyllostomatid bat, 54 Columnea florida Morton, 113-116 Comstock, John Adams, Biography, 57—60: bibliogra- phy, 60-68 Crabs, Range extensions of porcelain and hermit crabs, 56 ty an Diptera, New species of asilids, 43—47 Dugas, D. D., see Wilson, L. D Eckelbarger, K. J., and D. J. Reish: A first report of self-fertilization in the wood-boring Family Teredinidae (Mollusca: Bivalvia), 48-50; Ef fects of varying temperatures and salinities on settlement, growth, and reproduction of the wood boring peleypod, Lyrodus pedicellatus, 116-127 Ephemeroptera, New Baetodes, 52 Eucyllus Horn, Review, 80-91 Eupsophus, Review and definitions, 2—11 Fauchald, K., and B. W. Belman: A_ notophycid polychaete from California, 107-108 Fitch, J. E., and D. W. Gotshall: First record of the black scabbardfish, Aphanopus carbo, from the Pacific Ocean with notes on other Californian trichiurid fishes, 12—18 Fleharty, E. D., J. R. Choate, and M. A. Mares: Fluctuations in population density of the hispid cotton rat: Factors influencing a “crash,” 132- 138 Frog, Generic review of Eupsophus, 2-11 Gastropoda, New species, 106—107 Genoways, H. H., see Baker, R. J. Gibo, D. L.: An introduced population of social wasps, Polistes apachus, that has persisted for ten years (Hymenoptera: Vespidae), 53 Gotshall, D. W., see Fitch, J. E. Haig, J., see Brusca, R. C. Hemiptera, Aradidae, 91-93; Phymatidae, 93-95 Holothurian, New dendrochirote from Catalina Island, 145-150 Honduras, Riiadinaea godmani, 50-52: Coral snake. 139-145; New snake, 159 Hovore, F. T.: Three new sympatric Pleocoma from the southern Sierra Nevada mountains of Cali- fornia (Coleoptera: Scarabaeidae), 69-80 Hymenoptera, Persistent population of Polistes apachus, 53 Indian Ocean, New Pugnus, 106-107 Johnson, J. V., see Abbott, D. P. Jones, C. E. Jr., and P. V. Rich: Ornithophily extra floral color patterns in Coluninea flo Morton (Gesneriaceae), 113-116 Key to: Californian trichiurids, 17; Cercariae of 18 Californian trematodes, 39-42: spe r WwW coxia, 46: subspecies of Pleocoma / species of Eucyllus, 83: species of Californ gobiids, 108-112: species of Cucuriari 56 BULLETIN SOUTHERN CALIFORNIA Kilmer, F. H.: Late Pleistocene of 150-157 Kormilevy, N. A.: Two new species of North Ameri- can flat bugs (Hemiptera: Aracidae), 91-93; A new species of ambush bug from Arizona (Hemiptera: Phymatidae), 93-95 A new species of sea otter from the Northwestern California, Loomis, R. B., and J. P. Webb, Jr.: A new species of Microtrombicula (Acarina: Trombiculidae) from Hidalgo, México, 159-161 Lynch, J. D.: Generic partitioning of the South Amer- ican leptodactylid frog genus Eupsophus Fitzinger, 1843 (Sensu lato), 2-11 Lyrodus pedicellatus, 116-127 MacDonald, C. K.: A key to the fishes of the family Gobiidae (Teleostomi) of California, 108-112 Mares, M. A.: see Fleharty, E. D. Martin, L. M., John Adams Comstock, 57-60 Martin, W. E.: An annotated key to the cercariae that develop in the snail Cerithidea californica, 39-43 Metabolism, Polychaetous annelids, 32—39 Mexico, New Microtrombicula from Hidalgo, 159-161 Meyer, J. R., see Wilson, L. D. Micrurus nigrocinctus, 139-145 Mollusca, Fertilization in Teredinidae, 48-50 1883-1970, Neanthes arenaceodentata, Metabolism, 32-39 New taxa: Aradus nevadensis, 91-92; Baetodes sigil- latus, 52; Cryptosclerocheilus baffinensis, new genus and species 131; Cucumaria salma, 145-150; En- hydra macrodonta, 151-157; Eucyllus carinarostris, 89-90; Eucyllus cinereus, 89; Eucyllus saesariatus, 86-88; Lumbrineris fauchaldi, 128-131; Macro- cephalus similis, 94-95; Metapogon amargosae, 46— 47; Metapogon obispae, 47; Mezira tropicalis, 92-93; Microtrombicula jacalae, 159-161; Phyl- lodocella bodegae, new genus and species, 107—108; Pleocoma hirticollis reflexa, 75-78; Pleocoma mar- quai, 69-73; Pleocoma rubiginosa, 74-75; Pugnus maesae, 106-107; Wilcoxia martinorum, 43-44; Wilcoxia monae, 44-45; Wilcoxia painteri, 45; Wilcoxia pollinosa, 45—46 Ornithophily, Patterns in Columnea, 113-116 Otoliths, Trichiurids, 12-18 Pacific Ocean, First record of Black Scabbard Fish, 12-13 Pelsue, F. W., and E. L. Sleeper: A review of Eucyllus Horn (Coleoptera: Corcolionidae, Brachyrhininae, Peritelini), 80-91 ACADEMY OF SCIENCES VOLUME 72 Perrin, W. F., and E. L. Roberts: Organ weights of non-captive porpoise (Stenella spp.), 19-32 Pleistocene, New sea otter, 150-157 Pleocoma, New sympatric species, 69-80 Pliocercus euryzonus, 159 Polychaeta, Cryptosolerocheilus baffinensis, new genus and species, 127—132; Metabolism in Neanthes, 32— 39: Phyllodocella bodegae, new genus and species, 107-108 Porpoise, Organ weights, 19-32 Reish, D. J., see Abati, J. L.; see Eckelbarger, K. J. Rhadinaea godmani, 50-52 Rice, D. W.: First record of the giant seadevil Ceratias holbolli from California, based upon a specimen from the stomach of a Sperm Whale, 158 Rich, P. V., see Jones, C. E., Jr. Roberts, E. L., see Perrin, W. F. Rodents, Factors in population density of cotton rat, 132-138 Roth, B.: Keeling 106-107 A new species of Pugnus from Cocos- Islands, Indian Ocean (Gastropoda), Scalibregmidae, 128 Sea Cucumber, New species, 145—150 Sigmodon hispidus, 132-138 Sleeper, E. L., see Pelsue, F. W. Smith, J. D., see Birnie, D. Snake, Micrurus nigrocinctus in Honduras, 139-145; Pliocercus euryzonus in Honduras, 159; Rhadinaea godmani in Honduras, 50-52 South America, Review of Eupsophus, 2-11 Teredinids, Literature review of effects of tempera- ture and salinity, 118-121; Self-fertilization, 48-50 Vampyressa brocki, 54 Webb, J. P., Jr., see Loomis, R. B. Whale, Seadevil specimen from stomach, 158 Wilcox, J.: New species of robber flies of the genera Wilcoxia and Metapogon (Diptera: Asilidae), 43-47 Wilson, L. D., and D. D. Dugas: Pliocercus eury- zonus Cope: An addition to the snake fauna of Honduras, 159 Wilson, L. D., and J. R. Meyer: Rhadinaea godmani, an addition to the snake fauna of Honduras, 50-52; The coral snake Micrurus nigrocinctus in Honduras (Serpentes: Elapidae), 139-145 Yingst, J. Y.: A new species of rock dwelling den- drochirote holothurian from Catalina Island, 145— 150 F LIBRARY JUN 19 1973 NEW YORK “ BOTANICAL GARDEN DUODHERN CALIFORNIA ACADEMY OF SCIENCES BULLETIN Jolume 72 Number 1 ae CAS-A72(1) 1-56 (1973) APRIL 1973 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 OFFICERS Jules Crane, Jr., President Donald Bright, First Vice President Wayland D. Hand, Second Vice President Stuart L. Warter, Secretary Donald R. Patten, Treasurer Patrick H. Wells, Technical Editor James Dale Smith, Managing Editor Roberta S. Greenwood, Index Editor BOARD OF DIRECTORS 1971-1973 1972-1974 1973-1975 John J. Baird Wayland D. Hand Donald Bright Takashi Hoshizaki Donald R. Patten Timothy S. Hillebrand Elbert Sleeper Donald J. Reish William J. Morris David L. Walkington Andrew Starrett Richard E. Pieper Stuart L. Warter Patrick H. Wells David P. Whistler / 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. PATinualis Members eisai ence eee tis seca s rote Ha aliyciecispeled ater are be Siero ne Eee $ 8.00 StudenteMembers = tgrcierece earner aeciconectine aie cise eer oe ee SEO e ele ee Uae 5.00 Teifes Members) cisietsrteyar scree Sic s hoe os Ae ooo oa CSOT eee GIS ESS ra ae Oe 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. en eee ee BULLETIN OF THE VOLUME 72 JUNI SOUTHERN ACADEMY OF CALIFORNIA SCIENCES 1973 NUMBER 1] NEW TAXA OF BRACHYURAN CRABS FROM DEEP WATER OFF WESTERN PERU AND COSTA RICA JOHN S. GARTH! ABSTRACT: Five new species and one new genus of brachyuran crabs obtained by E. M. del Solar and E. Fernandez-B. are described and illustrated. Of these, Homolodromia robertsi, Acanthocarpus delsolari, Pilumnus fernandezi, and Trizocarcinus peruyianus are most closely re- lated to western Atlantic deep-water species, whereas, Delsolaria enriquei, a new genus and spe- cies, is a west American endemic. The family Homolodromiidae and the genera Homolodromia and Acanthocarpus are reported for the first time from the eastern Pacific. It is postulated that the analogous pairs of deep-water Amphi-american species have been separated since free communication existed at or near the depths at which they presently occur. The Bolivar Trench which extends across northern South America is suggested as the probable Miocene or earlier connection. Through the efforts of Enrique M. del Solar-C., technical adviser to IMARPE (Instituto del Mar del Perti), carcinological collections from a hitherto untapped source have lately found their way into museums in the United States and western Europe, among them the Allan Hancock Foundation (AHF), Los Angeles, the U.S. Na- tional Museum of Natural History (USNM), Washington, and the Royal Netherlands Museum (RNM), Leiden. These come from deep waters off Peru, and are the by-product of a search for edible shrimp and other fisheries products con- ducted by the research vessels “Wiracocha,” “Challua japic,” and ‘“‘SNP-1” under the direction of del Solar. Independent probing of deep waters off western Costa Rica by Hno. Eduardo Fernandez-B., Colegio de La Salle, San José, has similarly enriched the collections of the U.S. National Museum. Not only have a number of the brachyuran species proven to be new to science, but repre- sented among them are genera and families not previously known to inhabit the eastern Pacific. Four of the new species described herein have already been mentioned in Peruvian journals (del Solar, Blancas, and Mayta, 1970; del Solar, 1972) with identifications to genus only, and three have been illustrated (Chirichigno, 1970), but without specific names. Since the genera under which they were first reported may differ from the ones used here, and since the published illustrations are sometimes insufficient for species identifica- tion, citations to the above are given for clarifica- tion only, and are not intended to establish priority for the species in question, which dates solely from this publication. This paper is Allan Hancock Foundation Contribution No. 343. SYSTEMATIC ACCOUNT Family HOMOLODROMIIDAE Homolodromia robertsi, new species Figure 1, A-F Type: Female, holotype, AHF No. 719, from off Peru, lat. 03°48.5’ S, long. 81°18.4’ W. 800 m, 11 January 1971, mud bottom, E. M. del Solar, collector (Orig. No. B-260). Ovigerous female. paratype. USNM No. 141570, from off Pert, lat. 07°59" S, long. 80°22’ W, 800 m, 26 November 1971, hard bottom, Trawler “Challua japic,” E. M. del Solar, collector (Orig. No. B-376). Measurements (all in millimeters): Fem hol type: length, including rostrum, 42.5, w length of rostrum, 4.0, width of rostrum, 7.5, chelipe 1 Allan Hancock Foundation, University of South AANNT ern California, Los Angeles. California 9 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 1973 (coxa-ischium-merus, 35.4, carpus-propodus, 37.8), 73.2, chela, 28.0, dactyl, 13.5, height of palm, 7.2, walking legs, from first to last, 98, 90, 36, and 39. Diagnosis: Rostral and exorbital spines short and subequal, the latter more forwardly than outwardly directed. Median and lateral portions of cervical su- ture continuous. Carapace, chelipeds, and walking legs spinulous. Description: Carapace thick, swollen, pilose, sur- face spinulous, punctate, divided by cervical and branchial sutures into unequal thirds. Posterior branchial regions dilated, limited anteriorly by bran- chial furrow; anterior branchial regions similarly de- limited from gastric and hepatic regions by cervical furrow. Front with two stout, triangular horns reach- ing extremity of third article of antenna, outer margins sloping toward a similarsand forward-directed exor- bital spine. Rostral horns and outer orbital spines straight, tubular, the former flattened beneath, the latter spinulous externally. Eyestalks thick, short, cylindrical, two spinules at base; cornea terminal, of diameter equal to eyestalk, color yellowish brown. Supraorbital margin smooth, with a narrow raised border bearing one or two spinules. Lateral margins spinulous, constricted by the aforementioned grooves. Basal antennal article with concealed green-gland opening internally, three or four spinules externally, the outer two in line; second article with two spinules in line externally, a cluster of spinules anteroin- ternally, and a large spine anteroexternally; third (first flagellar) article with a subdistal spinule ex- ternally and two spinules in line internally. Outer maxilliped densely haired, especially along internal margin, merus broader than ischium, arcuate Outer margin spinulous; palp coarse, cylindrical. Chelipeds of female equal, pilose, margins of merus, carpus, and manus spinulous, prehensile margins of fingers dentate, dactylus fitting into a fork-tipped propodus. First two pairs of ambulatory legs long, slender, cylindrical, pilose, dactyli long and strongly curved. Last two pairs of legs short, elevated, chelate, chela of third leg directed backward, that of the fourth leg directed forward, dactyl fitting a fork formed by two or more propodal spines. Female abdomen with seven free segments, seventh segment cordiform, longer than preceding six. The male of the species is unknown. Remarks: Homolodromia robertsi differs from spinulous, the western Atlantic Homolodromia paradoxa A. Milne-Edwards, 1880, as follows: and exorbital spines, the latter more forwardly than outwardly directed: the median portion of shorter rostral NEW BRACHYURAN CRABS FROM EASTERN PACIFIC 3 the cervical suture continuous with the lateral por- tions; the carapace and the walking legs spinulous; and in having the legs of the last two pairs shorter, although this may be a character associated with the female sex. The new species differs from the East African Homolodromia houvieri Doflein, 1904, in having the carapace broader posteriorly, the furrows more clearly discernible, giving it a tightly laced appearance, the rostral and exorbital spines are. straight incurving (cf. Doflein, 1904, text-fig. 1), the eyestalks short and instead of thick, the cornea pigmented, the outer maxilliped thicker, the antennal peduncle and walking legs more spinulous, including the meri and propodi, which in H. bouvieri are without spinules. I take pleasure in naming this new species for Henry B. Roberts, Senior Museum Specialist, U.S. National Museum of Natural History, Washing- D.C., whose assistance in preparing its description is gratefully acknowledged. ton, Family CALAPPIDAE Acanthocarpus delsolari, new species Figure 2, A-F Mursia sp. Chirichigno, 1970, p. 41, text-fig. 85. Type: Holotype, female, AHF No. 703, from near Banco de Mancora, off Pert, lat. 03°43’ S, long. 81°03’ W, 250 m, mud bottom, 30 August 1970, E. M. del Solar, collector (Orig. No. B-204). Measurements (in millimeters): Female holotype: length, 25.3, breadth, 27.6, frontal width, 3.4, fronto- orbital width, 10.8, length of meral spine, 11.0, length of chela, 17.9, dactyl, 7.8, height of palm, 12.1. Over- all width, including meral spines, 60.5. Diagnosis: Carapace ovate, posterolateral spine of moderate length. Two meral spines, the inferior much the longer. Posterolateral margin non- tuberculate. A tooth on posterior margin and a conical tubercle on either side of sternal plastron. Tubercles of carapace arranged in longitudinal rows, most prominent posteriorly and laterally. Carapace regularly convex, opposite middle, surface uneven, protuberances ar- ranged in five longitudinal rows of which one is ridges Description: widest median, tubercles of lateral increasingly prominent posteriorly, tending to coalesce, the outer- most protruding beyond posterolateral margin, which Figure 1. Homolodromia robertsi, new species, female holotype: A, dorsal view: B, ventral view of frontal region; C, outer view of right chela; D, tip of left third walking leg: E, right outer maxilliped; F, abdomen. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 Figure 2. Acanthocarpus delsolari, new species, female holotype: A, dorsal view; B, frontal view, showing suborbital tubercles; C, outer view of right cheliped; D, inner view of same, showing stridulating ridge; E, abdomen and sternal plastron; F, left outer maxilliped. 1973 bears a moderately long, conical, more laterally than posteriorly directed spine. Surface covered with mi- nute granules and punctae. Front moderately wide, trilobate, separated from orbit by a shallow notch. Orbits large, margins hairy, superior fissure. ob- solescent. Anterolateral margin with four obscure tuberculiform teeth, followed by three smaller tuber- cles at widest part of carapace. Posterior margin arcuate, bearing a large tooth at middle and a smaller tooth on either side. Spine at outer angle of merus of cheliped approxi- mately two-fifths width of carapace; superior spine approximately one-fifth length of inferior. Hand with a seven-toothed crest above and an oblique, five- toothed crest on outer surface extending from base of dactylus to posteroinferior angle; posterior tooth largest, conical, separated, from other teeth by a con- siderable interval. Six or seven scattered tubercles forming an irregular line between upper and lower crests. Stridulating ridge on inner surface of manus composed of about 47 closely placed, oblique striae, which engage a series of coarser oblique tubercles on the suborbital margin to produce the vibratory grating described for Acanthocarpus bispinosus by Rathbun (1937:224). Outer maxilliped truncate-triangular, inner margin of ischium denticulate, a diagonal row of setae cross- ing exognath and merus of endognath. Ambulatory legs naked, unarmed, surface smooth and polished, carpi ridged, propodi laterally com- pressed, dactyli long, vertically compressed, incurving. A conical tubercle on either side of first sternite. The male of the species is unknown. Color in alcohol: Carapace reddish orange, deepest on anterior portion. A spot of orange surrounding each stridulating ridge. Remarks: The proposed new species appears to be more closely related to Acanthocarpus alexandri Stimpson, 1871 (Massachusetts to Windward Is- lands), type species of the genus, than to A. bispinosus A. Milne-Edwards, 1880 (Florida Straits to Windward Islands). Acanthocarpus delsolari resembles the former in having the cara- pace narrowing posteriorly; two meral spines, the inferior the larger; a tooth on the posterior margin; and a pair of tubercles on the sternal plastron. Furthermore, while the postlateral spine is longer than in alexandri, it is not as long as in bispinosus, nor as laterally directed. Acanthocarpus delsolari differs from both in having prominent branchial tuberculate ridges, the outermost actually over- hanging the posterolateral margin. I take pleasure in naming this species for E. M. del Solar, of Lima, Pert, who recognized it as belonging to a genus hitherto unreported from the American west coast. NEW BRACHYURAN CRABS FROM EASTERN PACIFIC 5 Family MAJIDAE Delsolaria, new penus Type species: Delsolaria enriquei, new species Description: vated and tuberculated, posterior lamellate Rostrum consisting of flattened Orbit small, a fissure above and below, the lower one open Preorbital spine and postorbital cup closely approxi mated, intercalary spine absent. Basal antennal article Carapace ovate-triangular, regions cle margin two horns. broadened basally, forming floor of orbit, a stout spine at anteroexternal angle. Maxilliped with ex- ognath as wide as endognath proximally, ischium widening distally, merus truncate distally, expanded anteroexternally, notched at anterointernal angle. Chelipeds of female slightly more robust than walking legs, manus compressed and carinated, fingers pointed. Walking legs short and stout, carpi tuberculated. Abdomen of female unknown. Relationship: The new genus appears to belong among those genera of the subfamily Pisinae in which the supraocular eave is in close contact with the postocular cup. However, the remarkable broadening of the basal antennal article suggests affinities with certain genera of the subfamily seven-segmented; male sex Majinae as well. Final placement and inclusion in a key must await the discovery of a male specimen. Delsolaria enriquei, new species Figure 3, A-F Type: Female, holotype, AHF No. 704, from north side of Banco de Mancora, Peri, 35 m, gravel, 9 December 1970, E. M. del Solar, collector (Orig. No. B-256). Measurements (in millimeters): Female holotype: length, including rostrum, 37.7, rostrum, 6.2, width at level of branchial spines, 29.6, at level of postero- lateral margin, 26.4, cheliped (ischium-merus 12.7, carpus-propodus 17.0), 27.7, chela, 13.8, dactyl, 6.6, height of palm, 4.7. Diagnosis: Gastric, cardiac, and branchial regions swollen and tuberculate. Lateral branchial tubercle prominent. Rostral, preorbital, and antennal spines of like size and sharpness. Maxilliped distally fringed with clavate setae. Description: Carapace ovate-triangular, one and one-quarter times longer than wide, highly convex medianly and laterally, elevations studded rounded tubercles, depressions smooth but microscop- ically punctate, sides declivitous, I lamellate. with posterior ma Front bifid, horns sharp, cleft shallow, U-shaped, sides parallel or slightly concave, a double row of curled setae on each; frontal, antennal, and preorbital spines of comparable size and sharpness. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 B IO mm E 5 mm Figure 3. Delsolaria enriquei, new species, female holotype: A, dorsal view; B, right lateral view; C, left chela, outer view; D, right outer maxilliped; E, ventral view of frontal region; F, abdomen. 1973 Preorbital spine with outer margin concave, separated by a narrow slit from postorbital cup into which eye retracts, leaving but a fraction of the cornea visible from above, a slight constriction behind postorbital cup. Gastric region broad, swollen, surmounted “by a dozen or more small tubercles of which two are median, the others paired; epibranchial regions sur- mounted by a single large tubercle, with lesser tuber- cles in advance and on outer slope; mesobranchial re- gions with three large tubercles forming a triangle, the anterior in line with the lateral and outer epibranchial, the interior in a line with the lateral and anterior car- diac tubercle; cardiac region most elevated, with four tubercles in a diamond with two in line, sides sloping from summit in all directions; sides of hepatic and branchial regions each with one or two small tuber- cles; intestinal region low, pinched, marked by a single median spine; posterior margin thin, broadly rounded, slightly sinuous, a notch marking off a small lobe at base of second walking leg. Basal antennal article broad, spine at anteroex- ternal angle visible from above. Pterygostomian re- gion with a double row of four rounded tubercles. Exognath of outer maxilliped as broad basally as endognath; ischium of endognath widening distally, invading merus internally, merus broadening an- teriorly, expanded externally, and notched internally to receive palpus; distal margins of merus and palpus fringed with clavate setae. Chelipeds of female slightly more robust’ than walking legs, merus with anterior and external margins cristate, Outer crest with a subterminal spine; carpus tuberculate; manus narrowing distally, crested above and below, fingers long, slender, pointed, downcurving, incurving, ribbed, and multidenticulate. Walking legs, of which most are missing from the type specimen, short, cylindrical, diminishing in length from first to last, carpi tuberculated, dactyli robust, setose, those of the last pair nearly as long as their propodi, tips incurving. Female abdomen with seven free segments, each with a low median tubercle. The male of the species is unknown. Family XANTHIDAE Pilumnus fernandezi, new species Figure 4, A—-F Pilumnus sp. Chirichigno, 1970, p. 55, text-figs. 128, 129. Type: Male, holotype, USNM No. 141571, Pacific side of Costa Rica, 80 m, E. Fernandez-B., collector (Orig. No. 13). Male, paratype, AHF No. 698, from off Punta Sal, Peru (lat. 03°59’ S), 180 m, on con- tinental shelf, 9 May 1969, E. M. del Solar, collector (Orig. No. B-20). Measurements (in millimeters): Holotype male: NEW BRACHYURAN CRABS FROM EASTERN PACIFIC 7 length of carapace, 35.5, width of carapace, including spines, 47.8, length of cheliped (coxa-ischium meru 29.0, carpus-propodus 38.3), 67.3, length of chela, 37.0, of dactyl, 21.0, height of palm, 18.7. Paratype male: length, 29.2, width, including spines, 37.9 Diagnosis: Carapace convex, four anterolateral spines, a subhepatic spine. Three spinules in line inside anterolateral spines. Chelipeds walking leg spinulous, color of fingers terminating short of palm Merus of outer maxilliped pentagonal. Tip of first male pleopod doubly recurved. Description: and Carapace one and one-third time broader than long, strongly convex anteroposteriorly, slightly convex from side to side; regions indicated only by a groove outlining the anterior mesogastric, an obtusely angular groove defining the hepatic region, and an H-shaped depression separating the urogastric from the cardiac region; sparsely hairy, with a row of three spinulous granules paralleling the anterolateral margins, and numerous lesser spinules. Front ad- vanced, consisting of two slightly oblique separated by a wide and deep U-shaped sinus, each lobe bearing three spinules; an antennal spinule stand- ing between front and orbit. Orbits broad, spinulous margined, of the spines on upper margin, One next antennal spine and one at about middle, two closed fissures above, three outer lower marginal spines and two inner, the latter on an advanced process bearing two spinules in a longitudinal line on basal portion. Anterolateral spines four, including exorbital spine, each with a conical base and slender tip, inclining successively more outward and upward. Carapace widest opposite fourth spine. Pterygostomian region granulate; a subhepatic spine with a second spinule beneath it. Sternum densely granulate. Merus of external maxilliped, owing to an inner projection, pentagonal, outer angle rounded, not pro- duced, surface with two shallow depressions, granulate, and _ hairy. Chelipeds unequal, the right larger than the left. spinulous, and hairy. Merus with five or six spines along superior margin, the two distalmost largest, curved, and separated by a U-shaped sulcus. Carpus with a prominent inner spine and outer groove as well as spines On upper and outer surface. Manus with two rows of three or four spines above, with lesser lobes spines arranged in rows on outer surface giving way to irregularly arranged spinate tubercles on lower por- tion. Fingers elongate, meeting with a slight gape. tips crossing; dactyl grooved above. spinulous at base, and with a low bicuspid tooth: pollex deflexed, swollen at base, with a tricuspid tooth followed by a large and a small tooth. Color of fingers terminating in an irregular line considerably short of palm. Minor less robust than major, complete! manus palm spinulous, fingers longer and slenderer, teeth and triangular, pollex grooved to tip. Walking legs slender, hairy, longest almost twice the length of carapace, and spinate on an BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME feos aun. mt ro pH Fm mm = ina a (| A TO mm AGN Hh eye Imm EB Sesion F Figure 4. Pilumnus fernandezi, new species, male holotype: A, dorsal view; B, right chela, outer view: C, left chela, outer view; D, left third maxilliped; E, abdomen: F, tip of first pleopod. 1973 gins of meri, carpi, and propodi; dactyli long, straight, slightly curved toward tips, nails amber. Male abdomen smooth-textured with seven ments. The segments beyond third decreasing regularly in width, seventh segment narrowly triangular. Male first pleopod with two rows of marginal setae, tip pointed, sharply recurved. Color in alcohol: Bright orange-red fading to salmon pink in a network leaving many whitish or cream-colored spots on carapace, chelipeds, and legs; color beneath extending from pterygostomian region onto maxillipeds and sternum. Fingers brown. Hairs golden, Variation: The paratype from Peru differs from the holotype from Costa Rica in lacking the superior spines on the orbits; these, however, appear to have been broken off on the specimen, which is rigid due to formalin preservation. No trace can be found of the three spinules that parallel the anterolateral margin on the right sides; on the left side the anterior two of these are present but reduced in size. The tips of the fingers of the major chela are worn in the paratype and do not cross as in the holotype specimen. Other- wise, the two compare favorably. Remarks: I take pleasure in naming this new species for Hno. Eduardo Fernandez-B., Colegio de La Salle, San José, Costa Rica. seg- Family GONEPLACIDAE Trizocarcinus peruvianus, new species Figure 5, A-G Goneplax sp. Chirichigno, 1970, p. 61, text-fig. 150. Type: Female, holotype, AHF No. 699, off Paita, Perti, 80 fathoms (144 m), 10 May 1969, epibenthonic net, E. M. del Solar, collector (Orig. No. B-106). Male, paratype, AHF No. 705, off Pert, depth not stated, 25 February 1970, E. M. del Solar, collector (Orig. No. B-106). Measurements (in millimeters): Female, holotype: length, 13.1, width, including spines, 19.1, frontal width, 5.5, fronto-orbital width, 16.0, length of right chela, 16.3, of dactyl, 8.7, height of palm, 5.1. Male, paratype: length, 19.9, width, including spines, 28.9, frontal width, 8.2, fronto-orbital width, 22.6, length of right chela, 25.9, of dactyl, 13.5, height of palm, 8.4. Diagnosis: Only two anterolateral teeth, including exorbital tooth. A sharp transverse ridge opposite lateral tooth. Chelae covered with matted hairs. An infraorbital tooth. No pterygostomian stridulating ridge. Male first pleopod long, straight, attenuated. Description: Carapace about two-thirds as long as broad, cardiac and posterior branchial regions crossed by a blunt transverse ridge, gastric and anterior bran- chial regions crossed by a sharper transverse ridge: these ridges dividing carapace into thirds, each in- clined at a different plane. Carapace uneven, espe- NEW BRACHYURAN CRABS FROM EASTERN PACIFIC 9 cially posterior two-thirds; mesogastric and cardiac regions clearly delimited, Posterior branchial regions swollen, cach with a blunt ridge parallel to the con posterolateral coarsely verging borders, Surface minutely granulate, and devoid of hairs Front bimarginate, upper margin straight, lower mar gin slightly sinuous, separated from subacute inner orbital angle by a notch, Orbit ticulate, two notches above and a closed fissure below tooth present. Anterolateral margin with only one tooth behind the flattened exorbital margin between exorbital and the more slender, cylindrical, and upturned lateral tooth arcuate and denticulate. Ridge on pterygos- tomian region paralleling epimeral suture blunt, non- punctate, broad, margin den the outer tooth; an infraorbital also tooth, striate, and lacking stridulating armature. Merus of external maxilliped subquadrate, antero- external angle broadly rounded, anterior margin slightly concave, surface matted anteriorly with to- mentum. Chelipeds subequal, the left slightly larger than the right; merus with a sharp spine on upper margin and a dense mat of hairs on internal surface; carpus with a cylindrical tooth on inner margin and a mat of hairs distally; chela with a dense hairy covering ex- ternally, leaving only a portion at base of fingers exposed; dactyls slender, elongate, incurving, irreg- ularly dentate, tips crossing. Ambulatory legs long, slender, carpi and propodi fringed with short hairs, dactyli straight or slightly incurving, with hairy margins. Male abdomen with seven distinct segments: third segment broadest, like second touching coxae of last pair of walking legs; third to fifth segments concave: base of sixth segment constricted; seventh segment narrowly triangular. Male first pleopod long, slender, cylindrical, taper- ing to attenuated tip, and armed with numerous short, conical setae and a few longer setae. Male second pleopod short, curved, tip ovate, concave. Remarks: Trizocarcinus peruvianus differs from JT. tacitus Chace, 1940, of the western At- lantic by possessing but two anterolateral marginal teeth instead of three, by having the outer surface of the chelae almost completely covered with hairy matting instead of only proximally so. and by having the infraorbital tooth present. Both species differ from T. dentatus (Rathbun), 1893, the type species from the Gulf of California, by having the carapace nearly smooth instead of hairy, and by lacking striae on the pterygostomian ridge, which, together with a lamellate ridge on the merus of the cheliped, comprise a stridulating organ in that species. The crisp ridge at the level of the lateral tooth is a further distinguishing dentatus. The female has been selected as the holotype 10 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME Figure 5. Trizocarcinus peruvianus, new species: A—C, female holotype; D—G, male paratype. A, dorsal view; B, outer view of right chela; C, ventral view of front; D, abdomen; E, F, first pleopod; G, second pleopod. 1973 because it has good locality and depth data, both lacking in the male, and is in perfect condition, the male has been dismembered and shows signs of having been dried and rehydrated. The male paratype has been used freely in the description, the male abdomen and gonopod being diagnostic of the species. DISCUSSION AND CONCLUSIONS It will have been noted from the foregoing account that the nearest relatives of the newly described eastern Pacific species, where known, are found in the western Atlantic. Thus, the species most closely related to Homolodromia robertsi is H. paradoxa A. Milne-Edwards, 1880, off Nevis, Leeward Islands, 356 fathoms; the species most closely related to Acanthocarpus delsolari is A. alexandri Stimpson, 1871, from Massachusetts to the Windward Islands, 45 to 208 fathoms; the species most closely related to Pilumnus fernandezi is P. diomedeae Rathbun, 1894, from the entrance to the Gulf of Mexico, 130 to 184 fathoms; and the species most closely related to Trizocarcinus peruvianus is T. tacitus Chace, 1940, off Barbados, 209 fathoms. More remarkable still, although this may be an artifact of collecting, with the exception of Pilumnus fernandezi, which occurs off Costa Rica, the newly described species have not been found in the Bay of Panama, nor have their Atlantic counterparts been found in the western Caribbean. All are species of the continental shelf and slope, most from below the 100-fathom line, yet not from such abyssal depths as the Peru-Chile Trench (cf. Garth and Haig, 1971) or the Cayman Trench, yet to be explored. I believe that these deep-water species represent an older and more fundamental relationship be- tween the two faunas than is indicated by the twin or geminate species found intertidally or subtidally on both sides of the Isthmus of Panama. This circumstance is of potentially great significance to marine zoogeographers, who look to evidence of major displacements of marine biotas as confirma- tion of geological and paleoclimatological changes. Thus, while it is possible that the pelagic larvae of deep-water brachyuran species were able to cross a shallow-water barrier until mid-Pliocene, the date of the last known the degree of morphological distinctness between isthmian closure, their species pairs, which is of a greater magnitude than now exists between the shallow-water species, suggests that effective interchange between the NEW BRACHIIYURAN CRABS FROM EASTERN PACIFIC I] now-isolated populations ceased long before Therefore it is postulated that the analogous pairs of deep-water Amphi-american species have been separated since free communication existed be tween them at or near the depths at which they presently occur. This would date them from Mio cene or earlier, when South America was truly an island continent, separated from North America not by a shallow shelf, but by waters of from 100 to 300 fathoms deep. Furthermore, their absence from the Bay of Panama and the western Carib- bean would indicate that this deep-water connec- tion was probably not across the present Isthmus of Panama, but across northern South America, perhaps at the Rio Atrato-Buenaventura, Colom- bia, level of the Bolivar Trench. The one new species that does not appear to fit into this category, and for which it has been necessary to erect a new genus, is Delsolaria enriquei. The family Majidae, to which it belongs, is of New World origin, with most of its numerous genera common to the east and west coasts of the Americas, and with parallel speciation on the two sides. There are several endemic west-coast genera, among them Loxorhynchus Stimpson, 1857, and Mimulus Stimpson, 1860, of western North Amer- ica, and Pisoides A. Milne-Edwards and Lucas, 1843, and Lophorochinia (Garth, 1969) of west- ern South America. These are either north or south temperate in distribution, and do not cross the Equator. Delsolaria may be placed provision- ally in the latter category until its extra-Peruvian distribution becomes known. ACKNOWLEDGMENTS I am indebted to Henry B. Roberts, U.S. National Museum of Natural History, Washington, for permis- sion to describe and illustrate Pilwmnus fernandezi, to Fenner A. Chace, Jr., U.S. National Museum, whose description of Trizocarcinus tacitus was fol- lowed in describing T. peruvianus, and to Daniéle Guinot, Paris Museum, whose studies on the Gone- placidae (Guinot, 1969) facilitated its generic place- ment. The manuscript was read in part (the Majidae) by D. J. G. Griffin, Australian Museum, Sydney, and in its entirety by Henry B. Roberts and by my asso- ciate, Janet Haig. A subvention from the Sociedad Nacional de Pesqueria of Lima, Peru, provided for the illustrations by Jerry J. Battagliotti. LITERATURE CITED Ghace; sb AS, Jr; 91940: eports on results of the Atlantis expeditions to the oO 12 BULLETIN SOUTHERN CALIFORNIA Indies, under the joint auspices of the University of Havana and Harvard University. The Brachy- uran Crabs. Torreia, Num. 4:1-—67, text-figs. 1-22 Chirichigno-F., N. 1970. Lista de crustaceos del Peru (Decapoda y Stomatopoda), con datos de su distribucién geografica. Inf. Inst. Mar Peru —Callao, 35:1—95, text-figs. 1-193. del Solar-C., E. M. crustaceos del Pert. 38:1-21. 1972. Inf. Inst. Mar Peru Addenda al catalogo de Callao, del Solar-C., E. M., F. Blancas-S., and R. Mayta-L. 1970. Catalogo de crustaceos del Pert. Lima, Peru. 1-46. Doflein, F. 1904. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898-1899. VI. Brachyura. (i-xiv) 1-314, pls. 1-46, text-figs. 1-68. Garth, J. S. 1969. A new genus and species of oxyrhynchous crab from the west coast of South America. Bol. Soc. Biol. Concepcion, 41:5—-7, text-fig. 1. Garth, J. S., and J. Haig. 1971. Scientific results of the Southeast Pacific Expedition. Decapod Crustacea (Anomura and Brachyura) of the Peru-Chile Trench. Anton Bruun Rpt. No. 6:1— 20, pls. 1-3. Guinot, D. 1969. Recherches préliminaires sur les ACADEMY OF SCIENCES VOLUME 72 groupements naturels chez les crustacés décapodes brachyoures. VII. Les Goneplacidae (suite). Bull. Mus. Hist. Nat. Paris, (2)41(2):507—528, pl. 2, text-figs. 33-82. Milne-Edwards, A. 1880. Reports on the results of dredging under the supervision of Alexander Agassiz, in the Gulf of Mexico, and in the Carib- bean Sea, 1877, ’78, °79, by the U.S. Coast Survey Steamer “Blake,” .. . VII. Etudes sur les Crus- tacés, 1°™° Partie. Bull. Mus. Comp. Zool., Harvard, 8:1—68, pls. 1, 2. Rathbun, M. J. 1893. Scientific results of explora- tions by the U.S. Fish Commission Steamer Alba- tross. XXIV. Descriptions of new genera and species of crabs from the west coast of North America and the Sandwich Islands. Proc. U.S. Nat. Mus., 16:223—260. 1894. Descriptions of a new genus and > four new species of crabs from the Antillean region. Proc. U.S. Nat. Mus., 17:83-86. 1937. The oxystomatous and allied crabs of America. Bull. U.S. Nat. Mus., 166:1—278, pls. 1-86, text-figs. 1-47. Stimpson, W. 1871. Preliminary report on the Crustacea dredged in the Gulf Stream in the Straits of Florida, by L. F. de Pourtales, Assist. U.S. Coast Survey. Part I. Brachyura. Bull. Mus. Comp. Zool., Harvard, 2(2):109-160. Accepted for publication September 10, 1972. A NEW SPECIES OF GERBIL FROM SOUTH WEST AFRICA WITH REMARKS ON GERBILLUS TYTONIS BAUER AND NIE THAMMER, 1959 (RODENTIA: DUANE ABSTRACT: West Africa. known for the subgenus. GERBILLINAE) A. SCHLITTER! A new species of Gerbillus (Gerbillurus) is described from Gobabeb, South This new species exhibits the maximum cranial size and bullar The skins of Gerbillus hypertrophy tytonis are described for the first time Gerbillus tytonis was known previously only by cranial elements taken from owl pellets. The Smithsonian Institution’s African Mammal Project had field parties collect small mammals and their ectoparasites in southern Africa from 1963 until 1969. Among the specimens collected were numerous Gerbillus. In southern Africa, this genus is presently in a confusing taxonomic state. There appear to be two groups of nominal species in the genus Gerbillus in southern Africa; a “paeba” group including the names listed under Gerbillus paeba by Roberts (1951) and subse- quent authors and a “vallinus” group including Gerbillus vallinus Thomas, 1918, and Gerbillus tytonis Bauer and Niethammer, 1959. These species have been included under the subgenus Gerbillurus Shortridge, 1942. Shortridge (1942:52) proposed the name Gerbillurus as a subgenus of Gerbillus to include only the species vallinus. In the original descrip- tion, Gerbillurus was characterized as having a long and relatively tufted tail, which in some examples is half as long again as the head and body, partially bare soles, and a triangular skull with inflated bullae. Although elevated to generic rank by Roberts, 1951, and Lundholm, 1955, Gerbillurus was retained as a subgenus of Gerbillus by Ellerman, Morrison-Scott, and Hayman, 1953, and the species vallinus was included in the genus Gerbillus by Meester, Davis, and Coetzee, 1964. Herold and Niethammer (1963:54, 56) concluded that Gerbillurus was more closely related to Tatera after comparisons of the enamel patterns of the lower first molars of young animals and the molar alveoli of different genera of gerbillines. They did not state conclusively, however, whether Gerbil- lurus should be recognized as a distinct genus or remain as a subgenus of Gerbillus. A review of the taxonomic status of Gerbillurus Shortridge, 1942, and the species in both the “paeba” and the “vallinus” groups is underway. ] ~ BS However, this preliminary paper was completed so that the new name would be available. “vyal- linus” group and the skins of Gerbillus (Gerbil- lurus) tytonis Bauer and Niethammer, 1959, are described for the first time. A new species is being proposed in the METHODS All measurements were taken with dial calipers and are in millimeters, weights are in grams and capitalized color terms are from Ridgway “Color Standards and Color Nomenclature” 1912. Hind foot measurements of specimens in the Smith- sonian Institution toenail. Total length and length of tail were taken on dorsal surface of specimens; the latter measurement with tail held perpendicular to body. Breadth of auditory bulla is the distance from the outside edge of the auditory meatus to the inner edge of the bulla. All specimens are deposited in the National Mu- seum of Natural History, Smithsonian Institution include (USNM), unless otherwise indicated. the Transvaal (TM). Pretoria: British Museum (Natural History) (BM), Lon- don; and the American Museum of Natural His- tory (AMNH), New York, were also examined. The new species may be known as: Specimens from Museum Gerbillus (Gerbillurus) setzeri. new species Figure 1 Holotype:—Young adult female, skin and_ skull, United States National Museum of Natural History 1 African Mammal Project, Smithsonian Institution, Washington, D.C. 20560 and Dept. Zoology, Univer- sity of Maryland. College Park 20742. l4 BULLETIN SOUTHERN CALIFORNIA no. 342253, from | mi E Namib Desert Research Station, Gobabeb, South West Africa; obtained 22 November 1963 by A. C. Risser, original no. 359. Specimens examined.—Seventy-four, as follows: South West Africa: | mi E Namib Desert Research Station, Gobabeb, 23; 8 mi E Namib Desert Research Station, Gobabeb, 1; Namib Desert Research Station, Gobabeb, 10 (1 TM); Swartbank [= Zwartbank] Mountain, 36 km WNW Gobabeb, 22: Tumas Moun- tain, 1; near Swakopmund, 2 (AMNH): Goanikontes, 3 (AMNH); 10 km E Hope Mine, 1 (TM); east of Gobabeb, | (TM); 8 mi E Hope Mine, 3 (TM); east of Hope Mine, 2 (TM); Swakopmund, 5 (TM). GAZETTEER Goanikontes 22° 40’ S, 14° 50’ E Gobabeb D233 4a Sil Set O Sine Hope Mine DBMS AES ol Se aS ei Sossus Vlei 24° 44’ S, 15° 18’ E Zwartbank Mtn. 23° 224 S; 14° 58" E Swakopmund 22° 41’ S, 14° 32’ E Tumas Mtn. PRO DE SOUS Sil? Je! Measurements.—Selected external and cranial mea- surements of the holotype are: Total length, 249; length of tail, 143; length of hind foot, 32; length of ear from notch, 15; occipitonasal length, 32.6; greatest breadth across zygomatic arches, 17.1; greatest breadth of braincase, 15.1; height of skull, 13.5; least interorbital breadth, 5.3; length of nasals, 12.8; oblique length of audital portion of auditory bulla, 11.8; crown length of maxillary toothrow, 4.5; greatest breadth of M*—M®, 5.1; length of anterior palatine foramina, 5.7; and length of posterior palatine foramina, 25. Comparative measurements of three species of Gerbillus (Gerbillurus) are given in table 1. Diagnosis——Upper parts near Light Pinkish Cinna- mon, with slight admixture of gray hairs; all hairs plumbeous at base. Circumoral, entire underparts, supraorbital and postauricular spots, and dorsal sur- faces of hands and feet, white; all hairs uniformly white to base. Sharp line of demarcation present be- tween dorsal and ventral colors of body. Tail rela- tively short for subgenus and bicolored, dorsal color same as color of back, ventral color white; tail with penicillate tip of Mouse Gray dorsal hairs on distal one-third. Color of back extending to hairs on ex- ternal surface of pinna; hairs of internal face of pinna white; flesh of pinna Cinnamon-Buff. Skull large for subgenus: upper toothrow relatively short and robust; audital and mastoidal portions of auditory bulla relatively large and well inflated ventrally and posteriorly; mastoidal portion of auditory bulla pro- jecting beyond occiput; external auditory meatus well inflated anteriorly and foramen of Huschke well de- veloped ventrally; anterior palatine foramina rela- tively short and wide; posterior palatine foramina long. Comparisons.—From the nominal species of ACADEMY OF SCIENCES VOLUME 72 the Gerbillus paeba group in southern Africa, Gerbillus (Gerbillurus) can be distin- guished by its larger size, both externally and cranially. The large size of the body and the hind feet serve at once to distinguish this new species from any of the smaller G. pacha. From Gerbillus (Gerbillurus) vallinus vallinus as known from the vicinity of Berseba, South West Africa, and Tuin, South Africa, this new species differs externally by having a shorter tail, longer hind feet, smaller ears and being dorsally paler in color. Cranially, the skull of G. setzeri is longer and broader, the bulla larger and more inflated, the breadth across M*—-M® narrower, and the least interorbital breadth narrower than G. v. vallinus. Gerbillus setzeri differs from representatives of G. vallinus seeheimi from Seeheim, South West Africa, and numerous localities in the northern Cape Province, South Africa, by the lack of black hairs that are suffused in the dorsal pelage and make up the tufted tip of the tail in G. v. seeheimi. Skulls of G. setzeri differ from G. vy. seeheimi in the same manner as from the nominate subspecies. Even though the breadth across M?—M? is less in G. v. seeheimi [5.1 (4.7—5.4) 25] than in G. v. vallinus [6.3 (6.1-6.5) 7], this distance is still greater than the same measurement in G. setzeri (Table 1). Representatives of Gerbillus tytonis taken at the type locality of G. setzeri can be distinguished by the smaller size of the body, shorter and nar- rower skull, less inflated auditory bulla, and markedly shorter posterior palatine foramina. The dorsal pelage of G. tytonis is darker in color than that of G. setzeri; the former is a reddish gold color and matches the red sand of the desert south of the Kuiseb River at Gobabeb whereas the latter is a paler color near Light Pinkish Cinnamon and more nearly matches the pale feldspar and quartz plains north and east of the Kuiseb River at Gobabeb. Statistical Analysis.—Initially, samples of males and females of each of the three species were compared to determine if any secondary sexual variation existed in mensural data. Weights were not included in these tests. These analyses re- vealed the absence of secondary sexual variation in all of the measurements tested in each of the three species; consequently the sexes were com- bined for subsequent tests of significant differ- ences between the species. In addition, compari- sons between Gerbillus vallinus vallinus and G. vy. seeheimi revealed these two subspecies were sig- setzeri 1973 1 NEW SPECIES OF GERBIL FROM AFRICA Figure 1. from left to right: South West Africa (USNM 342196). G. setzeri and lateral view of same skull. nificantly different only in the breadth across the third upper molars. Results of statistical comparisons between sam- ples of Gerbillus setzeri with samples of G. vallinus and G. tytonis are given in table 1. Both “t” tests and “F” tests were computed but SS-STP tests were not run inasmuch as only grouped single samples for each species were available. For all but one measurement, the coefficient of varia- tion was within the range considered normal for small mammals. In the case of the posterior palatine foramina, the high values of the coef- ficient of variation are a result of the difficulty ocular of taking the measurement. Use of an micrometer or a craniometer would undoubtedly Skulls of the three species of the Gerbillus vallinus group Gerbillus setzeri holotype (USNM 342253); G. South West Africa (USNM 304852): and G. Bottom row, left to right: (Scale 2x ) Top row, ventral views vallinus vallinus, Berseba. tytonis, Namib Desert Research Station, Gobabeb. dorsal view of holoty pe of result in lower coefficient of variation values The high value of the coefficient of variation for the breadth across third upper molars of G. val- linus is a result of combining the samples of the two subspecies. Two of the results of the “F” test are ably significant. In the comparison of ear between Gerbillus setzeri and G. val * test would indicate the variances of samples are significantly different at the 95 per cent level but the low “t” value (3.189) indicates a questionable difference. In the case of the com- parison of crown length of maxillary toothroy between G. setzeri and G. tytonis. the low value (2.981) again indicates a questionable dif- VOLUME 72 DRY INCI SCIE MY OF Ey ACAD BULLETIN SOUTHERN CALIFORNIA 16 9S = F0'F €€ 9s €€ OE (S'p-L'€) (Zr-S'€) (€p-9'€) br00 = IIb xxSL0'0 = P6'E ¢S0°0 = LO'r Ef) 6h = EST ve wee se EOE LS 67E pe 98'E S— EST (6°8-6'L) (0°6-0'8) (L6-1T'8) (0°9-7'S) (€9-4'S) (6'S-€'S) ++++090'0 = FE'8 +7€80'0 = 19°8 IIl'0 = $06 MED 8r0'0 = IS°s 9L0°0 = SL's €S0'0 = 9g9°¢ ait 6c =F SO OKC CCmee ONC erp O1'Z 6t ©6667 US = yee (871-071) (TEI-6 IT) (S°€I-LT1) (9'bI-F'€1) (O'SI-0'€T) (19I-l'rL) +2060'0 = +7 ++L€10 = 89 TI SILO = OU'ET SHD +x680°0 = P6'El xx8S1'0 = C71 6¢1'0 + 76'FI add) igs L861 pe SPrIl 9€ E€L'8 6€ soe TE = O8'T CCM mE SHIG (9°1-9'0) (¢t-+'T) (L7-8'T) (9°91-@'r1) (S'°9I-1'ST) (PLI-L’S1) «+LS0'0 = 801 «+8100 = 861 L90°0 = TET ddd xx 80 = 19°S1 xe SST'O0 = 89°CT €s1'0 = 7991 VZ4 96 SSP re lpr ve SSP ep 787 iS VANE SC CSHO (€9-1'S) (L’S-L'v) (1'9-0'S) (F'0E-T'87) (6° 1€-T'87) (9°7E-S 67) 0L0';0 = €F's ++8L0°0 = 61'S 6800 = Lrs ddT xx €STO = PE'6T xx €8T'0 F 06°67 Tee0 = Erle NOO SE Ilr ZTE Os «L6°6 L Ly'@ Lg Sas ECan O0it er 109 (¢S-S br) (S9-L'P) (€'S-6'P) (FI-Z1) (9I-F1) (91-71) +S90°0 = L8'r 6810 = LES cs0'0 = Los Wd xx081'0 + 96°TI +exCCT'0 = br'rl STO = 98'EI Na os 8sLz €€ gars 61 LL€ LS 9@P Ke SAGE br ELE (S't-0'r) (S'¥-L'€) (9'p-T'p) (9€-87) (re-0£) (Se-0€) +«*££0'0 = ECP «+CL0'0 = 80°F CLO'0 = €€'F LTO xaLLe0 * LE EE xxbIb'0 F 6LTE S9€0 = SPTE LH Lp 9€7 ve OS? ce 10°€ cS = 86 97 IVL er = 60'S (+r 0I-£'6) (0° 11-6'6) (771-901) (IrI-€11) (9S1-611) (SPI-€11) ++«890'0 = 08°6 ++160°0 + 7901 OTTO = 6E TT avi 8691 & 6 9TI «xT L8'€ + S8'SEl 6L6'1 = SE'LTI VL Gs _ Airs ve €LPr 6% OLE cc IL 97 ESF frp €l'b (O'€I-s'01) (FTI-€'01) (l€I-v'11) (OPz-S07) (997-S ITZ) (€97-L17) ++191'0 = 791 s+7810 + PE TI TLV0 = 67 TI N19 *«x09T'T F IL'S7 xOPS'P F S8'6E7T 9E8'T F CO'EET TL s1uojway y) SNUIP]DA 3) 1492Jas **). S10} 73) SHUTP]DA 3) 140ZJaS 9) ‘Ry[nq Asoypne Jo YyIpesag ‘MA_H pur ‘][nys Jo YsIeYy ysajwais ‘SHH ‘vulwesoJ ouNLied Jola}sod Jo yu] “4dd ‘vullUBIOJ due] Vd Jolajur jo ysue] “4d TJ ‘JN 1 aeyed jo yproq ‘Wd :MO1Y4IOO} AIv][IXeW JO YSUI] UMOID “LTO ‘eI[Nq AJoUpne Jo uo}.od [vyipne Jo yu, onbiyqo ‘gy ‘s[BSPU JO YJDUA] ISalvais ‘NTH :WNsjsol Jo YIpeosg “YH ‘yipvoig [VIUGIOLO}UI Jsvol ‘G]T ‘esvoureIg JO YJpvasq jsavais ‘G|H ssayoiw oyeVwosAZ ssolov yIpraiq “YZd ‘yisue] [eseuo}dI990 ‘NOO ‘Yyojou Wolly Iva Jo YyysUa] ‘NA ‘OOF pury Jo yIWus] “PH *]!e1 Jo yu ‘VL SYyIUs] [%}I0} “TI, :SMOTJOJ sv ov syUdWaINsvaW Jo SUONRIADIQGY “(a2dUapIjUOd JO [PAI] [0'0 = ++ PUL BdUSPIJUOD JO [2AI] CY’) = + ‘IS9} 4, JOJ ‘9UIpPIJUOS JO [OAR] [0'O = yy PUB DdUSPIFUOS JO [oA] CO'0 = » 482} .1,, JOJ) JUBOIFIUDIS av sIVOJA] “H PUV SNUI]]VA *H JO asOY} pu IMazZjJas “H Jo safdurvs dy} UdIaMJaq SddUdIAFJIP YSIYM UL Payvorpul oiv s}UdWaAINSvaA, ‘o[duuvs ay} Ul SuaIdads Jo JaquINU ay} PUL UONRIIVA JO JUAIOIJIOI dy} OUT] PAY] 94} puv ‘salaI}Xe 9Y} SUI] PUODAS ay} ‘Ss1O1Ia PAvpUL}s OA snuIW puv snd uvoW 94) SOPN[OUI JUSWaINSvaW YORA JO} SUI] ISINY OY] “BOY uUIOyINOS WoIy sivosk] “DH pure ‘snuip]VA “D2 ‘Llazjas Snjjigiay Jo sajduvs udsanjoq uonVUeA ‘| aTAvy 1973 ference between the variances of the two samples. In all other instances where the “F” test is sig- nificant, “t values are high (6.181—25.003). Remarks.—Gerbillus setzeri is apparently. re- stricted to the very pale feldspar and quartz gravel plains in the Namib Desert. In sharp con- trast to this distribution is that of Gerbillus tytonis, a species apparently found only on the shifting red sands south of the Kuiseb River. In Novem- ber of 1963, field collectors for the Smithsonian Institution took both species at Gobabeb in the red sands of the dry river bed and adjacent dunes south and west of the research station. At this time both species were rather common in_ the vicinity of the field station. According to the late Charles Koch (pers. comm., 1969) in March of 1963, unusually heavy rains fell in the vicinity of Gobabeb and the area to the west with subse- quent heavy flooding in the Kuiseb River. In the months following these rains, vegetative growth on the gravel plains was exceptionally lush ac- cording to Dr. Koch. Judging from the ages of the specimens taken by the field collectors, by November a good breeding season had been ex- perienced inasmuch as mostly juveniles, sub- adults and young adults were trapped. No old adults were included in either of the samples of G. setzeri from Gobabeb or downstream near Zwartbank Mountain, also situated on the bank of the Kuiseb River. During this period of higher population levels, individuals of Gerbillus setzeri apparently dispersed from the gravel plain across the river bed and into the adjacent red sand dunes to the south and west of Gobabeb. Extensive trapping at Gobabeb in December 1969 revealed Gerbillus setzeri to be found rarely (five speci- mens in about 400 trap nights) and only on the gravel plain, whereas Gerbillus tytonis was com- moner but found only on the red sands. Individ- uals of the Gerbillus paeba group were found ubiquitously distributed and common at Gobabeb in 1969. The single individual from Tumas Mountain was taken in fine sand with less gravel present; dry grass formed a rather uniform cover in this area. The locality 10 mi E Hope Mine was also visited and proved to be nearly identical to the gravel plains near Gobabeb and Zwartbank Moun- tain. Meester (1963:245) reported a male speci- men shot on the gravel plain east of Gobabeb. In the only sample of weights available for G. males than females setzeri, averaged heavier taken at Zwartbank Mountain. Eight adult males A NEW SPECIES OF GERBIL FROM AFRICA 17 averaged 39.5 (37.0-42.5); whereas nine adult females averaged 34,3 (29,5~—39.0). Meester (1963:245) reported an individual from east of Gobabeb as Gerhillus vallinus val- linus. A skin with broken skull in the Transvaal Museum (TM 12929) was taken east of Gobabeb by the Bernard Carp expedition. This specimen is probably the same one reported by Meester and is referred here to G. setzeri This new species is named after Dr. Henry W. Setzer in honor of his efforts in African mammal- ogy taxonomy of desert rodents. and in particular for his interest in the REMARKS ON GERBILLUS (GERBILLURUS) TYTONIS BAUER AND NIETHAMMER, 1959 Originally proposed by Bauer and Niethammer (1959:255) as a subspecies of Gerbillus vallinus, Gerbillus tytonis was described from a collection of skulls removed from owl pellets collected at Sossus Vlei, South West Africa. Davis (1968:4) regarded G. tytonis as worthy of specific status; he distinguished the very short posterior palatine foramina and the small bullae of tyronis. I concur with this separa- tion of G. tytonis from vallinus. In spite of the tytonis from vallinus by large amount of variation present in the length of the posterior palatine foramina of G. tyronis, this measurement is still significantly shorter than that of G. vallinus. Gerbillus smallest species of the “vallinus” group (Table 1). To my knowledge, G. tytonis is known in the only from skulls recovered from owl tytonis is the literature pellets; a description of the skins of this species based on adult specimens taken on 28 March 1966 at the type locality follows: Upper parts near Hazel; all hairs plumbeous at base. Entire underparts, small supraorbital and well-defined postauricular spots, and dorsal sur- faces of hands and feet, white; all hairs uniformly white to base. Sharp line of demarcation present between dorsal and ventral color of pelage. Tail relatively long for subgenus and bicolored, dorsal color same as color of back, ventral color same as dorsal but with admixture of white hairs: tail with variable penicillate tip of grayish hairs. White circumoral ring absent. Ears essentially bare and Cinnamon-Buff in color: narrow fringe of black pigmentation present on lateral margin of pinnae. The dark dorsal pelage, long feet, small size of 18 BULLETIN SOUTHERN CALIFORNIA body and skull, short posterior palatine foramina and small bullae serve to distinguish Gerbillus tytonis from G. vallinus. ACKNOWLEDGMENTS Special acknowledgment must be given to Henry W. Setzer for allowing me to study the Gerbillus collec- tion at the Smithsonian Institution; this collection from southern Africa provided the bulk of the ma- terial which resulted in this paper. Thanks are due Richard W. Thorington, Jr., for making funds avail- able for statistical analysis and for commenting on the manuscript. Gratitude is extended to the curators of the museums mentioned at the beginning of this paper for making specimens and facilities available to me during this study. Others who deserve thanks for helping make this paper possible are, obviously, the many persons who collected mammals for the Smithsonian Institution in southern Africa. In addition, a special debt of gratitude is owed J. A. J. Meester, University of Natal, Pietermaritzburg; D. H. S. Davis, formerly with South African Medical Ecology Centre, Johannes- burg; the late Dr. C. Koch and I. L. Rautenback, Transvaal Museum, Pretoria; and C. G. Coetzee, State Museum, Windhoek, for their efforts and sug- gestions. Staff photographers at the National Mu- seum of Natural History made the skull photographs and Mrs. Helen J. Hutchinson typed the manuscript. This paper is a result of research supported by U.S. Army Medical Research and Development Com- mand Contract No. DA-49-193-MD-2738. LITERATURE CITED 1959. Uber eine Sudwest-afrika. Bauer, K., and J. Niethammer. kleine Saugetierausbeute aus Bonn. zool. Beitr., 10:236—261. ACADEMY OF SCIENCES VOLUME 72 Davis, D. H. S. 1968. Rodentia, Gerbillinae: Genera Tatera and Gerbillurus. in: Meester, J. (ed.) Smithsonian Institution preliminary identi- fication manual for African mammals, 18(2): 1-5. Ellerman, J. R., T. C. S. Morrison-Scott, and R. W. Hayman. 1953. Southern African mammals 1758 to 1951: A reclassification. Trustees Brit- ish Museum (Natural History), London, 363 pp. Herold, W., and J. Niethammer. 1963. Zur sys- tematischen Stellung des sudafrikanischen Gerbil- lus pacba Smith, 1834 (Rodentia: Gerbillinae) auf Grund seines alveolenmusters. Saugetierk. Mitt., 11:49-58. Lundholm, B. G. 1955. Descriptions of new mam- mals. Ann. Transvaal Mus., 22:279-303. Meester, J. 1962. Some mammals from the Namib Desert. Ann. Transvaal Mus., 24:241-248. Meester, J., D. H. S. Davis, and C. G. Coetzee. 1964. An interim classification of Southern African mammals. Raneved, distributed with the assis- tance of the Zoological Society of Southern Africa and the C.S.IR., 75 pp. Roberts, A. 1951. The mammals of South Africa. Trustees “The Mammals of South Africa Book Fund.” Johannesburg, xlviii + 700 pp. Shortridge, G. C. 1942. Field notes on the first and second expeditions of the Cape Museums’ Mammal Survey of the Cape Province; and descriptions of some new subgenera and sub- species. Ann. South African Mus., 36(1):27—100. Accepted for publication November 28, 1972. POLYCHAETES FROM CENTRAL AMERICAN SANDY BEACHES KRISTIAN FAUCHALD! ABSTRACT: new genus are described. The polychaete fauna of Central America has been the subject of several studies; early literature was summarized by Monro (1928, 1933a, 1933b). Hartmann-Schroder (1959) described polychaetes in shallow-water quantitative samples from man- grove areas of El Salvador. Sandy beaches, how- ever, have been largely neglected until recently. The present collections were made under the supervision of Deborah M. Dexter of California State University, San Diego, over a period of two years for the purpose of comparing the total fauna of the sandy beaches of the Atlantic and Pacific Oceans of Central America. STATION LIST Several beaches were collected at different times, so the list contains location only. Because of the com- plex coastlines, the stations are listed alphabetically within each country. ATLANTIC OCEAN Colombia: Boca Grande, Cartagena, 10°30’ N, 75°40’ W Isla San Andres, 12°32’ N, 81°34’ W Pradomar Beach, Puerto Colombia, 11°08’ N, 75°09’ W Rodadero Beach, Santa Marta, 11°15’ N, 74°13’ W Costa Rica: Airport Beach, Limon, 9°58’ N, 83°01’ W Cahuita North, 9°45’ N, 82°52’ W Cahuita South, 9°44’ N, 82°50’ W Playa Bonita, Limon, 10°01’ N, 83°04’ W Puerto Viejo, 9°40’ N, 82°44’ W Panama: Pina Beach, 9°20’ N, 80°05’ W San Blas. 9°30’ N, 79°20’ W Shimmey Beach, 9°23’ N, 79°57’ W PACIFIC OCEAN Colombia: Playa Juan Chaco, N of Buenaventura, 4°10’ 77°30’ W Costa Rica: Boca de Barranca, Puntarenas, 9°58’ N, 84°45’ W N, Twenty-six species of polychaetes are reported from sandy beaches in Colombia, Panama, and Costa Rica from the Atlantic and Pacific Oceans. Two new species and one Jaco, 9°37’ N, 84°38’ W La Punta, Puntarenas, 9°59’ N, 84°52’ W Playa Cocal, Quepos, 9°26’ N, 84°10’ W Playa Espadilla, Quepos, 9°24’ N, 84°10’ W Playita Blanca, 10°34’ N, 85°42’ W Samara, 9°53’ N, 85°38’ W 10°19’ N, 85°50’ W Playas del Coco, Coco, Tamarindo, Panama: Naos Island, 8°53’ N, 79°33’ W San Carlos, 8°29’ N, 79°58’ W Venado Beach, 8°55’ N, 79°36’ W Family Sigalionidae Sthenelais maculata Hartman, 1939 Figure 1f—g Sthenelais maculata Hartman, 1939, pp. 64—65, pl. 15, figs. 176-187. Material examined: Boca Grande, 8 April 1971 (2): Naos Island, 30 June 1969 (2), 29 August 1969 (1): Jaco, 21 March 1971 (1); Playa Cocal, 28 February 1971 (2). Distribution: Sthenelais maculata was originally described on material from Peru, Panama, and Mexico. The present records cover Pacific beaches in Costa Rica and Panama and one beach on the Atlantic side of Colombia. Remarks: The present specimens fit very well with the specimens originally described by Hart- man (1939). The superior part of the presetal lobe is deeply bifid (Fig. 1f); the distal tooth of the composite falcigers (Fig. 1g) is strongly curved. Family Pisionidae Pisione remota (Southern, 1914) Figure 2a Praegeria remota Southern, 1914, pp. 61-64, pls. VII and VIII, figs. 15a—k. 1 Allan Hancock Foundation, University of South- ern California, Los Angeles, California 90007. 20 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 Figure I. Glycera abranchiata Treadwell: b, tall proboscideal organ, lateral view, * 385; < 385; Sthenelais maculata Hartman: falciger, parapodium 85, x 950. Pisione remota Hartman, 1968, pp. 181-182, 5 figs. Material examined: Playa Espadilla, 1 March 1971 (1); Playita Blanca, 9 February 1971 (8). Distribution: Pisione remota has been reported from shelf-areas off Ireland and on the west coast of the Americas; the present records are from the Pacific side of Costa Rica. Remarks: The present specimens agree with P. remota as characterized by Hartman (1968) and with Laubier’s (1967) review of the species of Pisione. The simple bifid hooks have a series of long, neatly organized hairs between the two teeth (Fig. 2a). This brush-border is usually illustrated as being considerably less extensive than it is in the present specimens. Paired bosses are present lateral to the base of the secondary tooth in all simple hooks. Pisionidens indica (Aiyar and Alikuhni, 1940) Pisionella indica Aiyar and Alikuhni, 1940, pp. 89- 107, pls. 1-2, 9 text figs. a, median parapodium, >< O38 posterior view, c, tall proboscideal organ, frontal view, « 385; d, short proboscideal organ, frontal view, > 385; e, short proboscideal organ, lateral view, f, parapodium 85, anterior view, x 95; g, composite Pisionidens indica Aiyar and Alikuhni, 1943, p. 120; Siewing, 1954, pp. 81-83, pl. 23; Day, 1967, p. 133, fig. 4.1.f-j. Material examined: Cahuita South, 1 April 1971 (2); Pradomar Beach, 21 April 1971 (1); Rodadero Beach, 26 April 1971 (22). Distribution: Pisionidens indica is known from India, South Africa (Day, 1967) and has been re- ported from El Salvador and Brazil (Siewing, 1954). The present records are from the Atlantic side of Colombia and Costa Rica. Remarks: The present specimens fit very well with the species as originally described and later revised by Siewing (1954). Family Amphinomidae Hermodice carunculata (Pallas, 1766) Hermodice carunculata Fauvel, 1914, pp. 113-116, pl. 8, figs. 22-27, figs. 31-32; Monro, 1933a, p. 4; Hartman, 1951, pp. 22-25, fig. 1. 1973 Material examined: Pina Beach, 24 July 1969 (1); San Blas, 19 August 1969 (1). Distribution: Hermodice carunculata was originally described from the West Indies; it has been reported extensively from warm water areas in the Atlantic Ocean and is apparently limited to this ocean. Remarks: Hermodice carunculata Eurythoe complanata with which it is often con- fused by non-specialists, in that both species are long-bodied fire-worms. ‘The two species differ most noticeably in the development of the caruncle. This structure is long and flexuose with poorly developed, usually completely hidden, lat- eral folds in E. complanata; it is wide and ovate, with very obvious lateral folds in H. carunculata. resembles Family Phyllodocidae Anaitides near multiseriata Rioja, 1941 Anaitides multiseriata Rioja, 1941, pp. 684-687, pl. 1, figs. 2-6; Hartman, 1968, pp. 237-238, 4 figs. Material examined: Playa Juan Chaco, 5 September 1971 (1); Venado Beach, 14 July 1969 (17). Distribution: Anaitides multiseriata is known from western Mexico and Southern California; the present records are from Pacific Panama and Colombia. Remarks: The present specimens agree with A, multiseriata in most characters, but differ in the color pattern. Anaitides multiseriata has a series of dark transverse bars with light-colored mid-dorsal spots; the specimens reported herein are nearly uniformly grey. In addition, A. multi- seriata has a characteristic H-shaped pattern on the prostomium; this pattern is absent in the present specimens. The dorsal cirri are slightly more pointed in the present specimens than in A. multiseriata. Further identification will have to await a complete re-evaluation of the specific characters used to separate the many species in the genus Anaitides. Family Nereidae Ceratonereis mirabilis Kinberg, 1866 Ceratonereis mirabilis Kinberg, 1866, p. 170. Ceratonereis tentaculata Kinberg, 1866, p. 170; Monro, 1933a, p. 45; Hartman, 1940, p. 218, pl. 35, fig. 47. Material examined: Naos Island, 30 June 1969 (1): Rodadero Beach, 20 April 1971 (1). Distribution: Ceratonereis mirabilis is widespread in warm waters and may be circumtropical; it has been reported more frequently from rocky shores than from sandy beaches. Remarks: Ceratonereis mirabilis has a distally bifid prostomium and very long antennae. The POLYCHAETES FROM CENTRAL AMERICAN BEACHES 21 Fa ‘} i Cc 5 Ee | = Ie) e A, =A £ Figure 2. Pisione remota (Southern): a, simple hook, x 1900; Notodasus dexterae, new species: b, anterior end, lateral view, » 16; c, hooded hook, lateral view, xX 950: d, hooded hook, frontal view, approx. xX 950; e, abdominal setigers 8-10, dorsal view, x 16; f, abdominal setigers 15-17, dorsal view. Sai6: dorsal cirri are greatly prolonged in posterior setigers giving the specimens a slightly ragged appearance. Nereidae, indeterminable Material examined: Rodadero Beach, 20 April 1971 (fragment). Remarks: This fragment cannot be further identified; it appears different from Ceratonereis mirabilis reported from the same beach. Family Glyceridae Glycera abranchiata Treadwell, 1901 Figure la-e Glycera abranchiata Treadwell. 1901, pp. 200-201, fig. 49: Jones. 1962, p. 183. figs. 41-48. 22 BULLETIN SOUTHERN CALIFORNIA Material examined: Naos Island, 31 July 1969 (8). Glycera abranchiata is known from The Panama and is the Distribution: Puerto Rico, Jamaica, and the Lesser Antilles. present record is from Pacific first record of the species outside the western Atlantic Ocean. Remarks: The present specimens resemble G. tesselata Grube (1863) in general body propor- tions and in the shape of the parapodia. The two postsetal lobes (Fig. la) are somewhat more dis- tinct in G. abranchiata than in G. tesselata and the two presetal lobes are somewhat longer in the former than in the latter. The proboscideal organs are of two different kinds. The most numerous kind (Fig. |b-c) is narrow and slender and has 13 to 14 ribs in an oblique pattern. The other kind (Fig. 1d-e) is considerably wider and show a broad flattened shape; this second kind lacks ribs. The distinctness of G. abranchiata was recog- nized by Jones (1962) who re-examined the type material. These types were also examined during the present investigation and were found to be as described by Jones. Hemipodus armatus Hartman, 1950 Hemipodus armatus Hartman, 1950, pp. 83-84, pl. 12, figs. 1-5. Material examined: Boca Grande, 28 April 1971 (3); Cahuita South, 1 April 1971 (1); Puerto Viejo, 2 April 1971 (1); Samara, 8 April 1971 (9); Tamarindo, 9 April 1971 (1 large, several small). Distribution: Hemipodus armatus is known from western Mexico in shallow-water: the present records come from sandy beaches on both Atlantic and Pacific sides of the isthmus of Panama. Remarks: The present specimens agree with H. armatus in that the tall proboscideal organs have numerous transverse ridges and in that the presetal lobes of median and posterior parapodia have slender digitate distal processes. Family Nephtyidae Nephtys singularis Hartman, 1950 Nephtys singularis Hartman, 1950, pp. 98-100, pl. 15, figs. 1-6. Material examined: Naos Island, 30 June 1969 (6); Jaco, 27 March 1971 (3); La Punta, 14 March 1971 (8); Samara, 8 April 1971 (2): Tamarindo, 9 April 1971 (28). Distribution: Nephtys singularis was described from Guatemala and western Mexico; all present records are from the eastern Pacific Ocean in some- what lower latitudes. Remarks: The present specimens agree with ACADEMY OF SCIENCES VOLUME 72 the original description. Interramal cirri are pres- ent from setiger 4 in nearly all specimens, but can be identified from setiger 3 in very large speci- mens. The superior neuropodial lobes are best developed in setigers 8-15 and are completely re- duced posterior to setiger 30 in all specimens. Family Onuphidae Americonuphis, new genus Type species: is A. magna (Andrews, 1891, as Diopatra); A. hartmanae (Friedrich, 1956) and A. reesei, new species also belong to this genus. Description: Americonuphis includes macropodous onuphids with the anterior five or more parapodia enlarged and armed with composite falcigers. More posterior parapodia have simple limbate and pectinate setae, and distally bifid subacicular hooks. The prostomium has five occipital tentacles, each has a multiannulate base and a long, tapering style. The peristomium has a pair of tapering cirri. The thick, long, enlarged parapodia are armed with composite falcigers; the setal shafts are limited to their respec- tive segments. The long, cirriform dorsal and ventral cirri taper distally. Farther back the ventral cirri are replaced by thick, glandular pads. Branchiae are pres- ent behind the modified segments and are continued through many segments; each has many filaments in a pectinate arrangement. The maxillary apparatus resembles that of other onuphids. Remarks: Americonuphis is most nearly allied to Rhamphobrachium in having five or more, instead of two or three, anterior parapodia greatly enlarged and armed with special setae. The two genera differ in that the former have composite falcigers instead of scythe-like curved spines in these setigers and in that the shafts of the setae are limited to one segment rather than continued through several segments. Americonuphis reesei, new species Figure 3a-e Material examined: Gulf of Panama, 27 m, mud, coll. E. Reese (1, Holotype); Naos Island, 3 July 1969 (1). Description: The large holotype has been removed from a thick, mud-walled tube and is broken into several pieces; the posterior end is absent. The an- terior fragment, with 80 setigers, measures 85 mm long and 8 mm wide; more posterior segments bring the total length to 230 mm and the width to 9 mm; the total number of segments exceeds 300. The an- terior ventrum and the parapodial bases are mottled with black pigment. The tube has an inner lining of a tough parchment-like material and is externally covered with a thick layer of mud. 1973 The small prostomium is a flat rectangular lobe, slightly longer than wide on top of the thick, biartic- ulate palps. A pair of tapering frontal antennae are present. The thick ventral palps are basally fused and are deeply divided distally; their bases are large and overlie the oral aperture (Fig. 3a). The ventral lip is formed by the thick, broad anterior edge of the peristomium. A pair of eyes is at the outer bases of the inner lateral occipital tentacles. The five occipital tentacles are arranged so. that three are in a row across the posterior end of the prostomium; the other two are in front of the inner lateral pair and at the sides of the prostomium, The ceratophores are annulated; each of the median and inner lateral Ones has six short and a much longer distal article; the outer lateral ceratophores have only five short articles in addition to the distal long one. The long ceratostyles are tapering; the median is the longest and reach setiger 11. The peristomium is longer than the succeeding segments; its cirri are long, smooth and tapering; each is nearly twice as long as the peristomium. The first five pairs of parapodia (Fig. 3a) are en- larged; the first are at the sides of the prostomium, and the others are increasingly ventral so that the two of a pair nearly meet medially and are separated by a quadrate mid-ventral pad. Parapodia are sub- biramous; the notopodia are represented by a fascicle of slender acicula projecting into the long, tapering dorsal cirri; the ventral cirri are similar, but only about half as long (Fig. 3b). The base of the first parapodium is transversely wrinkled as though capable of great lateral extension. Pre- and postacicular lobes are distally prolonged to extend beyond the setal tips. Setae number twelve to fourteen in a fascicle; each is a composite falciger (Fig. 3c) terminating on a bifid tip and a short hood. These falcigers are accom- panied by three to five simple, capillary setae re- sembling the imbedded notacicula. The second, third, and fourth parapodia are more modified and directed forward on the ventrum; their setae resemble those in the first setiger. The ventral cirri are cirriform in these setigers; thereafter they are replaced by thick pads. Branchiae are first present from setiger 6 with three filaments in a pectinate arrangement. The maximal number of branchial filaments is seven and is found on the twelfth branchial setiger (Fig. 3d). The dorsal cirri in these setigers are short and tapering. Parapodia behind the modified setigers have long, limbate setae and one or two yellow. subacicular hooks, first present from setiger 15; they are distally bifid. Pectinate setae are numerous: each is distally flared and terminated in many short dentitions (Fig. 3e). Acicula number two or three in a ramus: each is yellow, nearly straight and terminates in a slightly acute tip. The maxillary apparatus (seen in dissection) is well developed. Maxilla I is the forceps: maxilla II POLYCHAETES FROM CENTRAT AMERICAN BEACHES 23 right; TIE has four left has ten teeth left and twelve fourteen left and nine right; [VY has the right-hand side; V has Distribution: Americonuphis reesei is and is absent on one tooth on each side known from two localities in the Gulf of Panama muddy bottoms in 27 m, the one in other intertidally in a sandy bottom. Remarks: Americonuphis reesei differs from A 1956) in that it anterior modified setigers rather than six. live The composite falcigers are bifid in the former and hartmanae (Friedrich, has entire in the latter. In addition, A. reesei differs from A. magna in the number of teeth on the different jaw-pieces and reesei has about six, rather than twelve branchial filaments where the branchiae are best developed. Americonuphis hartmanae (Friedrich, 1956), new combination Rhamphobrachium hartmanae Friedrich, 63-65, fig. Sa—c. Distribution: 1956, pp. Playa de las Flores, El Salvador in an intertidal sandy beach. Remarks: The specimens described by Fried- rich (1956) appear to have been as large as those described for A. reesei. The first six parapodia are enlarged and ventrally directed; parapodia 7 and 8 are transitional; the remainder of the parapodia are not modified and are lateral in position. Eyes are absent and the median occipital tentacle reaches back to setiger 5; its ceratophore has eight rings; each of the inner lateral occipital tentacles has twelve or thirteen basal rings. The modified anterior parapodia have equally long dorsal and ventral cirri and postsetal lobes. These parapodia are armed with weak simple superior setae and about four dark brown, thick, distally curved composite hooks. From setiger 7 the postsetal lobe is short. Branchiae are first present from this setiger as a single filament: the maxi- mum number of branchial filaments is five-seven at setiger 12. Dorsal cirri are shorter than the branchial filaments; has a curved fleshy spur at the base. Anterior ventral cirri are long through seven setigers and are absent posterior to each setiger 18. Postmodified setigers have numerous limbate setae and a pair of light brown subacicular hooks. Pectinate setae are present from setiger 11 and number three or four in a fascicle. Diopatra obliqua Hartman, 1944 Diopatra obliqua Hartman, 1944a, pp. 57-61. pl. 2 figs. 24-36, pl. 16, figs. 331-333: Fauchald, 1968 pp: 9-10, pl. 2, fig. a. 24 BULLETIN SOUTHERN CALIFORNIA Material examined: Naos Island, 29 August 1969 (1). Distribution: Diopatra obliqua is common along the coast from Golfo de California, Mexico to Peru in the eastern Pacific Ocean. Remarks: —Diopatra obliqua hooded hooks in anterior setigers and strongly has bidentate oblique pectinate setae. Details of the first para- podia were described by Fauchald (1968). Diopatra splendidissima Kinberg, Diopatra splendidissima Hartman, 1944a, pp. 56-57, pl. 1, figs. 21-23; Fauchald, 1968, pp. 12-13, pl. Dll Sa) Material examined: (1). Distribution: Diopatra splendidissima is known from southern California to Equador in the eastern Pacific Ocean. Remarks: Diopatra splendidissima has biden- tate hooded hooks in the anterior setigers; the pectinate setae are transverse; each has a few coarse teeth. A description of the first parapodia was added by Fauchald (1968). Venado Beach, 14 July 1969 Family Lumbrineridae Lumbrineris monroi Fauchald, 1970 Lumbrineris monroi Fauchald, 1970, pp. 99-102, pl. 16, figs. e-i. Material examined: Naos Island, 30 July 1969 (1). Distribution: Lumbrineris monroi is known from western Mexico in shallow water; the present record is from a sandy beach in Pacific Panama. Remarks: The present specimen differs from the species as originally described in that the simple hooded hooks are present from setiger 12 rather than from setigers 18—24 and in that the posterior postsetal lobes are slightly prolonged. The specimen is similar in size to the holotype. Lumbrineris zonata (Johnson, 1901) Lumbriconereis zonata Johnson, 1901, pp. 408-409, pl. 9, figs. 93-100. Lumbrineris zonata Hartman, 1944a, pp. 146-147; Fauchald, 1970, pp. 112-113, pl. 18, figs. e-i. Material examined: Venado Beach, 14 July 1969 (Gil), Distribution: Lumbrineris zonata was described from Washington on the west coast of North America and has previously been reported as far south as Baja California. The present record is from the Pacific coast of Panama. Remarks: Lumbrineris zonata has hooded hooks present from the first setigers and the post- setal lobes are short in all setigers. The present ACADEMY OF SCIENCES VOLUME 72 specimens do not appear to differ from specimens found in southern California or off western Mexico. Family Orbiniidae Orbinia johnsoni (Moore, 1909) Aricia johnsoni Moore, figs. 30-33. Orbinia johnsoni Hartman, 1969, pp. 33-34, 4 figs. Material examined: Samara, 8 April 1971 (13). Distribution: Orbinia johnsoni is previously known from central and southern California and from Costa Rica, where the present record is from. Remarks: Orbinia johnsoni has the transition between thorax and abdomen between setigers 16-19 according to Hartman (1969). The pres- ent specimens all have this transition at setiger 18. Branchiae are present from setiger 15 in all speci- mens. The ventral row of papillae on the transi- tional setigers is rather poorly developed. 1909, pp. 260-262, pl. 8, Family Paraonidae Paraonides platybranchia (Hartman, 1961) Paraonis platybranchia Hartman, 1961, pp. 86—87. Paraonides platybranchia Hartman, 1969, pp. 73-74, 2 figs. Material examined: Naos Island, 30 June 1969 (154). Distribution: Paraonides platybranchia was origi- nally described from southern California; the present record is the first from other areas and may indicate a wider distribution in the Pacific Ocean. Remarks: The present specimens agree with P. platybranchia in that they have branchiae pres- ent from setiger 4 through setigers 29-30. They lack antennae and modified setae. Family Spionidae Dispio uncinata Hartman, 1951 Dispio uncinata Hartman, 1951, pp. 87-90, pl. 22, figs. 1-5, pl. 23, figs. 1-4; Hartman, 1969, pp. 105— 106, 3 figs; Foster, 1971, pp. 73-78, figs. 161-174. Material examined: Naos Island, 30 June 1969 (4); Shimmey Beach, 15 August 1969 (1). Distribution: Dispio uncinata is known from both coasts of North and Central America in shallow water and intertidal areas in sand. Remarks: Foster (1971, pp. 76-78) discussed the possible relationship between the present spe- cies, D. schusterae and D. remanei Friedrich (1956) from El Salvador. The three species are very similar, but synonymizing the three species must await a re-examination of the type materials of all three. 1973 Paraprionospio species indeterminable Material examined: Naos Island, 30 June 1969 (1). Remarks: The present specimen cannot be identified to species. It is small and rather. badly preserved; two pairs of pennate brachiae are present starting on the third setiger; scars from branchiae on other segments identified. Foster (1971, pp. 79-112) revised the species usually assigned to Prionospio distributing them could not be on several genera. The characters used were the number and = structure of the branchiae. This makes generic identification of poorly preserved material difficult on occasion, but is probably the best alternative available. Foster (1971, p. 112) also synonymized all species assigned to the genus Paraprionospio into a single species, P. pinnata (Ehlers, 1901, pp. 163— 164). It is not clear from Foster's article that she had material of all the different species involved. Synonymies based solely on descriptions are not acceptable to the present author; experience has shown that differences often exist between popula- tions of closely related species, and that these dif- ferences often have escaped description. It is thus necessary to compare materials directly, preferably of course type materials, but if such are not available, at least materials from the type locality or the immediate vicinity. Scolelepis agilis (Verrill, 1873) Nerinides agilis Hartman, 1956, p. 291. Material examined: Airport Beach, 19 March 1971 (28); Boca Barrancas, 13 March 1971 (73); Boca Grande 28 April 1971 (11); Cahuita South, | April 1971 (2); Naos Island, 30 June 1969 (52); La Punta, 14 March 1971 (418); Playa Juan Chaco, 9 May 1971 (22); Pradomar Beach, 21 April 1971 (1); Puerto Viejo, 2 April 1971 (4); San Blas, 18 August 1969 (4); Shimmey Beach, 10 August 1969 (101). Distribution: Scolelepis agilis is known from the western Atlantic coasts from New England to Central America; the present records are also from the Pacific side of Central America. Remarks: Foster (1971, pp. 59-63, figs. 118— 131) following Pettibone (1963, p. 92) synony- mized this species with the European S$. squamata (Miller, 1806, for complete reference see Petti- bone, 1963). It is unclear from Foster and Petti- bone whether they examined any material from European waters or used the current descriptions as the base for this synonymy. Until this point has been cleared up, it seems preferable to keep materials from the Americas separate from the POLYCHAETES FROM CENTRAT AMERICAN BEACHES European materials, even if the descriptions overlap. Spionidae, species indeterminable Material examined: Playita Blanca, 9 February 1971 (4), Remarks: The present specimens are very badly preserved and can be recognized as spionids only on the distribution of the setae hooks are bifid. The hooded Family Magelonidae Magelona riojai Jones, 1963 Magelona riojai Jones, 1963, pp. 9-14, figs. 22-35. Material 27 March 1971 (3). Distribution: Magelona riojai was originally de from Veracruz, Mexico, Florida, USA. The present record is from Jaco, Costa Rica on the Pacific side of Central America. Remarks: The present specimens fit very well with M. riojai; the anterior end is somewhat more truncate than as illustrated by Jones (1963, fig. 22) and the mucronate tips of the modified setae of setiger 9 are slightly slenderer than the ones illustrated by Jones. examined: Jaco, scribed and Family Cirratulidae Cauleriella alata (Southern, 1914) Chaetozone alata Southern, 1914, pp. 112-113, pl. 12. figs. 27a—d. Cauleriella alata Hartman, 1969, pp. 225-226, 3 figs. Material examined: Naos Island, 30 July 1969 (8), 29 August 1969 (17). Distribution: Cauleriella alata has been reported from sandy beaches from worldwide areas. The rela- tionship between the different populations is unknown and it is doubtful that they can be adequately char- acterized on morphological characters alone. Remarks: The present specimens agree with C. alata in that they have bifid hooks present from the first neuropodium and have a pair of deeply imbedded black eyes. The bifid condition of the hooks is less obvious than as originally described, but is distinct in all setigers. Cirratulus 2? cirratus (Muller, 1776) Cirratulus cirratus Hartman, 1969, pp. 245-246, 2 fi Material examined: Naos Island, 30 June 1969 29 August 1969 (2). Distribution: Cirratulus in relatively shallow water: mostly in soft bottoms. Remarks: The present specimens belong to the cirratus is cosmopolitan genus Cirratulus in that they have transverse rows 26 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 1973 of tentacles on the first setiger and acicular spines present from an anterior setiger. Neuropodial spines are first present from setiger 12-14 rather than from setigers 6-11 as in C. cirratus. This character is considered important in identifying the different species of Cirratulus, so the present specimens are tentatively assigned to C. cirratus. Family Flabelligeridae Piromis americana (Monro, 1928) Stylarioides capensis americana Monro, 1928, pp. 96— 97, fig. 16; Monro, 1933b, pp. 1057-1058, text fig. 6. Piromis americana’ Hartman, 1969, pp. 305-306, 3 figs. . Material examined: San Carlos, 4 August 1969, in tubes under rock at low tide (1). Distribution: Piromis americana is known from Panama to central California. Remarks: The present specimen agrees with those described by Monro, except that the neuro- setae are somewhat less distinctly striated. Family Opheliidae Armandia ? bioculata Hartman, 1938 Armandia bioculata Hartman, 1938, pp. 105—106, figs. $1—54; Hartman, 1969, pp. 323-324, 3 figs. Material examined: Naos Island, 30 July 1969 (2). Distribution: Armandia bioculata is known from Washington to California in sandy intertidal areas. The present records come from Panama in a similar environment. Remarks: Both specimens are incomplete pos- teriorly; they resemble in all identifiable char- acters specimens of A. bioculata from California. Euzonus (Thoracophelia) furciferus (Ehlers, 1897) Thoracophelia furcifera Ehlers, 1897, pp. 101-103, pl. 7, figs. 164-167. Euzonus (Thoracophelia) furciferus Hartman, 1959, p. 431. Material examined: Playa Espadilla, 1 March 1971 (2). Distribution: Euzonus furciferus was originally described from intertidal areas in Punta Arenas, Chile: the present record comes from a similar environment in Pacific Costa Rica. POLYCHAETES FROM CENTRAL AMERICAN BEACHES 27 Remarks: The present two specimens fit with the species as originally described, including the numbers of setigers in the different parts of the body, the shape of the branchiae and the number and shape of the anal cirri. Family Capitellidae Notodasus dexterae, new species Figure 2b-—f 1969 Material examined: (30 TYPE): Description: Naos Island, 30 July The holotype is an incomplete speci- men with 29 setigers that is 17 mm long and 1 mm wide. The anterior end is inflated; the rest of the body is cylindrical. The specimens are greyish yellow in color. The peristomium and the first five setigers are strongly areolated (Fig. 2b); the remainder of the thoracal setigers less so. All thoracal segments are biannulated; the abdominal ringed. The prostomium is a flattened conical structure: the peristomium is considerably wider and is an- teriorly inflated. All eleven thoracal setigers are biramous and the setal bundles emerge from deep pockets. The two first abdominal setigers have short, conical parapodia; both noto- and neuropodia are similar aad setae emerge from conical protuberances rather than from pockets as in the thorax. All later abdominal setigers (Fig. 2e) are similar: the neuropodia are long welts that are separated ventrally only by the ventral nerve cord. The neuropodia are long enough laterally to be visible from the dorsal side. The large. nephridial papillae above the neuropodia are on ‘he dorsal side, and medial to these papillae near the dorsal midline, are two slightly raised ridges which represent the notopodia. These ridges have setae only in the abdominal parapodia 3 to 14: further posteriorly, the notopodia are completely reduced. In these posterior setigers, (Fig. 2f), the nephridial papillae are connected across the dorsum by a ciliated ridge. All thoracal setigers and the first two abdominal setigers have pointed, capillary setae only. The re- maining abdominal setigers have hooded uncini only. Each uncinus (Figs. 2c-d) has a large main fang and a crest of nine smaller teeth. These teeth are arranged segments are single- so that there are five in a lower row, three in the next row and a single tooth above the others. The margin of the hood is smooth. 2 Figure 3. b, first parapodium, anterior view, Americonuphis reesei, new genus, new species: a, anterior end, ventral view. SAS Ce distal end of composite falciger from parapodium, lateral view, x 540: d, branchial parapodium, anterior view, 46: e, pectinate seta, full view, x 1090. TABLE I. Species I. SPECIES FOUND IN ATLANTIC BEACHES Pisionidens indica Hermodice carunculata Il. SPECIES FOUND IN PACIFIC BEACHES Pisione remota Anaitides near multiseriata Glycera abranchiata Nephtys singularis Americonuphis reesei Diopatra obliqua D. splendidissima Lumbrineris monroi L. zonata Orbinia johnsoni Paraonides platybranchia Magelona riojai Cauleriella alata Cirratulus 2cirratus Piromis americana Armandia ?bioculata Euzonus (Thoracophelia) furciferus Notodasus dexterae Chone minuta 28 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 Geographical distribution of polychaetes reported from Central American sandy beaches. Previously known distribution Circumtropical Western Atlantic, warm waters Cosmopolitan Eastern Pacific, warm waters Western Atlantic, warm waters Eastern Pacific, warm waters Eastern Pacific, warm waters Eastern Pacific, warm waters Eastern Pacific, warm waters Eastern Pacific, warm waters Washington-Baja California Eastern Pacific, warm waters Eastern Pacific, warm waters Western Atlantic, warm waters Cosmopolitan Cosmopolitan Eastern Pacific, warm waters Washington-Southern California Chile Eastern Pacific, warm waters Central and southern California Ill. SPECIES FOUND ON BOTH SIDES OF THE ISTHMUS OF PANAMA Sthenelais maculata Ceratonereis mirabilis Hemipodus armatus Dispio uncinata Scolelepis agilis Eastern Pacific, warm waters Circumtropical Western Mexico The Americas, warm waters Western Atlantic, warm waters Distribution: Notodasus dexterae is known from intertidal sands on an island off Panama. Remarks: The genus Notodasus was previously represented by one species; N. magnus which was described from deep water off western Mexico by Fauchald (1972:246-247, pl. 51, fig. a-c). Notodasus magnus has notopodia developed in all abdominal setigers and the neuropodium is absent in the first thoracal setiger whereas, N. dexterae lacks notopodia in posterior abdominal setigers and all thoracal setigers are complete. It gives me pleasure to name this species after Deborah Dexter of California State University, San Diego who collected this very interesting material. Family Sabellidae Chone minuta Hartman, 1944 Chone minuta Hartman, 1944b, pp. 280-281, pl. 23, figs. 50-52, pl. 24, figs. 59-60; Hartman, 1969, pp. 671-672, 5 figs. Material examined: Naos Island, 29 August 1969 (8); Playita Blanca, 9 February 1971 (1). Distribution: Chone minuta is known from central and southern California in rocky areas. The present records are from Costa Rica and Panama in sandy beaches, but as indicated above, the specimens cannot be separated from C. minuta as presently described. Remarks: The present specimens agree with C. minuta in that the spatulate setae are sharply mucronated; the radioloi are united very high up on the crown and the number of segments is similar to that described for the species from central California. DISCUSSION Twenty-six species were found in the present col- lections; nineteen were in material from the Pacific Ocean only (Table 1) and of these, four- teen have their total distribution limited to the eastern Pacific Ocean. Three species are pre- sumed to be cosmopolitan, but have not been 1973 found on the western Atlantic coasts of Central America. ‘Two species previously were recorded from the Caribbean Sea only, but we did not find them in our collections from the Atlantic coasts. ‘Two species have been found in beaches on the Atlantic coast only; one of these appears limited to warm water in the Atlantic Ocean; the other has a reported circumtropical distribution, but was absent from our eastern Pacific collec- tions. Five species have been found in sandy beaches on both sides of the isthmus of Panama; two were previously known only from the eastern Pacific, one had been reported only from western Atlantic areas, another has been known from both coasts of the Americas, and one is circumtropical. A total of 24 species were found in the Pacific beaches and only seven in the Atlantic beaches. This difference may appear puzzling since the eastern Pacific Ocean usually is considered im- poverished when compared to other faunal areas at similar latitudes (Ekman, 1953:39-44). This is demonstrable for hard-bottom faunas, e.g. coral reefs, but a close investigation of the lesser known soft-bottom areas may very well change our views of the faunal relations between the two oceans. Under some circumstances, the difference in numbers of species of polychaetes might be due to a sampling bias; this cannot be the case in this instance since the same investigators sampled beaches in both areas. Dexter (1972) discussed the community struc- ture of two Panamanian beaches, one on each side of the isthmus. Some of her material is treated above. Dexter demonstrated that the varia- tion in the physical parameters was greater on the Pacific coast than on the Atlantic coast due to upwelling and that this upwelling had been associated with a greater primary productivity on the west coast. In terms of the organisms present, Dexter’s survey shows that the whole fauna is distributed essentially as is the polychaete fraction with the numbers of species and the biomass very much higher on the Pacific than on the Atlantic coast. The species diversity is lower on the Pacific coast than on the Atlantic side. This lower species diversity is probably related to the lower stability in physical parameters on the Pacific coast. The higher numbers of species of polychaetes and the higher biomass probably are related to the higher productivity on the west coast. There is no contradiction between this re- sult and the previous findings of a depauperized fauna on the Pacific side. The earlier conclusion was based on the coral reef fauna and the tem- POLYCHALTES FROM CENTRAL AMERICAN BEACHES 29 perature variation on the Pacific coast is so great (Dexter, 1972:449) that only a limited number of reef species are able to survive. Species burrow sensi hard ing in sand and mud are apparently not as tive to temperature fluctuations as their bottom relatives. ACKNOWLEDGMENTS The author is very grateful to Deborah M. Dexter, Olga Hartman made the first description of Americonuphis reesei and allowed who collected the material. the author to use this as the base for the present description. The excellent drawings of this species were made by Anker Petersen. LITERATURE CITED Aiyar, R. G., and K. H. Alikunhi. 1940. On a new pisionid from the sandy beach, Madras. Rec. Indian Mus., Calcutta, 42:89-107. 1943. Change of the generic name Pisionella Aiyar and Alikunhi, 1940, into Pisionidens (Polychaeta). Current Sci., Banga- lore, 12:120. Andrews, E. A. 1891. Report upon the Annelida Polychaeta of Beaufort, North Carolina. Proc. U.S. Nat. Mus., 14:277-302. Day, J. H. 1967. A monograph of the polychaeta of Southern Africa. Publ. British Mus. (Nat. Hist.), 656: XXIV and XVI, 878 pp. Dexter, D. M. 1972. Comparison of the Commu- nity Structure in a Pacific and an Atlantic Pana- manian sandy beach. Bull. Mar. Sci., 22:448—462. Ehlers, E. 1897. Polychiéeten. haenischen Sammelreise. Hamburg, 148 pp. Hamburger Magal- Friedrichsen & Co., 1901. Die Polychaeten des magellanischen und chilenischen Strandes. Ein faunistischer Versuch. Festschrift fiinfzigjahrigen Bestehens des sellschaft der Wissenschaften zu (Abh. Math.-Phys.) | Wiedemannsche handlung, Berlin, 232 pp. zur Feier der Hundert- k6niglichen Ge- GOttingen. Buch- Ekman, S. 1953. Zoogeography of the wick and Jackson, Ltd.. London, XIV ar Fauchald, K. 1968. from western Mexico. Biol., 3:1-82. Onuphidae (Polychaet fron Allan Hancock Monogr. Mar. 30 BULLETIN SOUTHERN CALIFORNIA 1970. Polychaetous annelids of the fam- ilies Eunicidae, Lumbrineridae, Iphitimidae, Arabellidae, Lysaretidae and Dorvilleidae from western Mexico. Allan Hancock Monogr. Mar. Biol., 5:1-335. 1972. 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Polychaetous Aphroditidae to Pisionidae. cific Exped., 7: 1-156. annelids. Pi 25 Allan Hancock 1940. Polychaetous Chrysopetalidae to Goniadidae. Pacific Exped., 7:173-287. 1944a. Polychaetous annelids. Pt. 5. Eu- nicea. Allan Hancock Pacific Exped., 10:1—238. 1944b. Polychaetous annelids from Cali- fornia, including the description of two new genera and nine new species. Allan Hancock Pacific Exped., 10:239-310. 1950. phtyidae. 1-181. Goniadidae, Glyceridae and Ne- Allan Hancock Pacific Exped., 15: 1951. The littoral marine annelids of the Gulf of Mexico. Publ. Inst. Mar. Sci., Texas, 2:7-124. ACADEMY OF SCIENCES VOLUME 72 1956. Polychaetous annelids erected by Treadwell, 1891 to 1948, together with a brief chronology. Bull. Amer. Mus. Nat. Hist., 109: 239-310. ——. 1959, 1965. Catalogue of the polychaetous annelids of the world. Allan Hancock Founda- tion Occas. Paps., 23:1-628, 1959; Supplement and index, 1-197, 1965. 1961. Polychaetous annelids from Califor- nia. Allan Hancock Pacific Exped., 25:1—226. 1968. Atlas of errantiate polychaetous an- nelids from California. Allan Hancock Founda- tion, 828 pp. 1969. annelids from California. dation, 812 pp. Atlas of sedentariate polychaetous Allan Hancock Foun- Hartmann-Schréder, G. 1959. Zur Okologie der Polychaeten des Mangrove-Estero-Gebietes von El Salvador. Beitr. Neotrop. Fauna, 1:69-183. Johnson, H. P. 1901. The polychaeta of the Puget Sound region. Proc. Boston Soc. Nat. Hist., 29: 381-437. Jones, M. L. 1962. lids from Jamaica, The West Indies. Mus. Nat. Hist., 124:169-212. On some polychaetous anne- Bull. Amer. 1963. Four new. species of Magelona (Annelida, Polychaeta) and a redescription of Magelona_ longicornis Johnson. Amer. Mus. Novit., 2164:1-31. Kinberg, J. G. H. 1865. Annulata nova. Ofv. Vet. Akad. Stockholm, Foérh., 21:559-574. 1866. Annulata nova. Ofv. Vet. Akad. Stockholm, Fo6rh., 22:167-179. Laubier, L. 1967. Quelques annélides polychétes interstitielles d’une plage de Cote d’Ivoire. Vie et Milieu, Ser. A, 18:573-594. Monro, C. C. A. 1928. On the polychaeta col- lected by Dr. Th. Mortensen off the coast of Panama. Vidensk. Medd. dansk Naturh. Foren., 85:75-103. 1933a. The polychaeta errantia collected by Dr. C. Crossland at Colon in the Panama region and the Galapagos Islands during the ex- pedition of the S. Y. St. George. Proc. Zool. Soc., London, 1933:1—96. 1973 POLYCHAELTES FROM CENTRAL AMERICAN BEACHES 4] 1933b. The polychaeta sedentaria collected Rioja, E. 1941, Estudios anelidologicos Il. Dato by Dr. C. Crossland at Colon in the Panama para el] conocimiento de les fauna de poliqueto region and the Galapagos Islands during, the ex- de las costas del Pacifico de Mexico. Ann. Inst pedition of the S. Y. St. George. Proc. Zool, Soc., London, 1933:1039-1092, Moore, J. P. 1909. annelids from Monterey Bay and San Diego, California. Acad. Nat. Sci., Philadelphia, 61:235—295, Polychaetous Proc. Miiller, O. I 1776. seu Animalium Daniae et Norvegiae indigenarum Zoologica Danicae Prodromus characters, nomine et synonyme imprimis popu- larium. Havniae, XXXIL and 274 pp. Miscellanea Zoologica. Pallas, P. S. 1766. Hagae, p. 77. Pettibone, M. H. 1963. Revision of some genera of polychaete worms of the family Spionidae, including the description of a new species of Scolelepis. Proc. Biol. Soc. Washington, 76: 89-104. Biol, Mexico, 12:669-746 Siewing, R. 1954. Zur Verbreitung indica Aiyar and Alikunhi. Kieler Meeresfor 10:81—-83. on Pisioniden schungen, Southern, R. 1914. Clare Island Survey. Dublin, 31:1—160. Archiannelida and polychaeta Proc. Roy. Irish Acad Treadwell, A. L. 1901. The polychaetous annelids of Porto Rico. Bull. U.S. Fish. Comm., 20 181-210. Verrill, A. E. 1873. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, with an account of the physical characters of the region. Rept. U.S. Fish. Comm., 1871- 1872:295-778. Accepted for publication January 2, 1973. POLYCHAETOUS ANNELIDS COLLECTED BY THE R/V HERO FROM BAFFIN ISLAND, DAVIS STRAIT, AND WEST GREENLAND IN 1968 JAMES A. BLAKE! ABSTRACT: the eastern Canadian Arctic. AND Davip DEAN? Seventy-one species of polychaetes are reported from four benthic stations in Two new species were found and described by Blake, 1972. Myriochele oculata was found to be abundant off Baffin Island and is redescribed and com- pared with M. heeri; one new synonym is presented. Aspects of the faunal analysis are discussed and compared with other Arctic areas. Benthic ecology has been studied in east Green- land fjords (Thorson, 1933, 1934, 1944; Sparck, 1933; Madsen, 1936; Bertelsen, 1937; Degerbgl, 1937; and Ockelman, 1958), from scattered points along the west coast of Greenland (Vibe, 1939, 1950; Madsen, 1940; and Petersen, 1962) and the Baffin Island-Labrador coast (Packard, 1867, 1891; Soper, 1928; Ellis and Wilce, 1961; see Ellis, 1960 for further references). All of this benthic ecological work has been conducted in inshore areas (fjords, sounds, inlets, etc.); prac- tically nothing is known about the biology of the fauna inhabiting the continental shelves. Of par- ticular interest is the polychaete component. Our knowledge of the polychaete fauna of this general area has been supplemented by several workers. especially Packard (1863, 1867), Moore (1902. 1909), Thorson (1936), Treadwell (1937). Berkeley and Berkeley (1943). Wesenberg-Lund (1934, 1947, 1948, 1950a, 1950b, 1951, 1953). Vibe (1950), Grainger (1954), Pettibone (1956) and Curtis (1970). Thorson (1934) found that infaunal biomass generally decreased with increas- 1 Pacific Marine Station, University of the Pacific. Dillon Beach, California 94929. 2Ira C. Darling Center, =<52 Walpole, Maine 04573. University of Maine. 32 BULLETIN SOUTHERN CALIFORNIA ing depth, that lamellibranchs dominate the fauna of shallow water, and that polychaetes dominate in deeper water. Ellis (1960) substantiates Thor- son’s The polychaete the deeper waters of the Davis Strait and the Labrador findings. fauna of Sea is inadequately known. At least 134 species of polychaetes have been reported from the eastern Canadian Arctic and subarctic waters (Grainger, 1954) and from west Greenland (Wesenberg-Lund, 1950a). During August of 1968, the University of Maine participated in the shakedown cruise of the Na- tional Science Foundation’s new polar research vessel, the R/V HERO, to the eastern Canadian Arctic. Invertebrate studies were conducted by the University of Maine in the Labrador Sea and Davis Strait. Although the primary mission of the cruise was to test the vessel in Arctic waters, it was possible to sample 32 biological stations in Arctic and subarctic waters. Four of these stations were benthic and the remainder planktonic. Two brief accounts of the cruise and progress in analy- sis of the data have been published (Dearborn and Dean, 1969; 1970). Another benthic station (Number 33), sampled near the end of the cruise in the Gulf of Maine, is not included. The present paper reports on the benthic poly- chaetous annelids collected during the cruise. In subsequent papers other personnel will report on the echinoderms, molluscs, zooplankton, and phy- toplankton. Station data and polychaete species obtained at the benthic stations are given below. The four benthic stations include use of both standard and specialized gear. Standard gear included a Ponar type grab (0.1 m*), Peterson grab (0.1 m?), Rock dredge, Triangular dredge, Tri-net trawl, Mop tangle, and underwater camera. Specialized gear included a modified Beyer Epibenthic | sled (Holme, 1964) equipped with a Clarke-Bumpus closing device and a “mouse-trap” sampler (Muus, 1964). The Epibenthic sled was towed along the bottom to obtain a plankton sample from about 0.5 m above the bottom, while the “mouse-trap” was used to collect a 225 cm? sample of the top 1—2 cm of bottom substrate. Polychaetes were screened on board ship and preserved as soon as possible after collection. Propylene phenoxytol (0.15%) was used as a nar- cotizing agent and either formalin or Bouin’s fluid After fixation the ma- terial was preserved in 70% ethyl alcohol. Identifi- was used as a fixative. ACADEMY OF SCIENCES VOLUME 72 cation and sorting were done at the Ira C. Darling Center during 1968-1969. The collections remain at the Ira C. Darling Center, pending completion of the remaining in- vertebrate studies. At that time, the entire collec- tion will be transferred to the United States National Museum, Washington, D.C. RESULTS Seventy-one species of Polychaeta are distributed in 27 families; two species are new to science. Details of these species have been presented in another paper (Blake, 1972). The Polynoidae and Sabellidae are best represented in the collec- tions with seven and eight species, respectively. The names and numbers of polychaetes taken at the four benthic stations are presented below. A systematic list summarizes the species collected. LIST OF BENTHIC STATIONS BENTHIC POLYCHAETA Hero Cruise No. 3 AuGust 1968 STATION 20 East Angiak Island (Baffin Island Region); lat. 65°43’ N—long. 62°05’ W; 14 August 1968, 1230-1700 hours EDT (Zone 4); sediment analysis: a well-sorted (So— 1.9) brown-grey sticky mud with a mean particle size of 150u. Gear: Ponar grab (A and B); Mop tangle and Triangle dredge towed in tandem (E). Depth, ranged from 132-245 m. POLYCHAETA: Antinoella sarsi E(1); Pholoe minuta A(7); Nephtys incisa A(1), B(1); Sphaero- dorum gracilis A(1); Scoloplos armiger A(2), B(3); Paraonis gracilis A(1); Prionospio steenstrupi A(3); Chaetozone setosa A(30), B(46), E(1); Cirratulus ciurratus B(1); Scalibregma inflatum A(2), B(1); maldanids A(2); Myriochele oculata A(36), B(47); Myriochele heeri A(25), B(12); Melinna elisabethae A(4), B(12); Lysippe labiata A(1); Glyphanostomum pallescens E(1); ampharetids A(2); Polycirrus me- dusa E(1); Euchone papillosa A(5), B(1), E(1). STATION 26 Southern Baffin Bay; lat. 67°49’ N— long. 60°46’ W to lat. 67°38’ N—long. 60°38’ W; 16 August 1968, 2330-1200 hours EDT (Zone 4); bot- tom temperature 0.1°C; sediment, thick sticky mud. Gear: Ponar grab (A), Peterson grab (B), Niskin cast (D), Triangular dredge (E), mouse trap sampler (F), Epibenthic sled and Triangular dredge in tandem (G). The Ponar and Peterson grabs (A and B) failed to operate, but mud was retained on the corners and edges of the grabs. This mud was seived for polychaetes as was mud retained on the weights and shackles of the Niskin cast. Depth, ranged from 1920-1745 m. 1973 POLYCHAETOUS ANNELIDS FROM EASTERN CANADIAN ARCTIC $3 POLYCHAETA; — Kteone flava’ A(1), BOL); POLYNOIDALI Pelagobia longicirrata A(1 was caught on screen Of — feynoe oerstedi Malmgren, 1865; 28. 29 grab as it was drawn through the water column); fapisca extenuata (Grube), 1840: 28, 29 Aglaophamus malmgreni G(1); Scoloplos sp. ACL); = Harmothoe imbricata (Linne). 1767. 29 Haploscoloplos. sp. BCL); Aricidea suecica FOS); Phyllochaetopterus sp. E(1); Chaetozone setosa A(6), ~ Gattyana nutti Pettibone, 1955: 29. New Recorp B(1); Brada villosa G(1); Cryptosclerocheilus baf- Aystrolacnilla mollis (Sars), 1871; 28 finensis D(2), E(1), G1); maldanids ACL), ECS), G(2); Myriochele heeri A( 1); Ampharete arctica G(1); Jasmineira schaudinni E(2), G(6). STATION 28 Davis Strait; lat. 66°28’ N—long. 57° 26’ W to lat. 66°29’ N—long. 56°42’ W; 17 August 1968, 0300-0900 hours EDT (Zone 4); bottom tem- perature 2.58°C; sediment, soft mud composed partly Gattyana cirrosa (Pallas), 1766; 29 Antinoella sarsi (Malmgren), 1865; 20 SIGALIONIDAE Pholoe minuta (Fabricius), 1780; 20 Neoleanira tetragona (Oersted), 1845; 28 of large agglutinated foraminifera. Gear: Tri-net EUPHROSINIDAE trawl (D). Depth, ranged from S80-610 m. Euphrosine borealis Oersted, 1843; 29 POLYCHAETA: Laetmonice — filicornis D(4); Eunoe oerstedi D(2); Lagisca extenuata D(8&); Aus- LOPADORHYNCHIDAE trolaenilla mollis D(1); Neoleanira tetragona D(1); Sphaerodorum gracilis D(3); Glycera capitata D(1); Lumbrineris latreilli D(3); Lumbrineris fragilis D(1); Lumbrineris fauchaldi D(7); Neothria conchylega D(many); Maldanella davisi D(2); Ampharete arctica. D(7); Glyphanostomum pallescens D(2); Eupolymnia sp. D(1); Pista cristata D(3); Terebel- lides stroemii D(2). STATION 29 Off Little Hellefiske Bank (West SYLLIDAE Greenland); lat. 64°57’ N—long. 58°38’ W; 18 August Typosyllis fasciata (Malmgren), 1867; 29 1968, 0800-1115 hours EDT (Zone 4); bottom tem- Typosyllis armillaris (Miiller), 1771; 29 perature 1.4°C; rocky bottom with encrusting epifauna = Exogone verugera (Clap.), 1868: 29 of hydroids and bryozoans. Gear: Niskin cast (A); 9 Sphacrosyllis sp.; 29 Tri-net trawl (C); Triangular dredge (F and G). — Syllid; 29 Depth, ranged from 79-85 m. POLYCHAETA: Eunoe oerstedi A(2 retained on NEREIDAE Niskin weights), Lagisca extenuata C(1), F(3); Nereis pelagica Linne, 1758; 29 Harmothoe imbricata G(1), F(4); Gattyana cirrosa F(11), G(4); Euphrosine borealis C(4), F(6), G(2); NEPHTYIDAE Eulalia bilineata C(4), G(1); Eumida spp. F(1), Nephtys ? incisa Malmgren, 1865; 20 G(1); Typosyllis fasciata C(10), G(6), F(3); Aglaophamus malmgreni (Théel), 1879: 26 Typosyllis armillaris C(4); Exogone verugera C(7): Sphaerosyllis sp. C(4); syllid C(1); Nereis pelagica C(9), F(6), G(1); Glycera capitata C(3); Nothria conchylega F(few), G(1); Spiophanes kroyeri C(1), G(2); Pygospio elegans C(2); Polydora concharum F(1); Polydora websteri G(1); Pherusa plumosa Pelagobia longicirrata Greef, 1879; 26 PHY LLODOCIDAE Eteone flava (Fabricius), 1780; 26 Eulalia. bilineata (Johnston), 1840; 29 Eumida spp.; 29 GLYCERIDAE Glycera capitata Oersted, 1843: 28, 29 SPHAERODORIDAE Sphaerodorum gracilis (Rathke), 1843; 20, 28 F(1); Thelepus cincinnatus C,F,G(many); Euchone LUMBRINERIDAE analis G(1); Chone duneri C(many); Chone in- Lumbrineris latreilli (Adouin and Milne-Edwards), fundibulariformis C(7); Potamilla neglecta C(3): 1833: 28 Potamilla sp. F(several); Branchiomma sp. F(2); 9 Lumbrineris fragilis (Miiller). 1776: 28 Serpula vermicularis C(many); Spirobis spirillum — Lumbrineris fauchaldi Blake. 1972: 28 C(many); Spirobis granulatus C(many). ONUPHIDAE Nothria conchylega (Sars), 1835: 28, 29 SYSTEMATIC LIST OF POLYCHAETOUS ANNELIDS AND COLLECTION STATIONS ORBINHDAE Scoloplos armiger (Miiller). 1776: 20 APHRODITIDAE Scoloplos sp. juvenile: 26 Laetmonice filicornis Kinberg, 1855; 28 Haploscoloplos sp.: 26 34 BULLETIN SOUTHERN CALIFORNIA PARAONIDAE Paraonis gracilis (Tauber), 1879; 20, New RecoRD Aricidea suecica Eliason, 1920; 26 SPLONIDAE Prionospio steenstrupi Malmgren, 1867; 20 Spiophanes kroyeri Grube, 1860; 29 Pygospio elegans Clap., 1863; 29 Polydora concharum Verrill, 1881; 29, New Record Polydora websteri Hartman, 1943 juvenile; 29 CHAETOPTERIDAE Phyllochaetopterus sp.; 26 CIRRATULIDAE Chaetozone setosa Malmgren, 1867; 20, 26 Cirratulus cirratus (Miller), 1776; 20 FLABELLIGERIDAE Pherusa plumosa (Miller), 1776; 29 Brada villosa (Rathke), 1843; 26 SCALIBREGMIDAE Scalibregma inflatum Rathke, 1843; 20 Cryptosclerocheilus baffinensis Blake, 1972; 26 MALDANIDAE Maldanella davisi Wesenberg-Lund, 1948; 28 Maldanids; 20, 26 OWENIIDAE Myriochele oculata Zaks, 1923; 20, NEW RECORD Myriochele heeri Malmgren, 1867; 20, 26 AMPHAERETIDAE Melinna elisabethae McIntosh, 1922; 20 Ampharete arctica Malmgren, 1865; 26, 28 Lysippe labiata Malmgren, 1865: 20 Glyphanostomum pallescens (Théel), 1879; 20, 28 Ampharetids; 20 TEREBELLIDAE Polycirrus medusa Grube, 1855; 20 Thelepus cincinnatus (Fabricius), 1780; 29 Eupolymnia sp.; 28 Pista cristata (Miller), 1776; 26, 28 TRICHOBRANCHIDAE Terebellides stroemii Sars, 1835; 28 SABELLIDAE Euchone papillosa (Sars), 1850; 20, NEw REcorpD Euchone analis (Kroyer), 1856; 29 Chone duneri Malmgren, 1867; 20 Chone infundibulariformis Kroyer, 1856; 29 ACADEMY OF SCIENCES VOLUME 72 Jasmineira schaudinni Augener, 1912; 26, New RECORD Potamilla neglecta (Sars), Potamilla sp.; 29 Branchiomma sp.; 29 1851; 29 SERPULIDAE Serpula vermicularis Linne, 1767; 29 Spirobis (Dexiospira) spirillum (Linne), 1758; 29 Spirobis (Laeospira) granulatus (Linne), 1767; 29 DISCUSSION The bottom types and depths were different at each of the four benthic stations. These differ- ences were dramatically reflected in faunal com- position at each station. Of the 71 species of Polychaeta, only 10 occurred at more than one station. No one species occurred at more than two localities. The following species shared sta- tions: Stations 20 and 26, Chaetozone setosa, Myriochele heeri; Stations 20 and 28, Sphaerodo- rum gracilis, Glyphanostomum pallescens; Sta- tions 26 and 28, Ampharete arctica, Pista cristata; Stations 28 and 29, Eunoe oerstedi, Harmothoe imbricata, Glycera capitata, Nothria conchylega. The benthic infauna was richest at Station 20, where sandy mud was favorable for grab sampling. Polychaetes were mostly sedentary tube dwellers or burrowers. The dominant species were Chaeto- zone setosa, Myriochele oculata, and M. heeri. Few polychaetes were retained in qualitative dredge hauls. These samples yielded large num- bers of echinoderms, especially asteroids and ophiuroids. One conspicuous asteroid, Ctenodis- cus crispatus, is a deposit feeder. Molluscs were generally small and few in numbers. This sug- gests a community structure dominated by infau- nal polychaetes and deposit-feeding echinoderms. The most interesting polychaete collections, from a taxonomic standpoint, come from south- ern Baffin Bay (Station 26). This locality, in depths ranging from 1745 to 1920 m, yielded one of the two new species and several range extensions. The occurrence of an unknown spe- cies of Phyllochaetopterus is also of interest as there are no other records of this genus from Arctic waters. Benthic polychaetes were mostly sedentary deposit feeders. The most conspicuous animals were Cryptosclerocheilus baffinensis, Jasmineira schaudinni, maldanids, and some large alcyonarian corals. Grabs failed to operate in the thick sticky mud. All polychaetes at this station were obtained by qualitative dredge or 1973 from mud which adhered to corners of grabs, shackles, or weights. Grabs were not used at Station 28 (Davis Strait). Qualitative dredge hauls yielded 17 spe- cies of polychaetes. Eleven of these were erran- tiate, predatory forms, whereas only six were sedentary tube dwellers. ‘The sediment was largely composed of large agglutinated foraminifera, The most abundant polychaete was the onuphid, Nothria conchylega. Station 29 (Little Hellefiske Bank) occurred off western Greenland. ‘Thirty-one species of polychaetes were taken of which 14 were seden- tariate. Most of the sedentary species, however, were species which attached their tubes to hard surfaces. The bottom type at this locality was rock and supported large numbers of erect bryo- zoa;, Thelepus cincinnatus was the dominant poly- chaete. Grabs were attempted at this station but were jammed by rocks. Comparisons with other regions: The most extensively studied areas in Arctic and subarctic seas are the western coasts of Greenland, the western Pacific and more recently the Ellesmere Island fjords (Canadian Northwest Territories). The polychaete fauna off Baffin Island is poorly known as compared with that of western Green- land. The present collections, though sparse, sug- gest that polychaetes in shallow waters off Baffin Island have remarkable affinities to those occupy- ing similar areas in the western Pacific. No less than 89 percent of the species taken off Baffin Island also occur in the western Pacific (data derived from Ushakov, 1965). This is a greater percentage of shared species than occurs off western Greenland (70 percent shared with the Baffin Island region, data derived from Wesenberg-Lund, 1951). A comparison of the Baffin Island polychaetes with those found by Curtis (1970) in the fjords of Ellesmere Island (N.W.T.), shows that only 53 percent of the Baffin Island species were found by him. This enhances the view that the Baffin Island fauna is indeed rich and deserves more extensive study. The deeper waters of Baffin Bay also warrant further study. The failure of grabs to operate in the thick clay sediments was disappointing in light of the interesting polychaetes which were taken by qualitative samples. Several transects from the shallow waters of Baffin Island to the deeper waters of Baffin Bay would be desirable and will be planned for future research. predatory New records and range extensions: The ma- jority of polychaetes taken in this study are widely POLYCHAETOUS ANNELIDS FROM EASTERN CANADIAN ARCTIC $5 Several records distributed in Arctic and boreal seas species were taken which represent new and range extensions for the areas sampled Among the more significant are: Gattayana nutti and Polydora concharum, new to western Green land (previously known only from Labrador and northeastern America); Paraonis gracilis, Myrio- chele oculata, and Euchone papillosa, new to the Baffin other circumpolar localities); Jasmincira schau- Island Region (previously known from dinni, new to Baffin Bay and areas west of Green- land (previously known from abyssal depths near Spitzbergen, Iceland, and eastern Greenland). Reproductive biology: Five species were noted to be ovigerous. Eggs were observed in the coelom of Neoleanira tetragona and Euchone papillosa. Exogone verugera contained eggs in a brood sac attached to the abdomen. Spirorbis spirillum was brooding eggs in the tube, while S. granulatus was brooding eggs in the operculum. SYSTEMATIC NOTES The majority of the species encountered are rela- tively well-known and do not require descriptive Lumbrineris fauchaldi and comment. Crypto- sclerocheilus baffinensis were described earlier (Blake, 1972). Several other species do merit further comment. FAMILY CHAETOPTERIDAE Phyllochaetopterus sp. An anterior fragment from Station 26E (South- ern Baffin Bay, 1800 m) measures 10 mm in length. The anterior region consists of 9 seg- ments. There are one or two modified spines on each side of setiger 4. Each spine is brown and terminates in a pointed tooth, which has an exca- vation on the tip. The species resembles P. monroi Hartman, 1967, from the Strait of Magellan. This appears to be the first record of Phyllochaetopte- rus from Arctic seas. FaMiILty OWENIIDAE Genus Myriochele Malmgren. 1867 Synonym: Galathowenia Kirkegaard, 1959. Galathowenia africana Kirkegaard. is herein considered synonymous with M. oculata Zaks. The criteria used to distinguish G. africana from other oweniids are not sufficient to distinguish 36 BULLETIN SOUTHERN CALIFORNIA Figure 1. ACADEMY OF SCIENCES VOLUME 72 Myriochele oculata Zaks, a. head region in ventral view; b. anterior end in lateral view; c. spinous notoseta from setiger 4; d, e. bifid uncinus from setiger 4. it from M. oculata. Kirkegaard’s specimens have more pigment, but this in itself is not sufficient for either generic or specific separation. Myriochele oculata Zaks, 1923 (Bigs 1) Myriochele oculata Zaks, 1923, pp. 171-174, figs. 1-3; Annenkova, 1937, p. 184; Ushakov, 1950, p. 214; 1955, p. 348, fig. 128h-i; 1965, p. 325, fig. 128h-i. Galathowenia africana Kirkegaard, 1959, p. 67, fig. 17. NEW SYNONYM. Numerous specimens were taken at Station 20: length up to 25 mm. The anterior end is truncate, has a midventral cleft but has no cephalic bran- chiae (Fig. la,b). There are two brownish eyes located on either side of the head. Paired bands of brown pigment are present dorsal to the eyes (Fig. 1b). Occasionally some weakly developed pigment bands may occur ventral to the eyes. All pigment bands may be absent. The head is compressed when the animal is preserved within its tube. If preserved out of the tube, the head is slightly inflated. The first three segments have notopodial fasci- cles only; from the fourth, parapodia are biramous with pointed setae in the notopodia and long handled uncini in the neuropodia. Notosetae are long and spinous (Fig. Ic). Uncini are bifid with the two nearly equal curved teeth, one set slightly higher than the second (Fig. ld,e). The uncini are small, numerous, ar- ranged in transverse rows and are present in 1973 \ Figure 2. Myriochele heeri Malmgren, a. spinous notoseta from setiger 4; b, c. bifid uncinus from setiger 4. parapodia to the posterior end. The tube is com- posed almost entirely of olivine and mica. The species was previously known from the Sea of Japan, Bering Sea, Chuckchee, Kara, Barents, and White seas; also from west Africa (as Galathowenia africana). This is a new record then from Baffin Island. POLYCHAETOUS ANNELIDS FROM EASTERN CANADIAN ARCTIC 47 Myriochele heeri Malmpren, 1867 (Fig. 2) Fauvel, 1927, pp. 204-205, fig. 7Ih-m (Syn- onymy); Wesenberg-Lund, 1950a, pp. 103-104; 1950b, p. 46. Specimens were obtained from Stations 20 and 26; length up to 22 mm, width up to 1.5 mm. The body is thick throughout, and has up to 30 segments. The anterior end is prolonged and has an anteroventral oral opening. In some specimens the cephalic segment is contracted and the oral opening is more anterior; there are no eyes. Setigerous segments 1-3 have only notopodial fascicles; thereafter parapodia are biramous. The uncini are long handled and bifid with two teeth, arranged one above the other (Fig. 2b and c). The notosetae are distally multiserrated (Fig. 2a). The posterior end terminates in two weakly de- veloped lateral lobes. below the anal opening. The tube is composed A small cirrus protrudes almost entirely of quartz grains. The species is widely distributed in Arctic boreal seas. ACKNOWLEDGMENTS This paper is contribution No. 4/7 from the Ira C. Darling Center for Research, Teaching and Service and contribution No. 29 from the Pacific Marine Station. The study was supported by National Science Foundation Grant GA-1405. We are grateful to John H. Dearborn and Robert C. Bullock for invaluable assistance in collection and preservation of the ma- terial while on board ship, and for other help during and after the cruise. We are also grateful to the administration of the Pacific Marine Station for allow- ing the first author time to complete the manuscript. LITERATURE CITED Annenkova, N. P. 1937. [The Polychaete fauna of the northern part of the Sea of Japan.]} Issledovaniya Morei S.S.S.R., No. 23:139-216. (In Russian.) Berkeley, E.. and C. Berkeley. 1943. Biological and Oceanographical conditions in Hudson Bay II. Polychaeta from Hudson Bay. J. Fish. Res. Bd. Canada, 6(2):129-132. Bertelsen, E. 1937. Contributions to the animal ecology of the fjords of Angmagssalik and Kangerdlugssuaq in East Greenland. Meddelelser om Grénland, 108(3):1—358. 38 BULLETIN SOUTHERN CALIFORNIA Blake, J. A. 1972. Two new species of Polychae- tous Annelid worms from Baffin Bay and the Davis Strait. Bull. So. California Acad. Sci., 71(3)) 127-132. Curtis, M. A. 1970. Depth distributions of benthic polychaetes in Hare Fiord and Tanquary Fiord, Ellesmere Island, N.W.T. Mar. Sci. Cen. Manus. Rept., McGill University, No. 16:1-73. 1969. of the Arctic In- United Dearborn, J. H. and D. Dean. vertebrate Studies. Antarctic J. States, 4(5):194-195. Invertebrates from the Davis Strait Antarctic J. of the United 1970. and Labrador Sea. States, 5(5):192. Degerbol, M. 1937. A contribution to the investi- gation of the fauna of the Blosseville coast, east Greenland, with special reference to zoogeogra- phy. Meddelelser om Grénland, 104(19):1—36. Marine infaunal benthos in Arctic Inst. N. Amer., Ellis, D. V. 1960. Arctic North America. Tech. Pap. No. 5:53 pp. Ellis, D. V., and R. T. Wilce. 1961. Arctic and Subarctic examples of intertidal zonation. Arctic, 14(4):224-235. Fauvel, P. 1927. Polychetes sedentaires. Addenda aux errantes, archiannelides, myzostomaires. Fauna de France, 16:1—494. Grainger, E. H. 1954. Polychaetous annelids of Ungava Bay, Hudson Strait, Frobisher Bay and Cumberland Sound. J. Fish. Res. Bd. Canada, 11(5):507—528. Hartman, O. 1967. Polychaetous annelids collected by the USNS Eltanin and Staten Island cruises, chiefly from Antarctic seas. Allan Hancock Monogr. Mar. Biol., No. 2:1-387. N. A. 1964. Methods of Adv. Mar. Biol., Holme, benthos. sampling the 2:171-260. Kirkegaard, J. B. Africa. 1959. The Polychaeta of West Atlantide Rept. No. 5:7-117. Madsen, H. 1936. Investigations on the shore fauna of East Greenland with a survey of the shores of other arctic regions. Meddelelser om Grénland, 100(8) : 1-79. 1940. A study of the littoral fauna of northwest Greenland. Meddelelser om Grégnland, 124(3):1-24. ACADEMY OF SCIENCES VOLUME 72 Moore, J. P. 1902. Descriptions of some new polynoidae with a list of other Polychaeta from North Greenland waters (and Alaska). Proc. Acad. Nat. Sci. Philadelphia, 54:258—278. 1909. The polychaetous annelids dredged in 1908 by Mr. Owen Bryant off the coasts of Labrador, Newfoundland, and Nova_ Scotia. Proc. U.S. Nat. Mus., 37(1703):133-146. Muus, B. J. 1964. the meiobenthos. A new quantitative sampler for Ophelia, 1(2):209-216. Ockelmann, W. K. 1958. Marine Lamellibranchi- ata. Meddelelser om Grgnland, 122(4):1-256. Packard, A. S. 1863. A list of animals dredged near Caribou Island, southern Labrador, during July and August, 1860. Canadian Nat. Geol., 8:401-429. 1867. Observations on the glacial phe- nomena of Labrador and Maine, with a review of the recent invertebrate fauna of Labrador. Mem. Boston Soc. Nat. Hist., 1:210-303. 1891. two summer cruises to that region. Hodges, Publisher, New York, 513 pp. The Labrador Coast. A journal of N.D.C, Petersen, G. H. 1962. The distribution of Balanus balanoides (L.) and Littorina saxatilis, Olivi, var. groenlandica, Mencke in northern West Greenland. Meddelelser om Grgnland, 159(9): 1-42. 1956. Proc. Pettibone, M. H. from Labrador. 531-585. Marine polychaete worms U.S. Nat. Mus., 105: Soper, J. D. 1928. A faunal investigation of south- ern Baffin Island. Bull. Nat. Mus. Canada, (Biol. Ser. 15) No. 53:1-26. Sparck, R. 1933. Contributions to the animal ecology of the Franz Joseph Fjord and adjacent East Greenland waters. I-II. Meddelelser om Grénland, 100(1):1-38. Thorson, G. 1933. Investigations on shallow water animal communities in the Franz Joseph Fjord (East Greenland) and adjacent waters. Med- delelser om Grgnland, 100(2):1-70. 1934. Contributions to the animal ecology of the Scoresby Sound fjord complex (East Greenland). Meddelelser om Grgnland, 100(3): 1-68. 1973 1936. The larval development, growth, and metabolism of arctic marine bottom in- vertebrates compared with those of other seas. Meddelelser om Grénland, 100(6):1-155,. 1944. Marine Gastropoda Prosobranchiata. Meddelelser om Grénland, 121(13):1-181. Treadwell, A. L. 1937. Polychaetous annelids col- lected by Captain Robert A. Bartlett in Green- land, Fox Basin, and Labrador. J. Washington Acad. Sci., 27(1) :23-36. Ushakoy, P. V. 1950. [Polychaeta of the Sea of Okhutsk.] Issledovaniya Dal-nevostochnykh Morei S.S.S.R., No. 2: 140-234 (In Russian. ) 1955. [Polychaeta of the Far Eastern Seas of the U.S.S.R.] Opredliteli Po Faune, S.S.S.R., 56:1—445 (In Russian.) 1965. Polychaeta of the Far Eastern Seas of the U.S.S.R. English edition of the 1955 publication. Translated by the Israel Program for Scientific Translations, U.S. Department of Commerce, Washington, D.C., 419 pp. Vibe, C. 1939. Preliminary investigations on shal- low water animal communities in the Upernavik and Thule Districts (Northwest Greenland). Meddelelser om Gr@nland, 124(2):1-43. 1950. The marine mammals and the ma- rine fauna in the Thule District (Northwest Greenland) with observations on ice conditions in 1939-1941. Meddelelser om Grénland, 150 (6) 21-115. POLYCHAETOUS ANNELIDS FROM EASTERN CANADIAN ARCTIC v7] Wesenberg-Lund, EF. 1934 The Committee’s 2nd East Greenland expedition in 1932 to King Christian [X's Land Scoresby Sound Gephyrean and annelids. Meddelelser om Grgnland, 104 (14):1-38. 1947. Syllidae (Polychacta) from Green land waters. Meddelelser om Grénland, 134(6) 1-38. 1948. Greenland waters. 134(9):1-38. Maldanidae (Polychaeta) from West Meddelelser om Grgnland, 1950a. The polychaeta of West Green- land, with special reference to the fauna of Nordre Strémfjord, Kvyane-, and Bredefjord. Meddelelser om Grgnland, 151(2):1—171. 1950b. Polychaeta. tion, 4(14):1-92. Danish Ingolf-Expedi- 1951. Polychaeta. land, Vol. II(19):1-182. The Zoology of Ice- 1953. Polychaeta. 1-169. Greenland. 122(3): The Zoology of East Meddelelser om Grénland, Zaks, I. G. 1923. [A new polychaete species be- longing to the family Ammocharidae, viz, Myrio- chele oculata, n. sp.] Trudy Petrogradskogo obschchestva_ Estestvoispytatelei, 53(1), Proto- kolyzasedanii, Nos. 1-8:171-174. (In Russian.) Accepted for publication June 6, 1972. ABRASION IN THE MEASUREMENT OF WATER MOTION WITH THE CLOD-CARD TECHNIQUE JAMES E. Dory! ABSTRACT: of diffusion rates of materials into and out of the organisms. determined by measuring the weight lost from immersed CaSO, blocks. AND MAXWELL S. Dory?’ Water motion around aquatic organisms produces a significant enhancement Relative diffusion rates can be A comparison of weight lost from CaSO, blocks and sediment load along a vertical transect indicated that abrasion was not a significant factor in the loss of weight from the CaSO, blocks. Aside from mass-action effects, water motion around an algal thallus produces a steepening of diffusion gradients, thereby increasing the rate at which materials diffuse in and out (Ruttner, Naturwissenschaften 14:1237—1239, 1926). The relationship between current velocity and uptake of phosphate has been described by Whitford and Shumacher (J. Phycol., 1:78—-80, 1965) in some detail following their experiments on benthic hydrophytes. However, in most natural situations water motion cannot be adequately described on the basis of current, i.e., velocity, alone. Relative diffusion rates can be determined by measuring the weight lost from a sparingly soluble material exposed to various conditions of water motion. Independently both Muus (Sarsia, 34:61— 68, 1968) and Doty (Botanica Marina, 14:32-—35, 1971) used the dissolution rate of CaSO, objects to obtain integrated measurements of water mo- tion over selected time periods. Muus gave no indication that he was aware of diffusion, as he only stated a concern for velocity. His instruments consisted of CaSO, spheres with masses of approximately six gr. Each was mounted on a nail or metal rod. The spherical shape of these instruments allowed for uniform dissolution even though the direction of the water motion might vary significantly. However, the small size of the instruments and the nature of their mounting are inadequate for most field situations we have encountered. Our concern has been for measuring the en- hancement of diffusion rates by water motion over what they would be if the water were mo- tionless at the measurement site. The instruments used (Doty, 1971, fig. 1) for this, called clod- cards, have a mass of about 30 gr. and are roughly rectangular in shape with rounded edges and corners. A piece of nylon ““Nitex 300” plank- ton netting cast in each clod and, protruding from 40 the bottom and cemented to the card, adds dura- bility to the clod-card preparation. The large size allows exposure periods sufficiently long that short-term variations in water motion will not greatly affect the results. Since the motion often differs so greatly with tide in shallow water marine work the exposure periods are usually near 24 hours, i.e., for a one-day tide cycle. Another advantage of the clod-cards is that they are mounted on rectangular pieces of white matte- finished plastic sheeting. This greatly facilitates labeling the individual instruments and securing them in place in the field. For both of these instrument types, weight losses indicate rates of dissolution which have been assumed (Doty, 1971) to be essentially due to diffusion, and this would seem certain in less turbulent water. However, the criticism has been voiced that a significant part of the weight loss from such instruments placed in more turbulent water may be due to abrasion from water-borne sediments. To test this hypothesis, an experiment was designed in which a series of clod-cards would be exposed in parallel along with a series of sediment traps. The sediment traps used were constructed from 16 fluid-ounce aluminum beverage cans. The top of one can was cut off and the bottom used to form the main part of the trap. A 4 cm bottom portion cut from a second can was used to form a lid. A hole one-half inch in diameter was drilled in each end to allow water to pass through. Rubber stoppers (size 00) were used to prevent spillage of samples during handling and storage. ‘Los Angeles County Dept. Arboreta and Botanic Gardens 301 N. Baldwin Ave., Arcadia, California 91006. “Dept. Botany, Univ. Hawaii, Honolulu, Hawaii 96822. 1973 CLOD-CARD MEASUREMENT OF WATER MOTION 4] ttt ot oo or TIDE RANGE | 1 n Nk S j oc x ts f WwW x = x x x x x Gis 0 a/b eh Ps ting 12 +000 C-l 2-3-4 « -5 6-7-8 O-9 -~--10 --Il:«~-12 «~-13:«~-14 ~-I5 -16 -I7 -I18 -I9 -20 -2I CLOD CARD CHANGE («) 0 6 #2 18 24 30 36 42 48 54 60 66 72 78 84 90 96 102 108 114 120 126 SEDIMENT TRAPPED (¢) GRAMS Figure 1. Weight change of clod-cards and amount of trapped sediment plotted as a function depth. Data shown are from one 24-hour exposure period. Above the 4.2 m level, all sediment traps and some clod-cards were damaged by a boat in this run of the experiment. The positions of curves were determined by eye. The two types of instruments, the clod-cards and the sediment traps, were mounted at intervals along a 24-foot long, 1-inch by 2-inch wooden board. At each interval a sediment trap and a clod-card were fastened on the stringer with masking tape, so that they were opposite one an- other and at the same elevation. The distance between successive positions was varied to pro- vide a more intense measurement series in certain “critical” segments of the transect. Thus, the intervals were 3 inch for the bottom 2 ft, then 6 inch for the next 2% ft. The maximum interval of one foot was in mid-water; then, near the surface, the interval was again 6 inch. In use this string of instruments was positioned so the to- and-fro surging motion of the water was parallel to the long axis of the sediment traps and the short axis of the clod-cards. The wooden stringer with the instruments attached was then strapped vertically to a piling on Goleta pier near the Santa Barbara, California, campus of the Univer- sity of California. Only the more complete of two experiments is described here. After 24 hours the instruments were retrieved. The weight lost from each of the clod-cards and the amount of sediment in each of the traps was measured (Fig. 1) and their values compared. The lack of values at several high tide levels was due to a boat having destroyed the instruments there. The clod-card weight increases the higher intertidal area probably are due to salt deposited from the evaporating of splash or spray. The amount of water-borne sediment collected was found to be at the bottom and diminished rapidly at successive intervals further from the bottom. Significant amounts of water- borne sediments were found only in the bottom two meters of the transect. Weight losses from the clod-cards, on the other hand, were found to be greatest in the mid-water region and diminished in the bottom two meters. These results lead us to believe that abrasion is not a significant error-producing factor in the measurement of water motion enhanced diffusion in greatest when using clod-cards as the measuring inst The financial assistance of U.S. AT(04-3)- ments. Energy Commission contract gratefully acknowledged. ele) Accepted for publication June 5. 19 VARIATIONS IN CONCENTRATIONS OF MAGNESIUM AND STRONTIUM IN RECENT SHELLS OF TIVELA STULTORUM RONALD J. GRABYAN! Abstract: Ten specimens of Tivela stultorum from Pismo and Torrance Beaches were analyzed for magnesium and strontium by atomic absorption techniques. Magnesium con- centrations in the umbonal areas were consistently greater than those along the margins. Strontium concentrations remained relatively uniform throughout the individual shell. Maxi- mum concentrations were 353 + 21 ppm for magnesium and 2690 + 300 ppm for strontium. Minimum concentrations 114 +21 ppm for magnesium and 1770 + 300 ppm _ for strontium. Maximum additional magnesium possibly diffused into the shell material of two shells immersed in running sea water for three months was 134+ 10 ppm. This apparent re-equilibration with sea water suggests that diagenesis can proceed rapidly after death of were the animal. In recent years investigations of skeletal carbonates have been largely confined to relating shell com- position to chemical and physical environments. Several relationships have been presented con- cerning regular variations between different groups of organisms even down to a differentia- tion between species. If evidence proving that a constant variation within an individual exists, then added importance might be placed on constant chemical variations as useful information to taxonomic grouping and ecological evidence. The purposes of this study are to examine the changes in concentration of magnesium and strontium within the individuals of the pelecypod, Tivela stultorum; to compare two widely separated populations; and to investigate the possible diffu- sion of magnesium into the skeletal material of the clams upon contact with sea water after death. Studies concerning the deposition of elements under various conditions in the skeletal material of marine fauna may lead to important discoveries with respect to pollution of the environment. A summary of some of the previous work on this subject will give a background to some of the problems and hypotheses submitted. Chave (1954) concluded that three variables, mineral- ogy, water temperature, and phylogenetic level are the controlling factors of skeletal magnesium in calcareous marine organisms. Also salinity, depth of water, and age or size of the individual are important factors. He came to three general conclusions applicable to most calcareous marine organisms. First, aragonitic skeletons are lower in magnesium than are those composed of calcite. Second, in all groups a linear relationship exists 42 between the magnesium in the skeleton and the temperature of the water in which the organism lived. Third, the total amount of magnesium in the skeleton decreases with an increase in phylo- genetic level of the organism. Lowenstam (1954, 1964) has shown that there is a Sr/Ca variation that is temperature-dependent in organisms having the ability to vary their aragonitic content with temperature variations. Pilkey and Hower (1960) concluded that MgCoO, content of Dendraster tests appeared to be directly related to water temperature and salinity, and that different echinoid species living under similar temperature conditions may deposit substantially different amounts of MgCOs. Harriss (1965) determined that only in the case of magnesium is mineralogy a dominant con- trolling factor in trace element distribution. Nelson (1967) determined that a heterogeneous distribution of strontium, barium, and manganese within eight species of freshwater clam shells is related to annual growth and shell structure. He also concluded that sodium and magnesium concentrations are relatively constant throughout the shells. An important paper by Weyl (1967) demon- strated that carbonate minerals in sea water do not behave as homogeneous thermodynamic phases. He concludes that the mineral phase in- troduced does not come into equilibrium with the solution, but rather, the surface layers of the solid adjust themselves to the aqueous environment. ‘Dept. Geological Sciences, Univ. of Southern California, Los Angeles, California 90007. 1973 Figure 7. Plan view of a typical shell of Tivela stultorum showing the position on each shell used for analysis. ANALYTICAL METHODS Sampling method.—Five live clams were collected from each of two California beaches separated by 200 miles of coastline—Pismo Beach and Torrance Beach. All ten clams were collected “in situ.” The percent MgCO., and SrCO. were determined for various parts of each specimen by standard methods of atomic absorption flame spectroscopy using the Perkin-Elmer Model 303 Atomic Absorption Spectrophotometer. From each locality two right valves and two left valves were analyzed. In order to determine accurately any possible trend of variation throughout the shell, the valves were sampled continuously in two perpendicular directions: a) the anteropos- terior plane crossing the muscle scars, and b) the plane perpendicular to the growth lines cutting the beak and continuing directly between the muscle scars (Fig. 1). The beak area and denti- tion were excluded from the analyses due to the complicated structures present in_ these Thirteen positions were analyzed—seven down and seven across with the center position being common to both directions. On three valves four additional positions were analyzed. Each position areas. Was approximately “4 inches square and occupied the thickness of the valve at that particular posi- tion. Three clams were also analyzed for varia- tions within the thickness of the valve was cut parallel and equidistant to the interior a sample and exterior planes, and the interior material was compared with the exterior material. MAGNESIUM AND STRONTIUM IN SHELLS OF lIVELA 43 Analyses. —For cach sample three analyses performed per element. The percent variation of this technique was 13 percent for magnesium and 17 percent for strontium. The four standard: used for magnesium in this experiment were 0.2 ppm 0.5 ppm, 1.0 ppm, and 1.5 ppm. Three standards were used for the strontium analyses—5.0 10.0 ppm, and 15.0 ppm. ppm, RESULTS Magnesium and strontium analyses.—-An appar- ently regular and substantial content variation in Mg is displayed within each of the eight individual clams analyzed. This variation appears to be valid as significant differences in concen- tration exist with respect to analytical precision. The maximum concentration of magnesium noted in any of the clams analyzed was 353 + 21 ppm, whereas the minimum concentration was 114 = 21 ppm. No systematic variation in the strontium content of the shells was noted. The maximum concentration of strontium noted was 2690 = 300 ppm, whereas the 1770 + 300 ppm. The plane passing directly between the muscle minimum concentration was scars and remaining perpendicular to the growth lines from the beak area to the outer edge is referred to herein as the vertical plane. Similarly. the anteroposterior plane crossing the muscle scars is referred to as the horizontal plane. The varia- tions of magnesium in the vertical and horizontal planes of each clam analyzed are shown dia- gramatically by the curves in figure 2. A trend can be detected from the concentrations at the respective positions on the shells. An increase in concentration generally appears toward the cen- ter of the valve in the umbonal region, and a sharp decrease is evidenced by the decline near the outer margin. A comparison of the Torrance clams with the Pismo clams demonstrates that the Pismo clams do not vary as consistently as the Torrance population. A difference in the average mean concentration between the two pop- ulations does not seem to be significant. A regular variation also appears to exist in the horizontal plane. This variation is somewhat in agreement with the vertical plane distribution as the concentric growth lines appear to have proximately equal concentration along o spective margins. Also. they appear to giv symmetrical appearance consistent with the growth lines. Standard electron microprobe ¢ 44 BULLETIN SOUTHERN CALIFORNIA 350 250 PPM 350 8 > 250 5 6 150 ACADEMY OF SCIENCES VOLUME 72 On wo @ aun POSITION ON SHELL Figure 2. Variation of magnesium in the shell material in two perpendicular directions: A—anteroposterior (vertical) plane—Torrance populations; B—anteroposterior plane—Pismo population; C—horizontal population. yses were conducted on two clams, but no ap- preciable trends are noted. Additional analyses were conducted on three clams 3, 4, and 6. Four additional analyses on each gives increased support to the constancy of concentrations along the growth lines. A varia- tion between the nacreous layer and the prismatic layer of the shell is also measureable. It can be seen in table 1 that the nacreous layer of the shell has a concentration much less tian the prismatic layer. These values may be the most important values determined, for they may lead to an understanding of the cause of the entire TABLE 1. nacreous Comparison of concentrations in the and prismatic layers for magnesium in position 4 of the clams analyzed. SAMPLE NUMBER PARTS PER MILLION 3—-N 150+ 21 3—P 290 + 21 4_-N 180 + 21 4-Pp 250 + 21 8-N 170 + 21 8-P 270 + 21 N—Nacreous layer of the shell P—Prismatic layer of the shell plane—Torrance population; and D—horizontal plane—Pismo variation with respect to the skeletal mineralogy and skeletal construction. Magnesium deposition experiment.—Two addi- tional clams were collected and cut into sym- metrical halves, and one half from each clam was placed in circulating sea water in an intake passage of the main tank in Marineland for three months. These shells were then analyzed for magnesium in the same way as the previous sam- ples. Figure 3 depicts the relative uptake of mag- nesium into the shell material. Only one position on the shell of one of the clams did not reflect a significant uptake of magnesium. The maximum uptake of magnesium noted was 134 + 10 ppm. DISCUSSION Tivela stultorum grows continuously throughout its life as the shell becomes thicker and increases in diameter (Fitch, 1950). The prismatic layer is secreted by the outer face of the outermost of three lobes at the mantle edge whereas the nacre- ous layer which does not extend beyond the pallial line is secreted by the general surface of the mantle (Cox, 1969). Timmermans (1969) presents a detailed account of possible mechanisms 1973 350 250 PPM 150 50 | (2 3 4 5 6 POSITION ON SHELL Figure 3. Results of magnesium deposition experi- ment after three months in circulating sea water. A and B represent the halves of the shells put in sea water for prolonged exposure and those that were not, respectively. of shell calcification. She concludes that the epithelial regions of the viscera are responsible for secretion of the shell layers including the periostracum and the two calcareous layers in gastropods. An extensive literature exists on depo- sition of calcium in various genera and species of molluscs. The prismatic layer thickens from the beak radi- ally, and the nacreous layer thickens in the reverse direction (Fig. 4). Dark colored growth rings commonly present on the shell can be seen dis- tinctly in the Torrance population, but they are lacking in the Pismo population. Their origin is due to a decreased rate of growth in August of each year when the temperature is highest and the seas invariably most calm (Fitch, 1950). The clams from Pismo Beach display a slight deviation from the Torrance Beach clams in that their mantles overlap the pallial line and extend almost to the shell margin. The exterior layer (Fig. 4) is the prismatic layer, and the interior layer is the nacreous layer. The contact on the interior plane of the valve where the nacreous layer and prismatic layer abut, pinpoints the outermost position where the mantle was attached to the valve. It has been established by Pilkey and Hower (1960), that MgCOs content is directly related to water temperature and salinity, and that dif- ferent species may secrete substantially different amounts of MgCO, under similar environmental conditions. Harriss (1965), however, proposed that in the case of magnesium, mineralogy is the MAGNESIUM AND STRONTIUM IN SHELLS OF PIVELA j Sr=2400+ 300 Mg=270 +20 Sr=1900* 300 Mg=220%20 Sr= 2300 * 300 = + Mg=|70+20 Figure 4. Cross section of clam 8 depicting the relationship of concentration values in ppm at two key positions. dominant controlling factor in its distribution. He also stated that metabolic processes and crystal growth considerations are of secondary impor- tance with respect to magnesium, but of major importance in strontium distribution. This latter point was initially determined by Odum (1957). He stated that the main factor controlling the Sr/Ca ratio was due to biological differences be- tween groups of organisms. Thus, the main dis- crepancy in the literature exists between Pilkey and Hower (1960) and Harriss (1965) as water temperature coupled with salinity and mineralogy are the respective arguments for the dominant factor in MgCO,. deposition. In order to discuss the results of the analyses presented in this paper, the following point must be analyzed. Cox (1969) has shown that alternat- ing layers of conchiolin (a scleroprotein) and aragonite are present in the nacreous layer. The fact that organic compounds may have strontium and magnesium as constituents has been outlined by Dodd (1967). These considerations present a problem which must be solved in order to validate the results presented in this paper. If the conchio- lin contains magnesium or strontium which has entered the solution phase in the analyses, then the results of this study will be altered by that factor. It is possible, however, that a variation of mag- nesium or strontium may be substantiated if the problem is approached indirectly. The nacreous layer contains much less magnesium than does the prismatic layer, therefore, it would seem that in the outer margin of the shell where the nacreous layer does not exist (Fig. 4), there would be an increase in the magnesium present. Instead. ther is a significant decrease. Therefore, a variation in the carbonate material is still a strong possi bility. Figure 4 presents a cross-section of clam 46 BULLETIN SOUTHERN CALIFORNIA If position A con- in the 8 depicting this relationship. tains a substantial amount of conchiolin nacreous material, and since position B belongs entirely to the prismatic layer and contains no conchiolin, then if it is assumed that there is no magnesium in the conchiolin, or if there is, that it is equally distributed and of small concentra- tion, then the expected results should accord a higher amount of magnesium in position B than in position A. The results, however, present the exact opposite. Therefore, if there is no magne- sium in the conchiolin as assumed, there would seem to be a variation of magnesium in the aragonitic shell material. The variation of mag- nesium distribution may occur among the pris- matic layers, the nacreous layers, or both. Dodd (1965) has shown that both magnesium and strontium concentrations vary with respect to the different layers of the specimen, and that a positive correlation between strontium concentra- tion in the prismatic layer and temperature is evidenced while in the nacreous layer a negative correlation with temperature is present. He also has demonstrated that a probable seasonal varia- tion in strontium and magnesium concentrations is shown in the outer prismatic layer in the antero- posterior plane. He has suggested that the cause of the variation between different genera and even between different structural units of the same shell is under strong biochemical control. Nelson (1967) indicates that a variation be- tween two zones of nacreous layers does exist for freshwater clams. He studied the distribution of trace elements among the annual layers within each of two nacreous zones. He found hetero- geneous distribution of Sr, Ba, and Mn within eight species of these clams which he attributed to annual growth and shell structure. A more important aspect with respect to this paper is that he determined magnesium concentrations to be relatively constant throughout the shells. He also reported that there were inherent differences in the trace element composition of clamshells of different species. This study has revealed that strontium distri- butions do not seem to vary between the prismatic or nacreous layers of the shell or across the shell to any great degree. Therefore, there may be a similar concentration of strontium in the conchio- lin as in the aragonite or an increase of strontium in the nacreous aragonite as opposed to the pris- matic aragonite. In order to determine the magnesium and stron- tium in the conchiolin, it would be necessary to ACADEMY OF SCIENCES VOLUME 72 separate the organic material from the crystalline material. A procedure is outlined by Degens, Spencer, and Parker (1967). However, even with- out doing this, the data substantiate a variation of magnesium in the shell material. Of the variables that have been presented, it seems that salinity and temperature of the water may be responsible for variations across the shell as these factors affected the shell uniformly at the same particular instant in time. It is possible that different secreting cells of the organism are responsive variably with respect to changes in temperature or salinity. It is important to note that since the outer margin of the shell is com- posed of the youngest shell material, the concen- tration of magnesium at these positions might reflect a susceptibility of the organism to en- vironmental parametric deviations. As the shells of Tivela stultorum are 100 per- cent aragonite, it does not seem that the trace element distribution would be controlled by min- eralogy. Crystal growth considerations may serve as an important factor in the variation of minor elements since prismatic crystals are deposited perpendicular to the surface, and in the nacreous layer thin leaves of the mineral are oriented al- most parallel with the inner surface of the shell (Cox, 1969). This difference may reflect a per- cent difference in accepting magnesium in the respective components of the shell structure. Mag- nesium concentration is strikingly less in the nacreous layer than in the prismatic layer as de- picted by table 1. If this were a proven variable, the percent MgCO, would increase toward posi- tions 5, 6, and 7 on each shell. This analysis of possible variables for the distribution of magnesium in the shell material leaves only one additional known variable which may account for the regular variation of MgCO, noted in Tivela stultorum—metabolic processes. Considering the possible diffusion of magne- sium into the shell material, it is significant to note that the uptake of magnesium may be due to an equilibration of the surface layers of the clams as outlined by Weyl (1967) for carbonate materials in sea water. Ragland, Pilkey, and Blackewelder (1969) support this suggestion as they have shown that Sr/Ca ratios of fossil mol- lusc shells may be significantly altered by in- organic processes before recrystallization. Lerman (1964) concurs with this in that he implies that concentrations of minor elements in fossil biogenic calcites may be altered by diffusion with no ap- parent effects in the crystalline texture. 1973 The fact that this diagenesis does occur indi- cates that post-mortem analysis of shells must be cautiously evaluated until further study determines the degree to which shells have been chemically altered from their original state. CONCLUSIONS A variation in the distribution of magnesium in the shell material of Tivela stultorum exists, and a constancy of this variation across the shell is apparant. Strontium, however, appears to either remain constant throughout the shell or increase in the nacreous layer. The concentration of mag- nesium and strontium should be determined in the organic compounds of the nacreous layers of the shells in order to arrive at more conclusive evidence, however, indirectly a variation does ap- pear to exist independent of this factor. The mag- nesium content is much less in the nacreous layer than in the prismatic layer. Therefore, in spite of the unknown concentrations of magnesium and strontium in the conchiolin, there still appears to be a variation of magnesium in the shell, and this variation appears to stay constant along the growth lines, but decreases from the umbonal area of the valve to the outer margin. From these probable and assumed facts, the following possi- bilities may exist along with others unknown to the writer. The biological secreting mechanism along with temperature effects may be responsi- ble for the differences in magnesium concentra- tions in each shell. Mineralogical control and salinity are possibilities that must be explored further. Regardless of the variations of magne- sium in the aragonite lattice, however, there is an uptake of magnesium into the shell material of post-mortem shells placed in circulating sea water, and due to this fact, correlations of trace element concentrations of the shells of living animals with fossil shells should be regarded cautiously. ACKNOWLEDGMENTS The writer is indebted to W. H. Easton, who presented the problem, D. F. Palmer, and J. L. Bischoff for helpful criticisms and approaches to the problem. Special gratitude is given to D. S. Gorsline, T. L. Ku, and R. H. Osborne for helpful discussion. Thanks are also given to William Feyerabend and Mary Friend for drafting the figures. This is Contribution No. 263, Department of Geological Sciences, Univer- MAGNESIUM AND STRONTIUM IN SHELLS OF PIVELA 4 sity of Southern California, Los Angeles, ¢ fornia 90007, LITERATURE CITED Chave, K. E. 1954. Aspects of the biochemistry of magnesium 1. Calcareous marine organism J Geol., 62:266-283. Cox, L. R. 1969. General features of Bivalvia Pp. N2—N129 in Treatise on Invertebrate Paleon tology: Part N, v. 1 (of 3), Mollusca 6, Bivalvia. (R. C. Moore, ed.), Geol. Soc. America Inc. and Univ. Kansas, xxxviii + N489 pp. T., D. W. Spencer, and R. H. Parker. Paleobiochemistry of molluscan shell pro- Comp. Biochem. Physiol., 20:553-579. Degens, E. 1967. teins. Dodd, J. R. 1965. tium and magnesium in Mytilus. Geochimica et Cosmochimica Acta 29:385-398. Environmental control of stron- strontium in cal- J. Paleo., 41:1313- 1967. Magnesium and careous skeletons: a review. 1329. Fitch, J. E. 1950. The pismo clam. California Fish and Game, 36 (no. 3):285-312. Harriss, R. C. 1965. Trace element distribution in molluscan skeletal material I. Magnesium, iron, manganese, and strontium. Bull. Mar. Sci., 15:265-273. Lowenstam, H. A. 1954. Environmental relations of modification compositions of certain carbonate secreting marine invertebrates. Proc. Nat. Acad. Sci., 40:39-48. 1964. Sr/Ca ratio of skeletal from the Recent marine biota of Palau and from fossil Gastropods. Pp. 114-132 in Isotopic and Cosmic Chemistry. (H. Craig, S. L. Miller, and G. J. Wasserburg, eds.), Amsterdam, North- Holland, xxv + 553 pp. aragonites Nelson, D. J. 1967. Microchemical constituents in pre-Columbian clamshells. Pp. 185-204 in Quaternary Paleoecology, v. 7. (E. J. Cushing and H. E. Wright, Jr., eds.), Nat. Acad. Sci. vii + 433 pp. Odum, H. T. 1957. Biogeochemical deposition strontium. Inst. Mar. Sci. 4:39-114. Pilkey, O. H., and J. Hower. 1960. Th concentration environment on the 48 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES magnesium and strontium in Dendraster. J. Geol., 68:203-216. Ragland, P. C., O. H. Pilkey, and W. Blackewelder. 1969. Comparison of the Sr/Ca ratio of fossil and recent mollusc shells. Nature 224 (No. 5225) :1223-1224. Timmermans, L. P. M. 1969. Studies on shell VOLUME 72 formation in molluscs. Netherlands J. Zool., 19(4) :417-523. Weyl, P. K. 1967. The solution behavior of car- bonate materials in sea water. Stud. Trop. Oceanog., 5:178—228. Accepted for publication, June 20, 1972. RESEARCH NOTES SPECIES GROUPS OF NORTH AMERICAN CHORUS FROGS (GENUS PSEUDACRIS: FAMILY HYLIDAE) Seven species of chorus frogs are currently included in the North American genus Pseudacris: Pscudacris brachyphona, P. brimleyi, P. clarki, P. Je. ornata, P. streckeri, and P. triseriata. These seven species have been divided into different intrageneric groups by different authors. ‘The most recent designa- tion was that of Chantell (Amer. Midl. Nat., 80:381— 391, 1968), who divided the extant species into two groups: the ornata group containing P. clarki, P. ornata, and P. streckeri; and the nigrifa group con- taining P. brachyphona, P. brimleyi, P. nigrita, and P. triseriata. Based on the data reported in the present paper, I propose that this scheme be changed by transferring P. clarki from the ornata group to the nigrita group. During the course of a detailed study of the com- plete skeletons of the 32 species of tree frogs found in the temperate regions of the northern hemisphere, I examined entire and disarticulated skeletons of specimens from the seven species listed above. Each specimen was cleared in 2 percent potassium hydrox- ide, stained with alizarin red S, and dehydrated and stored in glycerine. The following number of speci- mens of each species were studied: P. brachyphona, 7; P. brimleyi, 2; P. clarki, 5; P. nigrita, 4; P. ornata, 7; P. streckeri, 3; and P. triseriata, 5. The following species groups can be established solely on the basis of osteological data; the ornata group containing P. ornata and P. streckeri; and the nigrita group containing P. brachyphona, P. brimleyi, P. clarki, P. nigrita, and P. triseriata. The ornata group can be diagnosed in the following way (the contrasting character state found in the nig- rita group is given in parentheses): 1) a solid articula- tion is present between the nasal bone and the facial spine of the maxilla (Fig. 1) (no nasal-maxillary artic- ulation); 2) nasals are elongate and deeply notched laterally (nasals sub-triangular and not laterally notched); 3) a flattened process projects from the anterior arm of the septomaxilla (no anterior septo- maxillary process); 4) more than 30 percent of the frontoparietals are involved in the frontoparietal- sphenethmoid articulation (less than 30 percent): 5) a solid articulation is present between the palatine bones and vomerine tooth patches (no palatine- vomerine articulation); 6) vomerine tooth patches are relatively large (vomerine tooth patches are rela- tively small); 7) anterior extension of vomer directed toward the anterior end of the maxilla (an- terior vomerine extension is directed toward posterior end of premaxilla); 8) more than 30 percent of the nigrita, is the the sphenethmoid (less than 30 percent); 9) the posterior anterior arm of parasphenoid overlaps lateral arms of the parasphenoid are less cent of the 20 percent); 10) the horizontal arm of the is than 20 per the length of anterior arm (more than Quamo al horizontal length/ vertical 50 horizontal reduced and length zontal length/ vertical length = more than 80 percent); 11) the quadrate i arm arm less than percent (well developed hori arm, arm arm calcified (quadrate is cartilaginous); 12) the clavicles are thickened, especially medially (Fig. 2) (clavicles thin and narrow); 13) coracoids are directed postero- medially (coracoids directed medially); 14) pectoral fenestra is relatively large and rounded (pectoral 15) deltoid ridge of humerus is pierced longitudinally on its fenestra relatively small and elongate); ven- tral surface by a groove and foramen (no groove or foramen); and 16) the ventral acetabular expansion the ilium is relatively small large expansicn). of (relatively The above osteological character states are found in adult individuals and, even though some of them represent differences in degree and not in kind, it is very easy to distinguish members of the two species PREMAXILLA SEPTOMAXILLA NASAL << MAXILLA — SPHENETHMOID FRONTOPARIETAL PTERYGOID PROOTIC QUADRATOJUGAL SQUAMOSAL EXOCCIPITAL PREMAXILLA VOMER PALATINE PARASPHENOID PTERYGOID OPTIC FORAMEN QUADRATE COLUMELLA WA —_—_——————————_ OCCIPITAL B 1. The skull of Pseudacris streckeri Figure in dorsal view (A) and ventral view (B). 50 BULLETIN SOUTHERN CALIFORNIA — CLAVICLE FENESTRA CORACOID Figure 2. Ventral views of the pectoral girdles of Pseudacris streckeri (A) and Pseudacris clarki (B). groups from one another when one examines their skeletons. In general, the skeleton in specimens from the group is more developed than that in specimens from the nigrita group. In the former, the anterior end of the head is compact with two solid lines of articulation. Dorsally, the nasals provide a bony bridge joining the lateral maxillae solidly with the medial sphenethmoid. Ventrally, the palatines serve as transverse braces connecting the maxillae, sphenethmoid, and vomers. The arrowhead-shaped sphenethmoid is proportionately larger in size in the ornata group and involves much of the medial edges of the nasals in an overlapping articulation. The prootic and exoccipital ossifications are extensive and occupy much of the otic capsule. By way of contrast, the cranial ossifications in the specimens examined from the nigrita group are not as extensive (Fig. 3). The posterior end of the nasal capsules is not as heavily supported by bone, nor do the ossifications in the otic capsules develop as extensively. ornata ACADEMY OF SCIENCES VOLUME 72 Figure 3. The skull of Pseudacris nigrita in dorsal view (A) and ventral view (B). An examination of the external morphology of specimens of the seven species of Pseudacris suggests the same intrageneric division indicated by a com- parative study of their skeletons. Species in the nigrita group are smaller in size and resemble one another in general appearance. Pseudacris ornata and P. streckeri are stout bodied forms whose limbs are also more massive than those in the species of the nigrita group. ANTHONY J. GAUDIN, Dept. Biology, California State University, Northridge, California 91324. Accepted for publication November 16, 1972. 1973 THE OS TRANSILIENS IN FOUR SPECIES OF TORTOISES, GENUS GOPHERUS Ray (1959), first found a heterotropic or sesmoid bone, the os transiliens, in a chelonian, Gopherus polyphemus. Legler (1962) discovered the presence of the os transiliens in Gopherus agassizi and Go- pherus flavomarginatus. Ray (1959) showed histologically that the os transiliens in Gopherus polyphemus was bone and not calcified cartilage. Differential stains of toluidine blue (for cartilage) followed by alizarin red S (for bone) were carried out on a series of 20 os transiliens taken from Gopherus agassizi and) Gopherus— ber- landieri (Evans, 1949; Willey, 1969). X-rays were taken of the os transiliens and com- pared to known samples of bone and cartilage in Gopherus agassizi as a check upon the determination of the presence of bone. A General Electric model L1AA-3A X-ray machine was used at 40 KVP and 50 milliamps using Kodak Industrial type M-2 X-ray film. Carapace length, skull length, and length and width of the os transiliens were measured on nine Gopherus agassizi, one Gopherus polyphemus, and two Gopherus berlandieri. Skull length was measured in a straight line from the anterior extremity of the snout to the tip of the supraoccipital crest with the skull resting on the articulated mandible. The greatest width and length of the os transiliens were measured. The length of the carapace was measured along the midline between vertical axes erected at the posterior border of the supracaudals and the anterior border of the nuchal. The movement of the os transiliens was examined on seven thawed and two live Gopherus agassizi, using a wire and x-ray analysis technique developed by Inman (1947). The os transiliens is a prow-shaped bone in the aponeurosis of the M. adductor mandibularis externus and is articulated in a joint capsule with a facet formed by the quadrate and prootic bones, the pro- cessus trochlearis oticum (Gaffney, 1972). This facet places the os transiliens slightly higher on its inner surface and somewhat lower on its outer lateral sur- face. The os transiliens appears to provide a gliding or smooth motion for the fan-shaped aponeurosis of the M. adductor mandibularis externus over the edge of the superior temporal fossa where it passes ven- trally to an insertion on the mandibular coronoid process. The os transiliens of all adult Gopherus agassizi (15) showed positive stains for bone. Five juvenile Gopherus agassizi showed positive stains for cartilage suggesting a replacement of cartilage by bone with maturity. The differential technique indicated the presence of a small bony os transiliens in one specimen of Gopherus berlandieri which is the first to be so noted. The x-rays confirmed the stain- ing data in all cases. staining also RESEARCH NOTES co co 05 TRANSILIENS (mm) Z 4 6 SRULL LENGTH (cm) length Skull transiliens length (mm) in Gopherus agassizi (dots), Figure |. (cm) relative to the left o Gopherus polyphemus (open circles), and Gopherus berlandieri (open squares). The relationship between the os and skull agassizi was found by the method of least squares to be Y = 1.42X—1.18 and for Gopherus polyphemus to be Y = 1.85X — 3.90. transiliens length for Gopherus at a smaller size in Gopherus agassizi than has been found in Gopherus polyphemus (Fig. 1). The size of the os transiliens compared to carapace length in Goplicrus Ossification of the os transiliens occurs agassizi (Fig. 2) shows that the larger tortoises have proportionally larger os transiliens (Table 1). In Gopherus agassizi the superficialis portion of the M. adductor mandibularis externus contracts and pulls on the broad part of the fan-shaped aponeurosis attached to the upper end or posterior portion of the os transiliens. This is adducted (inside the joint cap- sule) posteriorly on the process trochlearis pulling on the tendon attached in part to the front of the os OS TRANS. -mm 1S 2s CARAPACE LENGTH Figure 2. Carapace length (cm) relative to length of the left os transiliens (mm) for Goplierus agassiz (dots), and a _ single polyphemus (open circle), and Gop/ier (open square). The relationship be carapace length for Goplierus agassizi Wé unc 2 the method of least squares. The relationship is Y = 0.259X + 0.33. specimen each of G 52 BULLETIN SOUTHERN CALIFORNIA Taste 1. Correlation coefficients between carapace length, skull length and os transiliens; A, Gopherus agassizi and B, Gopherus polyphemus. (re) A B xy between os and skull length 0.801 0.940 between os and . carapace length 0.742 — between skul! and carapace length 0.927 — transiliens and in part to the belly of the superficialis portion of the M. adductor mandibularis. This tendon pulls, ventrally, on the coronoid process thereby pulling the mandible up which in turn closes the mouth. No evidence was found for any involvement of the outer lateral edge of the M. adductor mandi- bularis (Fig. 3), in Gopherus agassizi as had been suggested for Gopherus polyphemus by Ray (1959). Ray (1959) had also suggested (based solely upon preserved specimens) that the os transiliens might act as a means of providing a longer lever arm for the action of the jaw closing musculature or as some sort of buttress for the muscle to pull against. It appears instead, that the os transiliens serves only as a means of providing a smooth or gliding action for the jaw musculature over the superior temporal fossa as shown by x-ray analysis. It thus appears that the os transiliens is probably found in all members of the genus Gopherus and develops in cartilage which is subsequently replaced GP RAY GA Figure 3. Comparison of suggested muscle action of M. adductor mandibularis externus in closing the jaw as suggested by Ray for Gopherus polyphemus (GP) and the present study for Gopherus agassizi (GA). The dark area on the right side of the Gopherus agassizi skull shows the relative position of the os transiliens. The symbols used in the figure include: a. tendon of M. adductor mandibularis ex- ternus; b. posterior belly of M. adductor mandibularis externus portio superficialis; c. posterior belly of the M. adductor mandibularis externus portio profunda. ACADEMY OF SCIENCES VOLUME 72 by bone. It also appears that the os transiliens ossifies earlier in young individuals of Gopherus agassizi than in other species studied. The os transiliens seems to be a device to smooth the translation of the horizontal pull of the M. adductor mandibularis ventrally in pulling the mandible shut. I wish to thank Bayard Brattstrom and James Smith for their assistance in this study and for reviewing the manuscript. LITERATURE CITED Evans, H. E. 1968. Clearing and staining small vertebrates, in toto, for demonstrating ossifica- tion. Turtox News, 26(2):42—47. Gaffney, E. S. 1972. An illustrated glossary of turtle skull nomenclature. Amer. Mus. Novit., 2486: 1-33. Inman, V. T. 1947. Functional aspects of the ab- ductor muscles of the (Amer.), 29:607-619. hip. J. Bone Surg. Legler, J. M. 1962. The os transiliens in two spe- cies of tortoises, Genus Gopherus. Herpetologica, 18:68-69. Ray, C. E. 1959. A sesmoid bone in the jaw musculature of Gopherus polyphemus (Reptilia: Testudinina). Anatomischer Anzeiger, 107(1/5): 85-91. Willey, R. B. 1969. Staining shark skeletons with alizarin Red S. Turtox News, 47(2):50—52. RoBERT PATTERSON, Dept. Biology, California State University, Fullerton, California 92634. Accepted for publication November 27, 1972. RATE OF HEAT LOSS BY LARGE AUSTRALIAN MONITOR LIZARDS Bartholomew and Tucker (1964) have shown that there is a difference in the rate of heating and cooling in Australian monitor lizards. They implicated vasomotor and respiratory mechanisms to explain this difference. The availability of several large Lace Monitors, Varanus varius, some more than twice the size used by Bartholomew and Tucker, allowed the testing of one of these mechanisms. Four large monitor lizards were used in a series of heating and cooling experiments where core (cloacal and esophageal), deep (10-28 mm) and _ shallow muscle, and skin surface temperatures were monitored continually. Temperature readings were taken at ir- regular intervals using a Yellow Springs Instrument 1973 a COOLING 35 30 47, 25°C et 2—40"—+10" TEMP. °C 47, 10°C L, 0 \ 2 3 4 5 6 7 2 HOURS Figure 1. Body temperatures (cloacal) of Lace Monitor lizards, Varanus varius of various sizes upon cooling from 40° to 25°, from 40° to 10°, and from 25° to 10°C. Lizard 1 is the largest, 4 is the smallest; see text for measurements and weights. Co. Telethermometer Model 44TD, 0-50°C and thermister probes. All probes were calibrated against a US Bureau of Standards Certified thermometer. Deep cloacal and esophageal temperatures were taken with #402 vinyl, ensheathed probes; skin surface temperatures with banjo probes; and subdermal and muscle temperatures with various sizes of hypodermic probes. Autopsy showed no significant or extensive hemorrhaging about the hypodermic probes. Head length, snout-vent length, tail length (mm), and weight (gms), respectively, for the four lizards used in this study are as follows: No. 1, 133, 765, 1149, 10,400; No. 2, 125, 676, 1129, 7,850; No. 3, 98, 535, 835, 2,563; No. 4, 67, 365, 645, 510. Lizard number one was the largest in size and weight. Lizards 1, 2, and 3 were captured in Victoria and 4 was obtained in Queensland. The lizards had been kept in captivity for between four weeks and a year and were in good condition. Lizards 1 and 4 had heavy layers of ven- tral fat. The lizards were not fed for at least three days prior to tests. Each lizard was tied to a wire frame two feet above a table so that air circulated all around the lizard. Though restrained, the depth of breathing did not seem impaired. The test lizard was placed in a large constant temperature room that maintained a temperature with + .2°C. Blowers which circulated air within the room passed air over and around the lizard at an average rate of 640 ft/min. After reaching the temperature of one room, the rack, lizard with imbedded probes, and _ telethermometer Were quickly transferred to another constant temper- ature room. As expected (Fig. 1) the rate of cooling depends both upon the temperature differential between the lizard and the environment and with size; larger lizards losing heat slower than the smaller lizards. RESEARCH NOTES COOLING 35 40°—>1|0° + | TEMP, °C HOURS Figure 2. Cooling curve for Lace Monitor No. 1 from 40° to 10°C, showing differences in tail, leg, muscle, and cloacal temperatures. Smaller lizards, of course, heat up faster than large lizards regardless of whether they are placed in a warm ambient temperature or are under heliothermic basking conditions (Bartholomew and Tucker, 1964: Cowles, 1958; Bartholomew and Lasiewski, 1965). Figure 2 shows the cooling curve for Lace Monitor No. 1 transferred from 40° to 10°C. Tail and leg temperatures drop quickly to ambient temperatures, whereas deep body temperatures take more than 6 hrs to reach ambient. Esophageal temperatures were almost the same (or 0.2°C cooler) as cloacal temper- atures. Figure 3 shows similar data for lizard No. 2 transferred from 25° to 10°C. Tail temperatures drop rapidly, back muscle, back subdermal, and deep abdominal (not shown in Fig. 2) are intermediate between tail and cloacal temperatures. Shielded (with masking tape) back skin surface temperatures were similar to subdermal, while side-skin surface tem- peratures were intermediate between subdermal and tail temperatures. The rapid drop of the tail and leg temperatures to ambient and the slower drop of the core temperature to ambient (up to 7 hrs) suggests that when cooling the Lace Monitor lizard may be shunting blood from the appendages to the center of the body. In one experiment, the temperature of the tail was taken at several points along its length. All tail temp I dropped at about the same rate, suggestit effect is not just that of a counter-curre change system or just due to differences in the phys- ical dimensions of different parts of the The temperatures shown in 54 BULLETIN SOUTHERN CALIFORNIA COOLING 25°10 #2 2 TEMP, °C HOURS Figure 3. Cooling curve for Lace Monitor No. 2 from 25° to 10°C, showing the rapid drop in tail temperature and slow drop in cloacal temperature. a gradient of temperature from outside to core rather than any bands of temperature or any core and shield differences. Autopsy revealed that the peripheral body muscles of monitors are few and thin; a condi- tion similar to most other lizards. There was no dorsal or lateral fat, though ventral fat was present. Thus the skin of the monitor does not appear to be very effective in reducing heat loss. Slow cooling rates have been reported for several reptiles (Cowles, 1958; Cogger and Holmes, 1960; Bartholomew and Tucker, 1964; Mackay, 1964; Bartholomew and Lasiewski, 1965; Bartholomew, 1966; Myres and Eells, 1968; Stebbins and Barwick, 1968). Stebbins and Barwick (1968) showed that monitor lizards often spend the night in tree-hole retreats. This behavior plus the data presented herein suggests that in such retreats (where there is less wind than in the experimental condition) the body tempera- tures of the monitors would probably not reach ambient until morning. This would facilitate nocturnal digestion of foods eaten during the day as well as reducing the heating time of the lizard in the morning. Though most reptiles are ectothermic, a small amount of internal endothermic heat production is possible. This amount is usually so small that it is immediately lost to the environment, hence it does not contribute to body temperature. Surface-volume relationships would suggest that in large reptiles a small amount of heat production plus a reduced rate of heat loss could contribute to body temperature. Bartholomew and Tucker (1964) calculated that at 25°C a monitor lizard might add 2°C to its body temperature by endogenous heat production. The large lizard used in the studies reported herein was stimulated for five minutes under an ambient tem- perature of 25°C (body temperature 25°C). In this experiment, the lizard raised its body temperature 2°C (to 27°C, but no higher) due to the increased muscular activity. When stimulated at an ambient temperature of 10°C no rise in body temperature could be detected. Heat retention and a reduction in thermal con- ductance are both enhanced by larger body size and reduced surface area. Large Galapagos tortoises thus show little internal daily fluctuation in body tem- ACADEMY OF SCIENCES VOLUME 72 perature (Mackay, 1964). Dinosaurs presumedly would have had even less of a daily fluctuation in body temperature. Further, vasomotor control of rate of heat loss may be another physiological mechanism used by reptiles to control and modify their body temperature and may also be an important step towards the development and evolution of homeothermy and endothermy. This work was carried out during an NSF Senior Postdoctoral Fellowship (56017) in the Department of Zoology and Comparative Physiology, Monash University, Clayton, Victoria, Australia. I wish to thank James W. Warren and A. K. Lee for their kindnesses and Lucien Vandevelde, Mike Owen, and John Nelson for collection, use, care, and assistance with these large lizards. LITERATURE CITED Bartholomew, G. A. 1966. A field study of tem- perature relations in the Galapagos marine iguana. Copeia, 1966:241—250. Bartholomew, G. A., and R. C. Lasiewski. 1965. Heating and cooling rates, heart rate and simu- lated diving in the Galapagos marine iguana. Comp. Biochem. Physiol., 16:573—-582. Bartholomew, G. A., and V. A. Tucker. 1964. Size, body temperature, thermal conductance, oxygen consumption, and heart rate in Australian varanid lizards. Physiol. Zo6., 37:341-354. Brattstrom, B. H. 1965. Body temperatures of rep- tiles. Amer. Midl. Nat., 73:376—422. Cogger, H. G., and A. Holmes. 1960. Thermoreg- ulatory behaviour in a specimen of Morelia spilotes variegata Gray (Serpentes: Boidae). Proc. Linnean Soc. N.S.W., 85:328—-333. Cowles, R. B. 1958. Possible origin of dermal tem- perature regulation. Evolution, 12:347—357. Mackay, R.S. 1964. Galapagos tortoise and marine iguana deep body temperatures measured by radio telemetry. Nature, 204:355-358. Myres, B., and M. Eells. 1968. Aggregation be- havior in the boa constrictor. Herpetologica, 24:61-66. Stebbins, R. C., and R. E. Barwick. 1968. Radio- telemetric study of thermoregulation in a lace monitor. Copeia, 1968:541-547. BAYARD H. BRATTSTROM, Dept. Biology, California State University, Fullerton, California 92634. Accepted for publication December 1, 1972. INSTRUCTIONS FOR AUTHORS _ The BULLETIN is published three times each year (April, August, and November) and includes articles in English in any field of science. Non-members will be assessed a page charge of $40.00 per page. Manuscript submitted for publication should contain results of original research, embrace sound principles of scientific investigation, and present data in a clear and concise manner. The current AIBS Style Manual for Biological _ Journals is recommended as a guide for contributors. Consult also recent issues of the BULLETIN. Authors - should strive for directness and lucidity, achieved by use of the active voice. Special attention should be given to consistency in tense, unambiguous reference of pronouns, and logically placed modifiers. 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Special attention should be given to the description of new taxa, designation of holo- _ type, etc. _ The literature cited section should include six or more references; entries for books and articles should take these forms. McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54: 452-458. Brattstrom, B. H. 1969. The condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), Univ. California Press, xii + 635 pp. q If fewer than six references are cited they should be inserted in text as follows: (McWilliams, Insect mimicry, p. 216, 1970); (Holmes and Speak, J. Mamm., 54: 452-458, 1971): (Bratt- strom, The Condor in California, Pp. 369-382, in Vertebrates of California, 1969). Tables and figures (line drawings, graphs, or black and white photographs) should not repeat data con- tained in the text. 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Wells, Department of Biol- ogy, Occidental College, Los Angeles, California 90041. Evaluation of a paper submitted to the BULLETIN _ begins with a critical reading by the technical editor; several referees also check the paper for scientific con- tent, originality, and clarity of presentation. Judgments as to the acceptability of the paper and suggestions _ for enhancing it are sent to the author at which time he may be requested to rework portions of the paper considering these recommendations. The paper is then re-submitted and may be re-evaluated before final - acceptance after which it is sent to the managing editor. Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He should promptly and carefully read the proof sheets for errors and omissions in text, tables. illustrations legends, and bibliographical references. He marks corrections on the galley (copy editing and proof proce- dures in Style Manual) and promptly returns both galley and manuscript to the managing editor, James Dale Smith, Department of Biology, California State University, Fullerton, California 92634. Manuscripts and origi- nal illustrations will not be returned unless requested at this time. Printing charges accruing from excessive addi- tions to, or changes in, the proofs »ust be assumed by the author. Reprint orders are placed with the printer. not the Managing Editor. CONTENTS New taxa of brachyuran crabs from deep water off western Peru and Costa Rica. By John 8. Garth 1 A new species of gerbil from Southwest Africa with remarks on Gerbillus tytonis Bauer and Niethammer, 1959 (Rodentia: Gerbillinae). By Duane A. Schlitter Polychaetes from Central American sandy beaches. By Kristian Fauchald. .............. Wegertolots 19 Polychaetous annelids collected by the R/V Hero from Baffin Island, Davis Strait, and West Greenland in 1968. By James A. Blake and David Dean Abrasion in the measurement of water motion with the clod-card technique. By James E. pen and Maxwell S. Doty \ — Variations in concentrations of magnesium and strontium in recent shells of Tivela stultorum. By Ronald J. Grabyan RESEARCH NOTES Species groups of North American chorus frogs (Genus Pseudacris: Family Hylidae) oat Anthony J. Gaudin The os transiliens in four species of tortoises, genus Gopherus. By Robert Patterson ..... Rate of heat loss by large Australian monitor lizards. Index to Volume 71 COVER: A new species of brachyuran crab from deep water off Peru. Its genus, Acanthocarpus, which was not previously known from the west coast of the Americas, is characteried by the long, transverse spine on the — merus of each cheliped. Illustration by Jerry J. Battagliotti, University of Southern California. mw MweRN CALIFORNIA ACADEMY OF SCIENCES LIBRARY LLETIN JAN 28 1974 \/ -) i olume 72 Number 2 NEW YORK BOTANICAL GARDEN ROACAI9N(9\) 101.119 (1072) AUGUST 1973 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 OFFICERS Jules Crane, Jr., President Donald Bright, First Vice President Wayland D. Hand, Second Vice President Stuart L. Warter, Secretary Donald R. Patten, Treasurer Patrick H. Wells, Technical Editor James Dale Smith, Managing Editor Roberta S$. Greenwood, Index Editor BOARD OF DIRECTORS 1971-1973 1972-1974 : 1973-1975 John J. Baird Wayland D. Hand Donald Bright Takashi Hoshizaki Donald R. Patten Timothy S. Hillebrand George Callison Donald J. Reish Jules Crane, Jr. David L. Walkington Andrew Starrett Richard E. Pieper Stuart L. Warter Patrick H. Wells David P. 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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. a a BULLETIN OF THE SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 NOVEMBER 27, 1973 NUMBER 2 THE SYSTEMATIC STATUS AND DISTRIBUTION OF COSTA RICAN GLASS- FROGS, GENUS CENTROLENELLA (FAMILY CENTROLENIDAE), WITH DESCRIPTION OF A NEW SPECIES PriscittA H. STARRETT AND JAY M. SAVAGE! Apstract: The glass-frogs (Family Centrolenidae) of Costa Rica comprise 13 species all placed in the genus Centrolenella. A review of the fleischmanni group indicates that six spe- cies: C. fleischmanni, C. colymbiphylhum, C. chirripoi, C. talamancae, and C. vireovittata, a new species, from southwest Pacific Costa Rica, occur in the republic. Salient features for distinguishing the species include head structures and proportions, degree of tympanum development, finger webbing, color patterns and male calls. The nominal taxa C. chrysops Cope, C. decorata Taylor, and C. millepunctata Taylor, are placed in the syn- onymy of fleischmanni; C. reticulata Taylor is regarded as a synonym of C. valerioi. Detailed distributional analysis for all 13 Costa Rican species of the genus indicates that most forms occur in lowland or premontane slope evergreen forest. As many as six or seven species may occur at the same locality, although few species show consistent co-occurrence at many sites. No obvious ecologic factor explains the diversity and differences in species composition from site to site. The mosaic distributional pattern and a similar unique mosaic of basic and de- rived features distinguishing each species makes determination of relationships within species groups difficult. Within the fleischmanni line, chirripoi, talamancae, and fleischmanni appear closely allied. Centrolenella vireovittata resembles this stock in basic coloration and male call but is unique in the family in having a striped dorsum. Centrolenella valerioi and C. colym- biphyllum do not appear to be closely related to any other forms, although the two may be included with vireovittata as a subgroup based upon characteristics of the head and snout region. valerioi, C. Among the most characteristic inhabitants of riparian situations along small fast-moving the leaves of streamside plants. In many areas. including vacant lots within some of the large streams in tropical Middle America, are a series of small, delicate, pale green arboreal frogs allied to Centrolenella fleischmanni (Boettger) of the glass-frog family (Centrolenidae). We have dis- cussed elsewhere (Savage, 1967; Savage and Starrett, 1967) the salient features of Central American glass-frog species belonging to the prosoblepon and pulverata groups of the genus Centrolenella and take this opportunity in de- scribing a new Costa Rican form to review the members of the fleischmanni group in the region. Frogs of this group are small (22-27 mm in standard length in adults) elusive creatures with males that call from vegetation overhanging the stream by night and with both sexes hiding by day, well-camouflaged by their coloration, among nr cities, these frogs are extremely common among stands of ginger-lilies (Family Zingiberacae: especially Costus and Hedychium), in and around small streams or even rivulets of only a few millimeters in depth. Members of the group share the following features (egg color not known for all species): no vomerine teeth: white bones; a colorless parietal peritoneum so that the viscera are visible through the transparent ab- dominal skin, in life: white visceral peritoneum: liver confined by white hepatic peritoneum to form a compact bulb-shaped organ rather than Dept. Biological Sciences and Allan Hancock Foundation, University of Southern California, Los Angeles, California 90007. 58 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES the large three-lobed structure characteristic of most other centrolenids; dorsum pale green fad- ing to whitish-yellow in preservative; no humeral hooks; no enlarged forearm muscles in males; no free prepollex or prepollical spines: eggs green, in life, white in preservative (Starrett, 1960) and are usually attached to the under surfaces of leaves overhanging water courses. The green pigment in the eggs is obvious when they are in the oviduct and at the time of laying. As cleavage takes place, the developing embryo becomes whitish-yellow but the yolk retains a greenish cast for some time. Taylor (1958) in the most recent survey of the situation, recognized nine Costa Rican species belonging to the fleischmanni group. Together with the Mexican form C. viridissima (Taylor) this makes a total of ten nominal species for Middle America. Although we regard some of these forms as conspecific, the small size (19-27 mm in standard length), few distinguishing features and subtle differences between closely allied taxa, make identification of preserved material extremely dirSicult. Familiarity with the animals in life in the field is an absolute prerequisite to any at- tempt to clarify the status of these frogs, partic- ularly as the significant features of dorsal colora- tion and the distribution of guanine on the heart and viscera may best be determined in life. Fortunately we have collected and seen living examples of all nominal forms except C. chirripoi and talamancae. Nevertheless, we hold no belief that this review is a final word on the group in the region. It is however the best that can be done on the basis of preserved material and cur- rent field observations. Further resolution of pos- sible problems must await intensive field study with emphasis on male call characteristics. Such studies may reveal the presence of cryptic sympat- ric species at several localities. We encourage herpetologists to undertake further field study of the following problems: 1. The status of the fleischmanni population of southeastern Atlantic slope Costa Rica with moder- ately large yellow dorsal spots and an indistinct but evident tympanic apparatus; the relationship of this population may be with C. talamancae which has a more fully evident tympanum and more protuberant nostrils than the fleischmanni population. 2. The possible occurrence of a form with the morphology of fleischmanni but with a call similar to valerioi at Los Diamantes, Provincia de Limon, Costa Rica. 3. The presence of a diminutive (adult males 19— VOLUME 72 20 mm) form with a distinctive weak fleischmanni- like call from near San Isidro de El General, Pro- vincia de San Jose, Costa Rica. 4. The possible sympatric occurrence of two species of reticulated frogs, all here referred to C. valerioi, from the Peninsula de Osa, Costa Rica. 5. The possible sympatric occurrence of frogs sim- ilar to C. talamancae with colymbiphyllum and fleischmanni in the Cordilleras Tilaran and Central of Costa Rica. GENERAL CHARACTERISTICS The following features are common to the species of the fleischmanni group treated in this report: upper surfaces smooth; nostrils directed laterally; eyes directed forward at about 45° angle to body axis; eye membranes opaque; orbit round; pupil of eye horizontally elliptical; undersides of palms and fingers covered by low smooth tubercles; fingers with lateral fleshy margins; disks rounded to truncate, slightly wider than fingers; thenar tubercle obscure, narrow and elongate; palmar tubercle small rounded; subarticular tubercles moderate; no webbing between fingers I-II; underside of foot smooth; toe disks small, rounded; a weak elongate inner metatarsal tuber- cle, no outer; subarticular tubercles small; modal toe webbing formula I 11-2 If 0-2 III 1-2% IV 2%-1 V:; throat smooth; venter and_ posterior ventral areas of thighs granular, rest of under- sides smooth; choanae large, ovoid; ostia phar- yngia moderate; tongue oval, small; paired vocal slits and a single external vocal sac in males. In all species of this group for which eggs are known (colymbiphyllum, fleischmanni, tala- mancae, and valerioi) the eggs are green but become white upon preservation. Egg color is unknown for chirripoi and vireovittata. Other centrolenids from Costa Rica having green eggs are C. spinosa of the prosoblepon group and C. pulverata of the pulverata group. The eggs of C. albomaculata, C. euknemos, and C. granulosa contain much melanin and appear dark (black or brown) and white in the oviduct or shortly after laying. The eggs of C. prosoblepon are black and C. ilex probably has black eggs as well, since the oviducal eggs are black in the holotype. POPULATIONAL DIFFERENCES Head Structures and Proportions: Among the most distinctive qualitative features distinguishing among the members of this group are the outline of the head viewed from above, the snout profile, 1973 C Figure 1. species of glass-frogs of the fleischmanni group. Dorsal and lateral views, A. Centrolenella fleischmanni, B. C. talamancae, C. C. chirripoi, D. C. vireovittata, E. C. valerioi, F. C. colymbiphyllum. the structure of the canthal and rostral regions, including the degree of development of raised nasal protuberances, the extent to which the eye protrudes laterally, past the margin of the lip, and the relative proportions of the interorbital and loreal (eye to nostril) regions. When viewed from above the head outline may be semicircular with a truncate (Fig. 1E) or slightly indented (Fig. 1D), rounded with a slightly indented rostral area (Fig. IC) or truncate with an indentation between the nostrils (Fig. 1F) or rounded with a weakly pointed tip (Fig. 1A-1B). In lateral view the snout is rounded (Fig. 1B) or vertical (Fig. 1E) in pro- file. In some forms each canthus rostralis is strong and the elevated area between them forms a distinct platform terminating with the SYSTEMATICS OF THE COSTA RICAN GLASS-VROGS 59 F Diagnostic features of head structures and proportions in Central American nostrils (Fig. 1D-1F). In others the canthus is weak and no platform is developed (Fig. 1A-1C). The nostrils open through very slightly indicated fleshy areas (Fig. 1A) in some forms. In others the nasal region is markedly swollen and the nostrils open through elevated fleshy protuber- ances that lie on distinct raised ridges (Fig. 1B-1F). The eyes, when not retracted extend laterally only slightly beyond the lip margin in most forms (Fig. 1A—-1E) but in one species the eyes extend laterally far beyond the lip margin to provide a pop-eyed effect (Fig. 1F). The relative proportions of the loreal (eye to nostril distance) to least interorbital width may also be used to characterize samples. The term loreal short refers to samples in which the loreal 0 BULLETIN SOUTHERN ( A Pericardial Figure 2. pigmentation in live white parietal pericardium; ventral pericardium; ventral view of C. R. W. Merritt. latter, courtesy of distance is less than the interorbital: loreal long which the equals or exceeds the interorbital width. Tympanum: Dunn (1931) and Taylor (1954) pointed out differences in the degree of external is used for forms in loreal distance evidence of the tympanum as a basis for dis- tinguishing among centrolenid species. In most members of the pulverata and 1967; Savage and Starrett, 1967) the tympanic mem- centrolenids (i.e., prosoblepon groups in Costa Rica, Savage, brane is evident although covered by a thin layer of integument and the tympanic annulus is dis- Within the fleischmanni group, some species have the tympanum distinct with the tympanic annulus In others the tym- tinct and demarcates the tympanic area. clearly indicated as above. panum and annulus, while present, are concealed beneath a thick portion of the integument and In this discussion, the term tympanic apparatus distinct (Fig. 1C, 1E, 1F) (Fig. 1A) refer to each of these conditions respectively. In one form (Fig. 1B) the tympanum is evident but between previously described extreme conditions. are only faintly indicated. and tympanic apparatus concealed indistinct and is intermediate the {LIFORNIA AC view of colymbiphyllum. {1DEMY OF SCIENCES VOLUME 72 the valeriot; fleischmanni glass-frogs of group. A. B. Colorless parietal courtesy of W. E. Duellman; Centrolenella Former, Webbing: In all but one species considered in the present report there are no more than vestigial webs at the base of fingers between I-II-II, but the web between fingers III-IV extends at least to the level of the penulti- mate subarticular tubercle of both digits. The modal finger webbing formula for these forms fingers following the system of Savage and Heyer (1967) is I 3-3 If 3-3 Ill 1%-2 IV. In Cen- trolenella chirripoi, the webbing between fingers II-III is almost as extensive as between fingers III-IV and the species has a modal finger web- bing formula of: I 3—3 II 1%—2% III 2-2 IV. Peritoneal and Pericardial Pigmentation: In all members of this group the venter and parietal peritoneum are transparent in life so that the internal are visible. In addition, most of the digestive organs are covered by white vis- ceral peritoneum. The presence of the white peritoneum is absolutely correlated with the occurrence of a compact bulb-like liver in all specimens of this group studied in life or shortly after preservation. Although we have seen no fresh examples of Centrolenella chirripoi, on 7G: these forms have the bulb- organs hepatic talamancae, 1973 SYSTEMATICS OF THI like liver typical of the fleischmanni group and surely have similar peritoneal pigmentation, In the prosoblepon group of centrolenids, the liver is clearly three-lobed. Centrolenella pulverata agtees with members of the fleischmanni group in liver form and in having a white hepatic peritoneum. In most species the heart is enclosed in a similarly colored white sac (Fig. 2A) the parietal peri- cardium, so that the heart appears as a white organ in life. Several other species, including the new form, lack white pigment in the peri- cardial sac, so that the heart appears red (Fig. 2B) in life. Careful dissection of well-preserved animals will usually show whether the heart is protected by white pigment or not, although very old examples or those exposed to strong for- malin or light may lack the pigment even if it were present in life. Since the heart or pericardial sac is visible through the ventral skin in life, these differences are valuable field characters for identifying certain species and should be recorded at the time of capture. Dorsal Color Pattern: The available evidence indicates that at least two chemically distinctive pigments are involved in the green coloration of Central American centrolenids. In members of the prosoblepon group, the green dorsal color of the live frog comes from a pigment con- centrated in a dense system of chromatophores. After a short time in formalin or alcohol this pigment appears purple and the animal seems purple to lavender in color to the unaided eye. In frogs that have been in preservative for a long time or exposed to light, the purple changes to brown. The pigment that appears to be purplish after preservation in this group is here- after called pigment A. Dried skins of fresh examples of the prosoblepon group that have not been fixed in preservation change from green to purple after a short time. In most members of the fleischmanni group the green dorsal color of live frogs is produced by widely scattered chromatophores containing a pigment that turns to purple, or with a long period in preservative to brown, that we assume is pigment A, and a suffusion of green pigment that covers most of the dorsal surfaces. The latter pigment, hereafter called pigment B, is soluble in formalin and alcohol. In frogs of this group, the green rapidly disappears after preserva- tion and they appear a uniform yellowish-white to the unaided eye. Closer examinations reveal that the body is covered by widely scattered chromatophores that appear to be purple to COSTA RICAN GLASS-FROGS 61 Dried of fresh examples of this group that have not been brownish and contain pigment A, skins fixed in preservative resemble preserved examples, In this valerioi (Fig. 3D), pigment Bis one species of group, C. concentrated in the form of a broad reticulum mixed with melanophores. Upon preservation, the green pigment dissolves but the melanophores continue to indicate the extent of the A third pigment coloration of C. reticulum. may be involved in the pulverata. In life, the green coloration of this frog is formed by widely dorsal scattered chromatophores and a diffused green pigment that covers most of the skin area. In preservative the chromatophores change to purple and we assume they contain pigment A. The diffuse pigment dissolves in formalin or alcohol so that to a superficial view, the animal appears to be yellowish-white, but because of the fairly numerous chromatophores containing pigment A, the color has a pale lavender cast. Whether the diffuse soluble green pigment of pulverata corresponds to pigment B is questionable, since dried skins of fresh examples that have not been fixed in preservative remain a bright green with scattered purple chromatophores. Goin and Goin (1968) and Barrio (1965) provide a partial explanation for this situation through their studies of coloration in hylid and pseudid frogs. According to the Goins, the green color of hylids and probably some cen- trolenids that appear purple in preservative is a structural color. In this case three layers of chromatophores are involved. The deepest are melanophores, more superficial are guanophores and near the skin surface are the lipophores. In life, when visible light falls on the skin, the shorter wave length components (greens, blues. purples) are reflected back by the guanophores and the longer wave length components (reds, oranges, yellows) are absorbed by the melano- phores. As reflected the shorter wave lengths pass through the lipophores and the blues and purples are mostly absorbed while the greens are transmitted so that the skin appears green. On preservation, the fatty yellow materials in the lipophores are dissolved or destroyed and the reflected blues and purples are transmitted and the skin appears purplish. If this is in- deed the situation in centrolenids, pigment A is guanine. In centrolenids of the fleischmanni group pig- ment B_ is certainly biliverdin (Goin and Goin, 1968; Barrio, 1965) a green pigment almost 62 BULLETIN SOUTHERN CALIFORNIA VOLUME 72 ACADEMY OF SCIENCES Figure 3. B D Diagrammatic representations of basic dorsal color patterns in Centrolenella fleischmanni and its allies. The letters (A,B.D,) refer to the pattern types discussed in the text. that is accumulated in the skin rather than in chromatophores. On preservation the biliverdin- like pigment in centrolenids is dissolved or de- stroyed so that the skin appears pale yellow or white, although scattered purplish (pigment A) and black (melanophores) chromatophores may be present. Another diffuse green pigment besides biliverdin may be present in the skin of C. pulverata, since the pigment is not de- stroyed when the skin is dried. Barrio (1965) found biliverdin in bones, muscles, eggs, and lymph of several hylid and pseudid frogs. Although it has not been bio- chemically verified, the green pigment in the eggs of members of the fleischmanni group and in the bones of species of the prosoblepon and groups of centrolenids is probably biliverdin. It is interesting to note that in the fleischmanni group, the skin and eggs contain pulverata the green pigment, while in the prosoblepon group, the green pigment is concentrated in the In C. pul- verata, the green pigment is in both skin and bones, but not in the skin or eggs. bones but is much less concentrated than in the skin of fleischmanni and its allies or in the bones of members of the prosoblepon group. In life, all members of the fleischmanni group have a pale leaf green ground color with some light yellow markings. In preservation the yel- low disappears and the green fades until the frog appears almost uniform yellowish-white on casual observation. Closer examination under magnification usually shows that the green pig- mentation is still indicated by scattered contracted purplish or in examples that have been preserved for a long time, brownish chromatophores. Several patterns occur that distinguish several species (Figs. 3, 5): A—dorsum appears almost uniform green in life, but with many small yellow spots; in well-preserved specimens the light areas seen under a microscope correspond to the yellow spots, but because of the contraction of the dark chromatophores are larger in area than the spots in life; B—dorsum green in life with moderate-sized yellow spots; in well-preserved specimens the spots are indi- cated as in A above, with the same qualifications; C—dorsum green in life, with a distinct green mid- dorsal stripe bordered on either side by a_para- 1973 vertebral yellow stripe; remainder of dorsum and flanks with moderate-sized yellow spots; in well-pre- served examples the mid-dorsal stripe is indicated by contracted dark chromatophores and the light para- vertebral stripes by pigmentless areas; the remainder of the dorsum resembles the situation in pattern B above; D—in life the dorsal ground color appears to be a pale yellow to bright yellow-gold; superimposed on the light ground color is a regular broad reticulum composed of many black punctations suffused with a pale green; in some populations the green suffusion extends over the yellow areas surrounded by the re- ticulum so that they appear as a murky chartreuse; the large light spots apparently correspond to the yel- low spots in the other patterns (A, B, C); these light areas are usually at least one-half the eye diameter in pattern D and often are as large as the eye: in preservation the green is completely dissolved but the dark punctations of the reticulum are retained and the pattern is visible to the unaided eye; under magnifi- cation the reticulum is seen to be composed of a series of purplish to black chromatophores (brown in some examples that have been in preservative for a long time). Male Vocalizations: During the rainy season male centrolenids call nearly every evening. The calls seem to be related to both spacing of the males and courtship of the females. The calls of males in the fleischmanni group consist of single notes repeated after a pause of several seconds. Three distinctive calls are known for this group (Fig. 4). In C. valerioi, the note is of short dura- tion (0.2 sec.) and is a rather high pitched seet. In C. fleischmanni and the new species, C. vireovittata, the sound resembles a rising whistled Wheet, is lower pitched and lasts for 0.4—0.5 sec. The most distinctive call is the long trilled musical call (0.6-0.8 sec.) of C. colymbiphyllum. Unfortunately the calls of C. chirripio and C. talamancae remain unrecorded. Other Features: Taylor (1954) emphasized a number of other features as a basis for recogniz- ing species within the group. Review of his material and our extensive samples from through- out Costa Rica indicate that these characteristics vary individually or interpopulationally to such a degree that they cannot be relied upon for species recognition. Eye tunic color—in centrolenids, the outer surface of the eyeball, posterior to the cornea, is covered by a thin membrane that underlies the eyelids. In life this membrane or eye tunic con- tains a heavy concentration of guanine. Upon preservation as the green pigments of the integu- ment fade, the eye tunic pigment may be seen SYSTEMATICS OF THE COSTA RICAN GLASS-FROGS 63 through the upper eyelids as a definite white area. Taylor (1954) noted that in preservative some individuals lacked the white tunic area and he used this difference as a basis for separating several nominal species. The presumed differ- ences (white versus black eye tunics) are arti- facts, for upon preservation and depending upon the nature and time of death, strength of preserya- live, exposure to light and other factors, the white pigment may disappear and the area seen through the upper eyelid appears black from the color of the eyeball. In all the fleisch- manni group seen by us in life, the guanine is present in the eye tunic. members of After death some in- dividuals in any large sample lose the white color, while others retain it for years. Terminal Taylor (1954) de- scribed two pad conditions for this group: a) digital pads rounded, distinctly wider than adjacent part of digit; b) truncate or subtruncate, not or only a little wider than adjacent part of digit. Careful examination of series from single localities in- dicates that these features are variable within a population, with most individuals haying inter- mediate conditions. Thigh folds—three examples within Taylor's large series of fleischmanni have a pair of skin folds that run from the vent laterally and down- ward on the thighs. Five within our samples from widely scattered localities show similar folds formed by a wrinkle in the thigh integument. The folds are not present in living frogs and appear to be an artifact of preservation. caused by the simultaneous swelling of the skin and its separation from the underlying muscle and flexing of the thighs during the death throes. frogs STATUS OF NAMED MIDDLE AMERICAN FORMS Centrolenella_ fleischmanni The earliest available name for Costa Rican frogs of t,vs group is Hylella fleischmanni Boettger, 1893 (holotype: Senckenberg Museum 3760, Provincia de San Jose: San Jose, 1180 m). This form is represented in our collections by numerous specimens from San Jose and the immediate vicinity. Most of the animals called Centrolenella fleischmanni (Noble. 1924) Cen- trolene fleischmanni (Dunn, 1933). Cochranella fleischmanni (Taylor, 1951, 1952, 1958) and Centro- 1964) belong here, al- though it is difficult to separate poorly preserved ma- lenella fleischmanni (Goin, terial from the allied Centrolenella talamancae. Centrolenella fleischmanni may be briefly charac- 64 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES terized as follows: 1) head viewed from above, rounded with a weakly pointed tip; 2) snout truncate in profile, loreal short; 3) canthus rostralis weak, no elevated intercanthal platform; 4) nostrils open through very slightly indicated fleshy areas; 5) eye not protuberant: 6) tympanic apparatus concealed; 7) reduced webbing between fingers I-II—II, but web between fingers III-IV well-developed; 8) parietal pericardium white; 9) dorsum green with small to moderate yellow spots; 10) male call a single rising untrilled wheet, repeated after a pause. Cope (1894) described Hylella chrysops from Ala- juela and San Jose, Costa Rica and Taylor (1954) revived the name for fleischmanni-like frogs from near Sarchi, Provincia de Alajuela and Provincia de Cartago, Costa Rica. The sole nominal difference be- tween clirysops and fleischmanni, according to Taylor is the presence of a black eye tunic in the former and a white tunic in the latter. Since this difference is an artifact of preservation as pointed out above and since our extensive material indicates that only one species resembling fleischmanni occurs on the Meseta Central Occidental of Costa Rica, we regard the two forms as conspecific. The types of chrysops are lost and in order to assure the stability of our conclusion, we herewith designate Senckenberg Museum 3760, the holotype of Hylella fleischmanni, as the neotype of Hylella chrysops. Taylor (1942) differentiated Centrolenella viridis- sima (holotype: FM 100093, Mexico: Guerrero: Agua del Obispo) from C. fleischmanni of Chiapas on the basis of slight mensurational differences and eye tunic color (black in the former, white in the lat- ter). Duellman (1960) concluded that all Mexican material represented a single species, that the charac- teristics used by Taylor were variable in Mexico and overlapped to a considerable extent with series of Costa Rican fleischmanni, but tentatively recognized viridissima as distinct. We can find no basis for sep- arating viridissima from fleischmanni particularly as the Mexican and Costa Rican samples are connected by a continuous series of populations ranging through Guatemala, El Salvador, Honduras, and Nicaragua. The name viridissima is based upon a population hav- ing the parietal pericardium silvery white as in all fleischmanni and cannot be confused with the popu- lations here referred to C. colymbiphyllum, which have the pericardial sac colorless. Cochranella decorata Taylor, 1954 (holotypé “KU 36896, Costa Rica: Provincia de Cartago: Hacienda La Florencia, 4.8 km SW Turrialba, 800 m) was recognized as distinct solely on the basis of having prominent skin folds on the posterior thighs lateral to the vent. In addition to the holotype, Taylor re- ferred two examples (KU 36883-84, presumably from Costa Rica: Provincia de San Jose: San Jose, 1180 m) to this species. We have examined the types and five additional Costa Rican examples (Provincia de Alajuela: Candelaria de Naranjo, CRE 525, 2 specimens; nr. Carrizal, CRE 6037; Provincia de San VOLUME 72 Jose: La Palma, CRE 503, 2 specimens) with the thigh folds and conclude that the folds are artifacts of preservation, as noted above. All eight examples agree in every other respect with typical fleischmanni taken at the same localities and we refer C. decorata to the synonymy of the latter form. Cochranella millepunctata Taylor, 1954 (holotype: KU 36887, Costa Rica: Provincia de San Jose: La Palma, 1500 m) was distinguished from fleischmanni by its describer by having the terminal digital pads enlarged, distinctly wider than adjoining part of dig- its (versus digital pads truncate or subtruncate, not or only a littke wider than adjoining part of digit in fleischmanni). As noted above the shape of the terminal pads varies from rounded to truncate in samples of fleischmanni from single localities. The holotype and paratypes all from Costa Rica (topo- typic examples: KU 36883—86, 36888—-94; KU 36897 from Provincia de Alajuela: Sarchi and KU 23943 probably from Provincia de Cartago: Moravia de Chirripo) seem to represent the extreme rounded and expanded pad condition but other examples from the same localities have subtruncate to truncate pads and many are intermediate between the extreme condi- tions. For this reason we regard millepunctata as a synonym of C. fleischmanni. Centrolenella fleischmanni, as conceived of in this report, has a broad geographic and altitudinal range: lowlands and premontane slopes in humid areas from Guerrero and Veracruz, Mexico, south through Cen- tral America to western Ecuador and through north- ern Colombia and Venezuela to Surinam. Centrolenella colymbiphyllum In Costa Rica, C. fleischmanni is sympatric at a num- ber of localities with another small form that resem- bles it superficially both in life and in preservative. The earliest name for this second species is Centro- lenella colymbiphyllum Taylor, 1949 (holotype: KU 23812, Costa Rica: Provincia de Heredia: Isla Bonita 1200 m). Centrolenella colymbiphyllum may be dis- tinguished by the following series of characteristics: 1) head viewed from above truncate with an indenta- tion between nostrils; 2) snout truncate in profile, loreal long; 3) canthus rostralis strong, with an ele- vated intercanthal platform; 4) nasal region markedly swollen and nostrils opening through elevated fleshy protuberances that lie on distinct raised ridges; 5) eyes protruding laterally well beyond level of margin of lips; 6) tympanic apparatus distinct; 7) reduced webbing between fingers I-IJ-III, but web between fingers HI-IV well-developed; 8) parietal pericardium colorless; 9) dorsum green with small to moderate yellow spots; 10) male call a single sustained trill, repeated after a pause. Centrolenella colymbiphyllum occurs in Costa Rica in humid areas of the premontane zone on both slopes of the Cordillera de Tilaran, on the Atlantic slope of the Cordillera Central and extreme north- 1973 eastern extensions of the Cordillera de ‘Talamanca; the species has also been taken in Pacific lowland forests from the area just west of San Isidro de El General in the Golfo Dulce region of southwest’ Pa- cific Costa Rica. Centrolenella colymbiphyllum may have a substantially broader geographic range than suggested by Costa Rican samples. Specimens of fleischmanni allies lacking white pericardial pigment have been noted from Guatemala, Honduras, Panama, and Venezuela and some of these frogs may be referred to the present species after detailed study. Centrolenella chirripoi Taylor (1954) also described another distinctive spe- cies apparently related to both fleischmanni and colymbiphyllum, but closest to the former, as Cochi- ranella chirripoi (holotype: KU 36865, Costa Rica: Provincia de Limon: Rio Cocolis, near Suretka, 100 m). Examination of the holotype and eight topo- typic paratypes indicates that this form is a valid species that may be characterized as follows: 1) head viewed from above rounded with a slightly indented area between nostrils; 2) snout rounded in profile, loreal long; 3) canthus rostralis weak, but with an elevated intercanthal platform; 4) nasal region mark- edly swollen and nostrils opening through elevated fleshy protuberances that lie on distinct raised ridges; 5) eyes not protuberant; 6) tympanic apparatus dis- tinct; 7) webbing between fingers II-III-IV extensive: 8) color of parietal pericardium not known; 9) dorsum green with small to moderate yellow spots: 10) male call not known. Centrolenella_ valerioi Dunn (1931) described another distinctive fleisch- manni ally from Costa Rica as Centrolene valerioi (holotype: MCZ 16003, Costa Rica: Provincia de San Jose; La Palma, 1370 m). Because the original de- scription implied that the only two known examples, the holotype and a topoparatype (MCZ 16005), had a distinct narrow mid-dorsal stripe, Taylor (1954) failed to recognize the species among his materials. Re-examination of the type specimens of valerioi proves it to be conspecific with examples of a species of Centrolenella with a dorsal pattern of green re- ticulations on a light yellowish ground color. The “mid-dorsal green stripe’ mentioned by Dunn corre- sponds to the portion of the green reticulum that tends to coalesce down the back in this form. Dunn (1933) later associated the types of valerioi with ex- amples of the reticulate form from Panama: Cocle: El Valle de Anton and emphasized the “green chain markings” of the types. This species may be charac- terized by the following: 1) head viewed from above semicircular, with a truncate or slightly indented tip: 2) snout truncate in profile, loreal long; 3) canthus rostralis strong, with an elevated intercanthal plat- SYSTEMATICS OF THE COSTA RICAN GLASS-FROGS 65 form; 4) nasal region markedly swollen, with nostril opening through clevated fleshy protuberances that lie on distinct ridges; 5) cyes not protuberant; 6) tym panic apparatus distinct; 7) reduced webbing between fingers I-II-III, but web between fingers I-IV well developed; 8) parietal pericardium white; 9) dorsal pattern of large yellow spots surrounded by a strong reticulum of green pigment and dark punctations; 10) male call a single short seet, repeated after a pause. Cochranella reticulata Taylor, 1954 (holotype: Kt 32922, Costa Rica: Provincia de Cartago: near main bridge over Rio Reventazon, at Instituto Interameri- cano de Ciencias Agricolas, Turrialba, 591 m) is a strict synonym of C. valerioi. It was also recorded by Taylor from Costa Rica: Provincia de Cartago: Moravia de Chirripo; Provincia de Heredia: Cari- blanco; Provincia de Limon: Suretka; Provincia de Puntarenas: Golfito, Palmar. The reticulated species, C. valerioi, is known from the Atlantic foothills and slopes of the Cordillera Central of Costa Rica south along the margin of the Cordillera de Talamanca to the lowlands of the Valle de Talamanca near the Panama boundary; it also occurs on the Pacific versant in lowland sites from near San Isidro de El General south through south- western Costa Rica to El Valle de Anton in central Panama. Cochranella talamancae Taylor, 1952 (holotype. KU 4143; Costa Rica: Provincia de Cartago: Moravia de Chirripo, 1116 m) is an enigmatic form. Accord- ing to Taylor (1952; 1958) the species is closely al- lied to C. valerioi but re-examination of the type and three additional examples (KU 32936-38) referred to talamancae by Taylor (1958) do not support this concept. Salient features of this series are: 1) head viewed from above, rounded with a weakly pointed tip; 2) snout rounded in profile, loreal short: 3) canthus rostralis weak, no elevated intercanthal plat- form; 4) nostrils in protuberant fleshy swellings: 5) eye not protuberant; 6) tympanic apparatus indistinct: 7) reduced webbing between fingers I-II-III. but web between fingers III-IV extensive; 8) color of parietal pericardium unknown; 9) dorsum green with moder- ate yellow spots: 10) male call unknown. In most features, except 4 and 6, talamancae re- sembles C. fleischmanni populations from southeast- ern Atlantic Costa Rica. It is possible that the ap- parent distinctiveness of the nostrils and tympanum are the result of desiccation in preservative. If C. talamancae is valid we have been unable to locate additional examples in our extensive series and sur- mise that study of living material is needed to clarify its status. Tentatively, we recognize it as a distinct form known only from the type locality. NEW SPECIES FROM COSTA RICA Among the materials collected in Costa Rica during our fieldwork that began in 1957, is a 66 BULLETIN SOUTHERN CALIFORNIA (KHz) FREQUENCY me NW PUD NN Ow ACADEMY OF SCIENCES VOLUME 72 oe? 8 eres TIME INTERVAL = 0.1 SECONDS Figure 4. Sonograms of male vocalizations in glass-frogs of the fleischmanni group from Costa Rica. Left to right: Centrolenella valerioi, Provincia de Puntarenas: 3 km SW Rincon de Osa, November 22, 1968. C. fleischmanni, Provincia de San Jose: Curridabat, October 29, 1968: C. vireovittata, Provincia de San Jose: 0.5 km NE Alfombra, August 5, 1963; C. colymbiphyllum, Provincia de Puntarenas, 3 km W Rincon de Osa, August 22, 1969. small series of striped Centrolenella of the fleisch- manni group having a colorless pericardium. We first took this species in the Cordillera Costena between San Isidro de El General and the Pacific coastal village of Dominical in 1963. It occurs sympatrically with the reticulated species called valerioi in the present report, but because of Dunn’s (1933) reference to a green stripe down the back in the type of the latter, we presumed that the striped form belong to Dunn’s species. As pointed out above, the name valerioi is based on an example of the reticulated species and does not have a mid-dorsal stripe. The striped species cannot be referred to any other described form and may be known as: Centrolenella vireovittata, new species Holotype: LA 75141, (Fig. 5) an adult male, from Costa Rica: Provincia de San Jose: Canton Perez Zeledon: 0.5 km NE Alfombra: a place 16 km SW San Isidro de El General on the road to Dominical, 880 m; collected by Jay M. Savage and Norman J. Scott, July 17-18, 1963. Diagnosis: The new species is immediately distin- guished from all other centrolenids known from Mex- ico and Central America by its unique striped pattern (Fig. 5). No other striped species have been de- scribed in the family. Definitive Characteristics: 1) head viewed from above, semicircular in outline with a truncate or slightly indented tip; 2) snout truncate in profile, loreal long; 3) canthus rostralis strong, with an ele- vated intercanthal platform; 4) nasal region markedly swollen and nostrils opening through elevated fleshy protuberances that lie on distinct raised ridges; 5) eyes not protuberant; 6) tympanic apparatus distinct; 7) reduced webbing between fingers I-IJ-III, but web between fingers II-IV, well-developed; 8) pari- etal pericardium colorless; 9) dorsum green with small to moderate yellow spots and a mid-dorsal green stripe bordered by a pair of paravertebral yel- low stripes; 10) male call a rising whistled wheet. Size: The holotype (22 mm) and five adult male paratypes (CRE 688, 2647, 6101-02, 7220A—B) range from 21.5—23 mm in standard length. 1973 SYSTEMATICS OF Remarks: Calling males of this species were taken at the type locality on three occasions, July 17-18 and August 5, 1963 and June 1, 1964. The frogs were found on the upper sur- faces of leaves on low herbaceous bushes between 0.5—1 m above a shallow (5-7 cm deep) stream about 1.5 m wide. A sonograph of the call is shown (Fig. 4). The type locality is in a Premontane Rain- forest, following the classification of Holdridge (1967), along the Pacific slope of the Cordillera Costena that separates the narrow coastal plain from the interior valley of the Rio General. This place, near the small village of Alfombra, is also the type locality for five other nominal taxa of reptiles and amphibians: Anolis humilis mar- supialis Taylor, 1956; Dermophis occidentalis Taylor, 1955; Dipsas tenuissima Taylor, 1954; Hyla legleri Taylor, 1958; and Neusticurus apo- demus Uzzell 1966. Although the Anolis and Dermophis are representatives of wide-ranging species (A. humilis and D. parviceps, respec- tively) the Hyla and Dipsas are valid species restricted in range to Pacific southwestern Costa Rica and adjacent extreme southwestern Panama. The Neusticurus and Centrolenella are not known to occur elsewhere. At the type locality Centrolenella vireovittata was found sympatrically with C. valerioi of the fleischmanni group. Centrolenella_ fleischmanni and C. colymbiphyllum are known from a nearby locality Quebrada Salto, 6.4 km SW San Isidro de El General, near Palma on the road to Dominical, 750 m, on the interior slope of the Cordillera Costena. Centrolenella granulosa and C. prosoblepon of the prosoblepon group were also taken in virtual sympatry with C. vireo- vittata at the type locality. DISTRIBUTIONS Fleischmanni Group: The following lists include all Costa Rican locality records for the members of the fleischmanni group based upon material at the Uni- versity of Southern California and the samples now at the University of Kansas that formed the basis for Taylor’s (1958) review. Centrolenella chirripoi—LIMON: Suretka on Rio Cocolis, 60 m (Fig. 6). Centrolenella colymbiphyllum—ALAJUELA: Cari- blanco; 1 km SW and Cinchona; CARTAGO: 8.8 km NE and Tapanti, Rio Quiri; GUANACASTE: Finca San Bosco; PUNTARENAS: Golfito; Las Cruces: 0.5 km SE, ESE, SW, 1.75 km ESE, 2.6 km ESE Monteverde; 3 km W, 3 km SW Rincon de Osa; SAN THI COSTA RICAN GLASS-FROGS 67 ( ee MS) 9 (Gas WA Figure 5. Dorsal view of a male paratype (USC- CRE 6101) of Centrolenella vireovittata; scale equals | cm. JOSE: Quebrada Salto, 6.4 km S San Isidro de El General; 6-1450 m (Fig. 6). Centrolenella_ fleischmanni—ALAJUELA: Cande- laria de Naranjo; Rio Cariblanco and Sarapiqui road: nr. Carrizal; Cinchona; nr. La Florencia; 3.2 km E La Fortuna; San Jose, nr. Rio Jesus; Sarchi; Rio Segundo; nr. Tacares; Tesalia; 8 km N Zarcero; CARTAGO: Cartago; 1.5 km SW Chitaria: Floren- cia; Instituto Interamericano de Ciencias Agricolas, nr. Turrialba; Moravia de Chirripo; 1 km S Orosi; 1 km E Pacayas; 1 km NE and Tapanti: 1-2 km SE Santa Teresa; GUANACASTE: 1.9, 3 km E, 2 km NW, 1.2 km SW Tilaran; HEREDIA: 1.6 km ENE La Uvita; San Jose de la Montana; LIMON: 16 km SW (Surayo), 20 km SW (Alto Lari) Amubri; Los Diamantes; Guacimo; Pandora; PUNTARENAS: 1.6 km SW Aguabuena: Rio Coton: 0.5 km ESE, 0.5 km SW Monteverde; SAN JOSE: Alto Guadelupe: Rio Claro-Rio La Hondura; Curridabat; Desamparados: 3.2 km WSW Escazu: El General: Guadelupe; Que- brada Salto, 6.4 km S and San Isidro de El General; San Jose; La Palma; Quizarra; San Pedro, Ciudad Universitaria; 50-1680 m (Fig. 7). This species is known from humid lowland and premontane forests from Mexico throughout Central America through Colombia to northwestern Ecuador and eastward through Venezuela to Surinam. It is possible that more than one species is involved in the material as- signed to fleischmanni by Goin (1964), but only field studies give hope of clarifying their status. Centrolenella talamancae—CARTAGO: Moravia de Chirripo, 1116 m (Fig. 6). Centrolenella valerioQh—ALAJUELA: Cariblanco. Rio Cariblanco and Sarapiqui rd.; CARTAGO: 1.5 km S Chitaria; Florencia: Moravia de Chirripo; nor. bridge over Rio Reventazon, 4.5 km SE Turrialba: LIMON: Suretka; PUNTARENAS: 6.4 km SE and Golfito; Palmar; 1 km W Platanillo; 3 km W, 2.5 68 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES km SW Rincon de Osa, Quebrada Aguabuena; 4.8, 5.1, 5.6 km NW Villa Neilly; SAN JOSE: 0.5 km NE Alfombra; Quizarra; La Palma. Also in PAN- AMA: COCLE: El Valle de Anton; 6-1500 m (Fig. 8). Centrolenella vireovittata—SAN JOSE: 0.5 km NE Alfombra, 880 m (Fig. 6). Sympatric occurrences of two or eVen three species of the fleischmanni group at one locality is not un- common in Costa Rica. Sympatry is used in the sense that the individuals were collected at the same gen- eral locality at the same time of year. Detailed field studies will be required to demonstrate how various facets of the biology of these species may act to pre- vent interbreeding and to foster ecologic segregation at any particular site. Centrolenella colymbiphyllum, C. fleischmanni and C. valerioi have all been taken at ALAJUELA: Cariblanco 830 m: C. fleischmanni, C. talamancae, and C. valerioi are sympatric at CARTAGO: Moravia de Chirripo, 1116 m. Localities where two species have been taken sympatrically are: Centrolenella colymbiphyllum—C. fleischmanni: ALAJUELA: Cinchona, 1350 m; CARTAGO: Tap- anti, 1200 m; PUNTARENAS: vicinity of Monte- verde, 1400 m; SAN JOSE: Quebrada Salto, S San Isidro de El General, 750 m. Centrolenella colymbiphyllum—C. valerioi—PUN- TARENAS: Golfito 6 m; 3 km W Rincon de Osa, 25 m. Centrolenella fleishmanni—C. valerioi-CARTAGO: Instituto Interamericano de Ciencias Agricolas, 4-5 km SSE Turrialba, 602 m; SAN JOSE: La Palma, 1500 m:; Quizarra, 720 m. Centrolenella valerioi—C. vireovittata—SAN JOSE: 0.5 km NE Alfombra, 880 m. Centrolenella valerioi and C. chirripoi are almost certainly sympatric at LIMON: Suretka, 60 m. Prosoblepon and Pulverata Groups—In earlier pa- pers (Starrett, 1960, Savage, 1967; Savage and Star- rett, 1967) we have discussed the distinguishing char- acteristics and general distribution of members of these groups in Costa Rica and Panama. We take this opportunity to provide a full listing of Costa Rican localities in order that our views on the current systematic status and distributions for Costa Rican glass-frogs may be available in a single paper. Prosoblepon Group Centrolenella albomaculata—ALAJUELA: Sarchi; Tesalia; CARTAGO: 1.5 km S Chitaria; nr. Tur- rialba; GUANACASTE: 2 km NW, 1.2 km SW and Tilaran; LIMON: 20 km SW Amubre (Alta Lari); 1.6 km E, 1.6 km N and Los Diamantes; PUN- TARENAS: 6 km SE Golfito; Finca Loma Linda, 3.2 km SSW Canas Gordas; Rio Ferruviosa 6 km S Rincon de Osa; SAN JOSE: nr. Juntas de Rio Gen- eral; Quebrada Salto, 6.4 km S and San Isidro de El General; 4-1180 m. Also known from PANAMA: COCLE: EI Valle de Anton (Fig. 9). VOLUME 72 Centrolenella euknemos—SAN JOSE: 1.5 km S Alto La Palma; Rio Claro—Rio La Hondura; 1150- 1500 m. Also known from PANAMA: CANAL ZONE: Summit; DARIEN: Rio Jaque 1.5 km above Rio Imamado; Laguna; PANAMA: §S slope Cerro Campana; SAN BLAS: Summit Camp (Fig. 10). Centrolenella granulosa—ALAJUELA: nr. Concep- cion; La Garita; Tesalia; CARTAGO: 1.5 km SW Chotaria; Moravia de Chirripo; 5 km SE Turrialba; GUANACASTE: 3 km NW Arenal; LIMON: Los Diamantes; PUNTARENAS: 4.8 km, 5.1 km NW Villa Neilly; 3 km W Rincon de Osa; SAN JOSE: 0.5 km NE Alfombra; Quebrada Salto, 6.4 km S San Isidro de El General; 25-1116 m. Also known from PANAMA: BOCAS DEL TORO: 4.8 km W Almirante; CHIRIQUI: Progreso; DARIEN: Reversa de Pava, Rio Tuira; PANAMA: nr. Rio Chico Hydrographic Station. Villa (1972) tentatively identified specimens from Nicaragua as this form. We have seen examples (KU 85474) from NICARAGUA: MATAGALPA: Finca Tepeyac to confirm Villa’s records from the same departamento at La Cumplida and San Jose de la Montana (Fig. 10). Centrolenella ilee—ALAJUELA: nr. Concepcion; 8 km NW Ciudad Quesada; CARTAGO: 1.5 km SW Chitaria; LIMON: 16 km SW Amubre (Surayo); 250-775 m; PANAMA: no other data, FM 153710, first record for the republic (Fig. 9). Centrolenella prosoblepon—ALAJUELA: 0.3 km N Salto Angel; Cariblanco; nr. Carrizal; 1 km S and Chichona; La Fortuna; San Jose, nr. Rio Jesus; 4.8 km S Ciudad Quesada; Sarchi; 1.6 km, 8 km §S Zar- cero; CARTAGO: Cartago; Casa Mata; 1.5 km S Chitaria; Santa Cruz; Moravia de Chirripo; Pacayas; Tapanti, Rio Quiri; Santa Teresa; 3.2 km SE Tres Rios; HEREDIA: 4.3 km S Santa Domingo de El Roble; 1.6 km NW La Uvita; LIMON: 8 km SW Amubre; Los Diamantes; La Junta; Puerto Limon; PUNTARENAS: Aguabuena; Las Cruces; El Hele- chales; 0.5 km SE, 0.5 km SW Monteverde; 1 km W Platanillo; 5 km SSW Rincon de Osa, Rio Ferruviosa; SAN JOSE: 0.5 km NE Alfombra; Santa Ana, Rio Oro; Bebedero de Escazu; Rio Claro—Rio La Hon- dura; 1 km S San Cristobal Sur; 3.2 km WSW Escazu; San Isidro de Coronado; La Palma; Quebrada Salto, 6.4 km S San Isidro de El General; San Pedro; Ciudad Universitaria; Salitral; 20-1900 m (Fig. 11). The species is also known from southeastern Atlantic lowland Nicaragua and in humid situations on both coasts of western and central Panama south to north- western Ecuador. Centrolenella spinosa—ALAJUELA: 3.2 km E La Fortuna; Tesalia; HEREDIA: La Selva; LIMON: Bambu; Los Diamantes; PUNTARENAS: 2.5 km SW Rincon de Osa; 20-260 m. Also known from PAN- AMA: CANAL ZONE: Barro Colorado Island (Fig. 10). We also have seen specimens probably represent- ing this species from western Colombia and Ecuador. 1973 SYSTEMATICS OF THE COSTA RICAN GLASS-FROGS 69 © CHIRRIPO! @ COLYMBIPHYLLUM m TALAMANCAE oe VIREOVITTATA eS KILOMETERS Figure 6. Distribution of Costa Rican glass-frogs allied to Centrolenella fleischmanni; the dotted line indicates the 1500 m contour. KILOMETERS ] | | ut | Y = 86 85 84 83 Figure 7. Distribution of Centrolenella fleischmanni in Costa Rica: the dotted line indicates the 1500 m contour. 70 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES KILOMETERS Figure 8: Distribution of Centrolenella valerioi in Costa Rica; the dotted line indicates the 1500 m contour. © ALBOMACULATA a ILEX KILOMETERS Figure 9. Distribution of Costa Rican glass-frogs of the prosoblepon group; the dotted line indicates the 1500 m contour. VOLUME 72 1973 SYSTEMATICS OF THI COSTA RICAN GLASS-FROGS @ EUKNEMOS ‘D GRANULOSA @ SPINOSA 50 "KILOMETERS Figure 10. Distribution of Costa Rican frogs of the prosoblepon group: the dotted line indicates the 1500 m contour. 86 10 KILOMETERS 86 Figure 11, Distribution of Centrolenella prosoblepon in Costa Rica: the dotted line indicates the 1500 m contour. 71 ™s tw BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 50 ———| —— KILOMETERS Figure 12. Distribution of Centrolenella pulverata in Costa Rica; the dotted line indicates the 1500 m contour. Pulverata Group Centrolenella pulverata—ALAJUELA: Rio Ron Ron, 8 km N Ciudad Quesada; Tesalia; CARTAGO: Florencia; LIMON: 14.5 km SW Amubre; 1.6 km E and Los Diamantes; PUNTARENAS: 2.5 km SW, 3 km W Rincon de Osa; SAN JOSE: nr. Juntas de Rio General; Quebrada Salto, 6.4 km S San Isidro de El General. Also known from PANAMA: CHIRIQUI: Progreso; DARIEN: Rio Jaque, 1.5 km above Rio Imamado; Rio Siluganti. Villa (1972) tentatively in- cluded this species in the fauna of Nicaragua without further data. He apparently had seen a specimen (KU 85476) from NICARAGUA: MATAGALPA: Finca Tepeyac, which is unquestionably pulverata (Fig. 12). Sympatric occurrences of two to four species of the prosoblepon group at a single locality, often together with C. pulverata are common in Costa Rica. Al- though all species have not been taken at precisely the same locality, or on the same date, four species of the prosoblepon group (albomaculata, granulosa, prosoblepon, and spinosa) and pulverata occur to- gether in the area immediately southwest of Rincon de Osa, Provincia de Puntarenas (20-30 m) on the Pacific lowlands, although albomaculata has not ac- tually been taken in the same stream course with the others. All five of these forms occur in the same stream, at Los Diamantes, Provincia de Limon (260 m) on the Atlantic versant. Two species of the fleischmanni group (colymbiphyllum and_ valerioi) occur at the former locality making a total of seven sympatric glass-frogs at that place and one (fleisch- manni) at Los Diamantes for a total of six species. Other areas with high species numbers include: the Rio Lari canyon southwest of Amubre, Provincia de Limon (300-800 m), with C. albomaculata, C. ilex, C. prosoblepon, and C. pulverata and the type species of the fleischmanni group; the area just southwest of Turrialba, Provincia de Cartago (600 m) with C. albomaculata, C. granulosa, and C. pulverata and C. fleischmanni, and C. valerioi of the fleischmanni group; the region where the Rio Chitaria crosses the Turrialba to Puerto Limon road, 1.5 km S Chitaria, where C. albomaculata, C. granulosa, C. ilex, and C. prosoblepon occur with C. valerioi of the fleisch- manni group; at Tesalia, Provincia de Alajuela (€00 m) where C. albomaculata, C. granulosa, C. spinosa, and C. pulverata occur with C. fleischmanni and Quebrada Salto, Provincia de San Jose (750 m) where C. albomaculata, C. granulosa, and C. pul- verata are found with C. colymbiphyllum and C. fleischmanni. Additional records of sympatry within the prosoblepon and pulverata groups include: albomaculata—granulosa—P ANAMA: COCLE: El Valle de Anton, 800 m. albomaculata—pulverata—SAN JOSE: nr. Juntas de Rio General, 530 m. | ii | 1973 SYSTEMATICS OF THE COSTA RICAN GLASS-FROGS 73 albomaculata—prosoblepon—A LAJUELA: Sarchi, = =x = 970 m. = =o - Aus oa euknemos—prosoblepon—SAN JOSE: 1.5 km S sz=e seE=ze2aso as A : = 5 I St ep o Alto La Palma, 1500 m; Rio Claro—Rio La Hondura, =e ae a -—— — Z2z2~2 = > ed — Dept. Geography, Univ. California 92502. California, Riverside. 98 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 Figure 1. of southern California (refer to cover illustration). Guide to the mountain ranges, geomorphic provinces, and other points of interest 1, Sierra Nevada and Tehachapi Moun- tains; 2, Mt. Pinos-Tecuya Mountain—Plieto Hills area; 3, Santa Ynez Mountains; 4, San Gabriel Mountains; 5, Santa Susana Mountains and Oak Ridge; 6, Simi Hills; 7, Santa Monica Mountains; 8, Verdugo Mountains and San Rafael Hills (3-8, Transverse Range); 9, Fullerton Hills; 10, Puente Mountains; 11, Santa Ana Mountains (9-11, Peninsular Ranges). CV, Central Valley (San Joaquin Valley), note farmland; LA, Metropolitan Los Angeles; MD, Mojave Desert; OC, Orange County; OXP, Oxnard Plains; note farmland; SFV, San Fernando Valley; SV, Simi Valley. pear black. Concrete appears light grey. Intensity of reflectance in each spectral band as well as color composition serve as indicators of ground cover and its condition. High and low quality resi- dential areas can frequently be distinguished by the intensity of the red “signature” mixed with colors that represent streets and buildings. Tech- niques are available to enhance particular levels in intensity in each band to facilitate identifica- tion, as well as to automatically scan, identify and measure color composition and intensity levels in the total scene. Multi-spectrality clearly is a boon to the photointerpreter. Traditional photo-interpretative tools are widely 1973 A NEW RESEARCH AND MANAGEMENT TOOL 99 el’ 2S i Cll 28} Figure 2. Guide to other points of interest shown in cover illustration. 1, Mt. Pinos: 2, Old Fort Tejon; 3, Castaic Lake; 4, Fairmont Reservoir; 5, Hughes and Elizabeth Lakes: 6, Bouquet Reservoir; 7, Lake Palmdale; 8, Rosamond Dry Lake; 9, Rogers Dry Lake: 10, alluvial fans; 11, Mirage Dry Lake; 12, Mt. Baden Powell: 13, Mt. San Antonio-“Mt. Baldy”: 14, Morris Reservoir; 15, Castaic Reservoir; 16, Lake Piru; 17, Forestfire scar; 18, Lake Casitas; 19, Van Norman Reservoir: 20, Lake Sherwood in Hidden Valley: 21, Los Angeles International Airport; 22, Anaheim Back Bay; 23, Newport Back Bay: 24, Pt. Hueneme: 25, Pt. Mugu; 26, Pt. Dume; 27, Malibu Point, note bulldozer-cleared area for Pepperdine Univer- sity left of point; 28, Marina del Rey; 29, Palos Verdes Peninsula; 30, Los Angeles-Long Beach Harbor. Major faults shown in photo are: A, Garlock fault; B, San Andreas fault: C. San Gabriel fault (star indicates epicenter of 1971 Sylmar earthquake): D, Chino-Elsinore fault; E, Norwalk fault; F, Newport-Inglewood fault (star indicates epicenter of 1933 Long Beach earthquake): G, Malibu fault; H, Anacapa fault. Major river drainages shown are: LAR, Los Angeles River: SAR, Santa Ana River: SAnR, San Antonio River: SCR, Santa Clara River: SGR, San Gabriel River, VR, Ventura River. 100 BULLETIN SOUTHERN CALIFORNIA to interpret ERTS Using these tools, the more primary geomorphic and geologic Surrounding used images. features can easily be identified. the metropolis are two great mountain systems (Fig. 1), the Transverse Mountain Ranges on its north and northwest the Peninsular Ranges on its southeast side. Components of this system include the rugged San Gabriel Moun- tains with peaks up to 10,000 ft which wall in the northern edge of the metropolis; the Santa Monica Mountains which thrust boldly into the heart of the great city; and the Santa Ana Moun- tains which frame the portion of the metropolis lying in Orange County. Along the coastline, Pt. Dume and Palos Verdes are prominent headlands framing Santa Monica Bay. Most prominent among the geologic features are the major faults: The San Andreas, the Garlock, San Gabriel, and Big Pine Faults. These faults hint at the powerful natural forces that menace the metropolis to the south. Between the San Andreas and Garlock Faults lies the broad, angular expanse of the western Mojave Desert (Antelope Valley) only 35 miles northerly of downtown Los Angeles, where many geo- morphic as well as geologic features can be discerned. Among them are a series of alluvial fans, and the Rosamond and Rogers Playas (dry lake beds). The upper left corner of the scene contains the low-lying, southern end of California’s Cen- tral Valley. Dividing it from the elevated western end of the Mojave are the Southern Sierra Ne- vada and the Tehachapi Mountains. Hydrologic features are clearly shown, from major rivers down to many smaller streams and sides and tributaries. Among the major features the Los Angeles, San Gabriel, Santa Ana, and Santa Clara Rivers are identified. Other hydrologic features of note include the Los Angeles Aque- duct and a number of major reservoirs. Agricultural districts stand out boldly. Clock- wise from the top several are visible: the double- lobed irrigated district of the Antelope Valley, the dairy region in San _ Bernardino-Riverside Counties, the Irvine Ranch, the diverse, richly productive fields of the Oxnard Plain, and the southern San Joaquin Valley, California’s largest and most important agricultural region. The actual and potential applications of ERTS data are immense. The applications in education towards creating an awareness of the nature of ACADEMY OF SCIENCES VOLUME 72 the earth’s surface, its environmental character, and man’s use of it are obvious. Another power- ful application is in the area of scientific re- search involving fields such as geology, oceanog- raphy, hydrology, botany, pedology, climatology, and geography. A vast range of resource man- agement investigations are currently under way in agriculture, forestry, fisheries, mineral explora- tion, and water and air quality where a wide range of analysis and data handling procedures including monitoring capabilities are being ex- plored. Also under study is the use of ERTS data for urban and regional planning and manage- ment. One such project, funded by NASA, is a joint study by the Regional Planning Commis- sion of Los Angeles County, California, and the General Electric Company of Philadelphia, Pennsylvania, to evaluate ERTS-—1 data for its urban and regional planning application. This project stresses a semi-automatic electronic anal- ysis of multi-spectral information from ERTS. Results from this project indicate that ERTS—1 imagery will primarily yield new and unconven- tional data about the intensity of urban develop- ment, an often-used surrogate for environmental quality. Through ERTS synoptic coverage, de- sign concepts can be defined and intra-urban as well as inter-urban comparisons made. These results stem from viewing ERTS as a new re- search tool, the smallest scale available in a multi-stage process of urban and regional anal- ysis, and viewing multi-spectral analysis as a means of rapidly obtaining and identifying in- formation for object classes that are characterized by color. These results will not replace older forms of data used by planners nor readily con- form to them, but will give planners the capability to undertake old and new problems more ef- fectively. ACKNOWLEDGMENT This paper is based on data obtained from the Regional Planning Commission of the County of Los Angeles, California, and the General Electric Com- pany who are engaged in a joint project to analyze Earth Resources Technology Satellite (ERTS—1) data for its urban and regional planning applications. The joint project is completely funded by a National Aeronautics and Space Administration Grant. James Smith and Susan Payne prepared figures 1 and 2. Accepted for publication June 1, 1973. LATITUDE AND LITTER SIZE OF THE CALIFORNIA GROUND SOUIRREL, SPERMOPHILUS BEECHEY 1 JOSEPH A. CHAPMAN ABSTRACT: ' AND GORDON An inverse relationship between latitude and_ litter in the California ground squirrel (Spermophilus beecheyi). S. LIND? W to xISt litter size as found c Correlation analysis of size with climatic and physical factors revealed that the best correlation existed between litter size and the length of the freeze-free period, It is hypothesized that the larger litter sizes in southern localities may be a response to heavier predation brought about by an op- portunity for more ground squirrel activity outside the burrow at southern latitudes, where the freeze-free period is longer. The purpose of this paper is to report on the relationship between latitude and litter size of the California ground squirrel (Spermophilus beecheyi) and to suggest possible reasons for the relationships which exist. Lord (1960) reported direct correlation between latitude and mean litter size in several “nonhibernating prey species” of mammals. Other workers have reported similar relationships for additional mammals (Rowan and Keith, 1956; Burns and Burns, 1957; Blus, 1966). Lord (1960) suggested that mortality rates for populations of these species were higher in northern latitudes than in southern latitudes, and that higher productivity was essential to offset the higher mortality rates. Because of the shorter breed- ing season at higher latitudes, increased pro- ductivity was accomplished through an_ in- crease in the litter size rather than an increase in the number of litters. Spencer and Steinhoff (1968) rejected Lord’s hypothesis that increased litter size was a response to higher mortality rates. They proposed that “length of season and parental mortality related to repro- duction” were selective forces by which mean litter size was established in the different en- vironmental conditions encountered at the various latitudes. Lord (1960) failed to find a significant re- lationship between latitude and mean litter size in the one group of “hibernating prey species” of mammals (Spermophilus) which he examined. He postulated that the lack of relationship be- tween latitude and mean litter size among this group was caused by responses to the densities of breeding populations, which were related to the length of time that pressures of predation were avoided through hiberation. Spencer and a 101 Steinhoff (1968) suggested that their hypothesis would explain the absence of latitudinal variation in litter size in ground squirrels, because these usually have single litter Thus, the length of the breeding season would species a annually. have little effect on the annual production in these species. METHODS During June—August, 1967-68, California ground squirrels were collected in Benton and Douglas Counties, Oregon. Adult females were examined by necropsy and implantation sites in each uterus counted. Mean numbers of implantation sites per parous female were considerably lower (Table 1) than litter sizes reported for the species in general references on mammals (Hall and Kel- 1959; Asdell, 1964). Because of these dif- ferences, a search was conducted for information on mean numbers of embryos or implantation sites per parous female California ground squirrel. Records of litter sizes of California ground squirrels were obtained for nine other localities (Table 1). However, one of these records was not used in the correlation analysis because the mean litter size was believed to be abnormally large in response to an intensive control program son, (Jacobsen, 1923). Weather data were obtained (1959) and Sternes (1960). Estimated altitudes of the collection sites were taken from from Elford mean 1 Natural Resources Institute, Univ. Maryland, La- Vale 21502. =U. S. Bureau 97303. of Reclamation, Salem. Oregon 102 state highway maps. Four climatic and physical factors were considered as possible causes of the relationship between latitude and litter size: mean annual rainfall; 3) mean annual temperature; and 4) mean length of the freeze-free period (Table 1). Statistical tests used in the analysis of the data were taken (1960) and Dixon 1) mean altitude; 2) from Steel and Torrie and Massey (1969}. RESULTS AND DISCUSSION Mean litter sizes plotted against corre- sponding latitudes, and a line was fitted to these points by the least squares method (Fig. 1). An F-test indicated that the slope of the line differed significantly from a zero slope (P < 0.01). The calculated coefficient of determination (0.762) implied that more than 76 percent of the variation in litter size was associated with latitude (Table 2). An inverse relationship be- tween latitude and mean litter size in the Cali- fornia ground squirrel was evident. The addition of the quadratic term to the least squares line were TABLE 1. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 9 r= 0.762 w Y= 16,501~- O0.253X N ° tr ° 8 “ net ° E < a9 > es measure- ments tested (Table 1). Females were larger than the males in all of these measurements. In the remaining seven, males averaged larger in two (length of tail vertebrae and length of ear) and females were larger in five (total length, length of hind foot, length of forearm, postorbital constriction, and breadth across upper molars). Conclusions. —Of the 12 measurements analyzed, only length of tail exhibited enough individual varia- tion to warrant its deletion in analysis of geographic or interspecific variation in the genus Choeroniscus. Also, because of the difficulty in consistently taking the measurement, we also suggest elimination of post- orbital constriction. Specimens of Choeroniscus intermedius were found to exhibit significant secondary sexual variation in five of the 12 measurements studied. Therefore, it is clear that males and females should be ‘separated in anal- yses of variation within members of the genus. Fe- males were found to be the larger in 10 of the 12 measurements—similar to the situation found in sey- eral other groups of bats (see for example, Jones and Schwartz, 1967; Jones e¢ al., 1971; Peterson, 1965). Specimens examined. — TRINIDAD: Blanchisseuse, 5; Guayaguayare, 12; Las Cueves, 14; Maracas Val- ley, 1; San Rafael, 4. LITERATURE CITED Genoways, H. H., and J. K. Jones, Jr. 1971. Sys- tematics of southern banner-tailed kangaroo rats of the Dipodomys phillipsii group. J. Mamm., §2:265-287. Handley, C. O., Jr. 1966. Descriptions of new bats (Choeroniscus and Rhinophylla) from Colombia. Proc. Biol. Soc. Washington, 79:83-88. Jones, J. K., Jr., H. H. Genoways, and R. J. Baker. 1971. Morphological variation in Stenoderma rufum. J. Mamm., 52:244—247. Jones, J. K., Jr., and A. Schwartz. 1967. Bredin- Archbold-Smithsonian biological survey of Do- minica. 6. Synopsis of bats of the Antillean genus Ardops. Proc. U.S. Nat. Mus., 124(3634) :1-13. Long, C. A. 1968. An analysis of patterns of varia- tion in some representative Mammalia. Part I. A review of estimates of variability in selected measurements. Trans. Kansas Acad. Sci., 71: 201-227. RESEARCH NOTES 107 1969. in some representative Mammalia. Part Il. Stud An analysis of patterns of variation ics on the nature and correlation of measures of variation. Pp. 289-302, in Contributions in Mam malogy. (J. K. Jones, Jr., ed), Misc. Publ. Mu Nat. Hist., Univ. Kansas, 51:1—428. Peterson, R. L. 1965, A review of the bats of the genus Ametrida, family Phyllostomidae. Contrib. Life Sci., Royal Ontario Mus., 65:1—13. Smith, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 56:1—132. Sokal, R. R., and F. J. Rohlf. 1969. Biometry .. . W. H. Freeman and Co., San Francisco, xxi 4 1-776. HuGu H. Genoways, The Museum, Texas Tech Uni- versity, Lubbock, 79409; Ropert J. BAKER, and W. BERNARD Wyatt, Department of Biology and The Museum, Texas Tech University, Lubbock 79409. Accepted for publication June 1, 1973. A NEW SPECIES OF ELEUTHERODACTYLUS (AMPHIBIA: LEPTODACTYLIDAE) FROM ANDEAN ECUADOR Access to the Amazonian slopes of the Ecuadorian Andes has been difficult in the past except through the deep canyon of the Rio Pastaza (Banos to Puyo), and this route traverses little of the altitudinal transect on the eastern face of the Cordillera Oriental. The building of the Texaco-Gulf transandean pipe- line prompted the construction of a road along its course and field parties from the University of Kansas have worked the Amazonian slope portion of the transect as it was being developed. On one of his trips along this road, William E. Duellman secured two specimens of a moderate-sized frog beneath rocks in a stream. At first glance the frogs appeared to be a treefrog of the genus Hyla, but dissection of the digits and closer examination revealed them to represent a distinctive new species of Eleutherodac- tylus. Eleutherodactylus pugnax, new species Holotype—University of Kansas Museum of Natural History (KU) 146466, an adult female collected at Salto de Agua, 2.5 km NNE Rio Reventador. Napo Prov., Ecuador, 1660 m by W. E. Duellman, Bruce MacBryde, and John Simmons, 7 April 1972. Topoparatype—KU_ 146467. a male. collected syntopically. 108 Figure 1. Male paratype of Eleutherodactylus pug- nax, new species. From a kodachrome by William E. Duellman. Diagnosis.—A species of Eleutherodactylus unique in the following combination of characters: first finger shorter than second; all digits bearing broad discs; toes basally webbed with prominent lateral fringes; skin of venter coarsely areolate; ear absent (no tympanic annulus, cavum tympanicum, or columella); snout round in dorsal view, truncate in lateral profile, shorter than eye length; prevomerine dentigerous processes and teeth present. Only three other species of Eleutherodactylus now known lack ears—E. anotis Walker and Test (Vene- zuela), E. surdus (Boulenger) (high Pacific slopes of Andean Ecuador), and an unnamed species from the paramos of southern Ecuador. None of these has toe webbing or the prominent lateral fringes of the toes seen in E. pugnax. All three also have longer snouts (eye length equal to or less than eye-nostril dis- tance). The unnamed species is a small frog (adult females 20-21 mm SVL) with minute prevomerine processes and concealed prevomerine teeth. Description—Head as wide as body, wider than long; snout semicircular in dorsal outline, truncate in lateral profile (Fig. 1), short, eye length much greater than eye-nostril distance; upper jaw barely over- hanging lower jaw; canthus rostralis round, concave; nostrils near tip of snout, protuberant, directed laterally; loreal region flat; lips not flared; eyelids narrow, their width about equal to interorbital dis- tance; interorbital space flat, no frontoparietal ridges; frontoparietals complete, no fontanelle, not in con- tact with nasals; nasals narrowly separated; supra- tympanic fold thick, glandular; tympanum absent; BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 Figure 2. pugnax. Right, hand of holotype of Eleutherodactylus pugnax. Left, foot of holotype of Eleutherodactylus tongue longer than wide, weakly notched posteriorly, posterior edge free; choanae round, slightly larger than a prevomerine dentigerous process, not con- cealed by palatal shelf of maxillae; prevomerine dentigerous processes round, median and posterior to choanae, bearing 3-4 teeth along the posterior border. Skin of dorsum bearing flattened, ill-defined warts; flanks areolate; skin of upper eyelid covered with numerous wart-like ridges; skin of venter and about anus coarsely areolate; discoidal folds prominent; concealed surfaces of limbs smooth; upper surfaces of limbs smooth with scattered flattened warts; outer edge of forearm bearing a row of ulnar warts; three palmer tubercles, inner largest; numerous well- defined supernumerary palmer tubercles; subarticular tubercles round, non-conical, distal tubercles tend to be bifid; fingers bearing weak lateral fringes, lacking web (Fig. 2); all digits bearing discs which are wider than long; discs largest on fingers II, III, and IV; first finger shorter than second. No tubercles on heel or outer edge of tarsus; inner edge of tarsus bearing a fold along its distal one-third; two metatarsal tubercles, outer flat, ill- defined, one-fifth to one-sixth size of elongate inner (twice as long as wide, not compressed); few super- numerary plantar tubercles; subarticular tubercles smaller than those of fingers, elongate, simple, non- conical; toes bearing discs that are smaller than those of fingers; toes with prominent lateral fringes (Fig. 1973 2) and basal webbing; the webbing formula for the holotype (based on the formula given by Savage and Heyer, Beit. Neotropischen Fauna 5:111-131, 1968): I 2-2 If 2-3" WI 2%-4 IV 3%-2% V; legs of moderate length, heel of adpressed hindlimb reaches to tip of snout; when legs are flexed at right angles to the sagittal line, heels touch. The holotype is an adult female with convoluted oviducts and a few moderate-sized (2.0 mm in di- ameter) yellow eggs interspersed among many small (0.5-1.0 mm) white eggs. The paratype is an adult male with vocal slits and a sub-gular vocal sac; the testes are large and white and the thumbs are swollen at their bases. The measurements (in mm) are as follows; data for the holotype are given first, data for the paratype follow in parentheses: snout-vent length 30:8 (22.1), shank 17.0 (12.2), head width 12.2 (8.6), head length 9.7 (7.3), eyelid width 2.6 (1.7), interorbital distance 2.5 (2.2), eye length 4.2 (3.0), eye-nostril distance 2.8 (2.5). In preservative, E. pugnax is brown with darker brown blotches, interorbital triangle, and limb bars. The limb bars are about as wide as the interspaces. Dark brown canthal and supratympanic stripes and labial bars are present. The posterior surfaces of the thighs are brown with cream flecks. The venter of the female is dirty cream with numerous brown spots; the lower surfaces of the hindlimbs are brown with cream reticulation. The venter of the male is creamy-white with a few brown flecks (primarily on the lower venter); the undersides of the thighs are heavily speckled with brown. In life, BE. pugnax was described as “Dorsal sur- faces and flanks grayish-brown with dark brown markings. Venter gray with brown flecks. Iris red- dish-brown.” (W. E. Duellman field notes, 7 April 1972). Etymology.—Latin, pugnax, meaning fighter; in loose reference to the collector, William E. Duell- man. Natural history—The pronounced lateral fringing and basal webbing of the toes suggests that E. pugnax is a riparian species. The holotype and paratype were collected beneath rocks in a fast-moving stream in cloud forests by day. The male is reproductively active (swollen, non-spinous thumb; large testes) but the female probably is not. Relationships—Few mainland Eleutherodactylus species have toe webbing; those that do (e.g., E. anomalus, E. bufoniformis, E. fitzingeri) are mem- bers of the FE. binotatus group [first finger longer than second, skin of venter smooth, ear prominent (7.e., tympanic annulus large and externally visible) | and frequently live near streams. The West Indian Eleutherodactylus with toe webbing and prominent fringes are likewise strongly riparian in habit (Schwartz, Stud. Fauna Curacao and other Carib. Islands 24:1-62; Shreve and Williams, in Williams, et al., Bull. Mus. Comp. Zool. 129:291—342, 1963). I do not think that either of the more distinctive RESEARCH NOTES 109 features of 2. pugnax (earlessness, toe webbing) are reliable indicators of relationship. Eleutherodactylus member of the Cochran and Goin’s (Bull. United States Nat. Mus (288):1-655, 1970) Group If which I prefer to call the £E, group, defined by granulate or areolate skin on the first finger being shorter than the second. Fleuthero dactylus pugnax has no apparent within this group. William E. Duellman loaned the specimens, pro vided additional information about the locality and its habitat, and permitted me to use his kodachrome of the frog. pugnax 18 a Wusuigas having a venter and the close relatives JOHN D. Lyncn, Dept. Zoology, University braska, Lincoln, Nebraska 68508. of Ne- Accepted for publication April 5, 1973. A NEW SPECIES OF TREMATODE (TREMATODA: MONOGENEA) FROM THE GILLS OF DIAPHUS WATASEI JORDAN AND STARKS, 1904. Through the courtesy of B. Nafpaktitis, Department of Biological Sciences, University of Southern Cali- fornia, some monogenetic trematodes from the gills of a myctophid fish, Diaphus watasei Jordan and Starks, 1904 (identified by Nafpaktitis) were ob- tained for study. The fish were collected between 400 and 450 fathoms off the southeast coast of Africa by the Anton Brunn International Indian Ocean Expedition. Two of eighteen fish had a total of five parasites that are described here as a new species of Diclidophora Diesing, 1850. They were stained with Mayer’s paracarmine, cleared with methyl benzoate, and mounted in Canada balsam. Measurements are expressed in microns. Diclidophora sprostonae,. new species Diagnosis: With the characters of the genus. Length 2198, 1092 wide. Opisthaptor attached to ventral side of body, with 8 short peduncles and clamps (two lost from best extended specimen) 770 long, 182-266 wide. Clamps with muscular pads, 140- 160 long. 112-126 wide. Oral cavity followed by pharynx 65 long, 44 wide. Body tapers anteriorly. Esophagus 2-3 times length of pharynx. Intestinal crura confluent with ticula directed peripherally and medially. A ticulum extends into opisthaptor. Testes diver- posteriorly numerous in radiating cords, mainly postovarial between testinal crura. Convoluted vas deferens extends an- mid-esophageal teriorly to ventral genital pore at 110 BULLETIN SOUTHERN CALIFORNIA Figure I. Ventral view of Diclidophora sprostonae, new species; C, clamps; G, genital pore; I, intestine; O, ovary; OP, opisthaptor; P, pharynx; S, sucker; T, testes; U, uterus; V, vitellaria; VA, vas deferens; VR, vitelline reservoir. level. Cirrus diameter about 30, armed with six curved spines about 9 long, ovary ribbon-like, looped, median and pretesticular. Vitelline follicles small, numerous, in groups between intestinal diverticula and extending into opisthaptor. Vitelline reservoir adjacent to ovary, with expanded anterior end. Genitointestinal canal on right side. Seminal re- ceptacle not seen. Uterus median, tubular, from testicular level to common genital pore. None contained eggs. Host: Diaphus watasei Jordan and Starks, 1904. Habitat: Gills. Type Locality: Off southeast coast of Africa. Holotype: No. 639, deposited in the Hancock Parasitology Collection, University of Southern Cali- fornia. ACADEMY OF SCIENCES VOLUME 72 Figure 2. sclerites of A, Cirrus spines and B, clamp; (M, muscular pad) of Diclidophora spros- tonae, New species. DISCUSSION These worms were removed from the gills of fish that had been preserved for several years and most of them were poorly distended. Diclidophora spros- tonae differs from other speceis of the genus in the numbers of cirrus spines and in the fan-like arrangement of the testes. Several authors (Llewel- lyn and Tully, J. Fish. Res. Bd. Canada, 26: 1063-— 1074, 1969; Sproston, Trans. Zool. Soc. London, 25: 185-600, 1946, etc.) have mentioned the varia- tion in the number of cirrus spines or hooks in in- dividuals of the same species. All specimens of Diclidophora sprostonae possess six hooks. Per- haps a larger number of specimens might show variation in this character. Although all specimens 1973 appeared mature, none had eggs. Yamagutt (Sys- tema Helminthum, Vol. 1V. Monogenea and Aspido- cotylea. Intersci. Publ, New York, 1963) stated that members of the genus Diclidophora are pata- sitic on the fish families Gadidae, Merlucciidae, and Macrouridae. The present paper adds the family Myctophidae. Monogenetic trematodes are gener- ally considered to be host specific. W. E. Martin, Dept. Biological Sciences, Uniy. Southern California, Los Angeles, California 90007. Accepted for publication May 2, 1973. A NEW SPECIES OF STRAPAROLLUS (ARCHAEOGASTROPODA) FROM THE MISSISSIPPIAN OF NEVADA During a recent biostratigraphic study (Mount, 1972a, 1972b) of the larger invertebrates in the Up- per Paleozoic section of the central Pancake Range, Nye County, Nevada, I placed particular emphasis on the distribution of Mollusca in the Mississippian and Pennsylvanian rocks. Excluding the Cephalop- oda, mollusks of this age are poorly known in the Great Basin area of North America. In his pioneer- ing study of the Eureka district of Nevada, Walcott (1884) described several new bivalves and gastro- pods, however in later years so little attention has been given to this group that at the present time it is virtually useless for age determination and correla- tion. Recently, the stratigraphic positions of Wal- cott’s species have become better known as a result of the biostratigraphic work of Gordon (1971). It is the purpose of this paper to describe a new species of the gastropod Straparollus. Numerous fossil lists include reference to this genus, but it is generally represented by material too poorly preserved or too scanty to allow species assignment. References to the Department of Geological Sciences, University of California, Riverside, are hereafter abbreviated as UCR. Family Euomphalidae de Koninck, 1881 Genus Straparollus de Montfort, 1810 Subgenus Euomphalus Sowerby, 1812 Straparollus (Euomphalus) pancakensis, new species Figure 1 Straparollus (Euomphalus) n. sp. A: Mount, 1972b, p. 75, pl. 2, figs. 18, 19, 20. Diagnosis: Shell relatively small; whorls 5: spire slightly elevated; single high, broad, rounded spiral rib medially situated between the upper suture and outer shoulder. Description: Shell relatively small for the genus, subdiscoidal; spire slightly elevated, the first two RESEARCH NOTES II] Figure 1. new species. B. Basal view. C. Apertural view. Straparollus (Euomphalus) Holotype UCR 7101/1. v9 pancakensis, A. Top view. whorls flattened; moderately phaneromphalus, um- bilicus deep with nearly vertical sides, umbilical angle near 80 degrees. Whorls 5, with moderately rounded angulations on outer-upper and -lower shoulders, outer sides of whorl moderately convex, umbilical profile of whorl weakly convex to flattened. Upper sutures shallow and_ slightly impressed, passing around the previous whorl just below the upper shoulder. Protoconch unknown. Spiral sculpture con- sists of a single high, broad, rounded rib between upper suture and angulation on the outer shoulder. width of rib about one third that of whorl. slopes of rib slightly concave. Growth lines orthocline. extend- ing outward to rib on upper whorl, there bending into a sharply defined but shallow sinus: beyond rib growth lines curve slightly backward around whorl to lower suture forming a slightly projecting upper lip over the aperture. Holotype: UCR 7101/1; height 7.8 mm, diameter 20.8 mm, height of aperture 6.0 mm. 112 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Paratypes: UCR 7101/2; height 7.3 mm, di- ameter 20.5 mm, height of aperture 6.7 mm (shell incomplete). UCR 7101/3; height 5.2 mm, diameter 18.8 mm, height of aperture 4.1 mm (shell incom- plete). Type locality: UCR locality 7101; 6530 feet south and 675 feet west of long. 115° 50’ W., lat. 38° 55’ N.: at elevation of 7,240 feet On crest of ridge extending west from first hill north of hill 7865, Brown Summit 714 minute quadrangle (1968 edi- tion), Nye County, Nevada (same as Department of Geology, University of California, Los Angeles, lo- cality 6058-60). Stratigraphic Position: Ely Group, Moleen For- mation, unit 60 of Mount (1972a, 1972b); the fossils occur in platy, thin-bedded, dark gray, silty limestone 20 feet above the base of the formation and can be collected best from the abundant weath- ered platy slabs that cover the slopes. Age: In addition to the gastropod described here, the following fossils have been identified: SCYPHOZOA: Paraconularia sp. BRACHIOPODA: Composita cf. C. subquadrata Hall, Leiorhynchus carboniferum Girty, Ovatia sp. indet. GASTROPODA: Bellerophon (Bellerophon) sp. BIVALVIA: Aviculopecten cf. A. affinis Walcott, Ayiculopinna cf. A. consimilis (Walcott), Schizodus cf. S. semistriatus Girty. SCAPHOPODA: Plagioglypta sp. indet. TRILOBITA: Paladin sp. BLASTOIDEA: Pentremites sp. indet. CONODONTA: Cavusgnathus gigantus Gunnell, C. lautus Gunnell, C. spathus Dunn, Gnathodus aff. G. girtyi simplex Dunn, G. muricatus (Dunn), Spathognathodus cf. S. minutus (Ellison), Strepto- enathodus unicornis Rexroad and Burton. This fauna is judged to be Late Chester (Late Mississippian) in age and is assigned to the upper part of the Rhipidomella nevadensis Assemblage Zone of Sadlick (1954). This assignment is based primarily on the presence of the conodont Gnathodus aff. G. girtyi simplex which occurs in this zone in southern Nevada (Webster, 1969) and of the charac- teristic Mississippian brachiopods Composita cf. C. subquadrata and Leiorhynchus carboniferum. Also, about 175 feet higher in the same section Rhipi- domella nevadensis (Meek) is abundant in beds that also contain Mississippian species of the brachiopod genera Anthracospirifer, Cleiothyridina, Flexaria, In- flatia, and the bryozoan Archimedes. Discussion: Straparollus (Euomphalus) pancak- ensis is easily distinguished from all other species of the subgenus Exvomphalus by its more strongly pronounced revolving rib which is more medially located between the upper suture and outer shoulder. The only North American species that approaches it is S. (E.) umbilicatus (Meek and Worthen, 1860: 462) the Middle from Pennsylvanian Desmoines VOLUME 72 Series of Illinois (Knight, 1934: 151), the latter form having a much higher spire, larger number of whorls, narrower spiral rib, and more angulated whorl profile. Straparollus (Euomphalus) planodorsatus (Meek and Worthen, 1860: 462) from the Mississippi Valley is the only species of this subgenus recorded from the Chester Series and it is readily distinguished from the new form described here by the lack of a spiral rib, more rounded whorl profile, higher spire, and somewhat larger size. The new name proposed here refers to the Pan- cake Range, the type locality. LITERATURE CITED Gordon, M., Jr. 1971. Biostratigraphy and age of the Carboniferous formations, in Mississippian stratigraphy of the Diamond Peak area, Eureka County, Nevada. U. S. Geol. Survey Prof. Pap. 661:34—-S5. Knight, J. B. 1934. The gastropods of the St. Louis, Missouri, Pennsylvanian outlier; 7, The Euomphalidae and the Platyceratidae. J. Paleo., 8:139-166. Meek, F. B., and A. H. Worthen. 1860. Descrip- tions of new Carboniferous fossils from Illinois and other western states. Proc. Acad. Nat. Sci. Philadelphia, 12:447—472. Mount, J. D. 1972a. Late Paleozoic biostratig- raphy of the Pancake Range, Nye County, Nevada. M.S. thesis, Dept. Geology, Univ. Cali- fornia, Los Angeles, viii + 79 pp. 1972b. Late Paleozoic biostratigraphy of the Pancake Range, Nye County, Nevada. So. California Paleo. Soc. Spec. Publ., No. 1:1- 80. Sadlick, W. 1954. Mississippian-Pennsylvanian boundary in northeastern Utah. J. Paleo., 28: 506-507. (Abstr.) Walcott, C. D. 1884. Paleontology of the Eureka district. U. S. Geol. Survey Mon., 8:1—298. Webster, G. D. 1969. Chester through Derry conodonts and stratigraphy of northern Clark and southern Lincoln Counties, Nevada. Univ. California Publ. Geol. Sci., 79:1-121. Jack D. Mount, Dept. Geological Sciences, Uni- versity of California, Riverside 92502. Accepted for publication March 12, 1973. INSTRUCTIONS FOR AUTHORS - The BULLETIN is published three times cach year (April, August, and November) and includes articles in English in any field of science. 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Use numerals before units of measurements: 5 ml, but nine spines (10 or numbers above, such as 13 spines). The metric system of weights and measurements should be used wherever possible. Taxonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International Code of Botanical Nomenclature (Lawjouw et al., 1956), the International Code of Nomencla- ture of Bacteria and Viruses (Buchanan ef al., 1958), and the International Code of Zoological Nomenclature (Stoll et al., 1961). Special attention should be given to the description of new taxa, designation of holo- type, etc. The literature cited section should include six or more references; entries for books and articles should take these forms. McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54: 452-458. Brattstrom, B. H. 1969. The condor in California. 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CONTENTS The systematic status and distribution of Costa Rican glass-frogs, genus Centrolenella (Family Centrolenidae), with description of a new species. By Priscilla H. Starrett and Jay M. Savage The chiggers (Acarina, Trombiculidae) parasitizing the Side-blotched Lizard (Uta stansburiana) and other lizards in Joshua Tree National Monument, California. By Richard B. Loomis And Robert Se Stephenscticrs cschia cress iievwhs a lelslevelalsu syakelalers: ayoilelone, oitoleherelohelteley kak meee eRe renin Notes on bats from the Mexican state of Queretaro. By David J. Schmidly and Chester O. Martin Colubrid snakes of the genus Tantilla from Nicaragua. By Larry David Wilson and Jaime Villa Earth resources technology satellite: a new research and resource management tool. By Jene MekKnightsand\) Darryl Goehring) a. cccc 2'2)2.cnieo)* «felienerel ier enet lores aiels\o)-1 +1 ier ore eae Latitude and litter size of the California ground squirrel, Spermophilus beecheyi. By Joseph Ay Chapman and: Gordon iSong oc eate | a fosekel ala =) ere ets: SL (mm) 33°15’/55”N, 118° 13/50” W 214-234 Velero IIT 1151-40 5/7/40 5 2 2.4-2.4 (USNM 143010) 33°15’30’N, 118°15’/05”’W 190247 Velero IV 1848-49 12/6/49 I | 1.9-2.4 (AHF) 33°15/00"N, 118°11’35”W 265-283 Velero II 1188—40 29/9/40 | 1 2.1—2.3 (AHF) 33°14/15”N, 118°10'00”W 278-366 Velero IIT 1430-41 25/10/41 3 1 1 2.2-3.4 (AHF ) 33°00/45”N, 118°42’/00”W 91-274 Velero HL 1018-39 23/11/39 6 3 1.9-3.3 (AHF) 33°00/30”N, 118°32’20”W 95-112 Velero III 1239-41 22/2/41 1 2.9 (AHF) 33°00/20’N, 118°33/10”’W 110-155 Velero III 911-39 18/2/39 9 I I 1.7-3.6 (AHF) 32°59/30”N, 118°38’25”W = 216-384 Velero HI 1026-39 9/12/39 1 3.1 (AHF) 32°46/30’N, 118°21/15’W 143-391 Velero III 914-39 19/2/39 4 2 2.43.1 (AHF) 32°45’55”N, 118°26'10”W 100-126 Velero IIT 1012—39 9/11/39 7 3 2 1.6—2.6 (AHF; USNM 143008) 32°35’/00’N, 119°11’45”W 165-201 Velero III 1343-41 10/6/41 4 1 1.9-3.5 (AHF) 32°33”15°N, 117°22'05”W = 143-148 Velero III 1240-41 23/2/41 3 1.9-2.5 (AHF) West coast Baja California, Mexico 28°12/35”N, 115°33/15’W =121-148 Velero III 1251-41 26/2/41 1 17 (AHF) 28°12’/05”N, 115°33’20”W =: 130-174 Velero III 1010-39 20/9/39 1 1.4 (AHF) 28°05’45”N, 115°31/00"W 117-119 Velero III 1253-41 26/2/41 1 2.5 (AHF) 26°16/17”N, 113°41/03”W 99 Velero IV 1710-49 7/3/49 12 1 3 1.8—2.7 (AHF; USNM 143009) Gulf of California 28°00'02”N, 111°24’40”W 201 Velero III 735-38 29/3/37 1 2 2.4—2.9 (AHF) change. In accordance with the Rules of Zoo- logical Nomenclature (article 49), the name mertensii Cannot be retained for Holmes’ species even though it is now being transferred to yet another genus. The references of both Alcock (1905) and Gordan (1956) refer, in part, to Parapagurodes makarovi and, in part, to Pagurus mertensii [sensu lato]. The distributions recorded by Rath- bun (1904, 1910) and Schmitt (1921) of Kam- chatka, Alaska and British Columbia are er- roneous, and reflect the misidentification of this species by Benedict and Holmes. Taylor’s (1912) record of Parapagurus mertensii from southern British Columbia, which was repeated by Hart (1940), was most probably based on misidentifi- cations of specimens of Pagurus h. hirsutiusculus. Parapagurodes laurentae, new species Figures 4b, 9-11 ?Pagurus [sp]: Menzies and Miller, 1954. p. 153 (see discussion). HOLOTYPE—é#@ (SE = 3.5 PARATYPES.—See table 3. TYPE LOCALITY.—2% mi SE of Seal Rocks, Santa Catalina I., California, 33° 16’ 20” N, 118 15’ 20” W, Velero III station 1429-41, 159-174 m. DESCRIPTION.—Shield slightly longer broad, or less frequently. mm), AHF 4127. equalling width, length 30 Fig sta. d, BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 ure 9. Parapagurodes laurentae, n. sp., paratype, ¢@ (SL = 3.2 mm), “Velero III” 1429-41: a, shield; b, right cheliped (dorsal view); c, right cheliped (mesial view); left cheliped (dorsal view). Scales equal 1 mm (a) and 3 mm (b-d). 1973 DECAPOD CRABS FROM CALIFORNIA Figure 10. Parapagurodes laurentae, n. sp., paratype, ¢ (SL = 3.2 mm), “Velero II” sta. 1429-41, a,b, Ps: a, lateral view: b, mesial view.—c,d, P;: c, lateral view: d. mes view.—e, Px, lateral view. Scales equal 3 mm (a—d) and 1 mm (e). 132 BULLETIN SOUTHERN CALIFORNIA rarely slightly broader than long; anterolateral margins sloping; anterior margin between rostrum projections somewhat concave or oc- casionally straight; posterior margin truncate or roundly truncate; dorsal surface slightly inflated, convex, usually with few scattered tufts of short setae; anterolateral angle produced, blunt or with small spine or spinule. Rostrum moderately long, or less frequently, short, usually exceeding lateral projec- tions, depressed distally; triangular, usually slender, occasionally moderately broad, frequently with weak median keel; terminating acutely or subacutely, often with small spine and few moderately long setae. Lateral projections obtusely triangular, unarmed or occasionally with submarginal spinule. Ocular peduncles moderately short, one-half to two-thirds length of shield, stout; slightly inflated basally and with corneae dilated; dorsal surface with irregular row of tufts of short setae. Ocular acicles triangular or subtriangular, narrow or moderately broad, with dorsal surface frequently slightly con- cave; terminating subacutely with strong, submarginal spine; separated basally by one to one and one- half times basal width of one acicle. Antennular peduncles moderately long, exceeding ocular peduncles by three-fourths to entire length of ultimate segment. Ultimate segment with one or two short setae on dorsal surface distally; penulti- mate segment often with few short setae; basal seg- ment with small spine on dorsolateral margin, ventro- distal margin often with small spinule and tuft of setae. Antennal peduncles moderately long, exceeding ocular peduncles by one-fourth to three-fourths length of ultimate segment; with supernumerary seg- mentation. Fifth segment unarmed, usually with few stiff setae or bristles dorsally and ventrally. Fourth segment unarmed, with few tufts of short setae. Third segment with strong spine at ventro- distal margin, partially obscured by tuft of setae. Second segment with dorsolateral distal angle pro- duced, terminating in strong spine, dorsal surface and mesial margin occasionally with one or two small spines or spinules and tufts of short to long setae, lateral margin with scattered short setae; dorso- mesial distal angle with strong spine, mesial margin with short, stiff setae. First segment with small spine or spinule on lateral face distally; ventrodistal margin produced, with one to several spines later- ally. Antennal acicle moderately long, slightly shorter than or equalling length of ocular peduncles, and usually exceeding proximal third of ultimate peduncular segment; terminating in strong, acute spine; dorsal surface and mesial margin with tufts of stiff setae. Antennal flagella long, equalling or exceeding tip of right cheliped; naked or with inter- mittent, short bristles. Mandible without significant characters. Maxil- lule with proximal endite slightly tapered; endopodite with one bristle on moderately well-developed in- and lateral ACADEMY OF SCIENCES VOLUME 72 ternal lobe, external lobe moderately well-developed, not recurved. Maxilla with endopodite long, exceed- ing scaphognathite in distal extension, slightly re- flexed. First maxilliped with flagellum of exopodite short, slender. Second maxilliped with basis-ischium fusion incomplete. Third maxilliped with — basis- ischium fusion incomplete; basis with one to three strong teeth; ischium with crista dentata well-de- veloped, one accessory tooth; merus and carpus each with spine at dorsodistal margin. Sternite of mxp, With prominent spine and row of long setae on either side of midline. Right cheliped usually long, moderately slender, particularly in large specimens; surfaces usually with scattered tufts of long, stiff setae. Dactyl moderately long, two-thirds to three-fourths length of palm, occasionally equalling or slightly exceeding length of palm; set at slightly oblique angle to palm; terminating in small, corneous claw, slightly over- lapped by fixed finger; cutting edge with calcareous teeth on proximal one-half to two-thirds, replaced distally by row of small, corneous teeth; usually with prominent, longitudinal hiatus; dorsomesial margin with row of moderately strong, blunt, or, in small specimens, acute, tubular spines, dorsal sur- face slightly elevated in midline, usually with one or two strong, tubular spines and frequently short longitudinal row of small spines or tubercles not extending beyond proximal half; mesial and ventral surfaces unarmed. Palm usually elongate, one-half to two-thirds length of carpus, moderately slender, particularly in large specimens, somewhat inflated dorsoventrally; dorsal surface convex, with four or five, widely spaced, irregular rows of tubular spines, usually very prominent in large specimens, not ex- tending beyond proximal half of fixed finger; dorso- mesial margin with row of moderately short spines, mesial face proximally produced into prominent shelf, marginally armed with strong, tubular spines, dis- tally with irregular row of small spines or tubercles; dorsolateral margin with row of small spines, in- creasing in size and becoming tubular on fixed finger, lateral face usually unarmed; ventral surface with scattered small tubercles or spines. Carpus very elongate, considerably exceeding length of merus; dorsomesial margin with row of strong, slender spines, increasing in size distally, dorsal surface with row of moderately strong spines slightly mesiad of midline extending from distal margin proximally two-thirds to three-fourths length of segment, rarely extending to proximal margin, and with usually short row of small spines laterad of midline, distal margin usually with one or two subacute spines; dorsolateral margin not delimited or with irregular row of small spines or spinulose protuberances, lateral face unarmed or slightly spinulose; ventro- lateral margin with short row of small to moderately strong spines distally, ventral surface unarmed or with few low, occasionally spinulose protuberances; mesial face with scattered, often spinulose pro- 1973 DECAPOD CRABS FROM CALIFORNIA So -—_——a,d,e Figure 11. Parapagurodes laurentae, n. sp., a,b, d—g, paratype, ¢ (SL = 3.2 mm), “Velero III” sta. 1429-41: a, sternite, Ps; b, coxae and sternite P;: d, telson: e, branchial lamella: f, antennular peduncle; g, antennal peduncle—c, h-m, @ (SL = 3.4 mm), “Velero III” sta. 895-38: c, coxae and sternite, P;—h-m, same, mouthparts (left internal face): h. mandible; i, maxillule; j, maxilla; k, mxpi; 1, mxp2; m, mxp;. Scales equal 0.5 mm (a.d.e) and 1 mm (b,c,f-m). 134 BULLETIN SOUTHERN CALIFORNIA tuberances, ventromesial margin with irregular row of moderately strong spines, increasing in size dis- tally. Merus subtriangular; dorsal surface with ir- regular rows of spinulose pro- tuberances: mesial and lateral faces usually unarmed, margins each with low, occasionally ventromesial and ventrolateral single or double row of spines, strongest mesially. Ischium with row of small spines on ventromesial margin, mesial face often with prominent protuber- ance dorsally. Coxa unarmed. Left cheliped moderately long, usually reaching to base of dactyl of right, slender; surfaces usually with tufts of long, stiff setae. Dactyl long, one and one-half to more than twice length of palm; cutting edge with row of small, corneous teeth; terminating in broad, corneous claw; often with slender, longi- tudinal hiatus; dorsal surface usually unarmed, oc- casionally with few, low, spinulose protuberances proximally. Palm moderately short, one-fourth to one-third length of carpus; dorsal surface convex, with irregular, single or double row of tubular spines, usually not extending onto fixed finger, dorsomesial margin with row of strong, tubular spines, becoming obsolete distally on fixed finger; dorsolateral margin not delimited or with irregular row of small spines or spinulose protuberances, lateral, mesial, and ventral surfaces unarmed. Carpus elongate, usually exceeding length of merus, sub- rectangular; dorsomesial and dorsolateral margins each with row of moderately strong or strong spines; mesial and lateral faces usually with few, scattered, low protuberances, ventromesial, and ventrolateral margins each with row of small spines, or less fre- quently with only few low protuberances. Merus subtriangular; dorsal surface with two irregular, longitudinal rows of low protuberances; mesial and lateral faces usually unarmed, or occasionally with few low protuberances, ventromesial and ventro- lateral margins each with single or double row of moderately strong spines. Ischium with row of small spines on ventromesial margin. Coxa unarmed. Second and third pereiopods long, usually equalling or overreaching tip of right cheliped; dorsal and ventral surfaces usually with tufts of moderately long, stiff setae. Dactyls elongate or moderately elongate, equalling or somewhat exceeding length of propodi; in lateral view, curved ventrally; in dorsal view, usually straight; terminating in strong corneous claws; dorsal margins each with row of widely spaced, low protuberances; mesial and lateral faces unarmed; ventral margins each with row of strong, corneous spines, increasing in size distally. Propodi elongate, one and one-half to twice length of carpi; dorsal surfaces each usually with row of low pro- tuberances; mesial and lateral faces unarmed; ven- tral margins each with one or two small, corneous spines or spinules distally. Carpi moderately short, two-thirds to three-fourths length of meri; dorsal surfaces each usually with small spine at distal mar- gin; lateral and mesial faces unarmed; ventral ACADEMY OF SCIENCES VOLUME 72 margins each frequently with small, occasionally spi- nulose protuberance. Meri laterally compressed; dor- sal surfaces each with row of low, occasionally spi- nulose protuberances; lateral faces unarmed. mesial faces each with small spine on ventral margin dis- tally; ventral margins usually with row of small spines (P.) or unarmed (P,). Ischia and coxae un- armed. Fourth pereiopods usually weakly subchelate, oc- casionally not subchelate. Dactyls with very small, preungual process on lateral face; propodal rasp with one to three rows of corneous scales. Fifth pereiopods chelate; coxae symmetrical. Sternite of third pereiopods subsemicircular; an- terior margin with long, stiff setae. Males with well developed, short sexual tube on right, occasionally with vas deferens of left slightly produced. No paired pleopods, unpaired pleopods, pl; to pl;, biramous, weakly developed, occasionally pleopods absent. Females with paired gonopores; no paired pleo- pods, unpaired pleopods, pl: to pli, with both rami moderately well-developed, or infrequently weakly developed; pl; usually rudimentary, occasionally ab- sent. Uropods asymmetrical. Telson with posterior lobes generally symmetrical; separated by very shallow median cleft; terminal margins concave or slightly oblique, each with row of very small spinules and one to four small spines at exterolateral angles, lateral margins with few short setae; anterior lobes unarmed, margins with long setae. COLORATION.—Living color unknown. In pre- servative: Body and appendages straw-colored. DISTRIBUTION.—Southern California, Channel Islands, to west coast of Baja California, Mexico; Gulf of California; 16 to 475 meters. AFFINITIES.—Although Parapagurodes laurentae is most closely related to P. makarovi, its superficial resemblance to an undescribed species of Pagurus that has come to our attention, may cause mistakes in identification. In addition to the presence of sexual tubes in the males, the lack of spines on the distal portions of the dorsal surfaces of the dactyls of the chelipeds, the moderately long and relatively slender dactyls of the second and third pereiopods, and the lack of regular setation on the articles of the antennal flagella are characters which distinguish P. laurentae. ETYMOLOGY .—Parapagurodes laurentae is named in honor of Michéle de Saint Laurent, the French carcinologist who has contributed so greatly to pagurid systematics. DISCUSSION.—As previously mentioned, both species of Parapagurodes were often found in- fested with Stegophryxus sp. The questionable inclusion of Pagurus [sp.] in the synonymy of Parapagurodes laurentae is based on the fact that this record of Menzies and Miller (1954) is 1973 the only other report of this bopyrid genus from western North America. these authors would not recognize their pagurid It is improbable that have been able to host had it been P, makarovi, as this species has been well illustrated under the name Parapagurus mertensii (Rath- bun, 1904, 1910; Schmitt, 1921; Makaroy, 1938). In contrast, Parapagurodes laurentae is not only an undescribed species, but it does bear a strong, superficial resemblance to some species of Pagurus. Key to the Species of Parapagurodes Palm of right cheliped with dorsal surface unarmed proximally, and with scattered small spinules or spinulose tubercles distally and on fixed finger. Palm of left cheliped with dorsal surface unarmed or with minute spinules P. makarovi, n. sp. Palm of right cheliped with dorsal surface armed proximally with four or five irregular rows of widely spaced, strong, tubular spines, not ex- tending onto fixed finger. Palm of left cheliped with dorsal surface with single or double row of strong tubular spines _ P. laurentae, n. sp. ACKNOWLEDGMENTS The authors are indebted to M. de Saint Laurent, Muséum National d’Histoire Naturelle, Paris, for her advice in the preparation of this manuscript; to R. Ingle, British Museum, London, for providing syntypic material of Pagurodes inarmatus, to Henry Roberts, National Museum of Natural History, Smithsonian Institution, for the specimens of Bene- dict’s Eupagurus mertensii, and to Fenner A. Chace, Jr., National Museum of Natural History, Smithson- ian Institution, and Anthony J. Provenzano, Jr., Uni- versity of Miami, for their reviews of the manuscript. One of the authors (McLaughlin) also wishes to ex- press her thanks to the Graduate School of Arts and Sciences, George Washington University, Washing- ton, D. C., for permission to publish data extracted from her doctoral dissertation. This research was supported, in part, by research grant GB-7075X from the National Science Foundation. LITERATURE CITED Alcock, A. 1905. Anomura. Fasc. I. Pagurides. Part Il in Catalogue of the Indian decapod Crus- tacea in the collections of the Indian Museum. Indian Museum, Calcutta, xi + 197 pp. Benedict, J. E. 1892. Preliminary descriptions of thirty-seven new species of hermit crabs of the genus Eupagurus in the U. S. National Mu- seum. Proc. U. S. Nat. Mus., 15:1—36. DECAPOD CRABS FROM CALIFORNIA Brandt, F. 1851. dorff, et al, und Osten Sibiriens wahrend der und 1844, Zoologic, 2(1):77—148. Krebse. Jn Dr. A. T. Reise in den ati Midde versten yOrcc?r Jahre 1443 Dana, J. D. 1851. in orbis terrarum circumnavigatione, print from] Proc, Acad, Nat. Sci (1)5:267-272. Conspectus crustaceorum quae [Pre Philadelphia Doflein, F. 1900. tischen Meere. Die Dekopoden Krebse der ark- In F. Romer and F. Schaudinn, Fauna Arctica, Gustav Fischer, Jena, 1:313 362. Gordan, J. 1956. A bibliography of pagurid crabs, exclusive of Alcock, 1905. Bull. Amer. Mus. Nat. Hist., 108(3):253-352. Hart, J. F. L. 1940. Reptant decapod Crustacea of the west coast of Vancouver and Queen Charlotte Islands. Canadian J. Res., 18(3):86— 105. Henderson, J. R. 1888. Report of the Anomura collected by H. M. S. “Challenger” during the years 1873-76. Vol. 27 in Scientific Results Explor. Voyage HMS Challenger, Zoology, xi + 221 pp. Holmes, S. J. 1900. Synopsis of the California stalk-eyed Crustacea. Occas. Pap. California Acad. Sci., 7:1—262. Lenz, H. 1901. Ergebnisse einer Reise nach dem Pacific (Schauinsland, 1896-1897). Crustaceen. Zool. Jb. Syst., 14(5):429-482. McLaughlin, P. A. genus Pagurus 1972. The hermit crabs of the from northwestern North America, with a partial revision of the genus. PhD dissertation, The George Washington Uni- versity, Washington, D. C., University Micro- films, Ann Arbor, Michigan, xiv + 554 pp. McLaughlin, P. A. (in press). The hermit crabs (Crustacea: Decapoda: Paguridea) of north- western North America. Zool. Verhandel., Leiden. McLaughlin, P. A., and A. J. Provenzano, Jr. (in press). Hermit crabs of the genus Paguri . from the tropical western Atlantic. Pt. I. . Bull. Mar. Sci. Makarov, V. V. [Crustacés Shtakel’berg 1938. Rakoobraznyey. Anomura. Décapodes anomures]. Jn A. A. (ed.), Fauna SSSR. Akad nauk. 136 BULLETIN SOUTHERN CALIFORNIA Zool. Instit. ns. No. 16, 10(3): {English translation: Crustacea, Anomura. Jeru- Israel Program for Scientific Transla- Published for the National Science Foun- Washington, x + 324 pp. salem: tions. dation and Smithsonian Institution, D. C., 1962, iv + 278 pp.] Markham, J. (in press). Parasitic bopyrid isopods of the amphi-American genus Stegophryxus Thompson, with the description of a new spe- cies from California. Menzies, R. J.. and M. A. Miller. 1954. Isopoda. In R. I. Smith, ef al., Intertidal invertebrates of the central California coast. S. F. Light’s labora- tory and field text in invertebrate zoology. Univ. California Press, xiv + 446 pp. Owen, R. 1839. voyage; ... Behring’s Straits ... The zoology of Captain Beechey’s during a voyage to the Pacific and H. G. Bohn, pp. 77-92. Rathbun, M. J. 1904. Decapod crustaceans of the northwest coast of North America. Jn Crus- taceans. Jn Harriman Alaska Series, 10:[1]—190. {Reprinted 1910, Washington: Smithsonian In- stitution.] Saint Laurent, M. de 1968. Révision des genres Catapaguroides et Cestopagurus et description de quatre genres nouveaux. I. Catapaguroides A. Milne-Edwards et Bouvier et Decaphyllus nov. gen. (Crustacés Décapodes Paguridae). Bull. Mus. Nat'l. Hist. Nat. Paris, (2)39(5.6):923-— 954 and 1100-1119. 1969. Ibid. THI. Acanthopagurus de Saint Laurent (Crustacés Décapodes Paguridae). Bull. Mus. Nat'l. Hist. Nat. Paris, (2)41(3):731- 742. 1970a. Ibid. IV. Solenopagurus de Saint Laurent (Crustacés Décapodes Paguridae). Bull. Mus. Nat'l. Hist. Nat. Paris, (2)41(6):1448— 1458. 1970b. Ibid. V. Trichopagurus de Saint Laurent (Crustacés Décapodes Paguridae). VI. Conclusions. Bull. Mus. Nat'l. Hist. Nat. Paris, (2)42(1) :210-222. ACADEMY OF SCIENCES VOLUME 72 1972. Sur la famille des Parapaguridae Smith, 1882. Description de Typhlopagurus foresti gen. noy., sp. nov., et de quinze espéces Ou sous-espéces nouvelles de Parapagurus Smith (Crustacea, Decapoda). Bijd. Dierk., 42(2):97— 123. Saint Laurent-Dechancé, M. de. 1966. Remarques sur la classification de la famille des Paguridae et sur la position systématique d’/rodopagurus de Saint Laurent . . . Bull. Mus. Nat'l. Hist. Nat. Paris, (2)38(3):257-265. Schmitt, W. L. 1921. The marine decapod Crus- tacea of California ... Univ. California Publ. Zool., 23:1—470. Smith, S. I. 1879. The stalk-eyed crustaceans of the Atlantic coast of North America north of Cape Cod. Trans. Connecticut Acad. Arts Sci., 5:27-136. Stimpson, W. 1857. On the Crustacea and Echi- nodermata of the Pacific shores of North America. Part I. Crustacea. Boston J. Nat. Hist., 6(4) :444—532. 1858. Crustacea. Pars VII in Prodromus descriptionis animalium evertebratorum, [Preprint (December 1858) from] Proc. each Nat. Sci. Philadelphia, (2)10:225-252. 1907. Report on the Crustacea (Brachy- ura and Anomura) collected by the North Pacific Exploring Expedition, 1853-1856. Smithson. Misc. Coll., 49(1717):240 pp. Taylor, G. W. 1912. Preliminary list of one hun- dred and twenty-nine species of British Columbia decapod crustaceans. Contrib. Canadian Biol., 1906-1910:187-214. Williams, H. C. 1915. Crustacea, Decapoda. Larven. Part VI in K. Brandt and C. Apstein (eds.), Nordisches Plankton, Zoologischer Teil, 3:315-588. Accepted for publication February 20, 1973. COMPARATIVE STUDY OF THE CALCANEA OF MEMBERS OF THE URSIDAE AND PROCYONIDAE Howarp J. STAINS! Asstract: Calcanea of all of the bears and all but one genus of procyonids are com pared and described. These calcanea indicate a close similarity of members of these two families. Some arrangement into subgroups can be made in each of the families. In the Ursidae, the calcanea of Ursus maritimus, Tremarctos, and Helarctos are similar; and those of Ursus americanus, Ursus arctos, Selenarctos, Melursus, and Ailuropoda are similar. In the Procyonidae, the caleanea of Procyon and Nasua, Potos and Bassaricyon, and Ailurus and Bassariscus are similar. The degree of development of the trochlear process and the proximal extension of the trochlear shelf is the major difference between members of the two families. tuberosity. The relationship of members of the families Ursidae and Procyonidae seem to be close, at least on the same line of evolution as the Canidae. The disputed position of the giant panda is one indication of the close relationship of these two families. Some of the early workers (Raven, 1936; Gregory, 1936) conclude that the giant and lesser pandas are closely related based on soft anatomy, skull and dentition, and place both in a separate subfamily of the Procyonidae. Walker (1964) places the giant panda in the Procyonidae. Davis (1964), based on soft anat- omy, and Leone and Wiens (1956), based on serology, place the giant panda in the family Ursidae. Simpson (1945) and Romer (1966) discuss the relationship of these two families to the Canidae. Stains (in Anderson and Jones, 1967) summarizes the morphological charac- teristics and taxonomy of these families. Like studies of the teeth, skulls, myology, and blood, study of the calcanea of living pro- cyonids and ursids of the world indicate these two families are closely related. The object of this paper is to point out the intra-family simi- larities and differences in the structure of cal- canea of living members of these two families. There is no intent to imply evolutionary re- lationships simply because of the similarities or differences noted. A discussion of the features of the calcanea and a comparison of these fea- tures by genus within each family follows. The terminology follows that of Stains and Robinette (1970) and Stains (1959 and 1962) and is illustrated in figure 1. Measurements were taken in the manner illustrated in figure 2 and all are expressed in millimeters. Differences are also evident in the various articular surfaces and the greater All genera and species of the family Ursidae were available for study. Because of the similarity to the calcanea of Melursus, Ailuropoda me- lanoleuca is included with the bears rather than the procyonids. Of the Procyonidae, Nasuella was the only genus not available for study. Aside from island species of Procyon and Nasua., other questionable species not available were Bassaricyon alleni and Bassariscus sumichrasti. CALCANEA OF MEMBERS OF THE FAMILY URSIDAE Ursus maritimus—Polar Bear In general, the calcanea of Ursus maritimus are larger than those of other bears. There is some overlap, especially in length of calcaneum, with Ursus arctos (Table 1). Only five of 28 speci- mens of U. arctos have calcanea with widths as great or greater than the bones of two immature specimens of the polar bear, and only one speci- men of the brown bear has a heel bone as wide as one of five subadult and adult specimens of the polar bear. Proportionately, the width/ length index of the calcaneum in the polar bear (Table 2, W/TL) is slightly larger than in the brown bear. These two bears can be distinguished most easily by the proximo-distal length of the troch- lear process compared with the width of the calcaneum (Table 2, LTP/W). In the polar o 7 c f ' *Dept. Zoology, Southern Illinois Univ bondale, Illinois 62901. 137 138 GROOVE FOR TENDON OF ACHILLES CALCANEAL TUBEROSITY = w > 3 Fa = BODY > bp a < POSTERIOR m = ARTICULAR SURFACE PROXIMAL END x L 4 ul m = » = > a in 4 = OORSAL TABLE MEDIAL ARTICULAR a SURFACE w = ~] SUSTENTACULUM m TROCHLEAR PROCESS ANTERIOR ARTICULAR SURFACE OISTAL ENDO CUBOID FACET Figure I. Dorsal view of generalized caleaneum of a carnivore. bear this process is a smaller percentage of the width than in the brown bear. The bear closest to the polar bear, in the form of the trochlear process, is Tremarctos (Fig. 3e) where the trochlear process extends outward as a continuation of the posterior ar- ticular surface. In Ailuropoda (Fig. 3j) the trochlear process is short as in the polar bear but is not a continuation of the posterior articular surface and has the development of a notch between the posterior articular surface BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 DORSO-VENTRAL HEIGHT SUSTENTACULAR THICKNESS TOTAL LENGTH LENGTH OF TROCHLEAR | PROCESS Se OT AN Ge LENGTH OF POSTERIOR ARTICULAR SURFACE LEAST WIDTH OF canal Figure 2. Measurements taken on the calcaneum. and the trochlear process (Fig. 31). In other bears this process tends to go below to form a distinct shelf under the posterior articular surface. Tremarctos also is closest to the polar bear in an average width to length index (Table 2, W/TL). Selenarctos is next but has a distinct TABLE 1. Measurements of the calcanea of members of the Ursidae. Total length (mm) Width (mm) LPAS (mm) No. Range Avg. Range Avg. Range Avg. 1 5 100.6-111.3 103.5 69.7-77.4 74.2 30.5—38.8 35.1 2; 20 66.2—111.2 83.7 44.0-71.9 54.1 24.2-41.4 30.6 3 17 60.4— 75.6 66.5 33.5-46.1 41.1 17.5—25.1 22.0 4 14 58.1-— 71.5 64.1 37.8-44.6 40.9 20.6—27.9 24.0 5 10 44.3— 67.9 60.4 30.2-46.3 40.2 16.8—25.6 21.5 6 5 55.1— 70.8 63.4 33.2-43.6 39.0 17.8—24.0 DDS: iT 4 45.6— 52.8 49.8 30.6-35.1 33.6 18.1—20.6 19.7 8 1 50.1 32.0 17.8 ST (mm) D-VH LWB No. Range Avg. Range Avg. Range Avg. 1 5 17.5—22.2 19.6 50.4—57.6 53.9 21.1-25.3 232) 2 20 12.3-24.1 16.3 34.0-57.6 43.4 12.6—23.1 17.4 3 17 7.2-12.9 11.7 30.5—37.4 34.5 10.6-14.9 13.4 4 14 8.4-12.2 10.2 27.0-34.0 29.6 10.6—12.7 11.6 5 10 7.0=11.3 ON, 20.9-35.9 28.7 8.5-13.2 10.6 6 5 6.8-11.8 9.7 24.5-35.8 30.7 10.8—-13.7 11.9 7 4 8.7-— 9.8 9.3 22.0-26.4 24.3 7.3-— 8.4 Tell 8 1 6.5 21.1 8.0 *1, Ursus maritimus, 2, U. arctos, 3, U. americanus, 4, Ailuropoda melanoleuca, 5, Selenarctos thibetanus, 6, Melursus ursinus, 7, Tremarctos ornatus, 8, Helarctos malayanus, LPAS length of posterior articular surface, ST sustentacular thickness, D-VH dorso-ventral height, LWB least width of body (see Fig. 2 for measurement procedures). 1973 Figure 3. CALCANEA OF URSIDS AND PROCYONIDS 149 EEX wis Right calcanea of members of the Ursidae. a, Ursus maritimus; b, U. americanus: c, Melursus ursinus; d, Helarctos malayanus; e, Tremarctos ornatus; £, Selenarctos thibetanus; g, U. arctos; h, showing distance from outer edge of trochlear shelf to center of inner edge of posterior articular surface in Ursus americanus; i, notch between posterior articular surface and trochlear process in Ailuwropoda melanoleuca; j, Ailuropoda melanoleuca. shelf formed by the trochlear process 3f). Specimens examined: AMNH_ 14058 (yg). 15686-7 (yg), 22885, 75244-5, 100039 (AMNH 22885 illustrated, Fig. 3a). (Fig. Ursus arctos—Brown Bear The caleaneum of Ursus arctos is extremely variable in size within and between different age groups. In 20 adults the range is from 66.2— 111.2 mm (avg. 83.7) in length, and two young (greater tuberosity not completely ossified) range from 67.9-75.5 mm. In width of calcanea, adults range from 44.0-71.9 (avg. 54.1 mm), and six young range from 43.6—53.1 (avg. 48.4 mm). The larger specimens come from Alaska, the brown bears of western United States and are less than 85 mm smaller calcanea come from in length and less than 55 mm in width. The brown bear (Fig. 3g) can be distinguished from the polar bear (Fig. greater proximal expansion of the trochlear pro- 3a) because of the cess in the brown bear. In Ursus maritimus, the proximo-distal length of the trochlear process is from 40.8-48.1 (avg. 44.7) percent of the width and in the brown bear, 51.2—69.8 (avg. percent. This ratio will separate both and adult polar bears from brown bears. 59.6) young 140 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES TABLE 2. VOLUME 72 Indices for caleanea of members of the Ursidae. W/TL" LTP/W* D-VH/TL* LWB/D-VH* Species Range Avg. Range Avg. Range Avg Range Avg. Ursus maritimus 68-75 72 41-48 45 49-56 52 39-45 43 Ursus arctos 60-73 65 51-70 60 48-58 53 33-50 40 Ursus americanus 55-67 62 55-77 68 49-56 52 31-44 39 Ailuropoda melanoleuca 59-70 64 35-44 40 44-51 46 36-43 39 Selenarctos thibetanus 61-70 67 45-69 58 44-54 47 33-41 37 Melursus ursinus 60-62 62 59-69 64 45-51 49 36-44 39 Tremarctos ornatus 66-70 68 28-32 31 47-52 49 28-35 32 Helarctos malayanus = 64 — 51 — 42 —— 38 *W/TL, width divided by total length; LTP/W, length of trochlear process divided by width; D-VH/TL, dorso-ventral height divided by total length; LWB/D-VH, least width of body divided by dorso-ventral height (See Fig. 2 for measurement procedures). The calcaneum of the brown bear can be dis- tinguished from the black bear (Ursus ameri- canus, Fig. 3b) by the anterior articular surface being continuous with and broadly connected with the medial articular surface. In only four of 26 specimens of the brown bear are the two surfaces separate; in no case is the anterior articular surface absent. In the calcanea of 20 black bears, 10 lack the anterior articular surface and 10 have a small anterior articular surface, usually separate from the medial ar- ticular surface, although the two surfaces may be joined by a narrow band. There seems to be no relationship between age of the bear and the amount of juncture of the two surfaces in the grizzly although in all the young the two sur- faces are joined. Regardless of age, the best calcaneal character for the separation of the brown and black bears is in the size of the posterior articular surface. In the bones of the brown bear, the length of this surface is greater, and in the black bear is shorter (Table 1, LPAS). Using this charac- teristic, only two adults, of 26 young and adult brown bears, fall wtihin the upper limits of the range of the black bear, and only one of 17 adult black bears falls within the lower limits of the range of the brown bear. Except for size, the calcaneum of Selenarctos (Fig. 3f) seems to be the most difficult to dis- tinguish from Ursus arctos. The indices of width to total length are the closest of all the bears. If the calcanea of the larger Selenarctos are compared with those of small brown bears, the length of the posterior articular surface, total length, width of the bones, and the proportion of the length of the trochlear process to the width of the calcanea are closest in these two species. The thickness of the sustentaculum does sepa- rate the calcanea of Selenarctos from U. arctos (Table 1, ST). Ursus arctos also can be dis- tinguished from Melursus and Ailuropoda on the basis of the thickness of the sustentaculum. In addition, the length of the posterior articular surface in Melursus is less (Table 1, LPAS). The posterior articular surface also tends to be less in Ailuropoda. The calcanea of both Melursus (Fig. 3c) and Ailuropoda (Fig. 3j) have a dis- tinct notch at the outer distal corner of the posterior articular surface (Fig. 3i), a charac- teristic which separates the bones of these genera from those of all other bears. The calcaneum of the smallest specimen of Ursus arctos is greater in length and width than any specimens examined of Tremarctos or Helarc- tos. In addition, the heel bones of Tremarctos (Fig. 3e) and Helarctos (Fig. 3d) have a dis- tinct rolling of the outer edge of the posterior articular surface which U. arctos lacks. Specimens examined: AMNH 3767 (yg), 14054, 35590, 35887, 45150, 62553-4, 70163, 70330, 70448, 73694, 73696-7 (both yg), 125407, 129379 (yg), 135502, 149805, USNM ‘ 1973 9483 (yg), 12322, 21337 (yg), 123386, 174950, 199252, 252303, 283624 (yg), 292072, 301690 (USNM 123386 illustrated, Fig. 3g). Ursus Black americanus Bear The calcaneum of the black bear is much smaller than that of the polar bear and no overlap in measurements exists. Differences in the calcanea of the black and brown bears have cussed under Ursus arctos. As is indicated in table 1, one difference be- tween U. americanus and Selenarctos is in the length of the calcaneum. As will be discussed later under Se/enarctos, there seem to be two distinct groups of calcanea; the adults with the smaller calcanea of these two groups fall below the size of the smallest adult black bears, but the group with larger calcanea (length 66.1— 67.9 mm; width 40.6-46.3 mm) falls within the size of the black bears (Table 1). One measurement that separates adult U. americanus from all Selenarctos is the distance of the outer edge of the trochlear shelf to the center of the inner edge of the posterior articular surface (see Fig. 3h). In bones of the adult black bear, this distance ranges from 11.1—14.2 (avg. 12.7 mm) and in adult Selenarctos from 5.9-10.8 (avg. 8.4 mm) in five specimens. In five of 10 adult specimens of Selenarctos, the shelf of the trochlear process does not extend proximal enough to obtain a measurement. This is true of all three of the specimens of young animals. Seven young and very young black bears tend to have less development of the troch- lear shelf producing measurements within the range of the adult Selenarctos (range 5.7—13.9, avg. 11.9 mm). Ursus americanus seems to be most like Melursus in various calcaneal characteristics and indices (see Table 2). One character separates all Melursus (five adult specimens) from all U. americanus (24 adult and young specimens). The calcanea of Melursus (Fig. 3c) have a dis- tinct notch (Fig. 3i) on the outer distal corner of the posterior articular surface as does A ilurop- oda. In Ursus (Fig. 3b), the posterior articular surface blends into the trochlear process on the distal part of the calcaneum. The calcanea of Tremarctos (Fig. 3e) differ from those of U. americanus in the absence of a trochlear shelf, and Helarctos (Fig. 3d) differs in the extensive rolling of the outer edge of the posterior articular surface. In addition, both been dis- CALCANEA OF URSIDS AND PROCYONIDS 14] Tremarctos and Helarctos are smaller than adult black bears. The expanded medial and posterior articular surfaces, the shorter trochlear proces and the surfaces black notched posterior articular separate Ailuropoda (Fig. 3j) from the bear. Specimens examined: AMNH 24157, 35046, 45149, 90321 (yg), 90333 (yz), 90334, 98950, 100125, 120843, 128521, 131829, 131830 (yg), 149894 (yg), 164283, 164284 (yg); USNM 49664 (yg), 220264 (yg), 240613, 255072, 260642, 269812, 283630, 302901, 303193 (USNM 255072 illustrated, Fig. 3b). Ailuropoda melanoleuca—Giant Panda The giant panda possesses a calcaneum similar to those of bears in most respects. The trochlear process is a prominent projection on the calcanea of Ailuropoda (Fig. 3j}) and ursids but is knob- like and small in most procyonids. However. it does not form a shelf below the posterior articular surface in Ailuropoda as it does in most bears. Tremarctos (Fig. 3e) is similar to Ailuropoda in this respect. The trochlear process is not grooved either in the giant panda or in the ursids. It is grooved in all procyonids except for some individuals of Procyon lotor (absent in eight of 17 specimens, faintly grooved in the remaining nine). The proximo-distal length of the trochlear process is about one-fourth the total length of the calcaneum in Ailuropoda Tremarctos; it is longer in other bears. The calcaneal width-total length index (Table 2) of Ailuropoda lies within the range exhibited by the ursids, and is greater than that found in most procyonids. This index reflects the rela- tive larger size of the trochlear process of the giant panda and the ursids. The most distinct differences observed between and the calcanea of ursids and Ailuropoda involve the posterior and medial articular surfaces. These two surfaces are raised higher above the bulk of the calcaneum in the giant panda than in the bears. The posterior articular surface in the giant panda forms a well-developed ledge because of its position high above the body of the cal- caneum; this ledge is absent in the calcanea of ursids. The outer distal corner of the posterior articular surface forms a notch or groove in the calcaneum of Ailuropoda (Fig. 3j) and is sit to that in the calcaneum of Melursus (Fig. 3c). The posterior articular surface is longer com- 142 BULLETIN SOUTHERN CALIFORNIA pared to width in Ailuropoda than in the bears (flatter). This sigmoid in the caleanea of most procyonids except and is less curved surface is Potos and Nasua nasua. Unlike other members of the Procyonidae and Ursidae, the medial articular surface in Ailurop- oda is expanded to the extent that the bulk of the hidden “when viewed from the dorsal aspect. In addition, a prominent sustentaculum is notch is formed between the medial and anterior articular surfaces. This notch is present in Melursus (Fig. 3c), and Tremarctos (Fig. 3e) but tends to be obscured in these bears by the ventral expansion of the sustentaculum. Such a notch is not present in procyonids. The anterior articular surface of the calcanea in Ailuropoda varies from absent (six specimens), to a minute triangular articulation entirely sepa- rated from the medial articular surface (10 specimens), to a condition where the two sur- faces tend to be continuous (two specimens). This variation in the anterior articular surface is similar to the variation found in Ursus ameri- canus (Fig. 3b). In all procyonids the anterior articular surface, when present, is usually minute and is never continuous with the medial articular surface, although it is joined by a small waist in eight of 18 specimens of the genus Procyon. In general, the sustentaculum of Ailuropoda and the ursids is lower in position (more toward the cuboid surface) than in procyonids. The giant panda and the ursids lack the notch on the ventral side of the cuboid surface which is characteristic of procyonids. Of the species of bears, the calcanea of Ailurop- oda resemble most closely those of Melursus in shape of the greater tuberosity, the notch at the distal corner of the posterior articular surface, the notch between the medial and anterior ar- ticular surfaces, and many of the indices. Davis (1964) with only the genus Ursus and does not go into compares the calcanea of Ailuropoda great detail. Specimens examined: USNM 258423, 258425, 258834 (yg), 258835—6, 258838, 258644, 258984 ), 259027-9, 259075, 259078 (yg), 259400 ), 259401-3 (USNM 258644 _ illustrated, 5 SDE ag q (y (y gg Fi Selenarctos thibetanus—Asiatic Black Bear There seems to be at least two, and perhaps three, distinct sizes of calcanea within Selenarctos ACADEMY OF SCIENCES VOLUME 72 with no relationship to sex or age of the animals. The largest group (six specimens) has calcanea which range in length from 66.1 to 67.9 mm with the young of this group lacking the cap of the calcaneal tuberosity and still being 61.6 mm long. In addition, two of the adults measuring 66.6 to 67.0 mm still have traces of the symphysis of the calcaneal tuberosity, an indication that they probably are subadults. Seven adult specimens of the smaller group range in length from 44.3— 59.6 mm with two of the adults being extremely small (44.3 and 50.4 mm). The two young of this group possess caps and measure 48.6 and 52.4 mm. Perhaps this smaller group is composed of two subgroups. Both the larger and smaller groups are repre- sented by both sexes and both groups have a preponderance of specimens obtained from zoos, although one from the group with large cal- canea came from Japan and one from northern Burma; one of the small group also came from the Himalaya Mountains. If these localities are accurate, there simply may be a _ tremendous amount of variation similar to that found in Ursus arctos. Such distinction in size usually is an indication of species differences in the calcanea of most mammals. Thus there seem to be two, and perhaps three, distinct species present. The effect of living as caged animals may be the explanation of such variation, although in most caged animals the effect usually is rugoseness of the bone as is obvious in USNM 221064, a zoo specimen at least 15 years old. The differences between the calcanea of Selenarctos and Ursus maritimus, U. arctos, and U. americanus have been discussed. Selenarctos (Fig. 3f) differs from Melursus (Fig. 3c) and Ailuropoda (Fig. 3j) in lacking the notch at the outer distal corner of the posterior articular sur- face (Fig. 31). The calcaneum of Selenarctos tends to be wider than long when compared to Melursus (see the width-length indices in Table 2). The trochlear process in Selenarctos is not as well-developed as in Melursus. differs Ailuropoda in having a much longer trochlear process (Table 2) compared to the width of the (Fig. 3e) and Ailuropoda 3j) lacking the development of a trochlear Selenarctos from Tremarctos and bone; Tremarctos (Fig. shelf. Specimens examined: AMNH 23086, 35016, 35496, 70094 (yg), 70320, 80248, 90184 (yg), 100396 (2 animals same number), 114544 (yg), 1973 119476, 171285; USNM 221064 (AMNH 70320 illustrated, Fig. 3f). Melursus ursinus—Sloth Bear The heel bone of Melursus (Fig. 3c) can be distinguished from all bears except Ailuropoda (Fig. 3j)) in having a distinct notch located at the outer distal corner of the posterior articular sur- face (Fig. 31). The length of the trochlear pro- cess divided by the width of the calcaneum separates all specimens of Melursus from Ailurop- oda (Table 2). In general shape and expansion of the trochlear process, the calcaneum_ of Melursus is similar to Ursus americanus (Fig. 3b). Specimens examined: AMNH 22720, 22896 (malformed), 35602, 35898, 54464—-5 (AMNH 54464 illustrated, Fig. 3c). Tremarctos ornatus—Spectacled Bear The similarities of Tremarctos to Ursus mari- timus have been discussed. All members of the genera Ursus, Melursus, and Ailuropoda have longer calcanea than the largest 7Tremarctos ex- amined. Tremarctos can be separated from all of the bears on the basis of the ratio of the length of the trochlear process to the width of the calcaneum (Table 2, LTP/W). This index is less than that of other bears, Ailuropoda being closest. In overall size, the calcaneum of Helarc- tos is closest to Tremarctos. Specimens examined: AMNH 35480, 70164, 100010, 194309 (AMNH 70164 illustrated, Fig. 3e). Helarctos malayanus—Malayan Sun Bear Only one specimen of the sun bear was avail- able for examination. The caleaneum differs from the calcanea of all other bears in having the posterior articular surface prominently rolled over on the outer edge (Fig. 3d). There is a slight trochlear shelf present. The cuboid surface is at a great angle with the long axis of the bone. A flat groove leads from the proximal end of the posterior articular surface toward the greater tuberosity; this groove was not noted in other bears. A small anterior articular surface is pres- ent. Specimens examined and _ illustrated: AMNH 70089, Fig. 3d. CALCANEA OF URSIDS AND PROCYONIDS l4 CALCANEA OF MEMBERS OF FAMILY PROCYONIDAE THI Procyon lotor—Raccoon The calcanea of members of the genus Procyon lack a distinct knob on the trochlear process (Fig. 4a and b). The trochlear process tends to be broad, originating close to the cuboid surface and gently sloping from the point knob is commonly present on other procyonids, where the to a point below and beyond the posterior ar- ticular surface. In this feature, the calcanea of Nasua narica (Fig. 4e) and Bassaricyon gabhi (Fig. 4d) most nearly resemble Procyon. The heel bones of the two species of Procyon seem to differ in size (P. being larger) and in length of the posterior articular cancrivorus In addi- tion, the trochlear process of P. lotor has a less distinct groove, the dorsal edge of the cuboid facet in P. lotor has a distinct point, and there is an addition of another surface alongside the anterior articular surface (between this surface and the cuboid facet) in P. cancrivorus (Fig. 4b). Procyon differs from Nasua narica (Fig. 4e) in that the medial artcular surface is not folded over the proximal edge of the sustentaculum. The distal edge of the trochlear process is almost even with the cuboid surface in Procyon. In both species of Nasua (Fig. 4e and f), the distal edge of the trochlear process originates more proximally than in Procyon. The calcanea of Procyon are much longer than those of Bassaricyon (Table 3. TL). In addition, Bassaricyon is similar to Nasua in the position of the distal edge of the trochlear pro- cess. The tuber, or body, leading to the calcaneal tuberosity is more robust in Procyon than Potos. the cuboid facet is not turned dorsally in Procyon as in Potos (Fig. 4c), the medial articular surface is round in Procyon but is tear-shaped and wider than long in Potos, and the posterior articular surface is strongly sigmoid in Procyon but in surface (P. cancrivorus being longer). Potos tends to form a smooth curve. The distinct knob-like Bassariscus (Fig. 4h) and Ailurus (Fig. +g) dis- trochlear process of tinguishes the calcanea of these genera from Procyon (Fig. 4a and b). Specimens examined: USNM 575, 1386. (yg), 144069, 187907, 22243. young), 256031, 256057 (yg). 21136 256028-30 (all 259130-1 (both 144 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 Figure 4. young), 259132, 259137, 271097 (USNM 271097 illustrated, Fig. 4a). Procyon cancrivorus—Crab-eating Raccoon Comparison and characteristics of Procyon can- crivorus have been made under Procyon lotor. The calcaneum of the crab-eating raccoon tends to be larger than that of the common raccoon but otherwise is not too different. Specimens examined: USNM 6949, 49718 (USNM 49718 illustrated, Fig. 4b). Potos flavus—Kinkajou In general, the posterior articular surface of the calceaneum of Potos is long, narrow, and prac- tically in line or parallel with the body leading to the calcaneal tuberosity (Fig. 4c). In all other Right calcanea of members of the Procyonidae. crivorus; c, Potos flavus; d, Bassaricyon gabbi; e. Nasua narica; f, N. nasua; g, Ailurus fulgens; h, Bassariscus astutus. a, Procyon lotor; b, P. can- procyonids, this surface tends to be broader and at more of an angle to the body of the bone. The trochlear process is located proximally in Potos near the distal edge of the posterior artic- ular surface. In some specimens this process is poorly developed and virtually absent. Differences between the bones of Procyon and Potos have been discussed. The extreme dorsal turning of the cuboid surface and the formation of a tear-shaped medial articular sur- face which is wider than long, separate Potos from the remaining procyonids. The procyonid most closely resembling Potos in general features of the calcaneum is Bas- saricyon (Fig. 4d). The difference listed above, as well as overall size (Table 3), will distinguish the two easily. Specimens examined: AMNH 14073, 35192, 35352, 35434-5, 35766, 35916 (yg), 42302, 1973 TABLE 3. Measurements of Total length (mm) Width (mm) me No. Range Avg. Range Avg. 1 2 32.2—36.1 34.2 15.0-17.7 16.4 2 9 24.7-31.3 27.3 12.3-15.7 11.8 ee i) 21.0-27.5 24.0 9.7-13.3 12.1 4 10 23.7-29.7 27.3 12.1-16.1 14.2 5 4 26.0-27.7 26.8 13.3-18.1 14.8 6 12 24.3-31.1 27.1 12.3-16.8 14.8 Wf 2) 17.9-18.4 18.2 91-92 9.2 8 10 14.0-17.1 15.8 7.2—- 9:8 8.7 "1, Procyon cancrivorus, 9 8, Bassariscus astutus; see T ‘able 1 and Fig. 2 for 48216, 80223 (yg); USNM 49943, 241106, 241181, 255123, 257352, 258316; KU 68059 (yg), 69058 (USNM 258316 illustrated, Fig. 4c). Bassaricyon gabbi—Olingo On the basis of the trochlear process, Bassaricyon gabbi (Fig. 4d) is in the intermediate position between a broad, gently sloping condition and a distinct knob-like process. Other than Bassaris- cus, Bassaricyon possesses the smallest calcaneum. There is no overlap in size ranges with the larger species (Table 3). There is a slight ten- dency for the posterior articular surface to be sigmoid. The proximal edge of the medial artic- ular surface is folded over the sustentaculum (true of the calcanea of all procyonids except Procyon and Potos). The anterior articular surface is poorly devel- oped, as in Petos, and could be listed as absent. In most characteristics, the calcaneum of Bas- saricyon (Fig. 4d) resembles Potos (Fig. 4c). Specimens examined: USNM 305748-9 (US- NM 305748 illustrated, Fig. 4d). Nasua nasua—Red Coati The caleaneum of Nasua is intermediate in pro- cyonid characteristics. The two species examined (N. nasua, Fig. 4f, from South America and N. narica, Fig. 4e, from Central and North America) differ in the ratio of the length of the posterior articular surface to the total length (Table 4, LPAS/TL). The posterior articular surface tends to be more sigmoid in N. narica, and more a smoothly curved surface N. nasua. The groove on the edge of the trochlear process tends to be more distinct in N. narica. in Nasua narica has a more conspicuous groove on CALCANEA OF URSIDS AND PROCYONIDS the caleanea Procyon lotor, 3, Potos flavus, 4, measurements taken, l4 of members of the Procyonidae. LPAS (mm) D-VH LWB Range Avg. Range Avg. Range Avg 13.5-15.0 14.8 13.3-15.8 14.6 5.8~-6.6 6.2 8.4-11.4 10.0 9.8-13.5 11.1 3.5~5.3 4.5 8.0- 9.2 8.5 7.6-10.1 8.5. 2.6-3.6 3.1 8.8-11.3 10.2 8.9-10.8 10.1 3.3~—5.8 4.8 8.0— 9.0 8.5 10.2-10.6 10.4 3.9-4.2 4.0 9.4-11.9 10.8 9.5-12.8 11.4 3.0-4.5 3.7 6.1— 6.3 6.2 6.8— 7.1 7.0 2.5-2.6 2.6 5.6— 6.4 5.9 5.8— 6.6 6.2 2.1—2.8 2.5 Nasua narica, 5, NAR 6, Ailurus fulgens, 7, Bassaricyon gabbi the outer surface of the body, below and proximal to the posterior articular surface. The trochlear process tends to be more knob-like in Nasua_ nasua than in N. Most of the above characteristics are matter of degree, but there was no difficulty in separating the specimens available. Perhaps specimens from northern South America would be intermediate between the two groups (species) examined, indicating a single species. Since specimens from the intermediate region are not available, the differences noted are of enough magnitude to regard these individuals different species. More specific comparisons of the calcanea of Nasua have been discussed under the genera Procyon and Potos. Ailurus (Fig. 4g) and Bas- sariscus (Fig. 4h) tend to have smaller and more distinct knob-like trochlear processes and a tuberosity which tends to flare out and be- come more distinct from the body. The heel bones of Nasua diiffer from Bassaricyon (Fig. 4d) in being of larger size, having a longer trochlear process in relation to total length, and in the presence of a minute additional articular surface next to the narica. a as members of two anterior articular surface (this additional surface also is present on the calcaneum of Procyon cancrivorus). Specimens examined: AMNH 347, 42699. 100448 (yg): USNM 49644, 238671 (yg). 270366 (AMNH 347 illustrated, Fig. 4f). Nasua_ narica—White-nosed Coati Comparisons of Nasua narica with other species of procyonids and with Nasua nasua were made previously under a discussion of red coati. Specimens examined: AMNH 5904, 35656, 70078; USNM 983, 21982 (yg). 146 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 Taste 4. Indices for caleanea of members of the Procyonidae. W/TL Index* LTP/W Index* | D-VH/TL* LWB/D-VH* — LPAS/TL* | Species Range Avg Range Avg. Range Avg. Range Avg. Range Avg. Procyon cancrivorus 46-49 48 77-86 82 41-44 43 42-44 43 — 42 Procyon lotor 44—56 50 67-92 76 39-44 42 36-46 41 34-39 37 Potos flavus 47-56 Sl 46-66 58 32-38 35 32-46 36 33-39 38 Nasua narica 48—S4 52 54-71 65 35-40 37 37-54 47 34-41 38 Nasua nasua 48—54 5 61-77 67 37-40 39 37-40 39 30-34 32 Ailurus fulgens 50-59 54 33-49 42 39-46 42 27-36 32 35-44 40 Bassaricyon gabbi 50-51 51 52-53 53 39 39 35-38 37 oe 34 Bassariscus astutus 51-58 55 47-S7 52 37-41 39 36-45 41 35-40 38 * See “Table 2 and Fig. 2 for measurements taken. 257314, 271136, KU 63118, 68056 (USNM DISCUSSION 257314 illustrated, Fig. 4e). Ailurus fulgens—Lesser Panda Ailurus fulgens (Fig. 4g) and Bassariscus astutus (Fig. 4h) have the most knob-like trochlear processes of all the procyonids. In addition, this process begins higher above the cuboid surface. The distal edge of the trochlear process is even with the distal edge of the medial articular surface. In all other procyonids the distal edge of the trochlear process is distal (closer to the cuboid surface) to the medial articular surface. The calcaneum of Ailurus is much larger than that of Bassariscus and tends to have a greater flaring of the tuberosity. The calcaneal relationships of Ai/urus to other procyonids have been discussed under previous genera. Specimens examined: AMNH 119474, 119675, 146778; USNM = 252087, 252091, 252146, 256098, 258051, 258350, 258850, 305770-1 (USNM 252146 illustrated, Fig. 4g). Bassariscus astutus—Ring-tailed Cat is the In almost all features, the calcaneum of Bassariscus is a minia- ture of Ailurus fulgens (Fig. 4g). Specimens examined: AMNH 35909, 100094, 135964-6, 137029-30; USNM 1619, 49934; KU 58649 (AMNH 35909 illustrated, Fig. 4h). The calcaneum of Bassariscus astutus smallest of all the procyonids. The calcanea of bears are quite similar (Fig. 3, Tables 1 and 2). However, there are differences which make it possible to identify each species. Bears have a calcaneum with an obvious trochlear process. This process is expanded into a promi- nent trochlear shelf which in most cases (except Tremarctos and Ailuropoda) extends to below the posterior articular surface. The posterior articular surface is smooth, never sigmoid. The medial articular surface tends to be rolled over the proximal edge of the sustentaculum. The amount of rolling is extremely variable within all species with the angle of curl being from slight to 90°. There is a tendency for the medial and anterior articular surfaces to be joined; this is especially true of the bones of larger bears (U. arctos, U. maritimus, and Melursus). In the others, the two surfaces may or may not be joined, but seldom broadly. The sustentaculum varies in thickness in the different bears. Some question the generic separation of Tremarctos from Ursus and place Selenarctos midway be- tween Tremarctos and Ursus (Simpson, 1945). The placement of Thalarctos (the polar bear) and Euarctos (the black bear) in the genus Ursus is generally accepted today (see Stains, in Anderson and Jones, 1967). The generic separation of Melursus and Ursus is adopted by all authors (Simpson, 1945). Studies of the calcanea indicate a close similarity between Ursus maritimus and Tremarctos, between Ursus americanus and Melursus, and between Ursus arctos and Selenarctos. 1973 The calcanea of all procyonids are basically similar in appearance to those of bears; both groups having calcanea with an obvious. troch- lear process. There is less massive development of the trochlear process in procyonids thus a smaller ratio of width of the calcaneum to length (Tables 4 and 2, W/TL). There is little extension of the shelf of the trochlear process up to or beyond the distal edge of the posterior articular surface in the procyonids as in the bears. This process forms a distinct knob in some procyonids. Size and location of the trochlear process are the most obvious differences between various members of the Procyonidae (Fig. 4, Table 3). In most procyonids, except Potos and Nasua nasua, the posterior articular surface is sigmoid in shape. In Bassaricyon it tends to be slightly sigmoid. In no case is there a tendency for the surface to be rolled as in some ursids. The cuboid surface tends to be half-moon in shape caused by a notch on the ventro-lateral edge of the cuboid surface and tends to slope dorso- ventrally, especially in Potos (Fig. 4c). The anterior surface may be present, minute, or absent but seldom joined to the medial articular surface in the procyonids. CONCLUSIONS On the basis of the characteristics of the cal- caneum, bears can, in general, be grouped ac- cording to similarities and differences. Ursus maritimus and Tremarctos are most similar in general shapes and proportions of the heel bone. Ursus americanus and Melursus likewise are similar to each other. Ursus arctos, Selenarctos, and Ailuropoda seem closer to the U. americanus- Melursus group and Helarctos seems closer to the Ursus maritimus-Tremarctos group. The trochlear process in procyonids is not expanded to the extent found in bears. In most procyonids, the process is expressed in the form of a knob-like structure which tends to be grooved. Based on the shape of the trochlear process as well as other features of the calcaneum, the procyonids can be arranged in three subgroups: (lotor and (nasua and narica); 2. Potos—Bassaricyon; and 1. Procyon cancrivorus )—Nasua 3. Ailurus—Bassariscus. Both ursids and procyonids have an obvious trochlear process and, in most characteristics. CALCANEA OF URSIDS AND PROCYONIDS 14 Although the 17 calcanea of have similar calcanea. ference is great between member of the two families, the ratios, length of troch lear process divided by width (LTP/W), dorso j total (D-VH TL), and least width of body divided by dorso ventral height (LWB/D-VH), are similar (Tables 2 and 4). The length in ursids does produce a higher W/TI ventral height divided by length larger width in relation to ratio in the bears (Tables 2 and 4). The degree of development of the trochlear process and proximal extension of the trochlear shelf forms the major difference between members of the two families. Other tend to have broadly joined anterior and medial articular surfaces, the procyonids do not: the differences are: the bears bears have a gently curved posterior articular surface which in many of the sigmoid; and the greater tuberosity is bulbous procyonids is in bears and not bulbous in procyonids. ACKNOWLEDGMENTS David Johnson, then at the United States National Museum (USNM), Richard Van Gelder and Sydney Anderson of the American Museum of Natural History (AMNH), and E. Raymond Hall and J. Knox Jones, Jr., of the Museum of Natural History at the University of Kansas (KU) provided the specimens which made this study possible. (No. 2843, 1960) by the Philosophical Society possible museums to look at A grant American these and other made visits to fossil materials and recent specimens. LITERATURE CITED Anderson, S., and J. K. Jones, Jr. 1967. Recent Mammals of the World, A Synopsis of Families. The Ronald Press Co., New York. 453 pp. Davis, D. D. 1964. The Giant Panda, a Mor- phological Study of Evolutionary Mechanisms. Fieldiana: Zool. Memoirs, Vol. 3, 339 pp. Gregory, W. K. 1936. On the phylogenetic re- lationships of the giant panda (Ailuropoda) to other arctoid Carnivora. Amer. Mus. Novit.. 878:1-29. Leone, C. A.. and A. L. Wiens. 1956. Compara- tive serology of carnivores. J. Mamm.. 37:1i- 93 Raven, H. C. 1936. Notes on the anatomy of the 148 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES viscera of the giant panda (Ailuropoda melano- leuca). Amer. Mus. Novit., 877:1—23. Robinette, H. R., and H. J. Stains. 1970. Com- parative study of the calcanea of the Pinnipedia. J. Mamm., 51:527—541. Romer, A. S. 1966. Vertebrate Paleontology. Univ. Chicago Press, 468 pp. Simpson, G. G. 1945. The Principles and Classifi- cation and a Classification of Mammals. Bull. Amer. Mus. Nat. Hist., 85:1—-350. VOLUME 72 Stains, H. J. 1959. Use of the calcaneum in studies of taxonomy and food habits. J. Mamm., 40:392-401. 1962. classification. Osteological data used in mammal Syst. Zool., 11:127-130. Walker, E. P. 1964. Mammals of the World. The Johns Hopkins Press, Baltimore, Vol. 2:647— 1500. Accepted for publication March 12, 1973. THE SYSTEMATIC POSITION OF HELMINTHOGLYPTA TRASKII FIELDI PILSBRY, 1930 (GASTROPODA: STYLOMMATOPHORA) Barry RoTH! ABSTRACT: Dissection of the genitalia of the snail described as Helminthoglypta traskii fieldi shows that it belongs to the so-called Helminthoglypta ayresiana series, a small group of species of coastal, south-central California and the northern Channel Islands. Comparison with other members of the group suggests recognition of it as a distinct species, H. fieldi. distribution of the group. Helminthoglypta traskii fieldi Pilsbry, 1930, was originally described as a subspecies of Helmin- thoglypta_ traskii (Newcomb, 1861), a wide- spread and variable land snail with many named races in southern and Lower California. The latter is the type species of Charodotes Pilsbry, 1939, a subgenus principally characterized by the single, thick wall of the penis, in contrast to the double-walled penial structure of Helmin- thoglypta, sensu stricto. While defining the sub- genus, Pilsbry (1939) stated that he had dis- sected only three of its members: H. traskii, H. petricola, and H. proles. He assigned other species and subspecies, including H. traskii fieldi, to Charodotes provisionally, pending examination of their anatomies. In September 1972, I collected a series of near-topotypic H. traskii fieldi from under sand dune plants just north of the mouth of the Santa Ynez River, Santa Barbara County, Cali- fornia. The dune habitat here is continuous with that at the type locality, Surf, a station on the Pleistocene and post-glacial climatic conditions appear to have influenced Southern Pacific Railroad’s Coast Route less than a mile to the south. Adult specimens were dissected and showed genitalial characteristics re- lating them, not to Charodotes and the group of H. traskii, but to Helminthoglypta walkeriana (Hemphill, 1911), a species of coastal and sub- coastal San Luis Obispo County to the north. Details of shell sculpture and zoogeography sup- port this conclusion. Further data on range and habitat were obtained during field work in October 1972. Recognition of the taxon as a distinct species seems indicated. Helminthoglypta fieldi Pilsbry Figures 1 and 2 Helminthoglypta traskii fieldi Pilsbry, 1930:66-—67, pl. 5, figs. 2-4. Helminthoglypta traski fieldi Pilsbry, Pilsbry, 1939: 178, fig. 88e.—Webb, 1952:4—5, pl. 2, figs. 3, "Dept. Geology, California Academy of Sciences, San Francisco, California 94118. 1973 4, 8; pl. 3, figs. 10—-14.—Webb, 10, fig. 1.—Smith, 1970:42. 1954:15, pl. Helminthoglypta — walkeriana — (Flemphill), — Field, 1930:30 (?in part).—Chace, 1951:7 (record from Surf only). [Not Helix walkeriana Hemp- hill, 1911.) Type material: Lectotype (designated by Baker, 1962: 9), 151516, Academy of Natural Sciences, Philadelphia; paralectotype 327210; three paratypes 152464 in same institution. Type locality: Surf, Santa Barbara County, Cali- fornia, under ice plants and sage on the beach (Pilsbry). Original description: “The shell is more elevated than H. t. phlyctaena, the height 74 to 77 percent of the diameter (in phlyctaena 60 to 66 percent); umbilicus smaller, about 2 mm. wide; post-nuclear whorls are not papillose; last 24% whorls are spirally engraved with lines cutting the striae, strongly de- veloped on the last whorl. Color cinnamon-brown, with a darker, chestnut-brown band bordered on both sides with chamois. Peristome narrowly ex- panded, the columeller margin reflected. “Height 18 mm., diam. 24.5 mm, 6% whorls. “Height 20.7 mm., diam. 26.3 mm., 614 whorls.” (Pilsbry, 1930). This description was repeated, with minor changes, by Pilsbry (1939). ANATOMY The external anatomy appears to present only the features usual for the genus. The living animal is dark slate grey, the depth of color varying somewhat between individuals. The mantle col- lar appears lighter, being densely set with whitish mucus glands. The mantle over the lung is tan with sparse black maculation. Specimens for dissection were prepared by drowning followed by preservation in alcohol. A total of five specimens was dissected. In gross morphology the dissected reproduc- tive systems agreed substantially with the dis- section figured for the species by Webb (1954: pl. 10, fig. 1). The appearance of the lower organs in their everted state was well shown earlier by Webb (1952: pl. 2, figs. 3, 8). Table | presents measurements of various organs in the author’s dissections. The dart sac, atrium, and base of vagina are enveloped in membranous tissue associated with the mucus glands. The common duct of the mucus glands is extremely thin and fine. The penial complex is shown in figure 1. The basal section of approximately 4 mm is cavernous, with a single thin wall. This section terminates distally at an opaque, apparently muscular, con- SYSTEMATICS OF SNAILS 14 du ted rep Pas_e 1. Lengths (in mm) of sele i organs of Melminthoglypta fieldi Specimen | 2 3 j Shell diameter 23 21.5 22 1 Shell height 18 15.5 16 | Epiphallic caecum 22 28 28 24 23 Epiphallus + penis 26 30 30 28 22 Vagina i) 1.5 1.5 3 2 Dart sac + atrium 9 10 10.5 10 8.5 Common mucus duct 2:5 2 2.5 1.5 1.5 Spermatothecal diverticulum 57 67 65 58 51 Sp. div. origin from base of spermatothecal duct 14 striction. Beyond the constriction, the tube is nearly filled by a second, thick-walled inner tube. Longitudinal ridges (four large and five smaller, in the figured specimen) project into the cavity of this inner tube, almost filling it. The genitalia of Helminthoglypta traskii, H. walkeriana, and H. fieldi are all similar in general character and relative proportions of the organs (cf. Pilsbry, 1939: fig. 63b, H. walkeriana; fig. 84b, H. traskii). All three share a relatively large dart sac, short common duct of the mucus glands, and long spermatothecal diverticulum. These features also appear in Helminthoglypta ayresiana (Newcomb, 1861). which Pilsbry (1939) grouped supraspecifically with H. walker- iana as the “Helminthoglypta ayresiana series.” Helminthoglypta walkeriana and H. fieldi have in common a double-walled penis, a pronounced constriction of the penis anteriorly, and a very short vagina. The vagina of H. fieldi is propor- tionally the shortest observed in any Helmintho- glypta species. SHELL CHARACTERS In addition to the author's series of H. fieldi. material examined includes specimens in the Geology Department, California Academy of Sciences (including syntypes of H. walkeriana and “Helix 1911 and the private collections of Allyn G. Smith and S. Stillman Berry. Helminthoglypta fieldi exhibits the basic sculp- var. morroensis’ Hemphill, ture characteristic of both the H. tr and the H. ayresiana series: incised which obliquely intersect radial growth striaz, the 150 BULLETIN SOUTHERN CALIFORNIA pr 4 hi Figure 1. fieldi Helminthoglypta complex: ec, epiphallic caecum; ep, epiphallus; pe, penis; pr, penial retractor muscle; vd, vas deferens. Pilsbry, penial Transverse sections slightly enlarged from main drawing. raised, roughly rhomboidal surfaces thus defined being more or less produced into papillae. In H. traskii the degree of papillation is variable from race to race; H. t. phlyctaena, for example, the nearest geographic neighbor of H._ fieldi in the group, is conspicuously papillose on the spire, the papillae being reduced or absent on the last and penultimate whorls. Helminthoglypta walkeriana bears moderate to strong papillation, frequently eroded from the early whorls but dis- tributed generally over the body whorl and into the umbilicus. Helminthoglypta fieldi tends to develop groups of papillae only immediately be- hind the outer lip of some adult individuals. Scattered papillae, sometimes in linear groups paralleling growth lines, are rare on other parts of the shell, and many specimens have none at all. Occasional papillae occur in the umbilicus. The protoconch of H. fieldi (described from juveniles hatched to captive topotypic specimens, A. G. Smith collection No. 7954) is minutely granular in texture and more or less evidently ACADEMY OF SCIENCES VOLUME 72 Figure 2. shell. Helminthoglypta Author’s collection #229A; just north of mouth of Santa Ynez River, Santa Barbara County, California. fieldi Pilsbry, wrinkled radially, particularly on the dorsal sur- face. Protoconch and first neanic whorl are covered with short, evenly spaced, stiff perios- tracal bristles with recurved ends. These are rubbed off all metaneanic and older specimens seen, but a few round, boss-like bristle-bases sometimes remain as far down as the third or fourth whorl. Other authors have sometimes termed these bosses “papillae,” but they are periostracal in origin and not homologous to the papillose sculpture previously described for the adult shells. The H. fieldi protoconch largely resembles that described by Webb (1941) for H. dupetithouarsi, particularly in the presence of recurved bristles. These bristles readily catch on loose organic matter and may help keep the young snails from being blown or washed out of the clumps of vegetable debris in which they usually live—a matter of some gravity in the sand dune habitat. Apices of well-preserved adults of H. walker- iana show substantially the same characters of wrinkling, granulation, and the presence of mi- nute bosses. The apex of H. walkeriana morroensis is also similar, with the periostracal bosses more persistent on subsequent whorls. The shell form of H. fieldi is quite constant in all material examined (Fig. 2), with the ex- ception of specimens from one locality, dis- cussed below. Adult specimens (i.e., those with reflected peristome) 18.4 range in size from 1973 Figure 3. Helminthoglypta walkeriana (Hemphill), shell. Syntype, CAS Geology Type Collection 1272; “Near Morro, California.” mm-—27.2 mm in diameter and from 14.6 mm— 20.9 mm in height. Number of whorls ranges from 5.8—6.3, the mean being just slightly above six. Adult H. walkeriana (Fig. 3) show greater variation, ranging from 18.0 mm-—28.8 mm in diameter and from 13.6 mm—23.5 mm in height. Whorl-counts of from 5.3—6.2 were noted, the mean being 5.7; more than six whorls were found on specimens approaching or over 27 mm in diameter. Adult H. walkeriana morroensis have from 5.2—5.5 whorls. The relationship of whorl number to greatest diameter in these three taxa is shown by figure 4. The inner lip of H. fieldi covers less than half the umbilicus; half or more is commonly covered in H. walkeriana. Arcuation of the inner lip, cited as a distinctive character by Pilsbry (1939), is subject to individual variation. Specimens from sand hills on the north side of Oso Flaco Lake, southwestern San Luis Obispo County (A. G. Smith collection No. 8010 and in the author’s private collection), are in some Ways intermediate between H. fieldi and H. walkeriana. They combine the open umbilicus of H. fieldi, the large size and tumidity of H. walkeriana, and an intermediate degree of papil- lation. Whorls range from 5.6—6.2, with the mean being 5.8 (Fig. 4). This combination of characters seems confined to this local popula- tion, which may represent a past site of inter- breeding between the two species. SYSTEMATICS OF SNAILS 15] ECOLOGY AND RANGE Helminthoglypta fieldi has previously been re ported only from beach habitats in the vicinity of its type locality. with Munz (Mesembryan- (M. coastal isocoma (Haplopappus venetus), and bush lupine (Lupinus spp.) plants. Dead shells collected by the author a short distance inland from the Santa Ynez River mouth suggest that the snail may range up the river valley, possibly into the agricultural region west of Lompoc. The low hills directly behind the shore are composed of loose, sandy and There it is associated the Coastal Strand plant community of and Keck (1965), with themum chilense), Hottentot-fig sea-fig edulic), among the conspicuous diatomaceous soils; here occurs more shrubby vegetation, the Coastal Sage Scrub plant com- munity of Munz and Keck (1965), which might provide suitable snail cover as it does for H. walkeriana at Morro Bay. A lot in the California Academy of Sciences (No. 42776) extends the snail’s known range some 48 km north to “Pismo” (= Pismo Beach), San Luis Obispo County, also immediately on the coast. Inland, the author has collected it at Halcyon, 5 km southeast of Pismo Beach. A very juvenile specimen from a drainage gully on the northwest slope of Nipomo Mesa, south- east of Oceano, agrees in all details with hatchling H. fieldi described above. At Oso Flaco Lake, the snail inhabits the Coastal Strand community. Helminthoglypta walkeriana appears to be most characteristically a snail of the Coastal Strand and Coastal Sage Scrub plant communi- ties in the vicinity of Morro Bay. It has been found from near sea level at the south end of the bay to 45 m elevation in dunes atop the lowest marine terrace in Montana de Oro State Park. Inland, as in canyons of the San Luis Range, it is replaced by Helminthoglypta um- bilicata (Pilsbry, 1898); the latter species also appears more aggressive in colonizing disturbed areas. Field’s (1930) record of H. walkeriana from “the sandy beaches above Point Con- ception” certainly refers, at least in part. to the specimens which became the type lot of H. traskii fieldi later the same year. Hemphill’s (1911) original locality citations for H. walkeriana (“San Luis Obispo”) H. w. morroensis (“San Luis Obispo County”) were imprecise. The syntypes of the former in the California Academy of Sciences (Geology Type Collection Nos. 1271-1276) are labeled and 152 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 72 | (pare ae eee bo al onl "= | H.walkeriana A i 28 |__| H.w. morroensis A | H. fieldi - i ial Suir O A =| Pismo e@ O Tr. a a. m4 | Oso Flaco | = O -e— -—___—___ ——SSSSSS O al a A oa > lige = A; A kb | = gw, 4 € — O 0 = O [= DAG |e O w a e O 4 Ww = a O < rae A a Ph O a a O O ) 2 iP A A fa O O O Ee 0 pare x 6 Fe 22 ~ qasuy aqeaow Jo 2oonpay (s99uld Jo age -QA0lW JO JOIONPpPL [NJIEAO0d Sursop> day) sasduy 2)112W0} -nap JO 10101 prvaunog aawo} -nap JO 10}v}OI pPIeAUyT aawoinep Jo (uonow -o1d) qUaWaAOW pPIeEMIOZ auawolnap Jo Josseidaq ayiauoynep Jo ]UAWeAOW preMyoeg Biaoayo Jo 1ojINpqy jauy apquaow Jo asvq “] “N N ‘ienuy ajqvuaow yo aseq 9y} 1 oumONAs poeduys-uvy v UL ad2aAu0d YSIyA\ sdins snouipua; uo youne saqiy (C1) yitay paraaos surewal — ayliawojynep jo ulsreu yenusA “LL °N ayiawoynap jo aseq jesioppiw “[ “N ayiewoynap jo aseq jesiop JauUr “IPT[IIqryouo |, 30} -nap Jo aseq jemusa “L ‘N ayiewonep Jo aseq [esioppra “Ieyqiyjouoy *N ayliapasowia ]exod Jo Jepiog Jolejsod 19uUt “L O}IOWO) -nap jo [pea sno “LCN ayeuuid “y ‘yjVUs ayeuuld ‘a}iawoy -nep ut coeds Jsofvu s][J ‘aaisseus puv asivy “4 aawo nap jo sya [wy -UdA puv JoUU! ‘[esIop “A “A wasup ajqvaowd sujpysado 9/98 JV] ayeuuid BxXOO ‘paysiM] pur [e]UOZIIOY (C1) jo [JBM [esayey Jano “4 ‘NF aysoddo ‘[wolipuljAd “4 (Cae) ayeuuid ym pareaoo Apied “4 “a ‘AWARD [BXOD JO JAvd 19jNO * BxO9 jo [[eM Jesiop Jouul “4 “¢ ajeuuld ysayprus “4 SaJISNU ajLlauUojnap-VxO) ATOSAW OISNTALNI Sa[OSNUE SISUII]X® YyIA poa1eaoo ‘ayirajosoyua [exo jo [jem yesjusa o} parfdde Ajasopa ‘oyyuey ‘uy? “A “A BxOO ysno1y} sunt ajiiajosojua yexoo Jo [eM [eUsA “4 ‘q a}1a[9sojua ayeuuld ‘(9) YIM yexoo Jo ulsiew Jou “4 “q = — PedeA0d urewel ‘padeys-uey Sa]ISnuUd a}l4auojnap AJlda]ISOJUd [px0oD [altered = “yaeys snourpue} e yum pepraoid § Al[et} -uaa ‘padeys-uoqqu uly) “¢ (OI) YUM peteaod sulewel ‘4 ‘q goponpqy (81) soynppy CLI) ayawoynap [exoo JONOXy (91) ayawoynap [exoo Aolojuy (SL) ayiawoynap jexoo soladng (PT) ayldawoynep aytdaposoque [exo JouaxA (EL) o}l4auoynap d}LLa]DSO}Ua [PXOD JOLIO}UT (ZI) aytaposoqus [e1ooid -a}L1a[ISOJUA [VXOD IOLI9}SOd (11) uonoun{ uorjiasuy sioqly ajosnul Jo JUawWaduPIIE uIsLIO ‘ginjonas ‘adeys :ainjeNn aposny (panuyuo)) “T ATavL BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 DEUT 1 COX 2 ao COXENT rea, COXENT 1o-< eee I ‘4 = = aae 8 COXENT COXENT 1974 CHELICERAL MUSCLES OF An outline of the classification and anatomical arrangement of the scorpion cheliceral muscles is given in table | and figures 1-6. Hach muscle has been given a number (shown in the paren- theses) which is used in the discussion and descrip- tion to avoid lengthy nomenclature. DISCUSSION Scorpions are predaceous and extract juices from prey (Brunson, 1951; Stahnke, 1966). The role of chelicerae in feeding can be clearly understood from the work of Stahnke (1966). According to him . “The pedipalps moye the prey to the chelicerae which are’ small chelate appendages, their shearing jaws are used to pierce, cut and tear the soft and hard parts of the prey into minute pieces. The tiny particles produced by the chelicera are packed between the coxa of the pedipalps.” All these actions necessitate a free movement of these appendages in various directions. A casual study of the arthrology of the chelicera revealed that the coxal joint simply consists of arthrodial membrane and allows great freedom of movement. The adduction is brought about by three muscles, viz. (10), (3), and (6). The first two muscles are broad and with parallel fibers, whereas, the last is fan-shaped and exhibits pinnate structure. All these muscles are well-developed and powerful. Although they arise from different regions of the prosoma, their contraction results in an in- ward movement of the chelate end of the ap- pendage. The muscles (11) and (7) serve as abductors. Their contraction moves ends of the two chelicerae apart. Both these muscles are well-developed with parallel fibers. Inward and outward rotatory movements of the chelicerae are brought about respectively by (1) and (5) small but their pinnate nature suggests powerful action. muscles. These muscles are relatively The promotory and remotory movements of these appendages are brought about by the (2) THE INDIAN GROUND SCORVION 134 and (4) muscles, respectively, Both these muscle are narrow and tendinous. An inward flexion at the joint between the coxa and deutomerite segments of the chelicera result: These muscles show pinnate arrangement of the muscle from action of the (13) and (9) muscles fibers and are provided with tendons at their in sertion ends. As stated earlier, this joint allows rel atively lesser movement and as such requires the presence of powerful muscles for flexor move ment. The pinnate nature and the tendinous ma- terial of the (13) and (9) muscles are thus suitably The animal adapted to meet these functional demands. contraction of these muscles enables the to bring the chelate organ of its chelicerae much closer to the mouth. The muscle operating the extensor mechanism of these segments is relatively weak in action. A noteworthy feature of the muscles operating the chela of the chelicerae is that unlike the condi- tion in scorpion pedipalps, both adductor and ab- ductor sets of muscles are represented (Snodgrass. 1952). Compared with the abductor muscle (18). the adductor (17) is powerful. ACKNOWLEDGMENTS I wish to thank M. S. Dubale. Department of Zoology. Gujarat University, Ahmedabad, who suggested the problem and provided encouragement and guidance LITERATURE CITED Awati, P. R., and V. B. Tembe. Buthus tamulus (Fabr.). The Indian Scorpion: Mor- phology, anatomy and bionomics. Zool. Mono.. No. 2, Univ. Bombay, 68 pp. 1956. Brunson, R. B. 1951. An unusual meal. Turtox News, 29: 151. Cloudsley-Thompson. J. L. 1958. Spiders. Scor- pions. Centipedes, and Mites. Pergamon Press. 228 pp. Dubale, M. S., and A. B. Vyas. 1967. of the pedipalp of Palamnacus The myology fulviceps (C. Figure 1. Extrinsic muscles of chelicera muscles (other muscles removed). Figures 5 and 6. Intrinsic muscles of COXENT = Coxal _ entosclerite: (Dorsal Figures 3 and 4. coxa DEUT = Deutomerite view). Figure 2. Dorso deutomerite Intrinsic muscles of coxal entosclerite. segments. COX = Coxa. FING = Movable finger. and deutomerite segment: For identification of numbered muscles see table 1. l4 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Koch). Proc. 55th Ind, Sci. Cong. (Abstracts): tion and development of scorpions. Quart. J. 114. Micro. Sci., 68: Part IV. — 1968. The structure of the chela of Smodgrass, R. E. 1948. The feeding organs of Heterometrus sp. and its mode of operation. Archnida including mites and ticks. Smithson. Bull. So. California Acad. Sci., 67:240-244. Misc. Coll., 110:1-93. 1952, A text book of arthropod anatomy. 1969. Heterometrus sp. and the associated muscles. Proc. Nat. Acad. Sci. India, 39:169-177. On the endosternite of the scorpion Lankester, E. R., W. B. S. Benham, and E. J. Beck. 1885. On the muscular endoskeletal systems of Limulus and Scorpio with some notes on the anatomy and generic characters of scorpions. Trans. Zool. Soc. London, 11:311—390. 1924. Pawlowsky, E. N. Studies on the organiza- Comstock Publ. Assoc. Ithaca, 363 pp. Stahnke, H. L. 1966. Some aspects of scorpion be- haviour. Bull. So. California Acad. Sci., 65: 65-79. Venkateshwararao, P. 1967. Feeding apparatus in the scorpion Heterometrus fulvipes. Proc. India Acad. Sci., 68:1—9. Accepted for publication July 21, 1972. XIPHIORHYNCHUS KIMBLALOCKI, A NEW BILLFISH FROM THE EOCENE OF MISSISSIPPI WITH REMARKS ON THE SYSTEMATICS OF XIPHIOID FISHES Harry L. FIERSTINE! AND SHELTON P. APPLEGATE” AsstRAct: Xiphiorhynchus kimblalocki, a new species of extinct billfish from the Eocene of Mississippi, is described. This is the first record of Xiphiorhynchus outside of western Europe, and the material consists of a well-preserved rostrum, three partial vertebrae and two fin spine fragments. Xiphiorhynchus kimblalocki is compared with other living and extinct billfish and appears to be intermediate in morphology between the Xiphiidae and Istiophoridae. Various genera of fossil billfish are critically discussed and we suggest that the Blochiidae, Paleorhynchidae, and the “Cylindracanthus-group” should be placed in Xiphioidei /ncertae sedis until better evidence indicates that they are billfish. We speculate that Xiphiorhynchus is an extinct offshoot from an unknown pre-Eocene common ancestor between Xiphtiidae and Istiophoridae and is closer to the Istiophoridae than to the Xiphiidae. We also agree with earlier workers that the lineages of the Xiphiidae and Istiophoridae run back separately into basal Eocene times and that any common ancestry to each other and to the scombroids must have been prior to the Eocene and may have extended into the Cretaceous. Billfish remains have been described in rocks from the Cretaceous Age (Dixon, 1850) to the Pleisto- cene (Fierstine and Applegate, 1968). The exact taxon to which many of these remains belong has puzzled paleontologists because identifications are usually based on isolated skeletal parts, partic- ularly the rostrum. Detailed anatomical compari- sons of recent genera are lacking and the lack of information has led to nomenclatorial confusion and misidentification of the fossil forms. Various attempts have been made to synthesize the avail- able evidence and to make order out of chaos (Woodward, 1901: Leriche, 1905; Carter, 1927; Casier, 1946, 1966). Unfortunately, even the *Dept. Biological Sciences, California Polytechnic State University, San Luis Obispo, California 93401 and Research Associate, Dept. Vertebrate Paleontol- ogy, Natural History Museum of Los Angeles County, Los Angeles, California 90007. “Dept. Vertebrate Paleontology, Natural History Museum of Los Angeles County, Los Angeles, Cali- fornia 90007. 1974 latest monograph (Casier, 1966) has failed to apply the well-established nomenclature and bio- logical knowledge used by recent ichythyologists (Greenwood, Rosen, Weitzman, and Myers, 1966; Gosline, 1968; Howard and Ueyanagi, 1965; Morrow and Harbo, 1969; Nakamura, Iwai, and Matsubara, 1968; Robins and de Sylva, 1960, 1963). Thus, it is the object of this paper to describe a new species from the Eocene of Mississippi and to put at least a part of the fossil billfish problem in a more modern perspective. SYSTEMATIC DESCRIPTION CLASS OSTEICHTHYES Order Perciformes Suborder Xiphioidei Family Xiphiorhynchidae Genus Xiphiorhynchus, Van Benden, 1871 Xiphiorhynchus kimblalocki, new species Figures 1—4 Holotype: LACM 25575.1, a rostrum (Figs. 1, 2); LACM 25575.2, a partial anterior abdominal vertebra (Fig. 3); LACM 25575.3, a partial posterior caudal vertebra (Fig. 3); LACM 25575.4, a vertebral frag- C Figure 1. Rostrum of Xiphiorhynchus kimblalocki, new species (Holotype—LACM 25575.1): A. lateral view; B. ventral view; C. dorsal view. Scale equals 10 cm. NEW BILLFISH FROM EKOCENE OF MISSISSIPI] 1s Figure 2. Rostrum of Xiphiorhynchus kimblalocki, new species (Holotype—LACM 25575.1): A. cross- section 220 mm from distal tip: B. cross-section 170 mm from distal tip. Scale equals 15 mm. ment; LACM 25575.5, a partial fin spine (Fig. 4): LACM 25575.6, a partial fin spine (unfigured). Horizon and Locality: LACM locality 7003, Scott Co., Mississippi. Southwest side of Sherman Hill (Hill 618), NW% SW% Sec 16, TSN, R9E. Forest (?Hill) Quadrangle, U.S.G.S., 1950. The specimens were collected in the Shubuta Clay member of the Yazoo Formation, Jackson Group (Eocene). The Shubuta consists of green to greenish-gray calcareous to non-calcareous. glauconitic. fossiliferous. silty clays (DeVries, ef al., 1963). Selenite crystals are common. There is no indication that the fossil was collected near the base or near the top of the formation. The associated fauna consisted of a skull and cervical vertebrae of Zygorhiza Kochi, an extinct, primitive cetacean. The species is named in honor of Mr. Kim Blalock who collected the specimens and discovered the site. Diagnosis: The rostrum differs from other Yipii- orhynchus in its large size. rugose surface texture. lack of a central longitudinal nutrient canal at its distal end, and its diminutive alveoli. 16 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Figure 3. Vertebrae of Xiphiorhynchus kimblalocki, new species (Holotype): A. partial abdominal verte- bra (LACM 25575.2), lateral view; B. partial ab- dominal vertebra (LACM 25575.2), ventral view: C. partial caudal vertebra (LACM 25575.3), lateral view; D. partial caudal vertebra (LACM 25575.3), ventral view. Scale equals 25 mm. VOLUME 73 Figure 4. Fin spine of Xiphiorhynchus kimblalocki, new species (Holotype—LACM 25575.5): A. lateral view; B. anterior view. Scale equals 10 mm. Description: The greatest length of the rostrum is 580 mm; it is nearly circular in cross-section at its distal tip and it becomes progressively larger and more depressed in cross-section towards it proximal end. One centimeter from the distal tip, the rostrum measures 11 mm wide and 12 mm thick, and the proximal end (base) is 83.5 mm wide and 38.0 mm thick. The proximal one-half probably was slightly depressed and crushed during preservation. The dorsal surface is rugose at its distal end and longitudinally striated at its proximal end. Ventrally, the distal one-half is rugose. A poorly preserved alveolar layer covers the middle one-half of the ventral surface of the rostrum. The alveoli range from .25 to .4 mm in diameter. The proximal one- fourth of the ventral surface is not preserved. A cross-section of the rostrum (Fig. 2), cut 220 mm from the distal tip, revealed poorly preserved bone. Traces of matrix revealed a central longitudinal nutrient canal which is bordered laterally by a pair of smaller lateral longitudinal nutrient canals. A cross-section of the rostrum 170 mm from the distal tip (Fig. 2) revealed three right lateral longi- tudinal nutrient canals and two left lateral longitudinal nutrient canals. The central longitudinal nutrient canal had terminated prior to this section. The centrum (Fig. 3) of the anterior abdominal vertebra (LACM 25575.2) is 70 mm long and the anterior and posterior surfaces are nearly circular. 1974 NEW BILLFISH FROM The ratio of the height (60.4 mm) of the centrum, measured at its anterior surface, to its greatest length (70.0 mm) is 0.86. The surface texture is rugose and contains many pits and fossae. In lateral view, the ventral surface is gently concave; in ventral view the lateral surfaces are relatively flat and do not have a “pinched-in” appearance, The neural arch is broken at its base, but appears to be divided into a larger anterior zygopophysis and a smaller posterior zygapophysis. A large rib attachment area is present on each lateral side. The centrum (Fig. 3) of the posterior caudal vertebra (LACM 25575.3) is 92.5 mm long and the anterior and posterior surfaces are nearly circular. The ratio of the height (64 mm) of the centrum, measured at its posterior surface, to its greatest length (92.5 mm) is 0.69. The surface texture is rugose and contains many pits and fossae. In lateral view, the ventral surface is gently concave; in ventral view, the lateral surfaces are concave and present an hourglass outline. The zygapophyses and the neural and haemal arches are broken off at their bases. The two partial fin spines (LACM 25575.5 and LACM 25575.6) belong to a median (anal or dorsal) fin (Fig. 4). LACM 25575.5 measures 34 mm across its nearly complete base and 72 mm long. Although its distal tip is missing, it appears to have tapered rapidly to a point. When viewed from the side, it shows a slight posterior curvature. If the spine had been complete, we estimate that it would have mea- sured 112 mm long and 40 mm wide. LACM 25575.6 is a left or right half of a spine that measures 20 mm across its base and 138 mm in length. When viewed from the side, it shows no antero-posterior curvature. If the spine had been complete, we estimate it would have measured 40 mm across its base and 210 mm in length. DISCUSSION The rostrum of Xiphiorhynchus kimblalocki, like other species of Xiphiorhynchus (Leriche, 1905; Casier, 1966), is similar in morphology to those found in the Istiophoridae (Tables 1 and 2). It differs, however, in the number of nutrient canals (Table 2), in having smaller alveoli, and in having a more depressed cross-section at its base (Table 1). The longitudinal openings, revealed in trans- verse section (Fig. 2), can be called nutrient canals with some assurance, since histological exami- nation of the rostrum of a striped marlin (Tetrap- turus audax) revealed blood vessels within the canals (Vladimir Walters, Univ. California, Los Angeles, unpublished). The rostrum of the swordfish (Xiphias gladius). when compared to the rostrum of X. kimblalocki, EOCENE OF MISSISSIPPI / is more greatly depressed along its entire length However, measurements taken at the base of the rostrum (Table X. kimblalocki and X. gladius are nearly equally depressed, As noted 1) show that earlier, some of the depression at the base of the rostrum in X, kimblalocki may be due to defor the swordfish lacks alveoli and denticles, but it does mation during preservation. The rostrum of contain a central (? nutrient) canal and one pair of lateral nutrient canals. The rostra of Blochius and what we are calling the = Cylindracanthus-group Congorhynchus, (Aglyptorhynchus, Cylindracanthus, Glyptorhyn- chus, Hemirhabdorhynchus, ete.) are much smaller than the rostra considered above. They all prob- ably have numerous longitudinal nutrient canals (Carter, 1927; Casier, 1966; Fierstine and Apple- gate, unpublished). The two vertebrae of X. kimblalocki differ from those of other billfish (Table 3). The anterior abdominal vertebra is similar in size and shape to a third or fourth abdominal of a black marlin (Makaira indica). It differs, however, in rugose surface texture (the centrum of the black marlin is relatively smooth), the lack of a fossa on its ventral surface, and the shape and _place- ment of its rib attachment. The third vertebra of the black marlin lacks a pronounced transverse process and has a large oval scar for rib attach- its ment. The transverse process of X. kimblalocki is broken at its base, but it probably was large and quite pronounced. The third vertebra of X. well-developed transverse process and no obvious rib facet. In this respect, the centrum of X. gladius is similar to the centrum of X. kimblalocki. The third vertebra of the swordfish (like most other vertebrae in the vertebral column) is more lightly constructed, appears to be more weakly ossified, and the surface architecture is smoother than for the abdominal vertebra of X. kimblalocki. The caudal vertebra of X. kimblalocki differs considerably from any istiophorid caudal vertebra. It most closely resembles one from the posterior caudal region (approximately in the vicinity of the 21st vertebra). The height to length ratio of the centrum is much larger than the ratio for the three living istiophorid genera (Table 3). Thus, the caudal vertebra of X. kimblalocki is much gladius has a thin, more cube-like than those found in the Istio- phoridae. The height to length ratio of the centrum compares much more favorably with the 21st centrum of X. gladius. The probable place- ment of the neutral and haemal spines is similar. 18 BULLETIN TABLE |. Species Base Istiophorus platypterus 18 ——. — .82 (Sailfish ) 22 Makaira indica 46.5 : —— — ./0 (Black marlin) 66.0 Tetrapturus audax 22 —— = .59 (Striped marlin) 37.5 NXiphias gladius 39 fs = .44 (Swordfish ) 89 Xiphiorhynchus kimblalocki 38 = .46 83.5 but as with the anterior caudal vertebra, the surface texture and construction is much more rugose and massive in X. kimblalocki than in the swordfish. Casier (1966) described and figured from the London Clay, a vertebra that he identified as Xiphiorhynchus (?). This centrum is very similar in shape and preservation to the caudal vertebra of X. kimblalocki. Systematics of the Xiphioid Fishes.—Fierstine and Walters (1968) and Gosline (1968) indepen- dently concluded that the Istiophoridae and Xiphi- idae should be placed in the perciform suborder Xiphioidei apart from the Scombroidei. Both fol- lowed Regan’s (1909) and Gregory and Conrad’s (1937) view that the Xiphiidae and Istiophoridae extend into basal Eocene time and that any com- mon ancestry to each other and to the scombroids must have been prior to the Eocene and may have extended to the Cretaceous. These phylogenetic conclusions were partially based on the identifica- tions by paleontologists who we now believe were in error. The following discussion is a critical review of billfish phylogeny and _ classification based primarily on fossil forms. Woodward (1901) recognized that the Xiphiidae included all living billfish genera, as well as Xiphiorhynchus, Acestrus, and Brachyrhynchus. All the xiphiids were placed in the Division Scombriformes along with the scombrids, caran- gids, stromateids, and the extinct family Palae- orhynchidae. Woodward placed Blochius in the family Blochiidae in the division Blenniiformes. Regan (1909) placed the billfish in their own division, Xiphiiformes, within the perciform sub- order Scombroidei. In this division he separated the istiophorids, xiphiids, and xiphiorhynchids into SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Height to width ratios of the rostra of certain billfish. Middle Point Tip el cm) 13.5 8 —— = .80 — — 3 17.5 11 36.0 mie S83) Missing 43.5 15.0 552 — OW =s/2 225) ce 18 11 3) 228 60 39 35.3 11 —— — .62 = 2 their own families. In addition, Regan included the blochtids as a family within the Xiphiiformes. The placement of the Paleorhynchidae with the xiphioids has been labeled as dubious (Gosline, 1968). According to Danil’chenko (1960), the paleorhynchids have about 45—60 vertebrae, jaws which are very elongate with the lower jaw some- times longer than the upper, and ribs which com- pletely surround the abdominal cavity. Thus, the paleorhynchids have about twice the vertebral number and have jaws and ribs unlike any living adult xiphioid. Until the paleorhynchids have been thoroughly studied, we prefer to put them in the Xiphioidei Incertae sedis. Woodward (1942) stated that Blochius does not exhibit the character of a xiphioid. Specimens of Blochius have a low vertebral number (24) and they lack (or have very reduced) pelvic fins, a condition similar to that found in living sword- fish. However, blochiids are small (about 1 meter), have numerous dorsal and anal spines, have large scales, a round bill, and were living contem- poraneously with the various species of Xiphi- orhynchus and some istiophorids. Without ques- tion, the blochiids need additional study in order to determine their relationship and. until such re- search is accomplished, we prefer to put the Blochiidae in the Xiphioidei Incertae sedis. The relationship of the Cylindracanthus-group to the Xiphioidei is also highly questionable. This group is only known by fossil rostra; no other skeletal remains have ever been positively identi- fied. Encouraged by Woodward, Carter (1927) studied a Cylindracanthus rostrum from the Eo- cene of Nigeria. He showed that it was histo- logically similar to a fragment of a Blochius rostrum as well as to a dermal spine of an un- 1974 NEW BILLIEISH FROM TABLE 2. EOCENE OI VUISSISSIPPT 14 Rostral characters and chronological and geographical distributions of various billfish wenera Comparative Rostral Characters Size and Shape Taxon of Cross-Section BLOCHIDAE + Blochius small, round +- Cylindracanthus small, round group ISTIOPHORIDAR +- Acestrus unknown Istiophorus large, round Makaira large, round Tetrapturus large, round -+-Brachyrhynchus large, round PALEORHYNCHIDAE +-Enniskillenus unknown -+ Homorhynchus unknown +-Paleorhynchus unknown + Pseudotetrapturus (?) large, round XIPHIIDAE Niphias large, depressed --undescribed genus large, depressed XIPHIORHYNCHIDAE +-NXiphiorhynchus large, round ++ = extinct related living trunkfish (Ostracion). He con- cluded that the Cylindracanthus specimen was probably the bill of some extinct swordfish re- lated to Blochius. This relationship has been accepted by most other ichthyologists and pale- ontologists (Berg, 1940; Casier, 1946, 1958; Darteville and Casier, 1943, 1949: Gregory, 1951; Leriche, 1942; and Romer, 1966). Recently Casier (1966) divided the Cylindracanthus-group into two parts and questionably put one part in the family Blochiidae of the Order Heteromi and the remainder in family Xiphiidae of the Order Scombromorphi (= ?Scombroidei). No expla- nation was given as to why he thought there was a relationship to the Order Heteromi (= Nota- canthiformes). Woodward (1942) placed Cylin- dracanthus (= Coelorhynchus) in Incertae sedis and we agree with this decision. The transfer of the Cylindracanthus-group trom the Niphioidei proper removes all the pre-Eocene representatives. On the basis of the above discussion, the seven Nutrient Canals Number of Longitudinal Chronologic and Geographic Mange unknown Eocene, Europe Africa, Eu America to numerous Cretaceous, North Oligocene, Europe rope, unknown Eocene, Europe one pair Eocene, Europe, N one pair ; America to recent, one pair world-wide seas. one pair Eocene to Pliocene, Europe unknown Eocene, Europe unknown unknown unknown | Eocene to Oligocene, Europe central and one pair Oligocene, Europe to re- cent, world-wide seas. North America (2?) central and one pair Eocene, Eocene, Europe, N. America, Africa central and two pair genera: Acestrus, Brachyrhynchus, Istiophorus, Makaira, Tetrapturus, Xiphias, and Xiphiorhyn- chus are all that remain as members of the Xiphioidei proper. Beginning with the living genera, we can reiterate that the Xiphiidae (Xiphias) is structurally very different from the Istiophoridae (Istiophorus, Makaira, Tetrapturus). Xiphias has a poor fossil record. Leriche (1910) identified a vertebra from the Oligocene of Bel- gium as Xiphias rupelensis, and we agree with his identification since the specimen is very similar to the penultimate vertebra of X. gladius. As far as we know, all other fossil records are erroneously based on istiophorids or members of the Cylindra- canthus-group. Except for the rostrum, the skeleton of XY. gladius is rather weak and fragile so that preservation is probably poor. Not enough osteological information is known to distinguish between the rostra, skulls or verte- brae of the various recent or fossil istiophorids: identifications of fossil forms. therefore, exact 20 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Pabte 3. Height to length ratios of the centra of certain billfish. 3rd 21st Species Vertebra Vertebra Istiophorus platypterus 30.5 24.0 SU sas —— = .54 = osh7/ (Sailfish ) 57.0 66.5 Makaira indica 52.5 38.5 : — .84 ——. = 46 (Black marlin) 62.5 83.5 Tetrapturus audax 30 24 : ; ——_— .64 = 3h) (Striped marlin) 47 61 NXiphias gladius 43.0 54 cl = .69 — =.75 (Swordfish ) 62.5 We Abdominal Caudal Vertebra Vertebra NXiphiorhynchus kimblalocki 60.4 ae 64.0 70.0 92.5 which are usually fragmentary, are to be ques- tioned. Some of the fossil rostra may belong to Xiphiorhynchus and those identified as Brachy- rhynchus are probably congeneric with J/stioph- orus, Makaira, or Tetrapturus. This latter ob- servation was also noted by Woodward (1901). It seems, therefore, that Brachyrhynchus and the living istiophorids may form a continuum that dates from the Middle Eocene of Europe (Bruxel- lien). Acestrus is only known from the early Eocene (London Clay—Ypresien) and the remains con- sist of posterior crania. Casier (1966) feels that these crania may belong to one of the other bill- fish genera, but not Xiphiorhynchus. Acestrus, like the living billfish, has pronounced muscle fossae on the dorsal surface of the posterior part of the cranium, whereas fossae seem to be lacking in Xiphiorhynchus. Casier (1966) also emphasizes that the cranium of Acestrus is very similar to that of the extinct scombroid, Scombrinus. All known cranial fragments of Acestrus are much smaller (they measure 50-60 mm in_ length) than the counterparts in living adult billfish and they are about one-half the size of the crania of Xiphiorhynchus. Even though the exact taxo- nomic placement of Acestrus is uncertain at best, we prefer to keep the genus in the Xiphioidei proper until more is known. It is possible that Acestrus is an immature billfish. Xiphiorhynchus is intermediate in many respects to the Xiphiidae and Istiophoridae. Each frontal bone has ridges that radiate from a central point similar to the swordfish, whereas only the anterior ridges are pronounced in the Istiophoridae. The ratio of the length of the posterior part of the cranium (anterior edge of the supraoccipital to the posterior edge of the exoccipital) to the length of the anterior part of the cranium (anterior edge of the mesethmoid to the posterior edge of the frontal) is about 0.35 for Xiphiorhynchus priscus, T. audax, T. angustirostris, Istiophorus sp. and M. indica. A similar ratio for X. gladius is 0.13, thus, the swordfish has a much smaller posterior region of the skull than other billfish. The rostrum of X. kimblalocki appears to be broad at the base, similar to the swordfish, and it is round in the distal one-half, similar to the istiophorids (Table 1). Both the swordfish and Xiphiorhynchus have a central canal in their rostra, whereas the istiophorids have only lateral canals. However, X. kimblalocki lacks a central canal in the distal one-fourth of its rostrum. The abdominal vertebra of X. kimblalocki (Table 3) is similar in shape to the third vertebra of the black marlin (M. indica), yet the placement of the transverse process is similar to the third vertebra of the swordfish. In shape, the caudal vertebra of X. kimblalocki is similar to those of the swordfish. It seems, therefore, that Xiphiorhynchus is intermediate between the Istiophoridae and the Xiphiidae and gives evidence that the two living billfish families diverged from a common ancestor prior to the Eocene. However, chronologically, Xiphiorhynchus is not able to be the common ancestor since it has never been found prior to the Eocene. There is no evidence that it is a 1974 NEW BILLEISIT FROM surviving parental stock. Very recently one of us (S.P.A.) collected a 75 cm section of a swordfish- like rostrum from the Yazoo Clay of the Eocene of Mississippi (Fierstine and Applegate, unpub- lished). ‘This very depressed rostrum differs con- siderably from any rostrum known in’ Xiphio- thynchiidae and Istiophoridae and demonstrates that a swordfish-like animal was living contem- poraneously during the Eocene with the Xiphio- rhynchidae and Istiophoridac. At this time we speculate that Niphiorhynchus is an extinct off- shoot from a yet unknown common ancestor and is closer to the Istiophoridae than to the Xiphiidae. Thus, even though based on different evidence, we must return to the conclusions of Regan (1909) and Gregory and Conrad (1937) that both the Xiphiidae and Istiophoridae extend into the basal Eocene and that any common ancestry to each other and to the scombroids must have been prior to the Eocene and may have extended to the Cretaceous. Our proposed classification scheme for the billfish is as follows: Class Osteichthyes Order Perciformes Suborder Xiphioidei Family Istiophoridae: rhynchus, Istiophorus, Makaira, Tetrap- Acestrus, Brachy- turus. Family Xiphiidae: scribed genus. Family Xiphiorhynchidae: chus. Xiphioidet Incertae sedis. Family Blochiidae: Blochius. Family Paleorhynchidae: = Enniskillenus, Homorhynchus, Paleorhynchus, Pseudo- Niphias and unde- Xiphiorhyn- tetrapturus. Family unknown: Cylindracanthus-group. ACKNOWLEDGMENTS We wish to express our thanks to the various members of the Mississippi Gem and Mineral Society, Jackson, Mississippi for their cooperation in our field work. Particular thanks go to Kim Blalock who donated the type specimen to the Natural History Museum of Los Angeles County for scientific study. Armando Solis of the museum staff took all the photographs. Joseph Codespodi of Cal Poly printed the plates, and Leonard Bessom of the museum staff prepared the rostrum for study. Darlene Miller, Patricia Knapp, and Cydney Dawson of Cal Poly typed the many drafts of the manuscript. EOCENE OI VUISSISSIPPT a | LITERATURE CITED Berg; Le S recent and fossil Brothers. 517 pp rudy Zool, Instit 1940. Classification of fishe botl Ann Arbor, Michigan: Edward (translated into English from Akad USSR 5(2) Naubh Carter; J; T fish, 1927 Cylindracanthus Ihe rostrum of the fossil sword Leidy (Coclorh Eocene of Nigeria introduction by Sir Arthur Smith Nigeria, Occas, Paper nehu (with ar Woodward ) No. 5:1-15 Agassiz) from the Geol. Survey Casier, E. 1946. La faune ichthyologique de l'Ypresien de la Belgique. Mém. Mus. Roy. Hist Nat. Belgique., 104:3~—267 1958. Contribution a l'étude des poisson fossiles des Antilles. Mém. Suisses Paléont., 74(3):1-96. ——. 1966. Faune Ichthyologique du London Clay. British Museum (Natural History) Lon- don, 1:1—496. Danil’Chenko, P. G. 1960. Maikop deposits of the Caucasus. from Russian) Akad. Nauk USSR. 78: 1-247. Bony fishes of the (translated Trudy Paleo Instit., Darteville, E.. fossiles du and E. 1943. Les Bas-Congo et des Casier. poissons regions voisines (Primiere Partie ). Ser. A, 2:1—200. Ann. Mus. Congo Belgique.. 1949, et des regions voisines (Deuxiene Partie). Mus. Congo Belgique, Ser. A, 2:201—255 Les poissons fossiles du Bas-Congo Ann DeVries, D. A.. W. H. Moore, M. K. Kern. H. M. Morse, and G. E. Murray. 1963. Jasper County Mineral Resources. Bull. Mississippi Geol. Econ. Topogr. Survey, 95:1-101. Dixon, F. 1850. fossils of the Tertiary and London, 422 pp. The geology and Cretaceous formations of Sussex. Fierstine. H. L.. and S. P. Applegate. 1968. Billfish remains from Southern California with ren on the importance of the predentary bone. B So. California Acad. Sci... 67:29-39. and V. Walters. 1968. St and California Fierstine. H. L.. locomotion Mem. So. dies in anatomy of scombroid fishes Acad. Sci.. 3 -] 1 6:1-—S1. 22 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Gosline, W. A. 1968. Proc. U. S. Nat. The suborders of perciform Mus., 124(3647):1-78. fishes. Greenwood, P. H., D. E. Rosen, S$. H. Weitzman, and G. S. Myers. 1966. Phyletic studies of Teleos- tean fishes, with a provisional classification of living forms. Bull. Amer. Mus. Nat. Hist., 131: 339-456. Gregory, W. K. 1951. Evolution Emerging. Mac- Millan Company, New York, 1:1—736. Gregory, W. K., and G. M. Conrad. 1937. The comparative osteology of the swordfish (Xip/hias) and the sailfish (/stiophorus). Amer. Mus. Novit., 952:1-25. Howard, J. K., and S. Ueyanagi. 1965. Distribution and relative abundance of billfishes (Istio- phoridae) of the Pacific Ocean. Stud. Trop. Oceanogr. Inst. Mar. Sci., Univ. Miami, 2:1—134. Leriche, M. 1905. Les poissons Eocenes de la Bel- gique. Mém. Mus. Roy. Hist. Belgique, 3:49—228. 1910. Les poissons Oligocenes de la Bel- gique. Mém. Mus. Roy. Hist. Belgique, 5:231— 363. 1942. Contribution a létude des faunes ichthyologique marines des terrains Tertiaires de la plaine cotiere, Atlantique et du centre des Etats Unis. Le synchronisme des formations terti- aires des deux cotés de l’Atlantique. Mém. Soc. Geol. France (N.S.), 20(45):5-110. VOLUME 73 Morrow, J. E., and S. J. Harbo. 1969. A revision of the sailfish genus /stiophorus. Copeia, 1969: 34-44. Nakamura, I., T. Iwai, and K. Matsubara. 1968. A review of sailfish, spearfish and swordfish of the world (in Japanese). Misaki Mar. Biol. Instit., Kyoto Univ. Spec. Rep., 4: 1-95. Regan, C. T. 1909. tion of the scombroid fishes. Hist., Series 3, 8:66—75. On the anatomy and classifica- Ann. Mag. Nat. Robins, C. R., and D. P. de Sylva. 1960. Descrip- tion and relationships of the longbill spearfish, Tetrapturus belone, based on western north Atlantic specimens. Bull. Mar. Sci. Gulf Carib., 10:383-413. 1963. A new western Atlantic spearfish, Tetrapturus pfluegeri, with a redescription of the Mediterranean spearfish, Tetrapturus belone. Bull. Mar. Sci. Gulf Carib., 13:84-122. Romer, A. S. 1966. Vertebrate Paleontology. Chi- cago Univ. Press, Third Ed. 468 pp. Woodward, A. S. 1901. Catalogue of the fossil fishes in the British Museum (Natural History). London, 4: 1-636. 1942. Some new and little known Upper Cretaceous fishes from Mount Lebanon. Ann. Mag. Nat. Hist., Series 11, 9:537—568. Accepted for publication February 16, 1973. EVOLUTIONARY RELATIONSHIPS AND LIZARDS (FAMILY TEIIDAE, SUBFAMILY TEIINAE) WILLIAM Prescu! ABSTRACT: The evolutionary relationships of nine genera of macroteiid lizards, (Ameiva Cnemidophorus, Kentropyx, Teius, Callopistes, Dracaena are discussed based on an analysis of 25 osteological character states. Dicrodon, Crocodilurus, Tupinambis, and A phenetic scheme based on the average linkage technique and an evolutionary sequence diagram are presented. The result places the nine genera into two groups: A, Kentropyx, Teius, and Dicrodon, B, Callopistes, Tupinambis, Crocodilurus, and Dracaena Group B is considered the most derived group of genera; Dracaena being the most derived genus. An analysis of the fossil record of the macroteiids in association with present distributional patterns suggests that the ancestors of the macroteiids arose in North America and entered South America at the close of the Mesozoic Era. With the closure of the Panamanian Seaway in Early Pliocene, movement of Ameiva north and Cnemidophorus south, occurred. Sub- sequent geologic activity and contraction of the tropical climate, are responsible for the Ameiva, Cnemidophorus BIOGEOGRAPHY OF THE MACROTEIID present distribution of the genera. The lizard family Teiidae is composed of approxi- mately 30 genera and 140 species. The group ranges throughout the New World from the north- ern United States, through Mexico and Central America to southern South America: several genera occur in the West Indies. The members of the family are primarily terrestrial, diurnal forms but some are semi-arboral or semi-aquatic: many are semi-fossorial, being snake-like in appearance and habits. The family is comprised mostly of small lizards, the microtetids, but includes several genera of moderately large-sized lizards, the macroteiids; the largest of these, Tupinambis and Dracaena, reach a length of about a meter. Diets range from insectivorous to herbivorous: some are omnivorous and others © strictly carnivorous (Presch, 1970). Food items cover a spectrum including aquatic snails, birds, eggs, and other lizards. The purpose of this present study is to clarify the evolutionary relationships of one segment of the family Teiidae, namely the large teiid lizards Ameiva, Cnemidophorus, Callopistes, Dracaena, Kentropyx, Teius, Dicrodon, Tupinambis, and Crocodilurus. Few attempts have been made to place the various teiid genera into suprageneric groups. The accepted generic arrangement for the family Teiidae comes from the work of Boulenger (1884) in his systematic rearrangement of the Sauria. Prior to his work, the genera now referred to the Teiidae were placed in 27 separate families > aS by various authors (see Boulenger, 1884 for literature synonymy). Boulenger (1885) split the family into four groups based on the condition of the scales on the nasal region. The macroteiids differ from the microteiids (terms first used by Ruibal. 1952) in having the anterior nasal plates not separated by frontonasal scales, limbs well-developed and. as the name implies, being medium to large in size. This division formed Group | Boulenger’s (1885) key and the division has been referred by in recent workers (Uzzell, pers. comm.) as Group I of the Telidae (Ameiva, Callopistes, Cnemidoph- orus, Crocodilurus, Dicrodon, Dracaena, Ken- tropyx, Teius, and Tupinambis). Group II, HI, and IV of Boulenger’s key were distinguished by: anterior nasal plates separated by one or two frontonasal plates. five fingers all clawed: nostril pierced between the nasal and the first labial: no ear opening: nasal plates widely separated by a frontalnasal: ear exposed, inner finger, if distinct, clawless: respectively. These three groups contain the small teiid genera. This division has been referred to as the microteiids or Group II of the Teiidae: a group of small. secre- tive forms, several of which show a reduction in the limbs, climaxing in a limbless condition. This division includes: Alopoglossus, Anadia, Anoto- saura, Argalia, Arthrosaura, Bachia, Cercosaurus. 1Dept. Biological Sciences, California State Uni- versity, Fullerton. California 92634. 24 TaBLe |. Distribution of 21 BULLETIN SOUTHERN CALIFORNIA character states in the Teiinae. ACADEMY OF SCIENCES VOLUME 73 See text for explanation of character state description. , Pterygoid absent(b Premaxillary teeth Postfrontal-Postorbital Pterygoid flange Clavicular shape Clavicular hook present(a present(b absent(a postxiphisternal ribs prescaral vertebrate Second ceratobranchial absent (b Anterior teeth on maxilla absent(b Ectopterygoid contribution to inferior orbital formen concave(a present(a Shape frontal-parietal No. caudal vertebrate roof 60: more(a Supra-angular window present(a) absent(b present(a Condition fifth toe Scapular fenestra Cnemidophorus Kentropyx Dicrodon Teius Callopistes Tupinambis Crocodilurus Dracaena Colobodactylus, Colobosaurus, Echinosaurus, Ec- pleopus, Euspondylus, Gymnophthalmus, Hetero- dactylus, Iphisa, Leposoma, Macropholidus, Microblepharus, Neusticurus, Opipeuter, Panto- dactylus, Pholidobolus, Placosoma, Ptychoglossus, Prionodactylus, Proctoporus, Teuchocercus, and Tretioscincus. Burt and Burt (1931) divided the family into the specialized genera and the lower teiids. The former include: genera with one or more special- izations such as_ strongly compressed teeth (Dicrodon), the loss of digits (Teius and Bachia), the possession of weakened or vestigial limbs and serpentine form (Ophiogonomon, = Bachia, etc.), the loss of eyelids (Gymnophthalmus), the loss of ear openings (Heterodactylus), the separation of the prefrontals (Calliscincopus = Tretioscincus), the complete loss of the prefrontals (Proctoporus, Pholidobolus, etc.), loss of the frontonasal (Ophiognomon = Bachia), enlargement of dorsal granules to scutes or a series of polygons (A nadia, Euspondylus, Iphisa, etc.), keeled ventral scales (Kentropyx, Leposoma), and very small size (Microblepharus, etc.). The remaining genera, the lower teiids, were divided into two subgroups: (1) genera with an essentially superiorly bi- carinate tail; and (2) genera with a regular cyclotetragonal tail. The first group was com- prised of Dracaena, Neusticurus, Echinosaurus, and Crocodilurus. The second group (Callopistes, Tupinambis, and Ameiva) was considered the most primitive of the two and was indicated as the ancestral group from which the specialized teiids were derived. Since Boulenger’s work, there have been few papers on the systematic relationships of the large telids. Cope (1900) stated that his description of the general characters, distributions and systematic arrangement of the genera were based on Boulen- ger (1885). Burt and Burt (1931) attempted to summarize and clarify the generic and specific characters of some of the genera. Vanzolini and Valencia (1965) maintained the division of the family into macroteiids and microteiids, on exter- nal morphology only, and discussed the possible relationships of the genus Dracaena in this frame- work. A number of generic revisions have appeared in the last 55 years for members of Group I (macroteiids) and Group II (microteiids) (Uzzell, 1962, 1965, 1966; Vanzolini and Valencia, 1965; Vanzolini, 196la, 1961b, 1961c, 1969; Ruibal, TABLE 2. 1974 EVOLUTIONARY RELATIONSHIPS OF Characteristics Postfrontal-postorbital Quadrate process of pterygoid Dorsal squamosal process Pterygoid teeth Premaxillary tooth types Clavicular shape Clavicular hooks Interclavicular median process Scapular fenestra Divergent process series of caudal vertebrae or lack of caudal process Number of caudal vertebrae Number of presacral vertebrae and post-xiphisternal ribs Second ceratobranchial ‘Tooth type Supra-angular window Ectopterygoid contribution to inferior orbital foramen Parietal-frontal shape Fifth toe Primitive Condition separate (a) not expanded (b) present (c) present (d) conical (¢) expanded (f) absent (2) long (h) present (i) present (j) more than 60 (k) 26 vert., 12 post- xiphisternal ribs (1) present (m) conical, biconodont triconodont (0) present (n) much (p) concave (q) normal (r) MACROTE|MD LIZARDS 25 Evolutionary changes in certain characteristics of the skull and posteranial skeleton in the ‘Teiinas Derived Condition fused (A) expanded (B) absent (C) absent (D) biconodont, triconodont (EF) rod-like (F) present (Gs) short (H) absent (1) absent (J) less than 60 (K) 25 vert., 11 post-xiphi sternal ribs (L) absent (M) molariform or medially expanded (O) absent (N) little (P) other (Q) reduction in skeletal elements (R) 1952; Duellman and Zweifel, 1962; Peters, 1964: Schmidt, 1957; Burt and Burt, 1931; Barbour, 1915; and Montanucci, 1973). These have pro- vided a starting point for information on the evolutionary relationships of the genera involved. METHODS The study centers on a detailed analysis of the osteology of the nine genera of large teiid lizards. Both the cranial and postcranial skeleton was analyzed. The data on which the following dis- cussion is based was taken from Presch (1970). The reader is referred to this work for details of methodology and detailed descriptions of the osteological elements mentioned below. The dis- cussion is based on 21 osteological characters (Table 1) used by Presch (1970) to define and delimit the nine macroteiid genera. DISCUSSION Distribution of the character states in the nine macroteiid genera is summarized in table lL. Table 2 summarizes the character states and the presumed direction of change or evolutionary trends. Information from these tables is useful in determining the major phenetic patterns. Simi- larity matrices are presented in table 3. Total similarity of shared character states (in roman type) and percentage of shared character states are presented. Table 4 presents a summary of the derived character states, the number and distribution among the nine genera. Based on the percentage of total shared char- acter states, regardless of whether the states are primitive or derived, the following groups are expressed. Ameiva, Cnemidophorus, Dicrodon. Kentropyx, and Teius share the greatest phenetic similarity (77 percent). The greatest similarity with the other genera is 48 percent. Tupinambis, Callopistes, Crocodilurus, and Dracaena have a similarity of 66 percent intragenerically and only 30 percent with the former group of genera. Data in table 2 suggest that the Telinae (Presch, 1970) form two groups on the basis of the combination of shared character states: one group (Tribe Teiini) is comprised of Ameiva. 26 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 TABLe 3. Similarity matrices for Macroteiid genera. Roman type—number of shared character states; /talics— percent of shared character states. 1 2 3 | Ameiva and Cnemidophorus — 20 18 2 Kentropyx 95 — 7 3. Dicrodon 86 8] _ 4 Teius 76 72 90 $ Tupinambis 29 24 33 6 Dracaena 14 10 29 7 Crocodilurus 24 20 29 8 Callopistes 43 48 38 Cnemidophorus, Kentropyx, Teius, and Dicrodon. The second group (Tribe Tupinambini) is com- posed of Tupinambis, Callopistes, Dracaena, and Crocodilurus. The two groups have been defined formally as tribes (Presch, 1970). Ameiva, Cnemidophorus, and Kentropyx share the greatest similarity (95 percent) with each other but only 72 percent and 81 percent with Dicrodon and Teius. The latter two genera share a 90 per- cent similarity. These results indicate Ameiva, Cnemidophorus, and Kentropyx are more closely related to each other than any is to Dicrodon or Teius. In the Tupinambini, Tupinambis, Crocodilurus, and Dracaena share the greatest similarity (76-95 percent) with each other but only 52 and 76 percent with Callopistes. These phenetic relation- ships are illustrated in an average linkage cluster- ing method (Sokal and Sneath, 1963) phenogram (Fig. 1); TaBLeE 4. Summary of derived character states. Genus 4 5 6 7a RE 16 6 3 9 13 15 5 2 4 10 19 7 6 6 8 = 5 6 6 6 24 — 16 20 16 29 76 = 17 11 29 95 80 — iS) In an attempt to establish any evolutionary sequence for the macroteiid genera, weighting the osteological character states as either primitive or derived is necessary. There are several ways in which this has been done in the past (Kluge, 1967; Wake and Ozeti, 1969; Heyer, 1969; and Throckmorton, 1968). The absence of a skeletal element is regarded as a more specialized condition than its presence. The direction of change in serial elements, increase or decrease, may be determined by outgroup com- parison but with less confidence than when dealing with elements that are present or absent. Certain conditions are specialization but it is not always certain that such a specialization is not irreversible. Thus, in the evaluation of characters as primitive or derived, the greatest weight must be given to those characters which involve the complete loss of elements, less weight to those characters which involve some measure of judgment. For clarity, only the derived states (capital letters) are shown in the body of the table. Characters a/A b/B c/C 4/D e/E £/F 2/G h/H i/l j/). k/K 1/L m/M n/N 0/O p/P q/Q 1/R Ameiva and Cnemidophorus Kentropyx A A Teius A A aDwww Dicrodon Tupinambis SE oops) res} Dracaena Callopistes > @. Mion 1 A om yy Crocodilurus lEQGnG@y iG) | © m Q i nea Op BP R i P I L M P I Ki ie aN Ome I L P Q I Ke wes P 1974 Telus Dictodon Kontropys Ameiva Cnomidophoeus — Collopistor Tupinomtvs z Crocodilurus Orocoena 30. 40 50 60 70 B80 90 100 PERCENT SHARED CHARACTER STATE Figure 1, Phenetic relationships of the macroteiid genera. Phenogram constructed by average linkage method. Ideally the fossil history of the group should be employed in the determination of these states. However, all the specimens of fossil macroteiids are fragments of bone, mainly of the lower jaw, anterior tip of the skull, or the posterior portion of the brain case. Furthermore, these fossils are similar to modern forms and do not give any indication of early structural condition for the characters used herein. Eighteen of 25 characters (Table 2) can be evaluated on the bases of the preceding discussion as to their primitive or derived condition. From the list of primitive and derived char- Kentropyx ODicrodon Ameiva / Cnemidophorus i 4 2 ie} \4 12 10 8 6 Figure 2. EVOLUTIONARY RELATIONSHIPS OF Primitive Ancestor MACROTEMD LIZARDS 27 acter states, a hypothetical prototype macroteiid skeleton may be reconstructed using only the primitive condition for each character. By group- ing the derived character states in such a manner as to produce the least number of parallelisms, two groups are produced with a linear phylo- genetic sequence for each group. This is rep- resented in the form of a dendrogram (Fig. 2) which is superimposed on a background of ad- vancement indices. The hypothetical constructed would thus represent the ancestral con- dition of the group prior to the radiation of the modern genera. Comparison of this prototype with modern forms provides no such combination of all primitive characters in either fossil or modern genera. However, all the genera do ex- hibit either the primitive or derived state. There- fore, it seems reasonable to postulate a common ancestor from which two phyletic lines arose. Within Group A (hereafter referred to as the Teiini), Ameiva, Cnemidophorus, and Kentropyx are most similar osteologically. The shape of the macroteid prototype parietal-frontal roof is concave in Ameiva and Cnemidophorus, and convex in Kentropyx. Oste- ologically, there are no major differences. Teius and Dicrodon can easily be separated from Ameiva, Cnemidophorus, and Kentropyx by the increase in the jugal-postorbital-squamosal expanded teeth, lack of suture, transversely 0 2 4 a @ f Phylogenetic sequence of the macroteiid genera superimposed on an index of advancement. Capital letters represent derived character states. pterygoid teeth, and the ventral orientation of the retroarticular process of the mandible (Presch. 1970). reduction in the (Dicrodon with more than 60, Teius with less than Teius is separated from Dicrodon by the number of caudal vertebrae 60), loss of the divergent process series in the caudal vertebral and fusion and re- duction in ankle bones. Callopistes differs from the other Group B members (hereafter referred to as the Tupinam- in combining the of pterygoid teeth with biconodont and triconodont teeth on dentary and maxillary, premaxillary teeth simple in adults; second ceratobranchials present; clavicles sequence bini) presence without hooks; long interclavicular median pro- cess, reaching posteriorly to the level of the third sternal rib. Tupinambis, Crocodilurus, and Dracaena, have lost the second ceratobranchials; loss of pterygoid teeth: a reduction in the length of the inter- clavicular median process to the level midway between the second and third sternal ribs, the loss of biconodont teeth on the dentary and maxil- lary, premaxillary teeth being biconodont in Cro- codilurus and triconodont in Tupinambis. These three genera also share an increase in the ex- pansion of the quadrate in a_posteroanterior direction, beginning with Crocodilurus and Tupin- ambis and reaching its greatest expansion in Dracaena. The clavicles are simple in Tupinambis, with small hooks in Crocodilurus. Dracaena pos- sesses large clavicular hooks and a further re- duction in the interclavicle median process to the level of the second sternal rib. Vanzolini and Valencia (1965) proposed a sim- ilar phylogenetic relationship based on 13 external scale characters. Gorman’s (1970) interpretation based on chromosome morphology proposed a similar division: Group I with 2n = 50 chromo- somes; Cnemidophorus, Kentropyx, Dicrodon, and Teius with the same phylogenetic sequence as I have proposed. However, Group II with 2n = 34-38 chromosomes, though con- taining the same genera as I propose, was arranged in a slightly different phylogenetic sequence. Ameiva, BIOGEOGRAPHY OF THE MACROTEIIDS (SUBFAMILY TEIINAE) The fossil history of the macroteiids dates back to the Late Cretaceous. Estes (1964) described several lizards of modern morphology from the Lance Formation of Wyoming which he referred to the Teiidae. These include Chamops segnis, 28 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 most similar to Crocodilurus and Tupinambis, Meniscognathus ulymani bearing resemblance to the modern Kentropyx. The affinities of two other Leptochamops denticulatus and Hapto- sphenus placodon, to modern forms were unclear. Several vertebrae were also described and _ re- semble those of the modern genera Tupinambis forms, and Crocodilurus. The genus Chamops is considered by Estes (1964) to be closely related to Callopistes. 1 regard Chamops as a valid genus, but believe that it is more closely related to Crocodilurus than to Callopistes. Estes (1964) described Meniscognathus and Leptochamops as being Ken- tropyx- or Ameiva-like. It is difficult to distinguish between Ameiva, Cnemidophorus, and Kentropyx based on the fragments of fossil maxillary and dentary pictured by Estes. However, I agree that it resembles these three modern genera more so than any other macrotelid genera. Haptosphenus is referred to the family Teiidae, but shows a fused splenial, a characteristic of the family Xantusiidae. The teeth are pleurodont in the Xantusiidae but this does not agree with the teeth in Haptosphenus. Estes referred this genus to the Teiidae with the comment that further specimens may show a different familial relation- ship. Estes (1961) described Tupinambis teguixin from the Late Oligocene of Colombia and a new species of Dracaena (Dracaena colombiana) from the Late Miocene of Colombia. I have studied his diagrams and agree with his allocation of the specimens to modern genera. There is no doubt that Dracaena and Tupinambis were present in the Oligocene of South America. Estes and Tihen (1964) reported specimens of the genus Cnemi- dophorus from the Valentine Formation, Miocene- Pliocene boundary of Nebraska. Etheridge (1964) described a fossil dentary from the Late Pleisto- cene of Barbuda, British West Indies, referring it to Ameiva griswaldi. Ameiva chrysolaema and A. taeniura were reported from the Late Pleisto- cene of the Dominican Republic by Etheridge (1965) and A. thoracica was reported from the Pleistocene of New Providence (Ethridge, 1965b). Gilmore (1940) described two new genera and species of lizards, Polyglyphanodon sternbergi and Paraglyphanodon utahensis, from the Upper Cretaceous of South Dragon, Utah. After de- scribing the osteology of Polyglyphanodon stern- bergi, in detail, Gilmore (1942) erected a new family, the Polyglyphanodontidae, recognizing only that it was not related to either the [guanidae 1974 EVOLUTIONARY or Agamidae, and placed it in the Ascalabota division of the Sauria. In 1943 Gilmore described a new species, Paraglyphanodon gazini and placed the genus in the family Polyglyphanodontidae. Hoffstetter (1955) recognized Polyglyphanodon sternbergi as belonging to the family Teiidae. The genus Paraglyphanodon is currently under study but it probably belongs to the family Teiidae (Estes, pers. comm.). Estes (1969) described a new telid genus and species, Peneteius aquilonius, from the Cretaceous Hell Creek Formation, Montana. He believed it occupied a position midway between Polyglyphanodon and the modern genera Teius and Dicrodon. Late The presence of Tupinambis, Teius, Ameiva, and Cnemidophorus in the Oligocene and Draca- ena in the Miocene indicates an early origin for the macroteiid genera. It seems likely that the modernization of this group of genera took place not later than the Oligocene and quite probably much earlier. The presence in the late Cretaceous of Meniscognathus, a torm that Estes considers to be Ameiva-Kentropyx-like indicates that the ancestor for the Teiini were present by the middle or late Mesozoic. The distribution of the macroteiid restricted to South America with the exception of Ameiva and Cnemidophorus. It seems likely that the ancestor of the modern macroteiids reached South America sometime late in the Cretaceous or early Paleocene across the Pana- manian land-bridge (Savage, 1966) and under- went independent evolution during the period of isolation in South America from the late Paleocene to the early Pliocene. The ancestors of modern Ameiva-Cnemidoph- orus appears to have evolved in isolation in two areas north and south of the Panamanian portal throughout middle Cenozoic (Eocene-Miocene). Both diverged to a level of generic differentiation and some northern forms seem to have reached South America and several South American forms, Middle America, after the land connection was re-established in the Pliocene (Savage, 1966). In summary the family Teiidae is a strictly New World family, originating in the tropical to warm temperate areas of North America. The modern macroteiids are restricted to South America and genera is probably arose in South America in the early Tertiary. Cnemidophorus and Ameiva appear to have been derived from a common ancestor, and evolved in North America and South America, in isolation through much of the Cenozoic. RELATIONSHIPS OF MACROTEMD LIZARDS y rhe history of the macrotetid genera seems te parallel the development of the marsupials, cavio morph rodents, and primates in South America (Savage, 1973), The geological history of the South American land mass has played an important role in the present distribution of the macroteiid genera There is some discussion as to the exact time of separation of the North American land mass from South (1972) evidence of both Simpson (1940) and Hershkovitz (1969), reached the same conclusion as he did in America. Savage reviewed the and 1966 using mammalian distributions, based on the analysis of the Central herpetofauna of America. The landbridge between Cen- tral America and South America was submerged from the late Paleocene to the early Pliocene However, as pointed out above, the ancestors of the macroteiids were present by the close of the Mesozoic and probably crossed into South Amer- ica prior to the formation of the Panamanian Seaway. Those events which have shaped the present distribution of the macroteiids must then be restricted to the events which occurred from the beginning of the Cenozoic to the present time in South America. There are no complete studies on the geological history of South America, but there are several papers which review the major events that took place during the Cenozoic period (Kummel, 1961: Harrington, 1962: Lloyd, 1963). climatic changes during this time are generally known and summarized by Axelrod (1960). Though there are a great number of events which shaped the distribution of the macroteiids in South America, the most important geological occurrences are: 1) the uplift and formation of the Andean Mountain Range, beginning in the Paleocene, and ending in the Pleistocene, 2) the extensive flooding of vast areas of parts of the Amazonian, Parnaiba and San Francisco Basins and the extreme stability of the Guiana, Central Brazilian and Coastal Brazilian Shields (Harring- ton, 1962). The uplift and formation of the Andean moun- tain chain began in the Paleocene and lasted until the late Pleistocene (Kummel, 1961). The great- est uplift occurred during the late Pliocene or early (Haffer, 1967: 1962). The sea advanced several times during the Likewise. the Pleistocene Harrington, Cenozoic into the Amazonian. San Francisco Basins, and the southern Patagonian Basin, as well as areas in what is today northern Colombia and Venezuela. 30 BULLETIN SOUTHERN CALIFORNIA During this period of geological compression, folding and uplift, two areas apparently were little affected; the Guiana and greater Brazilian Shields. Harrington (1962), pointed out the stability of these shields, dating back to early Precambrian. These areas were not subject to major geological activity, nor were they ever flooded. They were relatively stable, geologically and sprobably places of relative environmental stability. The climate during the Cenozoic of South America has been summarized by Axelrod (1960) by inference from known Cenozoic fossil floras. The tropical and subtropical flora extended south to a latitude of 45°-50°. In response to lower temperature during the Eocene, the tropical cli- matic belt was reduced as indicated by the re- duction of the Neotropical Geoflora (Axelrod, 1960). During the middle to late Tertiary, dry climates developed along the east and west coast of South America, gradually confining the tropical flora to its present region (Axelrod, 1960). The fossil history of the macrotetids as outlined above, along with the extent of the tropical floras, indicates that this group has always been con- fined to areas of tropical to warm-temperate climate. This association with climate is repre- sented today with the exception of some of the northern species of Cnemidophorus. Callopistes and Dicrodon are dry-adapted groups which probably became restricted to the west coast of South America in the Late Pliocene when the main uplift of the Andes took place. Teius is found in an area of South America which has remained relatively stable and dry since the Late Oligocene. Its arrival in southern South America probably dates to the early Cenozoic and has remained there undergoing little change. Ameiva and Kentropyx today occupy a vast range in South America and are composed of about 32 and 7 species, respectively. It seems most probable that, during the entire Cenozoic, these two groups were repeatedly broken into fragmentary populations by the many geological events which occurred, undergoing speciation and expanding their ranges as conditions permitted. Throughout the Cenozoic, northwestern Colom- bia was repeatedly flooded, causing an extremely unstable environment, hostile to the survival of the macroteiids. With the closure of the last portion of the Panamanian Portal, in the early Pliocene, faunal interchange between North and South America became possible. Cnemidophorus, which evolved in North America, was able to move into South America, while South American ACADEMY OF SCIENCES VOLUME 73 forms of Ameiva were able to move north into Central America and Mexico. Cnemidophorus maintained its distribution in North America and was able to adapt to the change in climatic con- ditions (Axelrod, 1948) which occurred through- out the Cenozoic. Tupinambis and Dracaena are both humid- tropical forms. Dracaena is particularly abundant in tidal and marshy sections of the Amazon Valley and in the drier ground on the fringes of the swamps in southern Brazil. Tupinambis occurs over much of the forested areas of tropical South America. Fields (1957, 1959) has described the northern section of Colombia during the Miocene, from which Dracaena and Tupinambis are known, as being a wet-tropical climate. Fields describes part of this region today as having a semi-arid climate, a condition unfavorable to Tupinambis and Dracaena. Crocodilurus is also a wet-tropical form, in association with small, slow- moving rivers (Goeldi, 1902). It appears that these three genera occupied the wet-tropical areas of South America in the early Tertiary, but as the climatic conditions changed in the middle to late Tertiary these forms followed the contraction of the tropical flora and climate, their habitat being restricted both by the Andean uplift and, to a great extent, by the increasing aridity. found MATERIAL EXAMINED The following species of teiid lizards were studied from radiographs, as dry skeletons, or both. Each species is followed by the number of specimens examined. Teiinae: Ameiva: ameiva (12), auberi (2), bifrontata (9), bridgei (1), chrysolaema (5), dorsalis (2), erythro- cephals (2), exsul (2), festiva (22), fuscata (2), griswoldi (2), leptophrys (4), lineolata (4), maynardi (2), pleii (2), pluvianotata (2), polops (2), quadri- lineata (7), septemilineata (2), taeniura (4), undulata (12), wetmorei (4). Callopistes: flavipunctatus (9), maculatus (13). Cnemidophorus: angusticeps (3), burti (2), cali- dipes (2), ceralbensis (2), communis (1), costatus (13), cozumela (2), deppi (10), exsanguis (3), gularis (2), guttatus (3), hyperythrus (5), inornatus (2), labialis (1), lacertoides (1), lemniscatus (31), lineatissimus (2), maximus (1), motaguae (4), muri- nus (10), neomexicanus (3), ocellifer (2), sacki (1), septemvittatus (5), sexlineatus (2), sonorae (3), tigris (13), uniparens (2), velox (2), vanzoi (6). Crocodilurus: lacertinus (8). 1974 EVOLUTIONARY Dicrodon: guttulatum (20), heterolepis (3), holm- hergi (3). Dracaena: guianensis (12), paraguayensis (4). Kentropyx; altamazonicus (1), calcaratus (21), pelviceps (7), striatus (15), viridistriga (2), william- yoni (1). Teius: feyou (11). Tupinambis: rufescens (52), feguixin (55). LITERATURE CITED Axelrod, D. I. 1948. Tertiary Geoflora. Madro- 24(7) :433-—509, Evolution of the Bot. Rev., ——.. 1960. The evolution of flowering plants. Pp. 227-305 in The Evolution of Life. (Tax, ed.) Univ. Chicago Press, Vol. 1, viii -- 629 pp. Boulenger, G. A. 1884. of existing Lacertilia. (5) XIV:117-122. Synopsis of the families Ann. Mag. Nat. Hist., 1885. British Museum. Catalogue of the lizards in the 2nd Ed. Vol. I1:330-429. 1894. List of reptiles and batrachians col- lected by Dr. J. Bohls near Asuncion, Paraguay. Ann. Mag. Nat. Hist., 13(6):342-348. Burt, C. E. 1931. A study of the teiid lizards of the genus Cnemidophorus with special reference to their phylogenetic relationships. Bull. U.S. Nat. Mus., 154:1-286. Burt, C., and M. D. Burt. 1931. South American lizards in the collection of The American Museum of Natural History. Bull. Amer. Mus. Nat. Hist., 61(7):227-395. Cope, E. D. 1900. The crocodilians, lizards and snakes of North America. Ann. Rept. U.S. Nat. Mus., 1898 (1900) part 2:151—1294. Duellman, W. W., and R. G. Zweifel. 1962. A synopsis of the lizards of the sexlineatus group (genus Cnemidophorus). Bull. Amer. Mus. Nat. Hist., 123(3):159-210. Estes, R. 1961. South America. Miocene lizards from Colombia, Breviora, 143:1-11. 1964. Fossil vertebrates from the late Cretaceous Lance Formation Eastern Wyoming. Univ. California Publ. Geol. Sci., 49:1-180. 1969. lizards (Sauria: Relationships of two Cretaceous Teiidae). Breviora, 317:1-8. RELATIONSHIPS OF MACROTLHD LIZARDS if] Estes, R., and J. Tihen. 1964, Lower vertebrate from the Valentine Formation of Nebraska Amer. Mid]. Nat., 72:453-472 Etheridge, R. 19654. Pleistocene lizards from New Providence. Quart. J. Florida Acad. Sci., 2% 349-358, — 1965b. Fossil lizards from the Dominican Republic. Quart. J. Florida Acad. Sci., 28 83-105. Fields, R. W. 1957. the late Miocene of Colombia, South America Univ. California Publ. Sci., 32:273-404. Hystricomorph rodents from Geol. ——. 1959. Geology of the La Venta badlands, Colombia, South Univ. California Publ. Geol. Sci., America. 36:405-449. Gilmore, C. W. 1940. New fossil lizards from the Upper Cretaceous of Utah. Smithson. Misc. Coll, 99(16):1-3. —. 1942. Osteology of Polyglyphanodon, an upper Cretaceous lizard from Utah. Proc. US. Nat. Mus., 92(3148):229-268. 1943. Osteology of upper Cretaceous lizards from Utah, with a description of a new species. Proc. U.S. Nat. Mus., 93(3163):209- 214. Goeldi, E. A. 1902. Lagartos do Brazil (Capitulo do Monographia inedita “Reptils do Brazil”). Bol. Mus. Paraense, 3(3 and 4):499-S60. Gorman, G. 1970. Chromosomes and the system- atics of the family Teiidae (Sauria, Reptilia). Copeia, 1970:230-245. Haffer, J. 1967. Speciation in Colombian forest birds west of the Andes. Amer. Mus. Novit.. 2245:1-57. Harrington, H. J. 1962. Paleogeographic develop- ment of South America. Bull. Amer. Assoc. Petro. Geol.. 46(10):1773-1814. Hershkovitz. P. 1966. Mice, landbridges and Latin America faunal interchange. Pp. 725-751 in Ectoparasites of Panama. (R. Wentzel and V. T. Tipton eds.) Field Mus. Nat. Hist., vii + 861 pp. 1969. The evolution of mammals on southern continents. VI. The recent mammals of the Neotropical Region: a zoogeographic and ecologic review. Quart. Rev. Biol., 44:1-70. Nh Heyer, W. R. frog 1969. Biosystematic studies on the Leptodactylus. Unpublished Ph.D. Southern California. genus dissertation, Univ. Hoffstetter, R. 1955. Pp. 606-662 in (Masson et Cie eds.) Paris, Vol. 5., Squamates de type modern. Traite de paleont. 1113 pp. Piveteau, 1967. lizards and Nat. Hist., Kluge, A. G. gekkonoid Amer. Mus. Higher taxonomic categories of their evolution. Bull. 135(1): 1-60. Kummel, B. 1961. duction to Historical Geology. and Co., xii + 610 pp. History of the Earth: an Intro- W. H. Freeman Lloyd, J. J. 1963. Tectonic history of the south Central-America orgon. Mem. Amer. Assoc. Petro. Geol., 2:88—100. Montanucci, R. 1973. Systematics and evolution of the Andean lizard genus Pholidobolus (Sauria: Teiidae). Univ. Kansas Mus. Nat. Hist., Misc. Publ, 5921-52. Presch, W. 1970. The evolution of Macroteiid lizards (Family Tetidae): An osteological inter- pretation. Unpublished Ph.D. dissertation, Univ. Southern California. Ruibal, R. 1952. Revisionary South American Telidae. Zool., 106(11):477—529. studies of Bull. Mus. some Comp. Savage, J. M. 1966. The origins and history of the Central American herpetofauna. Copeia, 1966: 719-766. 1973. The isthmian link and the evolution of Neotropical mammals. Contrib. L. A. County Mus. Nat. Hist. In press. Simpson, G. G. 1950. Latin America. History of the fauna of Amer. Sci., 38:361-389. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Sokal, R. R., and P. H. A. Sneath. 1963. The principles of numerical taxonomy. Freeman, San Francisco., xi + 359. Throckmorton, L. H. 1968. Concordance and dis- cordance of taxonomic character in Drosophila classification. Syst. Zool., 17:355—387. Uzzell, T. 1962. Additional notes on teiid lizards of the genus Placosoma. Copeia, 1962:833-835. 1965. Teiid lizards of the genus Echino- Copeia, 1965:82-89. SauUra. 1966. Teiid lizards of the genus Neusti- curus (Reptilia, Sauria). Bull. Amer. Mus. Nat. Hist., 132(5):279-327. Vanzolini, P. E. 196la. Bachia: Especies Brasil- eiras E Conceito generico (Sauria, Teiidae). Pupeis Avulsos Dept. Zool., Sao Paulo, 14(22): 193-209. 1961b. Notas bionomicas sobre Dracaena guianensis no Paia (Sauria, Teiidae). Pupeis Avulsos Dept. Zool., Sao Paulo, 14(25):237-241. Vanzolini, P. E., and J. Valencia. 1965. The genus Dracaena, with a brief consideration of macro- teiid relationships (Sauria, Teiidae). Arquivos de Zool., 13:7—25. Vanzolini, P. E., and R. Reboucas-Spielez. 1969. On a large and surprising sample of Calliscin- copus aglis from Brazil with the invalidation of the genus (Sauria, Tetidae). Pupeis Avulsos Dept. Zool., Sao Paulo, 22(3):123-144. Wake, D. B., and Necla Ozeti. 1969. relationships in the family Salamandridae. peia, 1969:124—137. Evolutionary Co- Accepted for publication October 1, 1973. PARASITIC BOPYRID ISOPODS OF THE STEGOPHRYXUS THOMPSON WITH THE AMPHI-AMERICAN GENUS DESCRIPTION OF A NEW SPECIES FROM CALIFORNIA JOHN C, ABSTRACT: Stegophryxus liyptins, MARKHAM! heretofore longicarpus in New England, is recorded infesting ?. longicarpus in Georgia, P. annulipes North Carolina and Georgia, and P. bonairensis and P. miamensis in southern Florida known only as a. parasite of Pagurus The only known specimens of Stegophryxus thompsoni, the types, from an unknown host in Chile are redescribed. A new species from California and Baja California, S. liyphalus, is as a parasite of Parapagurodes laurentae and P. makaroyi. described These three species, the only members of the genus, are illustrated in detail and their distribution discussed Examination of shallow-water hermit crabs col- lected along the Atlantic coast from Florida to North Carolina has turned up several specimens of the athelgine bopyrid parasite Sregophryxus hyptius Thompson, previously known only from New England, thereby providing several new host records and a considerable extension of range for this species. Reexamination of the types of S. thompsoni Nierstrasz and Brender a Brandis from Chile, which had not been adequately described, became possible. Numerous parasites of hermit crabs taken off the coast of California and Baja California proved to belong to a new species. With these three species of Stegophryxus de- scribed, it became possible to review the genus in detail. The material examined was in or has been donated to the following museums: Allan Han- cock Foundation, University of Southern Cali- fornia (AHF); Rijksmuseum van Natuurlijke Historie, Leiden (RMNHL); Rosenstiel School of Marine and Atmospheric Science, University of Miami (UMML); United States National Museum of Natural History, Smithsonian — Institution (USNM); and Universitetets Zoologiske Museum, Copenhagen (ZMC). Stegophryxus Thompson 1902 Type-species, by original designation, Stegophryxus hyptius Thompson. Stegophryxus hyptius Thompson 1902 Figures 1-3 Stegophryxus hyptius Thompson, 1902, pp. 53-56, pls. 9, 10.—Richardson, 1904, pp. 59-60: 1905, pp. 532-535, figs. 578, 579.—Rathbun, 1905, p. 48.— ts Uy Sumner ef al., 1913a, p. 136; 1913b, p. 661 Kunkel, 1918, p. 236.—Reinhard ef al., 1947, pp 70-72.—Reinhard, 1949, pp. 17-31: 1956, p. 101 Reinhard and Buckeridge, 1950, p. 131.—Caullery, 1950, p. 97; 1952, p. 76.—Reverberi, 1952, p 292.—von Brand, 1952, pp. 256, 271, tab. 41; 1966, Noble, 1964, pp. 392, 393, figs. XVI-SA. SB, SC Smith, 1964, p. 105.—Kaestner. 1970, p. 463. Bourdon, 1968, p. 133.—Schultz, 1969, pp. 321 322, fig. 513 —Markham, 1972, p. 73. Stegophryxus hyptias—Miner, 1950, pp. 450. 453. pl. 145. Stegophrixus hyptius.—Nierstrasz and Brender a Brandis, 1931, pp. 197-198. Stegophryxus.—Baffoni, 1953, 1956, p. 93.—Kaestner, 1967, 425, 463. 447.—Reinhard. 1161; 1970, pp Pp. Pp. Material examined.—Infesting Pagurus bonairensis Schmitt. Bear Cut, Virginia Key, Miami, Florida, 9 February 1958, A. J. Provenzano, coll., 12. 142 USNM 101762. In Thalassia bed, shallow water, NW side Virginia Key, Miami, Florida. March 1969. J. C. Markham, coll... 12.1¢ (reference 4), USNM. Same locality, 8 November 1972, E. B. Hatfield, coll.. 25 19,14, AHF. Same locality, 12 February 1973, E. B. Hatfield. coll.. 192 (reference 2), 1¢, USNM. Shallow subtidal. off W. Arsenicker Key. Card Sound, Florida. 25 October 1972, E. B. Hatfield. coll. 12,1¢, UMML 32.4538. Infesting Thalassia bed, shallow water, NW Miami, Florida, 28 June 1972. E. P. A. McLaughlin. identification UMML 32.4539. Infesting Pagurus longicarpus Say. Tidepool. Sapelo Provenzano. In side Virginia Key. B. Hatfield. coll.. of 12. 124 Pagurus miamensis host. of Marine and Atmospheric 10 Rickenbacker Causeway. 1 Rosenstiel School Science, Univ. Miami. Miami, Florida 33149. 34 BULLETIN SOUTHERN CALIFORNIA Stegophryxus hyptius Thompson, reference A. Dorsal view. B. Ventral view. C. Left D. Right posterior ventral border of head. E. Right oostegite 1, interior. F. Same, exterior. G. Right pereopod 1. H. Left pereopod 7. I. Pleon, dorsal view. Scale indicates 1.0mm for A, B, 0.5mm for D-F, I, 0.2mm for C, G, H. Figure |. female. antenna 2. Beach, Georgia, August 1969. R. W. Heard, coll., one immature 9, 14, USNM. Woods Hole, Mas- sachusetts, date and coll. unknown, M. J. Rathbun, identification of host, 19, 1¢, USNM 54777. Infesting Pagurus annulipes Stimpson, all hosts identified by P. A. McLaughlin. Morehead Channel, North Carolina, 3 March 1967, R. W. Heard, coll., 392, 34, four cryptoniscan larvae, ZMC. Same lo- cality, 25 June 1970, R. W. Heard, coll., 42, 44, Figure 2. Stegophryxus hyptius Thompson, A-G, reference male. H. Second male. A. Dorsal view. B. Lateral view. C. Ventral view. D. Right antenna 2. E. Left antenna 1. F. Right pereopod 1. G. Left pereopod 7. H. Pleon and last pereomere, dorsal view. Scale at B, indicates 0.2mm for A-—C, 0.1mm _ for D-G; scale at H. indicates 0.1 mm for H. ACADEMY OF SCIENCES VOLUME 73 Figure 3. mature female, ventral view. B-—F, cryptoniscan larva. B. Dorsal view. C. Ventral view. D. Left antennae. E. Left pereopod 1. F. Left pereopod 7. Scale in- dicates 1.0mm for A, 0.1mm for B, C, 0.05mm for D-F. Stegophryxus hyptius Thompson. A. Im- two cryptoniscan larvae, RMNHL. Lacking collection data (probably North Carolina), 192, UMML 32.4540. Beaufort, North Carolina, 1969 or 1970, C. Kellogg, coll., 12, 1¢, AHF. Same, 12,14, both damaged, UMML 32.4541. Bulkhead Channel, Beau- fort Harbor, North Carolina, 22 June 1970, C. Kel- logg, coll., 19, 1¢, AHF. Same, 19, 1¢, UMML 32.4542. Beaufort Harbor, North Carolina, 1969 or 1970, C. Kellogg, coll., | damaged 9, AHF. Remarks.—Thompson (1902) presented a de- tailed description and excellent drawings of both sexes, an immature female and a cryptoniscan larva of Stegophryxus hyptius. Richardson (1905) reproduced Thompson’s descriptions and draw- ings of the adults. There is thus no need to re- describe this species in detail, and a diagnosis should suffice. For comparison, I have presented drawings of both sexes collected in Florida (Figs. 1, 2), an immature female from Georgia (Fig. 3A) and more detailed drawings of a cryptoniscan from North Carolina (Figs. 3B—-F). The characters which distinguish the females of S. hyptius (Fig. 1) are a head twice its width, a first oostegite which is bluntly pointed, a highly asymmetrical brood pouch which extends far back on the right side, a pleon which is less than half the total body length and bears very narrow lateral plates and pleopodal rami and enlarged prominent uropods. Males of S. hyptius (Fig. 2) are distinc- tive in having conspicuous eyes, the head clearly 1974 separated from the pereon, and the pleon markedly narrower than the last pereomere, roughly triangu- lar and ending in a rounded point. The immature female of S. hyptius can be distinguished by its lack of coxal plates, its clongate pleomeres and very prominent uropods. Two of the host Pagurus annulipes, in addition to bearing mature pairs of Sfegophryxus hyptius on the abdomen, also bore cryptoniscan larvae on the thorax. On one P. annulipes there were two cryptoniscans, one attached to the carapace, the other to a pereopod; on the other P. annulipes, three cryptoniscans were clinging to the carapace and a fourth was inside the branchial chamber. These larvae, one of which is pictured (Figs. 3B-F), agree in all respects with those which Thompson (1902) described, though he found them with the male in the brood chamber of the female Stegophryxus rather than on the host’s body. I have sent some of these larvae to Jarl-Ove Stromberg of the University of Lund, who plans to examine them with a scanning electron microscope. One pair of Stegophryxus hyptius from Pagurus bonairensis was examined while still alive. The female was clinging so tightly to its host’s pleopods that one pleopod was torn off in removing the parasite. Its only movement was a constant beat- ing of the anterior oostegites to aerate the larvae filling the closed brood pouch. The male, rather than being in the female’s brood pouch, as is typical, was crawling all over its body. The color of the female was transparent to whitish except that dorsally the gut was red-orange and_ the ovaries yellowish with developing eggs, and there were a few chalky white chromatophores scattered along the sides. The male was transparent except for black eyespots, a brownish gut and a network of interconnecting white lines in all pereomeres and the posterior third of the pleon. A second pair Was examined freshly preserved. The female had a uniform glossy white background with brick- red eyes, a pale yellow central dorsal region on the pereon and a few chalky white chromatophores scattered near the edges of the pereon; develop- ing embryos visible through the oostegites bore rows of punctate red chromatophores. The ac- companying male was also glossy white with black even eyespots, an orange gut and scattered black and white chromatophores. Previous records of S. hyptius are only from Rhode Island and Woods Hole, Massachusetts. At the latter locality it is so common that Reinhard (1949) used it in a classical study to demonstrate BOPYRID PARASITES AND NEW SPECIES FROM ¢ ALIFORNIA sex determination in the Bopyridae, and Reinhar et al, (1947) made important discoveries about the effects it had on its host, which was alw Pagurus longicarpus. Vhe new records thus add a considerable extension of range, down to south ern Florida, and several new host records, all species of Pagurus. Of the parasites examined, 17 were still with their hosts. Five of these hosts were males, 12 females, the females being hosts significantly more often. Stegophryxus thompsoni Nierstrasz Brender a Brandis 1931 Figures 4—5 and Nierstrasz and Brender a 198, figs. 87-89 Stegophrixus thompsoni Brandis, 1931, pp. 196 Material examined.—From unknown hermit crab Valparaiso, Chile, exact locality and date unknown, Th. Mortensen Pacific Expedition, coll. Holotype allotype ¢, both damaged, ZMC. Redescription of holotypic female (Fig. 4).—Body length 8.0mm, maximal body width 3.7mm, head length 1.6mm, pleon length 3.8mm, distortion of body axis 50 degrees. line uncertain (Fig. 4A). Head antenna | Body badly damaged. so out- twice as wide. Segmentation of (Fig. 4B) of five long as uncertain, antenna 2 Figure 4, thompsoni Nierstrasz and Brender 4 Brandis. holotype female. A. Dorsal view. B. Right antenna 2. C. Right maxilliped. D. Right posterior ventral border of head. E. Right oostegite 1. interior. F. Same, exterior. G. Left pereopod 2. H. Right pereopod 6. I. Pleon, ventral view. Scale indicates 1.0mm for A, C. E, F, I, 0.4mm for B. 0.25mm for D, 0.5mm for G, H. Stegophryxus 36 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES ’ al C Stegophryxus thompsoni Nierstrasz and A. Dorsal view. D. Left antenna Figure 5. Brender a Brandis, allotype male. B. Ventral view. C. Left antenna 1. 2. E. Right pereopod 1. F. Right pereopod 7. Scale indicates 1.0mm for A, B, 0.2mm for C-—F. segments. No eyes. Maxilliped (Fig. 4C) with very irregular outline. Posterior ventral border of head (Fig. 4D) with long pointed lateral projection, ir- regular blunt process medial to it. Pereon surrounding head, first two pereomeres obliterated centrally, first five surrounding head, sixth longest. First oostegites (Figs. 4E, F) rounded anteriorly, with dentate internal ridge, sharply angled falcate posterolateral point. Most of other oostegites damaged or missing, so size, shape and position un- known. First five pereopods with prominent coxal plates, sharply reflexed, reduced in size (Fig. 4G); last two pereopods lacking coxal plates, more or less straight, larger (Fig. 4H). Last two pereomeres (Fig. 41) with medial ventral tubercles near anterior mar- gins. Pleon nearly as long as pereon, gradually tapering posteriorly, of six pleomeres. Lateral plates and pleopodal rami (Fig. 41) all about the same size, ovate in outline; evidently pleopods and lateral plates lost from fourth and fifth pleomeres by damage. Uropods uniramous, oval, each nearly as large as whole last pleomere, extending far posteriorly from that pleomere. Tiny posterior point between uropods. VOLUME 73 Redescription of allotypic male (Fig. 5)—Body length 2.3mm, maximal body width 0.8mm, head length 0.3mm, pleon length 0.6mm. Sides of body nearly parallel, all segments clearly separated (Figs. SA; B). Head semicircular, narrower than pereon. No eyes. Antennae (Figs. SC, D) of three and five seg- ments, respectively. Pereon broadest, but only slightly so, across fourth pereomere. All pereomeres separated by slight antero- lateral indentations. Pereopods (Figs. SE, F) gradu- ally smaller posteriorly. Pleon nearly as broad as last pereomere, barely longer than wide, broadly oval in outline, greatest width about 1% of distance back from anterior edge. No indication of segmentation. The male has broken into two pieces between the third and fourth pereomeres, and one side of the pleon is damaged, so the drawings presented are partial reconstructions. Stegophryxus hyphalus, new species Figures 6—8 ?Stegophryxus sp.—Menzies and Miller, 1954, pp. 141, 153. Stegophryxus n. sp.—McLaughlin and Haig, 1973, pp. 119, 134-135. Material examined.—Infesting Parapagurodes lau- rentae McLaughlin and Haig (part of type series) collected by Allan Hancock Foundation Velero III among Santa Barbara Islands, California, and near Abrejos Pt., Baja California. Sta. 914-39, 32°47’N, 118°22’W, 143-319m, 19 February 1939, 19, 1é, AHF; Sta. 981-39, 33°35’-35°08’N, 119°01’W, 139- 159m, 29 May 1939, 19, 16, AHF; Sta. 1012-39, 32°46’N, 118°25’W, 100-126m, 9 November 1939, one immature 9, AHF; Sta. 1028-39, 33°18’N, 118° 16’W, 152—229m, 10 December 1939, 12, 16, ZMC: Sta. 1239-41, 33°01’N, 118°33’W, 95-112m, 22 February 1941, 19, AHF; Sta. 1429-41, 33°17’N, 118°16’W, 159-174m, 25 October 1941, 592, 14, AHF; Sta. 1710-49, 26°17’N, 113°41’W, 99m, 7 March 1949, one immature 9, USNM; Sta. 1937-50, 34°02’N, 119°27’W, 69-79m, 24 March 1950, one immature 9, ZMC. Infesting Parapagurodes makarovi McLaughlin and Haig (part of type series) collected by U.S. Bureau of Fisheries Steamer Albatross. Sta. 2946, in Santa Barbara Islands, California, 33°58’N, 119°31’W, 274m, 7 February 1889, 19, USNM. Sta. 3184, off Carmel, California, 36°27’N, 122°00’W, 141m, 3 April 1890, 12, 14, USNM. Collected by Velero III among Santa Barbara Islands, California, and near San Benito Islands, Baja California. Sta. 911-39, 39°O00’N, 118°33’W, 110-155m, 18 February 1939, 12, 18, AHF; Sta. 981-39, 33°35’N, 119°01’W, 139-159m, 29 May 1939, 79 (including one imma- ture), 34, AHF; Sta. 983-39, 33°44’N, 119°10’W, Figure 6. view. C. Left antennae. Stegophryxus hyphalus, new species, holotype female. A. Dorsal view. D. Posterior ventral border of head. BOPYRID PARASITES AND NEW SPECIES FROM CALIFORNIA B. Ventral E. Right oostegite 1. exterior. F. Same, interior. G. Left pereopod 1. H. Left pereopod 6. Scale indicates 2.0mm for A. B. E, F, 1.0mm for D, 0.5mm for C, G, H. 128m, 29 May 1939, 19, AHF; Sta. 1018-39, 33° OLN, 118°32’W, 110-247m, 23 November 1939, one immature 2, RMNHL; Sta. 1020-39, 33°03’N, 118° 40’W, 247-274m, 24 November 1939, 29, 1¢. AHF: Sta. 1026-39, 33°00’N, 118°38’W, 216-384m, 9 De- cember 1939, holotype 9, AHF 3928, allotype ¢. AHF 3928a, two other 2, one other ¢, AHF; Sta. 1119-40, 28°13’N, 115°34’W, 159-174m, 19 Feb- ruary 1940, 12, UMML 32.4543; Sta. 1173-40, 33° 17’N, 118°14’W, 197-201m, 20 August 1940, 19, AHF; Sta. 1393-41, 33°47’N, 119°59’W, 196—-229m, 26 August 1941, 19, 14, RMNHL. Collected by Velero IV near Santa Barbara Island, Sta. 2062-51. 33°33’N—34°33’N, 119°04’W, 256-296m, 18 October 1951, 12, 16, UMML 32.4544. Description of holotypic female (Fig. 6)—\Body ~ length 6.1mm, maximal body width 3.2mm. head length 1.2mm, pleon length 2.3mm, distortion of body axis 42 degrees. Body nearly oval in outline (Figs 6A, B). Head roughly cordate in shape. about 1.5 times as long as broad. Antennae I and 2 (Fig. 6C) of 3 and 5 segments. respectively: each segment smaller than that proximal to it. No eyes. Posterior ventral border of head (Fig. 6D) with raised central portion consisting of a blunt medial point and two laterally directed narrow rounded points: each side bearing a long tapering point laterally with irregularly shaped blunt projection medial to it. Pereon longer than broad. First five pereomeres concave anteriorly, more or less concentrically sur- rounding head: first two pereomeres incomplete in 38 BULLETIN SOUTHERN CALIFORNIA af Q \ J Aa am { ( \s- REX (4 yA ec ) | a } f PY LY IIE Stegophryxus hyphalus, new species, im- A. Dorsal view. B. Ventral view. Figure 7. mature female. 1.0mm _ indicated. center: last two pereomeres concave posteriorly, nar- rower than others; sixth pereomere by far the longest. First oostegites (Figs. 6E, F) rounded anteriorly, without conspicuous internal ridge in circular middle region, produced into broadly falcate posterolateral point; first two pairs of oostegites arching over head; fifth oostegites by far the largest, covering half of ventral surface, both produced into posterolateral pockets of equal length, posterolateral borders lined with several long setae. Pereopods of first five pairs (Fig. 6G) surrounding head, sharply reflexed; last two pereopods (Fig. 6H) far posterior, longer and straighter than others. Pleon about *5 as long as pereon, of six pleomeres. Numerous tubercles on ventral surface of first two pleomeres. Sides of pleomeres 1—5 produced into foliaceous lanceolate lateral plates identical to branches of biramous pleopods. Final pleomere bear- ing small bulbous uniramous uropods. Variations: Some of the other females examined show a few slight variations from the type. In several cases, the uropods are somewhat longer; in others, the tubercles on the first two pleomeres are either longer and flatter or absent. Ten of the 33 females examined are immature to varying degrees, so re- marks on their development are possible. At first, the body is nearly symmetrical, the small oostegites of equal size in each pair, gradually larger from the first to the fifth; the pleon, which is as long as the pereon and not abruptly narrower than it, extends straight back. In the earliest postlarval stages, only the exopodites of the pleopods are developed, the endopodites being small flaps and the lateral plates suggested by tiny bumps. As development progresses (Fig. 7), the branches of the pleopods become equal, while the corresponding lateral plates are still smaller. Other conspicuous differences of the immature fe- males from adult ones are large coxal plates at the bases of all pereopods and much larger uropods. ACADEMY OF SCIENCES VOLUME 73 Description of allotypic male (Figs. 8A—E).—Body length 2.6mm, maximal width 0.6mm, pleon length 0.7mm. Sides of pereon nearly parallel, head and pleon abruptly narrower (Figs. 8A, B). Head much wider than long, fused with first pereomere. Antennae | and 2 (Fig. 8C) of three and seven segments respectively, both bearing many setae on distal segments. No eyes. Pereon broadest, but only slightly so, across second and third pereomeres. All pereomeres separated by lateral incisions. Pereopods (Figs. 8D, E) all of same size, dactyli progressively smaller and carpi larger posteriorly. Pleon about 4% length of body, tapering to blunt point. No segments, but undulating margins indi- cating traces of segments. Appendages lacking. Ventral surface (Fig. 8F) bearing tiny paired slitlike markings. Variations: Several males have their heads dis- tinctly separated from their pereons, while the pleons of some are quite wide relative to pereonal width. In some cases (Fig. 8G) both of these variations occur. Etymology.—The specific name hyphalus is the Latinized form of a Greek word meaning “sub- merged” in reference to the consistent occurrence of this species in much deeper water than S. hyptius. Remarks.—Stegophryxus hyphalus differs from the other two species of the genus in several re- spects. The female is similar to that of S. hyptius, from which it differs in lacking eyes at all stages, having a much more symmetrical brood pouch, smaller uropods in the adult, more prominent pereopodal coxal plates, shorter pleomeres, and smaller pleonal lateral plates in immature forms. The adult female of S. hyphalus differs from that of S. thompsoni in having a relatively shorter head and pleon, more broadly pointed first oostegites, less conspicuous pereopodal coxal plates, narrower pleonal lateral plates and pleo- podal rami, and much smaller uropods. Males of S. hyphalus differ from those of S§. Ayptius in lacking eyes, having pereomeres more strongly separated, and pleon longer, more pointed, and with more undulate margins. The male of S. hyphalus may be distinguished from that of S. thompsoni by its shorter head and pointed pleon. It is entirely possible that all three species have eyes in both sexes; eyes have been observed only in live or freshly preserved specimens of S. hyptius. The only previous record of a Stegophryxus from the coast of California is that listed by Menzies and Miller (1954), who identified the host as a “Pagurus sp.” Their key to species in- dicates that they did indeed have a Stegophryxus; most likely S. hyphalus. Unfortunately, they list no locality or source for this record. 1974 Figure 8. 7. F. Pleon, ventral view. G. Dorsal view. C, D, E, 0.4mm for F. DISCUSSION With the description of three species of Srego- phryxus, it now becomes possible to provide a generic description, thus: Female. Body longer than broad, bent to left so right side is longer: head roughly pyriform with subparallel sides, pro- duced into point anteriorly, indented postero- dorsally; first three pairs of oostegites arching far over head, others forming voluminous but tightly closed brood pouch; fifth oostegites much the largest, covering at least half of pereon BOPY RID PARASITES AND NEW SPECIES FROM ¢ Stegophryxus hyphalus, new species. Dorsal view. B. Ventral view. C. Right antennae. D. Right pereopod 2. (LIFORNIA second male. A. E. Right pereopod A-E, allotype male. G. Scale indicates 1.0mm for A, B, G. 0.2mm for ventrally and produced posteriorly into lateral sacs, that on right side extending slightly to greatly posteriorly. First five pereomeres bent around and enclosing head; sixth pereomere much the largest, concave both anteriorly and _ posteriorly. First two pereopods anterior to head. second two beside it: first five pereopods clustered forward, sharply reflexed, last two far back. more or less straight. Pleon of six pleomeres. First five pleo- meres bearing biramous pleopods and _ lateral plates ranging from linear-lanceolate to subovate by species, all from common peduncle so pleopods 40 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 appear “triramous.” Terminal pleomere with pagurids and diogenids collected off Chile and uniramous bulbous uropods, ranging, by species, Peru and reports that none bore any abdominal from button-like structures to large clubs. Male. parasites. As observed in a discussion of Para- Body length about three times width, sides nearly parallel except for head and pleon. Head more than twice as wide as long, smoothly rounded anteriorly, usually set off from pereon, or at least lateral indentations indicating separation. All pereomeres separated by deep lateral incisions. Pleon from '5 to 's total body length, a single fused piece which is narrower than last pereomere, broadest behind anterior edge, rounded to pointed posteriorly. No pleonal appendages. Three other genera are closely related to Stegophryxus, their females being distinguishable from those of Stegophryxus by several characters. In Stegias Richardson, pereomere 5 is longest, the last oostegites are not produced into posterior sacs, the pleon is relatively much shorter and broader, and the fourth and fifth pleomeres lack lateral plates. There are no markedly elongate pereomeres in females of Anathelges Bonnier, the body is nearly as broad as long, and the brood pouch symmetrical. In females of Pseudostegias Shiino, the brood pouch does not extend beyond the head, and only the first four pleomeres bear lateral plates. The males of these four genera are more difficult to separate along generic lines. The distribution and host selection of the species of Stegophryxus are quite peculiar. First, this is evidently the only bopyrid genus found on both sides of the Americas and nowhere else. Stegophryxus hyptius, throughout its range, in- fests those species of Pagurus which occur on intertidal or shallow subtidal sandy bottoms, even though this range, in covering latitudes from about 26°N to 42°N, goes from an essentially tropical environment to nearly arctic environ- ment. The distribution of S. hyphalus is com- pletely different. It infests both known species of the pagurid genus Parapagurodes from a depth of 69 to 391m; and thus occurs nowhere close to the intertidal zone. Though the recorded latitudinal range of S. hyphalus is from 26°17’N to 36°27’N, there is little difference in tempera- ture at the depths in question. The host and collec- tion depth of §. thompsoni are unknown. In their description of this species, Nierstrasz and Brender a Brandis (1931) report: “Valparaiso. Kréyer. Ein Weibchen mit Mannchen auf Pagurus sp.” The label with the types bears the notation “af Pagurid,” i.e., “from [a] pagurid,” so the genus (and possibly even the family) of the host is uncertain. Janet Haig has kindly examined several thelges spp. (Markham, 1972), there seems to be little host specificity among athelgine bopyrids. Therefore, it would be impossible to judge what the host of the types of $. thompsoni might have been even if more specimens should turn up. ACKNOWLEDGMENTS Edward B. Hatfield collected and provided to me a number of parasitized hermit crabs from the Miami area. Richard W. Heard provided hermit crabs which he and Charles Kellogg had collected in Georgia and North Carolina, while the latter provided collec- tion data from some of them. Torben Wolff allowed me to examine the types of S. thompsoni. Janet Haig examined hermit crabs from Chile and Peru in the Allan Hancock Foundation collections, furnished col- lection data on the hermit crabs from California and granted me permission to describe their parasites. Patsy A. McLaughlin identified these crabs and made their parasites available to me as well as identifying many of the crabs collected along the Atlantic coast. Thomas E. Bowman and Lowell P. Thomas critically reviewed the manuscript. This is publication No. 1734 of the Rosenstiel School of Marine and Atmo- spheric Science, University of Miami, and No. 1 of the Arch Cape Marine Laboratory, Arch Cape, Oregon 97102. LITERATURE CITED Baffoni, G. M. 1953. Modificazione metaboliche dell’ epatopancreas di “Callianassa_ laticauda” nella castrazione parassitaria. Atti Accad. Naz. Lincei, 14:436—442. Bourdon, R. 1968. Les Bopyridae des mers Eu- ropéennes. Mém. Mus. Nat'l. Hist. Nat. (Paris), (ns., A), 50(2):77-424. Caullery, M. 1950. Le parasitisme et la symbiose. Second Ed. C. Doin et Cie., Paris, 358 pp. 1952. Parasitism and symbiosis. Sidgwick and Jackson, London, xii + 340 pp. Florkin, M. 1960. Ecology and Metabolism. Pp. 395-410 in The Physiology of Crustacea. (T. H. Waterman, ed.), Academic Press, New York, Vol. 1, xvii + 670 pp. Kaestner, A. 1967. ogie. Bd. I: Lehrbuch der speziellen Zool- Wirbellose, 2. Teil, Crustacea. 1974 BOPYRID PARASITES AND NEW SPECIES FROM CALIFORNIA Second Ed. Gustav Fischer. Stuttgart, pp. i-viii, Reinhard, BE. G., and FP, W. Buckeridge. 1950. Th 849-1242, effect of parasitism by an entoniscid of secondary sex characters of Pagurus longicarpu ———. 1970. Invertebrate zoology. Vol. IL, J, Parasit., 36;131—-138 Crustacea. (H.W. Levi and L. R. Levi, transl.), John Wiley and Sons, New York. xi -+- 523 pp. Reinhard, E. G., T. von Brand, and S. F. MeDuffic 1947, Observations on the fat content of hermit Kunkel, B. W. 1918. The Arthrostraca of Con- crabs parasitized by a bopyrid. Proc. Helminth necticut. Bull. Connecticut Geol, Nat. Hist. Soc, Washington, D.C,, 14:69-73 Sury., 26:1-261. Reverberi, G. 1952. Parassitismo, iperparassitismo Markham, J.C. 1972. Four new species of e sesso nei crostacei. Pubbl. Staz. zool. Napoli Parathelges Bonnier, 1900 (Isopoda, Bopyridae ), 23:284-295. the first record of the genus from the western Atlantic. Crustaceana, Suppl. 3:57-78. Richardson, H. 1904. Contributions to the natural : history of the Isopoda. Proc. U.S. Nat. Mu McLaughlin, P. A., and J. Haig. 1973. On the 27(1350):1-89. status of Pagurus mertensii Brandt with the de- scription of a new genus and two new species _ tgs. A monograph on the isopods of from California (Crustacea: Decapoda: Paguri- North America. Bull. U.S. Nat. Mus.. 54:liii dae). Bull. So, California Acad. Sci., 72: 113-136. 727 pp. Menzies, R. J., and M. A. Miller. 1954. Chelifera and Isopoda. Pp. 138-159 in Intertidal inverte- brates of the central California coast, S. F. Light’s laboratory and field text in invertebrate zodlogy. (R. L. Smith, F. A. Pitelka, D. P. Abbott, and F. M. Weesner, revisers), Univ. California Press, Smith, R. I. 1964. Keys to marine invertebrates of Berkeley, and Los Angeles, xiv -+ 446 pp. the Woods Hole region. Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Massachusetts, Contrib. no. 11:1—208. Schultz, G. A. 1969. How to know the marine isopod crustaceans. Wm. C. Brown Company, Dubuque, Iowa, vii + 359 pp. Miner, R. W. 1950. Field book of seashore life. G. P. Putnam’s Sons, New York, xv + 888 pp. Sumner, F. B., R. C. Osborn, and L. J. Cole. 1913a. NierstiaszeeEioe. and G. A. Brender A Brandis. A biological survey of the waters of Woods Hole 1931. Papers from Dr. Th. Mortensen’s Pacific and vicinity. Section I. Physical and zoological Expedition 1914-16. LVII. Epicaridea II. Bull. U.S. Bur. Fish., 31:1-441. Vidensk. Medd. Dansk naturh. Foren. Koben- havn, 91:147—226. —. 1913b. Ibid. Part II, section III. A cata- logue of the marine fauna. Bull. U.S. Bur. Fish.. Noble, E. R., and G. A, Noble. 1964. Parasitology. 31:547-794. The biology of animal parasites. Second Ed. Lea and Febiger, Philadelphia, 724 pp. Thompson, M. T. 1902. A new isopod parasitic on the hermit crab. Bull. U.S. Fish. Comm., 21: Rathbun, M. J. 1905. Fauna of New England. 5S. 53-56. List of the Crustacea. Occas. Paps. Boston Soc. Nat. Hist., 7:1-117. Von Brand, T. 1952. Chemical physiology of endo- parasitic animals. Academic Press. New York. Reinhard, E. G. 1949. Experiments on the deter- x + 339 pp. mination and differentiation of sex in the bopyrid Stegophryxus hyptius Thompson, Biol. Bull. 96: 9 __ . 1966. Biochemistry of parasites. Academic 7—3i. Press, New York, x + 429 pp. 1956. Parasitological reviews. Parasitic castration of Crustacea. Exp. Parasit., 5:79-107. Accepted for publication October 20, 1973. COMPARATIVE STUDY OF THE LANCETS AND SHEATHS OF SOME ACULEATE HYMENOPTERA DENNIS M. PoorReE! ABSTRACT: A comparative study was made of the stinging apparati of 102 species of aculeate Hymenoptera from the following families: Scoliidae, Vespidae, Eumenidae, Pompilidae, Sphecidae, Andrenidae, Halictidae, Megachilidae, Anthophoridae, and Apidae. Analysis was confined to the presence or absence of barbs: number, arrangement, and types of barbs. The sheaths were compared as to the types of setae and the presence or absence of segmentation. The purpose of this investigation was to compare the anatomical features of the lancets (first val- vulae) and sheaths (third valvulae) of the stinging apparati of some aculeate Hymenoptera. The dif- ferences found may be of importance in system- atics, phylogeny, and contribute information on sting anatomy. Morphological studies have been made on the stinging apparati, but with little emphasis on the piercing structures and sheaths. Dasymutilla occi- dentalis (Mutillidae) possesses six barbs on each lancet (Hermann, 1968). Polistes exclamans and P. annularis (Vespidae) possess small barbs on their lancets (Hunt and Hermann, 1970). In 1967 Hermann and Blum reported that the presence or absence of barbs on the lancets of formicids had little significance because there is no constant pattern. Using high magnification, Smith (1970) noted that Myzinum maculatum (Tiphiidae) pos- sesses companiform sensilla (sensory pores) and serrulea (subdivisions of the barbs into small pro- jecting teeth). Sheath segmentation was noted by Hazeltine (1967), but little practical application was made of it as far as length ratios of the segments and setae types present. METHODS Stings were extracted from preserved specimens, dehydrated in three solutions (80, 95, and 100 percent) of ETOH, cleared in toluene, and mounted on slides using piccolyte as a mounting medium. RESULTS AND DISCUSSION Only one species, Anthophora curta, of the 102 observed lacked barbs. Barbs have been cate- gorized into five types (Fig. 1A—E): acute: saw- toothed: rounded; knobbed: and reduced. Barbs with an angle of less than 75 degrees were classi- fied as acute; those with an angle of greater than 75 degrees were classified as saw-toothed. Single saw-toothed barbs were prevalent in pompilids. Rounded barbs were confined to scoliids and the tribe Sphecini (Sphecidae). Reduced barbs were found in specimens of Andrena prunorum. Knobbed barbs were confined to the tribe Am- mophilini. In some genera, Auplopus, Ceratina, Euody- nerus, Lasioglossum, and Stenodynerus, the num- ber of barbs remained constant. In others the number of barbs varied, but a typical number of barbs for those genera was evident. In some spe- cies the number of barbs varied from the typical number, but the range never exceeded plus or minus one barb. The Apis mellifera queen had a reduced number of barbs as compared to the workers, possibly a survival adaptation. The queens of Vespula pen- sylvanica and V. sulphurea possessed one more barb than the worker. This may be due to in- creased size and lack of sting autotomy. The barb type, size, and number found in Apis workers and in V. pensylvanica and V. sulphurea were com- parable, suggesting that some other mechanism (possibly a difference in muscular attachment between the sting and the abdomen) must be re- sponsible for the sting autotomy found in the Apis workers. Barbs were arranged on the lancet in a linear fashion and were of one type per species with the following exceptions: Tachysphex ashmeadii, in which the first six barbs at the distal end of the lancet were acute with the remaining five prox- barbs smoothed off imal into minute knobs; “P.O. Box 97, Armona, California 93202. 1974 LANCETS AND SHEATHS OF HYMENOPTERA E Figure 1, Lancets of some aculeate hymenopterans. A, Acute barbs of Apis mellifera worker 450: B, Saw-toothed barbs of Ancistrocerus abdiatus cytainus 535; C, Rounded barbs of Campsomeris tolteca 450; D, Knobbed barbs of Ammophila aberti 450%: E, Reduced barbs of Andrena prunorum 535. Prionyx foxi, in which the first five barbs at the distal end of the lancet were rounded with the remaining six proximal barbs smoothed off into minute knobs: Centris rhodopus, in which the most common barb arrangement, from distal to proximal, was one acute barb followed by two adjacent acute barbs and finally another acute barb; Centris pallida, in which the most common barb arrangement, from distal to proximal. was one acute barb followed by two sets of two ad- Figure 2. Sheaths of some aculeate hymenopterans A, Constricted sheath of cytainus 85%; B, Basal segment twice the ler Ancistrocerus abdiatu the second segment, Anoplius americanus ambiguu 50; C, Basal segment % the length of the second segment. tolteca 35%: D, First and second segments of approximate equal lengths. Sce/- iphron caementarium 50: E, I of Coelioxys obtusiventris 50 Campsomieris Unsegmented she: jacent acute barbs: Clypeadon utahensis. in which there were two acute barbs at the distal end. two sets of two adjacent acute barbs, and three acute barbs proximally. Most specimens studied from the sub- family Halicinae had the same number of barbs and the same barb arrangement: Two acute at the distal end. two sets of two adjacent barbs, and one acute barb proximally. Sheaths were found to be of three basic types: Constricted, two-segmented, and unsegmented Annulated constrictions (Fig. 2A) were found the sheaths of Eumenidae and were less nounced and appeared as a light band with mini- mal constriction in Vespinae. 44 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Taste lt. Lancet and sheath comparison. The number in the columns below represents the number of lancets of those species observed. SHEATH Types (ST): A, constricted; B, first segment twice as long as the second segment: C, second segment twice as long as the first segment; D, first and second segments of approximately equal lengths; FE, unsegmented. Bars Types (BT): A, acute barb; B, saw-toothed barb; C, rounded barb; D, knobbed barb; E, reduced barb. Number of Barbs per Lancet Species Examined ST BT (i) Sl 2, 3) (4 (Si SiG er eee Se Oe Oe eee) SCOLIIDAE Campsomeris tolteca (Sauss.) CG © 7 NS) 2 Campsoscolia alcione Banks (e (e gy) il POMPILIDAE Pepsinae Macromerini Auplopus architectus (Say) Auplopus architectus metallicus (Banks) jeome~] ee} NN Pompilinae Pompilini Anoplius americanus ambiguus (Dahlbom) B Anoplius relativus (Fox) B Episyron quinquenotatus hurdi Evans B Tachypompilus unicolor Banks B > ww n SPHECIDAE Larrinae Larrini Motes aequalis (Fox) o) > i) Tachytini Larropsis tenicornis (F.Sm.) Tachysphex ashmeadii Fox Tachysphex cockerellae Rohw. Tachysphex sp. Tachytes rufofasciatus Cress. GISOuwgi@ S>rrge ASS tv Nyssoninae Bembicini Bembix amoena Handl. Bembix melanaspis Parker Philanthinae Philanthini Aphilanthops hispidus Fox Clypeadon californica (Boh.) Clypeadon utahensis (Baker) Philanthus pacificus arizonae Dunning whe} >> Na Ne) OE) >rrer ) oy aS Sphecinae Ammophilini Ammophila aberti (Hald.) Ammophila azteca Cam. Ammophila sp. Podalonia luctuosa (F.Sm.) Podalonia sp. whokonene) whohonene) an Sceliphronini Chalybion californicum (Sauss. ) Sceliphron caementarium (Drury ) oO >a Ss) ht Sphecini Chlorion aerarius Patton (e Fernaldina lucae (Sauss. ) (C (S 2 Prionyx foxi (Boh. & Menke) D 1974 LANCETS AND SHEATHS OF HYMENOPTERA TABLE |. (Continued ) Number of Barbs per Lancet Species Examined wap! BT ON D2) 3 a Ses Go es eo Or liecd 2 Sphecini (continued ) Prionyx parkeri (Boh, & Menke) D ( Sphex ashmeadi (Fernald) ( ( af Sphex ichneumoneus (L.) (S (@ 4 nN EUMENIDAE Ancistrocerus abdiatus cytainus (Cam. ) Ancistrocerus spilogaster (Cam. ) Ancistrocerus tuberculiceps sutterianus (Sauss. ) Eumenes crucifera Prov. Euodynerus annulatum (Say) Euodynerus foraminatus blandinus (Rohw.) Euodynerus foraminatus scutellaris (Sauss. ) Euodynerus hidalgo vierecki (Cam.) Euodynerus pratense (Sauss. ) Leptochilus propodealis Boh. Pterocheilus morrisoni Cress. Stenodynerus acarigaster Boh. Stenodynerus microstictus (Vier.) >> PPP > Pb > PF Pw i) i) BRNEANOCAN >>rrrrrrrrrrrre >> Nw N PY > Stenodynerus toltecus (Sauss. ) VESPIDAE Polybiinae Epipona taiva Guerin Mischocyttarus flavitarsis (Sauss. ) m mm >> an \o 2 30 Polistinae Polistes annularis (L.) Polistes apachus Sauss. Polistes carolinus (L.) Polistes dorsalis californicus Boh. Polistes exclamans Vier. Polistes fuscatus aurifer Sauss. Polistes fuscatus variatus Cress. Polistes metricus Say eo mcomesmesmesmeomes| es) to Vespinae Vespula maculifrons (Buysson) Vespula maculata (L.) Vespula pensylvanica (Sauss.) queen Vespula pensylvanica (Sauss.) worker Vespula sulphurea (Sauss.) queen Vespula sulphurea (Sauss.) worker tn pS w& to tn >rererre ANDRENIDAE Andreninae Andrena cerasifolii Ckll. Andrena prunorum Ckll. mm m > an i) HALICTIDAE Halictinae Agapostenom sp. Augochlora cuprea (Sm.) Augochlorella pomoniella Ckll. Halictus farinosus F.Sm. Halictus ligatus Say Lasioglossum mellipes (Crwtd.) Lasioglossum sisymbrii (Crwfd.) AmMmMmmm mM -rrrerey > + 46 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Species Examined MEGACHILIDAE Lithurginae Lithurgus apicalis (Cress. ) Megachilinae Anthidiini Anthidium collectum Huard Callanthidium illustre (Cress. ) Megachilini Ashmeadiella opuntiae (CkIl.) Chalicodoma occidentalis Fox Coelioxys obtusiventris Crwfd. ANTHOPHORIDAE Anthophorinae Anthophorini Anthophora curta Prov. Emphorini Diadasia australis (Cress. ) Diadasia australis californica Timb. Diadasia enavata (Cress. ) Diadasia laticauda Ckll. Diadasia rinconis Ckll. Eucerini Melissodes sp. Nomadini Nomada formula Vier. Centridini Centris pallida Fox Centris rhodopus Ckll. Xylocopinae Ceratinini Ceratina acantha Prov. Ceratina punctigena Ckll. Xylocopini Xylocopa brasilianorum varipuncta Patton Xylocopa virginica L. APIDAE Apinae Apini Apis dorsata Fab. Apis florea Fab. Apis indica Fab. Apis mellifera L. queen Apis mellifera L. worker Bombini Bombus californicus F.Sm. Bombus edwardsii Cress. Bombus morrisoni Cress. Bombus sonorus Say Bombus vandykee (Frison) (Continued ) ST mm m ies) OlOs@Oi@i@® leolmeoMesico mcs! nam mm BT >>> >>rrrpy >> >> Prrpre >P Pry VOLUME 73 0 N 1 Number of Barbs per Lancet Poon 7 8 Oo on TD é) 5) 2 2 il, il 2 2 PhS 20 2 14 4 6 8 2 Ai aS ee 18 2 4 2, 4 26 36048 36 8 45 3 45) 11 15 26 1 IS, VG 8 12 4 8 9 1 1974 LANCETS sheaths, the basal segment was slightly longer than the second segment. Sheaths with two segments (Fig. 2B-D) were observed in species of sphecids, pompilids, scoliids, and in the tribes Emphorini and Eucerini (Antho- phoridac). Unsegmented sheaths (Fig. 2) were present in all other species. Two-segmented sheaths were of types. Sheaths with the basal segment twice the length of the second segment (Fig. 2B) were con- fined to the pompilids. Sheaths with the second three basic segment twice the length of the basal segment (Fig. 2C) were found in scoliids and in the tribes Philanthini (Sphecidae) and Emphorini (Antho- phoridae) and scattered throughout other groups. Two-segmented sheaths with the segments approxi- mately equal in length (Fig. 2D) were found in the tribes Bembicini and Ammophilini (Sphecidae ) and scattered throughout other groups. In the five species of Diadasia examined, the sulci were at approximately a 20 degree angle to the axis of the sheath, a more acute angle than on the other segmented sheaths with the second segment twice the length of the basal segment. Barbs and sheath characteristics, for all species examined are sum- marized in Table |. Plumose setae on the sheaths were present in all apoid species except Lasioglossum sisymbrii, Nomada formula, and Apis workers. The Apis workers had setae that were reduced in_ size. Simple setae were present in all other species. In some species, only one specimen was studied. It would be desirable to examine more specimens, including some from different geographical pop- ulations, to determine if the anatomical features remain constant for those species. In addition, more species within certain groups (genera, tribes, AND SHEATHS OF HYMENOPTERA et cetera) should be studied to determin: results obtained for each species in thi hold true for its entire group. ACKNOWLEDGMENTS I wish to thank Roy R. Snelling of the Natural His tory Museum, Los Angeles County for his identifica tion of specimens, advice, and review of this paper LITERATURE CITED Hazeltine, W. E. 1967. noptera and comparative morphology of and female genital segments of Bombinae (Hy- Female genitalia of Hyme- male menoptera: Apidae). Res. Bull. No. 833, Purdue Univ., 36 pp. Hermann, H. R., Jr. 1968. The Hymenopterous Poison Apparatus IV, Dasymutilla occidentalis (Hymenoptera: Mutillidae). J. Georgia Ent. Soc., 3:1—10. Hermann, H. R., Jr., and M. S. Blum. 1967. The Morphology and Histology of the Hymenopterous Poison Apparatus IIT, Eciton hamatus (Formic- idae). Ann. Ent. Soc. Amer., 60:1282-91. Hunt, A. N., and H. R. Hermann, Jr. 1970. The Hymenopterous Poison Apparatus X, Polistes annularis (Hymenoptera: Vespidae). J. Georgia Ent. Soc., (4):210-16. Smith, E. L. 1970. Evolutionary Morphology of the External Insect Genitalia 2. Hymenoptera. Ann. Ent. Soc. Amer., 63:1—27. Accepted for publication August 3, 1973. A NEW SPECIES OF CLYMENELLA (POLYCHAETA: MALDANIDAE) FROM TOMALES BAY, CALIFORNIA James A. BLAKE ' AND JERRY D. KUDENOV ” Apstract: A new species of Clymenella is described from shallow subtidal sediments in Comparisons are made with related Clymenella species. Tomales Bay. Four specimens of an unknown species of Clymenella were collected at a depth of about one meter during a low tide in White Gulch, Tomales Bay, California on 22 October 1969. The speci- mens could not be identified as any species of Clymenella known from California (Hartman, Atlas of the sedentariate polychaetous annelids from California, Allan Hancock Foundation, Los Angeles, 1969) and were determined to represent a previously undescribed species. This is contribu- tion number 33 from the Pacific Marine Station. Clymenella californica, new species Figures 1 and 2 Material examined: White Gulch, Tomales Bay, California, 22 October 1969, from loose silt sediment, depth about one meter; four specimens, holotype and three paratypes, deposited in the collections of the Allan Hancock Foundation, University of Southern California. Description: The specimens have from 22 to 27 setigerous segments with one preanal asetigerous seg- ment in addition to the peristomial and _ pygidial segment. The holotype has 27 setigerous segments, measures 200 mm in length and 2 mm in width. Individual posterior segments are greatly prolonged and maximally measure up to 17 mm in length while those from anterior setigers measure 2.5 to 3 mm. The anterior end terminates in an elongated ce- phalic plaque which has an anterior projecting palpode, followed posteriorly on each side by raised lateral margins which surround the plaque and merge posteriorly (Figs. la-lb). The margin is interrupted by small notches near the posterior end of the plaque. Two parallel nuchal grooves lie alongside an elevated nuchal ridge which extends through the mid-region of the plaque. Clusters of eyespots occur on the anterior lateral edge of the prostomium (Fig. la). The first three setigers lack collars, but bear fas- cicles of long capillary notosetae and one or two acic- ular neuropodial spines (Fig. 2a). Setiger 4 has a conspicuous smooth collar on its anterior border. Notosetae are capillaries, but the neurosetae consist of about fifteen non-bearded rostrate setae, each of which has a main fang and four secondary teeth (Fig. 2b). Setiger 5 and succeeding setigers bear notopodial capillary setae and bearded rostrate neuro- setae (Fig. 2c). The number of secondary teeth gradually increases from four to five. The setal fas- cicles gradually shift from dorsolateral positions in anterior segments (Fig. la) to lateral positions in posterior segments (Fig. Ic). The anal funnel terminates in a ring of 22 nearly equal cirri (Figs. 1c-Id). The anus is at the apex of a raised cone. Color of the first three setigers is iridescent pink; ! Pacific Marine Station, Univ. of the Pacific, Dillon Beach, California 94929. ° Dept. Biological Sciences, Univ. Arizona, Tucson, Arizona 85721. TABLE 1. Differential characteristics of California Clymenella species. Characteristics C. californica C. complanata Number of setigers Length 200 mm Transverse nuchal groove absent across plaque Marginal flange around present plaque Segmental collar setiger 4 Habitat 22-27 setigers and 1 asetiger silty sediments in Tomales Bay 21 setigers and 4 asetigers 90 mm present absent setigers 4 and 5 shaley rock crevices of the open coast 1974 jFigure 1. Clymenella californica, new species: setigers 4-7 have deeply pigmented red areas alter- ‘nating with the lighter pink bands; setigers 8-13 are ‘pale brown to white (white when sexually mature): ‘the remaining segments exhibit color gradients from light to dark orange. The entire body surface is covered with numerous lines and folds. _ The tube is rigid, J-shaped, and formed of cemented ‘sand grains. Distribution: Tomales Bay, California, Remarks: Clymenella californica differs con- siderably from C. complanata (Hartman, 1969) which is known only from San Mateo County and Point Conception in California where it inhabits shaley rocks from the intertidal to shallow sub- tidal. See table 1 for differentiating charac- teristics. On a global basis, Clymenella californica appears closest in morphology to C. cincta (St. Joseph, 1894) from France. It differs from C. cincta in having its collar margin entire instead of incised. The nuchal slits of C. californica are NEW POLYCHAETE FROM CALIFORNIA a, anterior end in lateral view; b, cephalic plaque in dorsal Clymenella californica, new species: a. seta from Figure 2. acicular seta from setiger 1: b. rostrate setiger 4: c, rostrate seta from setiger 21. SO BULLETIN SOUTHERN CALIFORNIA parallel, but transverse in C. cincta. Non-bearded rostrate neurosetae present in C. californica are not reported for C. cincta. The number of seti- gers is not Known. Clymenella californica is superficially confused with Axiothella rubrocincta (Johnson, 1901), a common maldanid in north Pacific bays and estuaries. The generic differences involve the presence of thick acicular neuropodial spines in ACADEMY OF SCIENCES VOLUME 73 setigers 1-3 of Clymenella where Axiothella has rostrate neurosetae. Biology: The functional morphology of feeding in Clymenella californica, Axiothella rubrocincta, and Praxillella affinis pacifica has been studied and will be published elsewhere (Kudenov, M.S.). Eggs were present in the coelom of some speci- mens of C. californica. Accepted for publication February 14, 1974. RESEARCH NOTES NOTES ON THE ARBOREAL ACTIVITIES OF PEROMYSCUS BOYLIT IN INYO COUNTY, CALIFORNIA Arboreal activities in various species of Peromyscus have been known for many years (Baker, 1968), Al- though the arboreal tendencies of Peromyscus boylii have been noted by Bailey (1932), Cahalane (1939), Long (1961), and Storer, Evans, and Palmer, (1944), comprehensive studies of this aspect are unknown to the author. Studies of home ranges, population densities, and other aspects of their life histories were made by Drake (1958), Long (1961), and Storer ef al., (1944). During the summer of 1971, while doing field work in the Owens Valley of California, the author estab- lished a base camp at Cottonwood Creek Camp- ground, 11 miles north and 4 miles west of Olancha, Inyo County. In the evenings, mice were observed in the canyon live oaks (Quercus chrysolepis). An attempt to study the arboreal habits of these mice, subsequently identified as Peromyscus boylii rowleyi, was initiated. The dominant tree in Cottonwood Canyon was the canyon live oak. There were also a few scattered pinyon pines (Pinus monophylla). Cottonwoods (Populus fremontii) were present only along the immediate stream side. Shrubs were common in the areas between the oaks; these were, in order of decreasing abundance: basin sagebrush (Artemisia tridentata), wingscale (Atriplex canescens), and rabbit brush (Chrysothamnus nauseosus). Two canyon live oaks were selected as the site of study. A Sherman live trap was placed on each of the main branches and five traps were placed at the base of the trees. The traps were baited with rolled Oats in the evenings and emptied each morning. Trapping in the oak trees was continuous from 13 August to 26 August 1971. A total of 14 nights with 196 trap-nights was recorded. Also, a single trapline, Taste 1. Recapture data of Peromyscus boylii rowleyi (tree captures). Age classes are indicated as: A (adult); S (subadult): J) (juvenile). Females M ales Times ee eae ke Captured A Ss J A S yj 1 Sl yD 2 i = 1 OX WG) 5 ee 3 it Olen 4 1 - - eae ee 5 =-— =- = Se er 6 SSS Th ageleemet T+ SS = Ue a (20 trap stations set 25 feet apart with th traps per station) was set on the canyon floor trapline maintained for July 1971) in a shrub area, approximately from the two oak Live caught mice were toe clipped (Burt was four night ( 100 feet trees. 1940) Sex and age data were recorded for each mouse; it was released upon the same branch from which it was originally captured. Ages of mice mined from pelage color as follows: adults, full-adult color; subadults, showing some adult color but with post-juvenile molt with dull grey pelage apparent and no indication of molt. In the trees, 34 P. b. Adult and subadult female often captured than were any males or juvenile fe- males (Table 1). Ground trapping for four days yielded nearly equal numbers of male and female mice of all age classes. Capture data for ground trapping was as follows: day 1, seven males and six females; day 2, five males and four females; day 3, four males and four females; day 4, four males and four females. Sex ratios for both tree and ground trapping are recorded in Table 2. A statistically significant difference (x* = 3.98, 1 DF, < 0.05 level) was found to exist between the sex ratios of tree and ground trapped mice. Females are apparently more prone to foraging in trees than are male mice, resulting in more captures of females than males. Though the combined number of fe- males captured was greater than males, there was not a significant difference in the total sex ratio eS 2206 IE DE >10!05))- A season of increased activity (June-August) was recorded for the brush mouse in the Sierra Nevada of California by Storer et al., (1944). They believed that this increased activity caused a decrease in the food supply, thus forcing the mice to forage in trees. Long (1961) indicated that brush mice in Kansas were deter incomplete; and juveniles, rowleyi were captured 95 times. mice were more TABLE 2. Sex ratio data for both trapping areas. The total number of mice for each age class is recorded. Ratio is recorded as males per female. Ad. Subad. Juv. Totals TREES Males 7 1 2 10 Females 10 9 5 24 Ratio 0.70 0.11 0.40 0.42 GROUND Males 7 10 3 20 Females 7 9 2 18 Ratio ile 1.50 1.11 52 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES seemed to prefer acorns as a food item. In the present study, the fact that female mice were more commonly taken in the oak trees than were males be due to competition with the males on the This competition may force the females to utilize an alternative food source in the trees. I thank Ross Hardy, Don Patten, and Lan Lester for their critical review of the manuscript. may ground. LITERATURE CITED Bailey, V. 1932. Mammals of New Mexico. N. Amer. Fauna, 53:1-412. Baker, R. H. 1968. Habitats and distribution. Pp. 98-126 in Biology of Peromyscus. (J. A. King, ed.), Amer. Soc. Mamm., Spec. Publ. No. 2, xiii + 593 pp. Burt, W. H. 1940. Territorial behavior and popu- lations of some small mammals in southern Michigan. Misc. Publ. Mus. Zool., Univ. Michi- gan, 45:1-S8. Cahalane, V.H. 1939. Mammals of the Chiricahua Mountains, Cochise County, Arizona. J. Mamm., 20:418—440. Drake, J. J. 1958. The brush mouse, Peromyscus boylii, in southern Durango. Publ. Mus. Michi- gan St. Univ., Biol., ser., 1:99—132. Long, C. A. 1961. Natural history of the brush mouse (Peromyscus boylii) in Kansas, with a description of a new subspecies. Univ. Kansas Publ. Nat. Hist., 14:99-110. Storer, T. I, F. C. Evans, and F. G. Palmer. 1944. Some rodent populations in the Sierra Nevada of California. Ecol. Monogr., 14:165—-192. JoHN O. Matson, Dept. Biology, California State University, Long Beach (Present address: Natural History Museum of Los Angeles County, California 90007 ). Accepted for publication October 8, 1973. NOTES ON SOME BATS FROM A CAVE ON PENINSULA PARAGUANA, VENEZUELA Cueva del Guano is a limestone cave located on the Peninsula Paraguana, 58 km N and 34 km W of VOLUME 73 as N LIMESTONE ; SINK \ } Wipe ‘ | i H ' Me are et i RRB : ie i x 1 ; a) y aoe ) : s Va ) / / A \ if T=31/.5°C \ 1 cm = 3'meters RH= 85-88% / T=32.5°C ao T= 32°C RH= 89% Figure 1. Diagrammatic sketch of Cueva del Guano showing the positions and sizes of the four rooms with temperatures and relative humidities. Coro, Falcon, Venezuela. The entrance to the cave is about 12 meters below the surface of the ground, in a limestone sink (Fig. 1). The authors visited the cave to obtain bats on July 10, 11, 22, 23, 27, and 31, 1968. Temperatures and relative humidity readings were recorded for each room on July 22, 23, and 31 (Fig. 1). The out- side temperature and relative humidity varied from 28.5°-31° C and 58—66.5 percent, respectively. Representatives of five species of bats were cap- tured in the cave. These were: Pteronotus davyi, P. parnellii, Mormoops megalophylla, Leptonycteris curasoae, and Natalus tumidirostris. This represents the first record of L. curasoae from the Venezuelan mainland of South America. In the cave, 75 speci- mens (65 4 and 10 92) of L. curasoae were obtained. Also, 34 specimens (7 ¢ and 27 @) of this species were netted in the xeric thorn forest, which was the — dominant vegetation of the peninsula (the net area was about 9 km S of the cave). These specimens are now deposited in the United States National Museum of Natural History (catalog numbers: 444758-444819 and 491749-491794); eventually a share of the collection will be returned to Venezuela. This species was first reported (as L. nivalis) from the Colombian mainland of South America by Marinkelle and Grose (Nature, 218(5140):487, 1968). Subsequently, other specimens were reported from a second locality, also in Colombia (Marinkelle and Cadena, Mammalia, 36:50—58, 1972). 1974 The maximum number of bats in)’ Cueva del Guano was estimated to be between 45,000 and 50,000 individuals. ‘This estimate was determined from a photographic slide, from which we were able to count the number of bats (mostly P. davyi) roost- ing per sq m on the ceiling in room A. ‘This room contained the greatest number of bats, 65 per sq m or approximately 30,000 individuals. Overall, L. curasoae and P. davyi were the most abundant, while P. parnellii, M. megalophylla, and N. tumidirostris were considerably fewer in number. On July 10 and 11, L. eurasoae was estimated to be about five times as abundant as P. davyi. How- ever, On subsequent visits, individuals of P. davyi were more numerous than they had been, L. curasoae being considerably less abundant. In fact, on 27 July only one L. curasoae was taken, and by 31 July they were not observed at all. It is interesting to note that P. davyi was also recorded to show temporal alternations of abundance with other bats in a Mexican cave (Dalquest and Hall, J. Mamm., 30:423—427, 1949). It is possible that L. curasoae and P. davyi range over large areas, alternately roost- ing at different localities. Leptonycteris curasoae, P. davyi, and P. parnellii were apparently roosting together on the ceiling in rooms A and B. Mormoops megalophylla were taken only from small crevices in the ceiling or on the walls in rooms A, B, and C. Natalus tumidirostris was the only species found in room D. In this room we recorded the highest temperature and humidity for the cave (Fig. 1). Natalus tumidirostris relative was also captured in room B, from small crevices felt, hands, to be higher than in the center of the room. Natalus tumidirostris was reported to prefer the driest where the humidity and temperature to our parts of warm, humid caves in Trinidad (Goodwin and Greenhall, Bull. Amer. Mus. Nat. Hist., 122: 187-302, 1961). However, in that paper the actual temperatures and humidities were not recorded. There- fore, a direct comparison is not possible. It may be that N. tumidirostris is forced into the more humid areas of Cueva del Guano because of crowding by the other species of bats. We thank Charles O. Handley, Jr., Ross Hardy, Jerry R. Choate, and Andrew Starrett for critically reading this paper at various times during its prep- aration. This paper is a contribution of the Smith- sonian Venezuelan Project, supported by the United States Army Medical Research Command (DA-49-193-MD-2788). and Development JoHN O. Matson, Natural History Museum of Los Angeles County, Los Angeles, California 90007, and Frep P. Brown, Jr., Rural Retreat, Virginia 24368. Accepted for publication August 8, 1973. RESEARCH NOTES TANTILLA TAENIATA (BOCOUR’ ADDITION TO THE SNAKE FAI OF EL SALVADOR Tantilla taeniata (Bocourt), a relatively wide ing member of the southeastern Oaxaca, genus, is known to occur México to central Nicaras (Wilson and Meyer, Herpetologica, 27:11—40, 1971 Wilson and Hahn, Bull. Florida St. Mus., 17:93-150 1973; Wilson and Villa, Bull. So. California Acad Sci., 72:93-96, 1973). This species has not been recorded from El Salvador, however, and this paper documents its occurrence there. Tantilla known specimens (12 to date), and it appears that almost taeniata is from relatively few every specimen collected since the last review is con- tributing markedly to the expansion of the diagnosis of this species. The present specimen is no exception. The specimen of 7. taeniata from El Salvador (University of Utah 4716) was collected at an eleva- tion of 1000 m on Volcan de Conchagua, 4 km S La Union, Depto. La Unién. It is a female with the following characteristics: 143 ventrals (this count is probably + 1 scale from the actual number because several ventrals are missing; I made the count by shifting to the lowermost dorsal scale row to skirt that point); tail incomplete; dorsal scale rows 15—15—15: anal plate single (all Tantilla, of course, characteristi- cally have a divided anal plate); supralabials 7—7 (3rd and 4th entering orbit); infralabials 6—6, first pair not in medial contact, 4 in contact with anterior chin shields; preocular single and in contact with postnasal: postoculars 2—2; temporals 1+ 1. The dorsum is brown and the pale stripes are tan edged with dark brown. The middorsal pale stripe occupies the middorsal scale row and the adjacent halves of the paravertebral rows, and the lateral pale stripe occupies the adjacent halves of rows 3 and 4. Both stripes continue onto the tail. The anterior end of the middorsal stripe is separated by 4 scales from the nape band. The first dorsal scale row is divided into dark brown upper and cream lower halves. The venter is immaculate cream. The tan nape band is complete, covers the tips of the parietals and 1% scales posterior to the parietals, and crosses the pos- terior half of the last supralabial. Pre- and postocular pale blotches are present. I have presented a relatively detailed description of the scutellation and color pattern of this specimen because it is peculiar in one respect and I need to justify my allocation of it to T. taeniata. The color pattern of UU 4716 is like that of all other specimens of T. taeniata that I have examined, especially with respect to the diagnostic characteristics of the species. viz., a pale middorsal stripe occupying the middorsal and adjacent halves of the paravertebral rows, a pale lateral stripe occupying the adjacent halves of rows 3 and 4, first scale row divided into dark upper and pale lower halves, and a complete nuchal collar crossing the last supralabial. 54 BULLETIN SOUTHERN CALIFORNIA The ventral count (143), however, even though approximate, is well below the range for other known females of this species (158-178). This number of ventrals falls within the known range for females of T. jani (141-153) known from low and moderate elevations of the Pacific versant from eastern Oaxaca, Mexico to Guatemala. In view of the fact that the ventral count of UU 4716 falls within the range for female 7. jani and in view of the geographic inter- mediacy of this specimen (Wilson and Meyer, 1971), it is conceivable that T. taeniata and T. jani may be conspecific. I feel it premature, however, to make the necessary nomenclatural changes, as no other speci- mens are known from areas between the ranges of the two species. In addition, variation in scutellation in 7. taeniata remains poorly known and there are distinct pattern differences between the two (Wilson and Meyer, 1971). For the time being, I will continue to consider T. taeniata and T. jani distinct species. The specimen from El Salvador bears some re- ACADEMY OF SCIENCES VOLUME 73 semblance to the holotype of Leptocalamus trilineatus Peters, the status of which has remained enigmatic since its description (Wilson and Meyer, 1971). The holotype of L. trilineatus, which I haye examined previously, is a female with 149 ventrals, a count close to that of UU 4716, but there are important pattern differences between the two and they do not appear to belong to the same species. The holotype of L. trilineatus has a middorsal pale stripe confined to the middorsal row, the upper edge of the pale lateral stripe is not outlined with dark pigment, and there is no continuation of the middorsal and lateral stripes onto the tail. The name Leptocalamus trilineatus still has questionable status. Larry Davip WILson, Division of Intercurricular Studies, Miami-Dade Community College, South Campus, Miami, Florida 33156. Accepted for publication January 31, 1974. INSTRUCTIONS FOR AUTHORS t The BULLETIN is published three times cach year (April, August, and November) and includes ar _ English in any field of science. Non-members will be assessed a page charge of $40.00 per page. 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CONTENTS 4 A new species of polychaetous annelid (Arabellidae) parasitic in Diopatra ornata (Onuphidae) i from Southern California. By Raymond R. Emerson. oo ci. 22 2 sci « « © «+1610 oletonecnlanneneyers 1 Descriptions of two new species of Euschoengastia (Acarina: Trombiculidae) from California 7 and Baja California Norte, Mexico. By William J. Wrenn and Ronald E. Somerby ........ 5) ‘ 1 The cheliceral muscles of the scorpion Heterometrus fulvipes. By A. B. Vyas ........+.+0--+>- 9 i Xiphiorhynchus kimblalocki, a new billfish from the Eocene of Mississippi with remarks on the systematics of Xiphioid fishes. By Harry L. Fierstine and Shelton P. Applegate ......... 14 - Evolutionary relationships and biogeography of the macroteiid lizards (family Teiidae, subfamily Melinae). By William Presche 5 0.6/5): cy0)e tics eue e's Ble Sieve ois tele s 10 '6 7 poke orlekeie aero a eR Renae 23 Parasitic bopyrid isopods of the amphi-American genus Stegophryxus Thompson with the descrip- tion of a new species from California. By John C. Markham ............cececeesvoees 33 Comparative study of the lancets and sheaths of some aculeate hymenoptera. By Dennis M. Poore 42 A new species of Clymenella (Polychaeta: Maldanidae) from Tomales Bay, California. By James A. Blake; and Jerry: Di RUdenov cide ees oo enw ssi eireye love sale siiaies'0 fos the ets) AOTC RR RIOR 48 RESEARCH NOTES Notes on the arboreal activities of Peromyscus boylii in Inyo County, California. By John O. TUL IRNO) Tete O RECA MITA BEARDS Tne MODE TOO C ODDO DEBIT DO Oooo CODdL COONS oil Notes on some bats from a cave on Peninsula Paraguana, Venezuela. By John O. Matson and Fred | LITTON Dae bana 6 CUOMO OS Saal Ob ot Gulto Gs Oreo os an PIM a HIAG HOO Ooo, 6.600,00,00.000 Sy 4 Tantilla taeniata (Bocourt): an addition to the snake fauna of El Salvador. By Larry David q WGUSON) De ieiscst ee oye oy vcokeveisveleboreta eka ha cared Secusy ayn yey Sion aheneual eats oy svar ROR Benoa cowdoo &b-o 53 ( : IndexsitomViolumep/2eer ie sere edd e gen slggleeihins nebo uu Ueno 55) ee ae fs Ceonzo DOOR COVER: An extinct Diatom, Kittonia elaborata Grobe and Strut, from Eocene sediments of the Indian | Ocean (magnified 500x). Photo by Dr. W. W. Wornardt, Union Oil Research, Brea, California. NCES Z 4 = — i = ~ = ’ Number 2 74 SE 19 AUGU Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 OFFICERS Jules Crane, Jr., President Donald Bright, First Vice President Wayland D. Hand, Second Vice President Richard E. Pieper, Secretary Donald R. Patten, Treasurer James Dale Smith, Technical and Managing Editor Roberta S. Greenwood, Index Editor BOARD OF DIRECTORS 1972-1974 1973-1975 1974-1976 Wayland D. Hand Donald Bright George Callison Donald R. Patten John J. Baird Kristen Fauchald Donald J. Reish Jules Crane, Jr. Takashi Hoskizaki Andrew Starrett Richard E. Pieper James Dale Smith Patrick H. Wells David P. Whistler Stuart L. Warter 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. Anmnuale Mem berspesciescsnistesace cinco ores av freee sooo SURI Boo ote bade oh Riot) 6 Lose a SSOVe SPSL CLT eae $ 8.00 StruidentelMem bers icc hier versicicicclass
o> ty = Bey oa San Francisco de Macanao; 2, Cerro Tragaplata; 3, La Estancia; 4, Paraguachi; 5, Salamanca; 6, Cerro Matasiete: 7, La Asuncion; 8, Guatamare; 9, La Aguada; 10, Hacienda Ochenta; 11, El Vallée; 12, Porlamar: 13, Laguna Marites. The 200 meter contours also are indicated. and to Susan Payne for preparation of illus- trations and typing the manuscript. We are in- debted to the following institutions and persons for the loan of critical comparative material (abbre- viations preceding the names of institutions are used throughout the text to identify the source of specimens): AMNH, American Museum of Nat- ural History (Karl F. Koopman); BMNH, British Museum (Natural History) (J. E. Hill); IZT, Instituto de Zoologia Tropical, Facultad de Cien- cias, Universidad Central de Venezuela, Caracas (Juhani Ojasti and Edgardo Mondolfi); KU, The Museum of Natural History, The University of Kansas (J. Knox Jones, Jr.); LACM, Natural History Museum of Los Angeles County (Donald Patten); MCZ, Museum of Comparative Zoology, Harvard University (Barbara Lawrence); MLS, Museo de Historia Natural, La Salle, Caracas, Venezuela (Omar Linares); RMNH, Rijksmu- seum van Naturlijke Historie, Leiden, Netherlands (A. M. Husson); TCWC, Texas A and M University, Texas Cooperative Wildlife Research Collections (W. B. Davis and Dilford C. Carter); TTU, The Museum, Texas Tech University (H. H. Genoways); and USNM, National Mu- seum of Natural History (C. O. Handley, Jr.). We also thank Donald Patten, Natural History Museum of Los Angeles County, for identifi- cation of specimens of Artibeus. All specimens are deposited in the Museum of Natural History, The University of Kansas, unless otherwise noted. All measurements employed in this study are defined by Smith (1972:6) and were taken with dial calipers, calibrated in twentieths of a millimeter and were recorded to the nearest tenth of a millimeter. Measurements of embryos are given as crown-rump lengths. ACCOUNTS OF SPECIES FAMILY EMBALLONURIDAE Peropteryx macrotis trinitatus Miller, 1899 Specimens examined (24)—Cave SW El Valle [Musso (1962:167) indicates this cave is surely Cueva del Piache], 1 (MCZ): no specific locality, 23 (RMNH). Additional records—Guatamare, near El Valle (Hummelinck, 1940:69); Cave near La Asuncion, Carretera a Porlamar; Cueva Chaure. Cerro Atagua (Musso, 1962:166). This species apparently is one of the more common bats in the mesic regions of Margarita Island. All of the individuals reported by Musso (1962:166) were found in caves. In a cave near La Asuncion, he encountered Saccoprteryx le living with a large colony of P. mm. trini tatus. Measurements of the material examined by 60 TABLe 1. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Selected cranial and external measurements of Noctilio leporinus from the Antilles and northeastern South America. Superscript numbers indicate a sample size that differs from those given in left-hand columns. Number Greatest Condylo- Length of and length of basal Zygomatic Mastoid maxillary Length of sex Forearm skull length breadth breadth toothrow mandible Noctilio leporinus mastivus, Puerto Rico (Anthony, 1925:16-17) 5 86.7° 27.3 20.3 19.6 19.1 (85.0-88.0) (26.6—28.5) — (19.8-20.7) (18.0—20.7) — (18.5-19.4) 5 Q 83.9" 25.5 18.9 eo, 18.0 (80.0-85.8) (24.9-26.3) os (18.6-19.1) (16.8-17.8) — (17.7-18.4) Noctilio leporinus mastivus, Dominica, Lesser Antilles 13 88.9 25.6 24.9 20.0 18.5 10.6 18.6 39 87.9 24.8 23.8 19:3 17.0 10.0 17.6 (86.4-88.7) (24.6-24.9) (23.6-23.9) (19.1-19.5) (16.9-17.2) (10.0-10.1) (17.5-17.7) Noctilio leporinus mastivus, Trinidad* 464 85.6 27.4% 24.7! 19.6 19.0* 10.5 18.7! (82.7-88.0) (26.7-28.5) (18.7—20.2) (10.0-10.8) 3 Q 86.1% 24.8, 26.9° 23.5, 24.6° 19.4 16.9, 17.0° 10.4 17.4, 17.7° (82.9-89.7) (19.3-19.5) (10.1-10.7) Noctilio leporinus mastivus, Margarita Island, Venezuela 1 Q 85.8 24.1 2325) 18.5 17.5 10.0 17.3 Noctilio leporinus mastivus, Estado Sucre, Venezuela 1 87.9 26.3 24.9 20.1 19.9 10.8 18.6 5 Q 87.2 D2, 24.0 19.0 17.0 10.3 17.7 (84.1-92.1) (24.8-25.7) (23.5-24.4) (18.7-19.4) = (16.3-17.8) (10.1-10.5) (17.6-17.8) Noctilio leporinus leporinus, Surinam (Husson, 1962:68 ) 56 79.0 25.0 PE}-3} 18.4 16.8" 10.4 17.8 (77.3-81.2) (23.3-26.2) (22.8-23.7) (17.0-19.1) (14.9-18.4) (10.3-10.5) (17.4-18.3) 6 Q 78.8 24.5° 22.6 17.8 1572 10.0 17.3 (78.5-82.1) (24.1-24.9) (22.0-23.2) (16.8-18.3) (15.1-16.0) (9.1-10.2) (17.1-17.6) * Includes measurements of two males and one female from Goodwin and Greenhall (1961:219). us and those published by Musso (1962) agree with those given by Goodwin and Greenhall (1961:216) for four specimens of P. m. trinitatus (including the holotype) from Trinidad. Saccopteryx leptura (Schreber, 1774) Specimens examined.—None. Additional records—Hacienda Ochenta; Cueva near La Asuncion, Carretera a Porlamar (Musso, 1962:167); El Vallé (Pirlot and Leén, 1965:369). Pirlot and Leon (1965:369) reported this species as very abundant in the forest, on the higher hills of the eastern portion of the island. Individuals were found in groups of two to five living in holes in rocks and fallen trees, and between the buttresses of Ceiba pentandra. Musso (1962:167) found this species roosting with a large colony of Peropteryx macrotis trinitatus in a cave near La Asuncion. FAMILY NOCTILIONIDAE Noctilio leporinus mastivus (Vahl, 1797) Specimens examined (1).—1% km N San Francisco de Macanao, 75 m, 1. A nonpregnant, lactating female, represents the first record of this large fish-eating species from Margarita Island. It was captured at dusk in a mist-net set across a small arroyo which drained into a small artificial pond. Lesser nighthawks, Chordeiles acutipennis, were cap- tured at this time as they foraged over the pond. The vegetation around the pond was low, thorny scrub. Koopman (1968:2) discussed the subspecific status of Noctilio leporinus with particular refer- ence to N. |. mastivus, and Davis (1973) has recently reviewed the species leporinus. The selected external and cranial measurements given 1974 in table 1 clearly show that the Antillean bats are much larger in size than those from Surinam, which is the type locality of N. /. leporinus as restricted by Thomas, 1911:131. Both Koopman (1968) and Davis (1973) proposed that the antillean population should be referred to the race N. 1. mastivus. In comparing cranial and external measurements of our specimen from Margarita Island and several from the adjacent mainland, we found them to agree more closely in size with the Antillean bats than those from Surinam, Therefore, we concur with Davis (1973) in applying the name N. /. mastivus to the populations of large fish-eating bats from northeastern Venezuela. Specimens used for comparison are as follows. —BRAZIL: Cruzeiro do Sur, Acre, | (LACM). CUBA: Km. 7, between San Juan de Las Lajas and La Ruda, Habana, 2 (TCWC). DOMINICA: Clarke Hall Estate, 100 ft, St. Joseph Parish, 1 (KU); Mouth of Layou River, sea level, St. Joseph Parish, 3 (KU); no specific locality, 2 (1 BMNH, 1 USNM). TRINIDAD: Fyzabad, 4 (3 KU, 1 TCWC); Peral, 4 (TCWC). VENEZUELA: 1% km NW EI Pilar, Sucre, 6 (KU). FAMILY MORMOOPIDAE Pteronotus parnellii fuscus J. A. Allen, 1911 Specimens examined (31).—Cueva Atagua, Cerro Atagua, 5 (MLS); Cerro Matasiete, 2 km N, 2 km E La Asuncion, 305 m, 1 (USNM); El Vallé, 50 m, 2; Cueva Honda del Piache, SE El Vallé, 2 (RMNH); La Aguada, 3 km S La Asuncion, 53 m, 12 (USNM); Cueva El Convento, 1 km N, 1 km W San Fran- cisco de Macanao, 100 m, 9. Parnell’s mustached bat appears to be rather common on Margarita Island. The ecological limits of this species are not well known. It was encountered in both xeric and mesic habitats on the island. At Cueva El Convento, near San Francisco de Macanao, P. p. fuscus was collected in close association with Leptonycteris curasoae. It is not known whether individuals of P. parnellii were roosting singly or as a segregated cluster within the colony of Leptonycteris as both species were obtained with a single discharge of a shotgun. The bats were roosting in a dome-like depression in the ceiling, about ten feet above the floor of the dry, guano-filled cave. One wall of the roosting chamber was open to the outside by BATS FROM MARGARITA ISLAND, Vio { way of a verti | eWay aii lighted, The and contit indibles of / keletal from the room was dim! of guano indicated a Jo: | VO Mm among the the cave by bats. were found weathered owl pellets removed floor. One female, captured on 16 July, was pri with one embryo that measured 27. Ty oO femal taken on 12 November, showed no signs of reproductive activity. In analyzing the geographic variation of P parnellii, Smith (1972:77) from Margarita Island and the Caribbean versant assigned specimens of Venezuela to the subspecies P. p. Although individuals of P. Margarita fall well within the range of variation of mainland populations of P. p. Juscus. parnellti from fuscus, they show a slight reduction in overall size. Populations of P. p. rubiginosus from the Amazon Basin are considerably larger in cranial and external size (Smith, 1972). Specimens Trinidad appear to more closely resemble P. p. of parnellii from rubiginosus than P. p. fuscus. Mormoops megalophylla tumidiceps Miller, 1902 Specimens examined (4).—Cueva Honda del Piache, SE El Vallé, 3 (RMNH); Cueva El Convento, 1 km N, 1 km W San Francisco de Macanao, 100 m, I. Additional record—El Vallé (Pirlot and Leon, 1965:369). The ghost-faced bat does not appear to be an abundant species on Margarita Island and is common only locally on the Venezuelan main- land. One of our specimens is a right mandible found in the owl pellet material from Cueva El Convento. The geographic variation of this species is discussed by Smith (1972). Selected external and cranial measurements of two males from Cueva Honda del Piache are as follows: length of forearm, 55.6, 55.2: condy- lobasal length, 14.4, 14.1; zygomatic breadth. 9.8, 9.6; rostral breadth, 5.5, 5.4; length of maxillary toothrow, 8.1, 7.8. FAMILY PHYLLOSTOMATIDAE Micronycteris megalotis megalotis Gray, 1842 Specimens examined (3)—El Vallé, 2 (MCZ): Cueva El Convento, 1 km N, 1 km W San Francisco de Macanao, 100 m, 1. Additional records.—all from Museo (1962:168). 68 BULLETIN SOUTHERN CALIFORNIA Cerro Tragaplata; La Estancia, 2 km W Paraguachi; Salamanca; San Francisco de Macanao. According to Musso (1962:168), this species is widely distributed on the island. Selected ex- ternal and cranial measurements of one male and one female, respectively, from El Vallée are as follows: length of forearm, 32.8, 33.7; zygomatic breadth, 8.7, 8.6; rostral breadth, 4.8, 4.8; length of maxillary toothrow, 6.7, 6.9; condylobasal length, —, 15.7. These measurements agree with those given by Sanborn (1949) and Husson (1962) for the subspecies M. m. At Cueva El Convento, this species was repre- sented by a right lower jaw found in the weathered remains of owl pellets. megalotis. Phyllostomus discolor discolor Wagner, 1843 Specimens examined (6).—El Vallée, 50 m, 6. Pirlot and Leon (1965:369), first reported this bat on Margarita Island. Their material also was from El Vallé, the only locality on the island from which this species has been recorded. The ecological preferences of P. discolor seem to be for mesic habitats and for this reason the species may be restricted to the northeastern portion of the island. Of five females mist-netted on July 15 and 16, one contained one embryo measuring 17. Two others were not pregnant but were lactating. A male taken at the same time had testes measuring 5 mm in length. These bats were captured as they foraged in a peach and mango grove. Valdez (1970), after studying the geographic variation of Phyllostomus discolor, employed the name P. d. discolor for populations east of the Andes. Power and Tamsitt (1973) have ques- tioned the validity of the recognition of intra- specific names in this species. In the absence of an extensive study of geographic variation in northern South America, we have assigned our topelemd: Selected external and cranial measurements of material discolor. the four females (average with extremes in paren- theses) and two males, respectively, from El Vallé are as follows: length of forearm, 61.3 (60.1—-62.2), 62.0, 63.3; condylobasal length, 26.4 (25.3-26.6), 25.8, 26.9; zygomatic breadth, 15.4 (15.0-16.0), 15.4, 15.6. These measurements agree with those given by Sanborn (1936:98) for specimens from Brazil, Venezuela, and French ACADEMY OF SCIENCES VOLUME 73 Guiana, and by Goodwin and Greenhall (1961: 238) for material from Trinidad. Glossophaga longirostris longirostris Miller, 1898 Specimens examined (26).—El Vallé, 50 m, 24 (3 MCZ); 114% km N San Francisco de Macanao, 75 ina P25 Additional records —La Estancia, 2 km W Para- guachi; Salamanca; Cueva del Piache; Hacienda Ochenta; San Francisco de Macanao (Musso, 1962: 169): El Vallé (Pirlot and Leon, 1965:369). G. M. Allen (1902:96) reported on three specimens from a large cave near El Vallé which he referred to the species Glossophaga soricina. One was an adult male, another a young (sex unknown) individual with deciduous teeth, and the third specimen was an adult female. He noted that the latter seemed distinct from the other two and thought that it might represent an undescribed species. We have examined Allen’s three specimens and found that only the female is an adult. We concur with Koopman (1958: 437) who assigned all three bats to Glossophaga longirostris. Pirlot and Leén (1965:369) report two male G. soricina from El Vallé. We have not examined these specimens, but suspect that these, too, are but immature individuals of G. longirostris. Adult G. longirostris are easily distinguished from G. soricina on the basis of greater length of forearm. In addition, the upper incisors of longirostris are nearly equal in size and are equally procumbent. The outer incisors of sori- cina are smaller than the inner pair and somewhat less procumbent. Eight of fourteen females mist-netted in a peach and mango grove at El Vallé on 15 July, carried one embryo which averaged 17.0 (12.0— 20.0) crown-rump length. The remaining six females showed no sign of reproductive activity. Table 2 shows the means and extremes of several selected external and cranial measurements of G. longirostris from selected localities. Our material from Margarita Island is well within the range of variation of the mainland G. I. longiros- tris. These measurements also widely overlap those given for the Antillean race G. I. rostrata. Whether or not this indicates a close relationship between these populations is not clear at this time. However, this may indicate that the source for the Antillean populations may have been directly from the mainland rather than from the mainland by way of Trinidad and Tobago. Until | 1974 BATS FROM MARGARITA ISLAND, VENEZUELA TABLE 2. Selected cranial and external measurements of Glossophaga loneirosiris from the Antill eastern South America. Superscript numbers indicate a sample size that differs from those given in the left-hand column Breadth of Length « Condylobasal Breadth of Mastoid rostrum at maxillar Number Forearm length braincase breadth canines toothrov: Glossophaga longirostris elongata, Curacao (Miller, 1913:429) 20 37.3 22.9 8.7 £1 (35.6-39.0) (21.8-23.2) (8.49.0) — (8.0—8.4 Glossophaga longirostris longirostris, Margarita Island, Venezuela 9 37.8 21.2 8.8 9.2 4.0 7.9 (36.4—29.6) (21.0-21.6) (8.5-8.9) (8.89.5) (3.7-4.3 ) (7.7-8.2) Glossophaga longirostris longirostris, Estados Sucre, Venezuela 12 37.4 21.2 8.8 9.4 4.) 7.8 (36.2-39.4) (20.4—22.0) (8.5-8.9) (9.1-9.6) (3.9-4.2) (7.5-8.0) Glossophaga longirostris longirostris, La Guaira, Venezuela (Miller, 1913:429) 10 37.2 DAT; 9.1 8.1? (35.4-38.6) (21.0-22.4) (8.89.4) — — (7:8=8'8)) Glossophaga longirostris major, Trinidad 5 40.2° 21.5 10.1° Sale 4.2 8.1 (39.8-41.0) (20.8—22.2) (10.0—10.3 (9.0-9.2) (4.1-4.3) (7.9-8.4) Glossophaga longirostris rostrata, Grenada 9 37.7 21.5 8.8 9.4 4.1 7.9 (37.i-38.5) (20.8-22.0) (8.69.0) (9.2-9.7) (3.84.4) (7.88.0) Glossophaga longirostris rostrata, St. Vincent 10 37.4 Die 9.4 US) (36.4-38.8) (20.6-21.9) — (9.3-9.6) 3.9 (7.5-7.9) this relationship is better understood we tenta- tively assign our material to G. /. longirostris. Leptonycteris curasoae Miller, 1900 Specimens examined (88).—El Vallé, 50 m, 5; Cueva El Convento, 1 km N, 1 km W San Francisco de Macanao, 100 m, 81; 1% km N San Francisco de Macanao, 75 m, 2. Additional record—The holotype and type series of L. c. tarlosti from El Vallé (Pirlot, 1965:6-7). Individuals of this apparently widespread and common species were captured in mist-nets and shot in caves. The majority of our material resulted from two visits (July and November) to Cueva El Convento, near San Francisco de Macanao. The cave is described in the account of Pteronotus parnellii fuscus, a species found living in direct association with this colony of Leptonycteris. When first discovered in July the colony was composed of approximately 4000 females nursing nearly full-grown young. The young bats had unfused phalangeal epiphyses and were covered with sparse greyish, juvenile pelage. In addition, these young were in varying stages of losing the upper deciduous canines and lower incisors, the third upper molars and the permanent lower incisors were nearly completely erupted, and all cranial sutures were unfused. The forearms of 15 young bats averaged 53.3 (52.0-55.2) in comparison with 53.4 (51.3-55.8) of 61 adult females. No adult males were captured on the July visit. In November, the composition of the colony had changed. There were no young individuals with unfused wing bones, and adult males breeding condition (testes 6-8 mm in length) were present. Of the 34 females captured in November seven were pregnant: each with one small embryo which averaged 6.3 (3.1—11.3). The smallest embryos had the limb buds only slightly differentiated, whereas these structures relatively well the embryos. Skeletal elements and complete skulls in were developed in largest 70 BULLETIN SOUTHERN CALIFORNIA Figure 2. Top, right lateral view of the rostrum of an adult female Leptonycteris curasoae (KU 118206) showing the hyperdontia of the canine. Bottom, occlusal view of same. of Leptonycteris also were encountered in the owl-pellet material from the floor of the cave. Hyperdontia in the genus Leptonycteris was encountered only in the species L. sanborni by Phillips (1971:76). In our series of L. curasoae ACADEMY OF SCIENCES VOLUME 73 female (KU 118206) with a unilateral duplica- tion of the upper right canine (Fig. 2). Another (KU 118193) exhibited a_persis- tent left upper deciduous canine in the space adult female between the permanent canine and first premolar; all teeth were fully erupted and partially worn. Our sample also indicates that incisors are appar- ently lost frequently during life. In two individuals upper premolars had been lost and in another the second right upper molar was lost. Pirlot (1965:607) evidently was unable to com- pare his material Margarita Island with specimens from Aruba, Curacao, or Bonaire when he described L. c. tarlosti. Table 3 shows selected external and cranial measurements (average with extremes in parentheses) of specimens of L. cura- from Margarita, Aruba, Curacao, and Bonaire islands. As can be seen, there is a great deal of overlap, especially between individuals from Margarita and Aruba. Although individuals from the oceanic islands Curacao and Bonaire appear to have somewhat narrower crania, a meaningful interpretation of any geographic trend is difficult with so few specimens. There appears to be little evidence to support the recognition of the population from Margarita Island as a distinct race. The species is widespread along the coastal lowlands of northern Colombia and Venezuela (Marinkelle and Cadena, 1972:53; Matson and Brown, 1974) and we believe that an analysis of geographic variation will show that mainland and insular populations are undifferentiated. We therefore consider Leptonycteris curasoae to be from sodae from Margarita Island, we found one adult monotypic. TABLE 3. Selected cranial and external measurements of Leptonycteris curasoae. Superscript numbers indicate a sample size that differs from those given in the left-hand column. Breadth Breadth oe Length of : Condylo- Length Length of Zygomatic of across maxillary basal mandibular Number Forearm breadth braincase canines toothrow length toothrow Margarita Island, Venezuela 12 53.4 ede 10.17 4.9" 9.4 26.4 10.2 (51.3-55.2) (10.7-11.4) (9.9-10.4) (4.8-5.0) (9.09.7 ) (25.4-26.9) (9.9-10.6) Aruba Island 2 54.4° 11.4, 11.6 10.2, 10.5 Saal 9.6, 9.7 27.0, 27.6 10.7, 10.6 (53.7—-54.7) Curacao Island 5 54.3 10.9 10.1* 4.9 9.4 26.6* 10.4 (51.4-55.7) (10.7-11.3) (9.8-10.2) (4.5-5.1) (9.2-9.6) (26.0-27.2) (10.2—10.8) Bonaire Island 1 5255 —- 10.2 Sy 9.5 10.6 26.8 1974 TABLE 4, Selected external and cranial measurements of Artibeus planirosti BATS FROM MARGARITA ISLAND, VENEZUELA from northeast and Trinidad. Superscript numbers indicate a sample size that differs from those given in the lefth Breadth of I Condylobasal Zygomatic Mastoid rostrum at maxil Number Forearm length breadth breadth canine tootl El Vallé, Nueva Esparta, Venezuela 5 55.9 24.0 16.6 14.7 7/5) 9.7 (52.6-57.4) (23.7-24.4) (16.2-17.0) (14.4-14.9 ) (7.1-7.6) 9.4—10.0 Cumana, Sucre, Venezuela 22 58.6 24.2 Woe 15.2 7.6 9.7 (55.7-61.0) (23.7-25.1) (16.4—17.9) (14.8-15.6) (7.2-8.2) (9.4-10.1) 5 km E San Antonio del Gulfo, Sucre, Venezuela 17 59.6 24.6 17.4 15.2 7.8 10.5 (56.3-62.9) (23.8-25.4) (16.6-18.1) (14.6-15.8) (7.5-8.1) (9.6-10.3) 1.5 km NW El Pilar, Sucre, Venezuela 22 59.8" 24.6 17.4 IS'2 7.9 10.0 (57.1-62.9) (24.0-25.4) (16.7—18.2) (14.6-15.7) (7.4-8.5) (9.7-10.5 ) Various localities on Trinidad 8 57.1° 24.3" tle 14.8" 1/ 9.9° (54.8-58.9) (23.6—25.1) (7.5-7.9) (9.6-10.2) (16.7-17.9) Carollia_ perspicillata perspicillata (Linnaeus, 1758) Specimens examined (2).—Las Piedras, 2. Additional records——La Estancia, 2 km W_ Para- guachi (Musso, 1962:170). Two males were captured in a mist-net as they foraged along a dry arroyo bordered by _ tall broad-leafed gallery forest composed primarily of the fig tree, Ficus yoponensis. Musso (1962: 170) collected this species from under the roof of an old abandoned house. Only two other species of bats, Artibeus planirostris and A. lituratus, were netted at this locality. The forearms of the two males captured 1S July measured 42.2 and 40.5. We follow Goodwin and Greenhall (1961), Husson (1962), and Pine (1972) in assigning our specimens to Carollia perspicillata perspicillata. Artibeus planirostris trinitatis Andersen, 1906 Specimens examined (18).—Las Piedras, 8; El Valle, 50 m, 10. Additional records—Hacienda Ochenta (Musso, 1962:170); El Vallé (Pirlot and Leon, 1965:369). This bat is one of the most common species in the wetter portions of the island and the adjacent mainland. At Las Piedras, the fruits of the native fig (Ficus yoponensis) were found (14.4-15.6) in the nets along with both Artibeus planirostris and A. lituratus. Some flying individuals also were observed carrying a single fruit of this fig. Two of five females caught at Las Piedras on 15 July each carried one embryo measuring 22.: and 28.8, respectively. Two other females were lactating and a third showed no signs of repro- activity. In addition, two females El Vallé on 15 July were pregnant with embryos that measured 27 and 28. We follow Patten (1974) in applying the specific name planirostris to these specimens of collected at Artibeus from Margarita. Although our material of this species from Margarita averages slightly smaller in external and cranial size than speci- mens from the adjacent Venezuelan mainland and Trinidad (Table 4), we do not feel justified in recognizing this population as a_ distinct geographic race. We therefore assign our material to Artibeus planirostris trinitatis. Artibeus lituratus palmarum J. A. Allen and Chapman, 1897 Specimens examined (6).—Las Piedras. 1: El Valle. 50 m, 5. Additional records—Salamanca (Musso. 1962: 170): El Vallé (Pirlot and Leon. 1965:369). This large Artibeus does not appear to be common on Margarita Island. Musso (1962) first 72 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES reported the species on the island on the basis Pirlot Leon (1965) specimens of their and lituratus in of one adult male. reported 17 collection from Margarita. One of our specimens was captured in a mist-net set across a dry boulder-filled arroyo bordered by fig Artibeus planirostris also was captured at this site. At El Vallé, A. lituratus was netted in a peach, mango, and coconut grove. External and trees. cranial measurements of our material agree with those given by Goodwin and Greenhall (1961: 261) for specimens from Trinidad. One of two females captured on 15 July at El Vallé carried a single embryo measuring 25. The testes of three males obtained the same night at El Vallé measured 5, 6, and 7. Desmodus rotundus rotundus (E. Geoffroy-St. Hilaire, 1810) Specimens examined (1).—1% km N San Francisco de Macanao, 75 m, 1. Additional records——El Vallé 1965:360). (Pirlot and Leon, Although this species is abundant on the Venezuelan mainland, its presence on Margarita Island was not reported until relatively recently by Pirlot and Leén (1965). Our specimen, an adult male, was caught in a mist-net set over a dry wash. The specimens reported by Pirlot and Le6n (1965) were captured in a peach, mango, and coconut grove near El Vallée. In July 1966, the Food and Agriculture Organization of the United Nations reported several cases of paralytic rabies in cattle from the areas of San Antonio and “Ochenta.” Cave fumigation was employed by the Venezuelan Ministerio de Agricultura e Cria in an effort to eradicate bats on the island early in 1967. Selected external and cranial measurements of the individual from near San Francisco de Macanao followed by those (average with ex- tremes in parentheses) of four males from the adjacent mainland (Cumana and vicinity) are as follows: length of forearm, 59.1, 55.4 (54.1— 57.6); condylobasal length, 19.9, 20.4 (20.0- 20.7); zygomatic breadth, 11.6, 11.6 (11.2-12.0); length of maxillary toothrow, 3.3, 3.3 (3.2-3.5). These measurements agree with those given by Goodwin and Greenhall (1961) and Husson (1962) for the nominal subspecies rotundus. Diaemus youngi youngi (Jentink, 1893) Specimen examined (1).—El Vallé, 50 m, 1. VOLUME 73 Additional records.—Sierra del Copey; Cueva del Piache (Musso, 1962:171); El Vallé (Pirlot and Leon, 1965:6). The avian vampire was first reported from Margarita Island by Musso (1962:171). In the region around Cerro Copey, Musso noted that Diaemus was common throughout the forest and that flying individuals were easily identified by their large white wing tips [A. planirostris and A. lituratus also possess white wing tips]. Seven individuals of D. youngi reported by Musso were from a large colony which he encountered in a hollow tree trunk. Goodwin and Greenhall (1961:271) described the peculiar oral glands of Diaemus which are located in the corners of the mouth. These glands were observed on the specimen from Margarita as well as several individuals from the adjacent mainland. When disturbed, the bat opens its mouth and the two glands evert, nearly filling the oral cavity. Each gland is bulbous in shape and measures about 2-3 mm in diameter when fully everted. The mouth glands of our July-taken specimen appeared to be quiescent because the odor described by Goodwin and Greenhall (1961:271) was not noted. Several February-taken males from the mainland had very active glands that squirted a fine stream of liquid, which had a “mustelid- like” odor, when these individuals were disturbed. Selected external and cranial measurements of our specimen from Margarita Island, those of three males (average with extremes in paren- theses) and one female from the adjacent main- land, and the holotype of D. youngi (taken from Husson, 1962:199) are respectively: length of forearm, 52.2, 52.1 (51.1-53.3), 52.3, 50.9; condylobasal length, 21.3, 21.4 (21.2—21.8), 21.4, 21.1; zygomatic breadth, 13.9, 14.0 (13.9-14.1), 13.3, 13.3; breadth of the braincase, 12.5, 13.0 (12.9-13.1), 12.9, 13.0; length of maxillary toothrow, 3.2, 3.6 (3.5-3.6), 3.5, 3.0. Thomas (1928:288-—289) proposed the recognition of a separate race, D. y. cypselinus, from Pebas, Pert, based on one female specimen. Later, Sanborn (1949:282-—283) reported an additional specimen identified as cypselinus from Yarinacocha, Peru. Comparison of the above-listed measurements with those given by Thomas (1928) and Sanborn (1949) suggests to us that the Peruvian subspecies In view of the paucity of material, we tentatively retain is only slightly larger than youngi. the name D. y. cypselinus as a weakly defined 1974 race. The specimens from northeastern Venezucla and Margarita Island are referred to Diaemus youngi youngt. Other Venezuelan specimens of Diaemus that we examined in this study include: Tacal, 11 km SSW Cumana, Sucre (KU) 1; Chara Santa Rita, 3 km SE Cumana, Sucre (KU) 1; 2 km S Cachipo, Monagas (KU) 2. FAMILY VESPERTILIONIDAE Rhogeessa minutilla Miller, 1897 Specimens examined (2).—El Vallé, 1 (USNM); un- specified locality, 1 (USNM). Additional record.—Laguna de las Marites (Musso, 1962:171). The three specimens listed above are the only examples of Rhogeessa minutilla from Margarita Island, the type locality. The individual reported by Musso (1962:171) was captured in a man- grove swamp. Miller (1897:139) described the first specimen from the island as a full species, which it remained until Goodwin (1958:7) relegated it a subspecies of R. parvula. Along with the material from Margarita Island, Goodwin assigned specimens from Trinidad, northern Venezuela, and eastern Colombia to this race. LaVal (1973) has recognized Rhogeessa minutilla as a distinct species and assigned to it specimens from Mar- garita Island, northwestern Venezuela, and ad- jacent Colombia. Tentatively, we follow LaVal’s arrangement with some reservations. We believe that when the relationships of these bats in northern South America are fully understood, minutilla will prove to be, at most, a geographic race of R. tumida. FAMILY MOLOSSIDAE Molossus molossus molossus (Pallas, 1766) Specimens examined (2).—El Vallé, 2 (MCZ). Additional records—Guatamare, near El Vallé (Hummelinck, 1940:72); Juan Griego Salamanca (Musso, 1962:172). This species is probably much more common than available specimens indicate. Musso (1962: 172) reported collecting his material (all males) from a bell tower of a church. One of us (Smith) observed this species at dusk as they emerged from under the eaves or out of the rain spouts of houses in Porlamar. BATS FROM MARGARITA ISLAND, VENEZUELA We follow Husson (1962:251) in ay specific name molossus to our specim species which were reported earlier Allen (1902:91) under the name obscura: i melinck (1940:72) and Musso (1962 ported their specimens under the species majo Examination of material from the Lesser Antlli mainland South America, Central America, and México leads us to believe that enough inter population variation exists to warrant the use of the trinomial at this time, although the species is still badly in need of a thorough review. The specimens from Margarita Island closely resemble typical material of M. m. from St. Lucia, Grenada, St. Vincent (all in MCZ), Trinidad (TTU), and a large series from Dominica (KU). Also material from adjacent parts of the mainland (Jusepin, Monagas, and San Antonio del Golfo, Sucre) seem to be assign- able to this subspecies (Table 5). A series of specimens from Maripa, Bolivar, are somewhat smaller in size than typical molossus but this may be simply a result of local variation (Jones, Smith, and Turner, 1971:23-24). Further delineation of the range of M. m. molossus on the mainland and the relationship of the Maripa specimens will have to wait until more material is available; however, it is sufficient at present to say that M. m. molossus occurs on the mainland of northeastern South America. molossus ZOOGEOGRAPHIC COMMENTS Koopman (1958) has discussed the zoogeography and distribution of bats on islands off the northern coast of South America. We take this opportunity to update Koopman’s data based on recent publications and re-examine his conclu- sions in light of this new information. The islands in the following list are those considered by Koopman (the first number, in parentheses, represents species reported by Koopman, 1958: the second number refers to species presently known from the island followed by the references upon which this number is based): Aruba, (3) 4 (Husson, 1960); Curacao, (6) 8 (Husson. 1960); Bonaire (3) 6 (Husson, 1960): Marga- rita, (7) 16 (this paper); Trinidad, (44+) 63 (Goodwin and Greenhall, 1961. 1962. 1964: Genoways, Baker, and Loregnard, 1973): Tobago. (16) 17 (Goodwin 1961): Grenada, (11) 12 (Jones and Phillips. 1970). We estimate the bat fauna of northern Venezuela and Greenhall. 74 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Trance 5. Selected external and cranial measurements of Molossus molossus from the Antilles and north- eastern South America. Superscript numbers indicate a sample size that differs from those given in the left-hand column. Breadth Condylo- — Lengthof —-Breadth Number Zygomatic of basal maxillary of rostrum and sex Forearm breadth braincase length toothrow at canines Dominica 7 38.3 10.5° 8.5° 14.8 529) 4.3 (37.2-39.2) (10.3-10.9) (8.4-8.7) (14.5-15.3) (5.76.0) (4.1-4.4) 8 2 37.8 10.0 8.3 14.1 5.6 4.0 (36.2-39.0) (9.8-10.1) (8.0-8.6 ) (13.8-14.6) (5.5-5.7) (3.9-4.1) St. Vincent 15 (023) — 10.2 8.7 14.6 Si 4.2 2 2(?) — 9.8, 9.9 8.8, — 13.8, 14.4 SIoSh5:5)57/ 4.0, 4.0 Grenada 29 — 9.7, 10.2 8.3, 8.6 13.7, 14.3 Bi a7 3.8, 4.1 Trinidad 10 3 38.1 10.8 8.9 15.1 6.1 4.5 (37.0-39.2) (10.4-11.1) (8.79.2) (14.7-15.4) (5.9-6.2) (4.24.6) 10 9 37.6 10.1 8.4 14.3 5.8 4.2 (35.5-39.2) (9.9-10.5) (8.0-8.7) (13.8-14.7) (5.46.2) (4.0-4.4) Margarita Island, Nueva Esparta, Venezuela it Bs — 10.0 8.6 14.6 5.9 4.1 19 38.6 — 8.6 14.9 5:9 4.0 San Antonio del Gulfo, Sucre, Venezuela 1 2 36.9 11.0 9.1 15.5 6.0 4.4 19 5:2 — 8.8 14.3 ay) 4.2 Jusepin, Monagas, Venezuela Sits 38.9 10.4 8.5 15.1 6.1 4.4 (37.3-39.9) (10.2-10.5) (8.4-8.6) (14.8-15.4) (6.0-6.3 ) (4.14.6) ay) 38.4 10.1 8.4 14.5 5.9 4.2 (37.3-39.5) (9.8-10.4) (8.3-8.7) (14.3-14.7) (5.8-6.0) (4.0-4.3 ) Maripa, Bolivar, Venezuela ay 33.8 10.0° 8.4 1397) 5.6 3.9 (5.45.7) (32.7-34.9 ) (9.8-10.1) to comprise 50-60 species based upon field investigations by Smith and previous records; sixty species of bats were reported from Surinam by Husson (1962). The only change in Koopman’s (1958) rank order of islands based on the number of chirop- teran species inhabiting them is Margarita, which is now known to have more species than Grenada. This could have been predicted by the fact that Margarita is a much larger and higher island as well as being a continental island rather than an oceanic island. (8.2-8.6) (13.5—13.9) (3.8-4.0) Among the 16 species which presently comprise the chiropteran fauna of Margarita Island there are two species of emballonurids, one noctilionid, two mormoopids, nine phyllostomatids, one vespertilionid, and one molossid. Within the family Phyllostomatidae, the subfamilies are repre- sented as follows: Phyllostomatinae, 2; Glosso- phaginae, 2; Carollinae, 1; Stenoderminae, 2; Desmodontinae, 2. Rhogeessa minutilla Miller, 1897 and Leptonycteris curasoae tarlosti Pirlot, 1965 were described from Margarita Island; neither is currently regarded as unique to the 1974 PISC CARN. SANG FOL. GLE INSECT. FRUG. AER. NECT. BATS FROM MARGARITA ISLAND, VENEAUELA —io.9) 30-34 35-43 494—54 Des, rot Mic, nic D Phy. ste Car. per. 0 Phy. dis Vam. hel Art, pla Art. cin Des. rot Phy. dis Art. pla Rhy. nas Noc. lab Sac. lep Pte. par Ept. bra Mor. meg Mol. mol Eum. gla Per. mac Sac. lep Mol. mol. Figure 3. Composite, two-dimensional niche matrix for the 16 bat species from Margarita Island and the 39 species actually captured on the adjacent Venezuelan mainland. The trophic role values are those employed by Wilson (1973) or only slightly modified. The model values for forearm length are those used by Fleming ef al., (1972). Each of the 49 cells of the matrix is divided into two parts, the upper represents species (abbreviated scientific names) and trophic role values for the Venezuelan mainland and the lower represents those for Margarita Island. At the left-hand margin appear the importance values (IV) for each trophic role in each of the two regions. The importance values are derived by dividing the total trophic values by the number of bat species in each fauna and expressed as a per- centage. Crosshatching represents “niches” that are apparently not occupied by the known chiropteran fauna of these regions. Dots represent “mainland niches” that are apparently unoccupied by the chiropteran fauna of Margarita Island. The dotted cells for aerial insectivores from Margarita may represent sampling biases. island. All of these bats are Neotropical in Margarita Island are the three small. insectivorous their affinities and all are found on the adjacent families Natalidae. Thyropteridae. Furipteridae. Venezuelan mainland. and the frugivorous Sturnirinae (Phyllostomati- Notably absent from the chiropteran fauna of dae), all of which occur in the chiropteran 76 BULLETIN SOUTHERN CALIFORNIA fauna of the adjacent mainland. In addition it should be noted that no more than two species of each major taxonomic group are known from the island. The latter observation is particularly striking when considering the wide diversity of the Phyllostomatinae and Stenoderminae on the adjacent mainland (Fig. 3). Only in the case of Artibeus planirostris and A. lituratus is a genus represented by more than one species on Margarita Island. However, of the genera represented on the island, the following are represented on the mainland by two or more sympatric species: Peropteryx, Noctilio, Pteronotus, Phyllostomus, Glossophaga, Ar- tibeus, Rhogeessa, and Molossus. The apparent absence of some species of vespertilionids and molossids on Margarita may be an artifact due to the fact that these bats are not readily captured and extensive collecting in and around their potential roost-sites such as attics of buildings, rain spouts, and elsewhere, was not conducted. With the use of other collecting techniques we would anticipate the addition of other species of these two insectivorous families to the Margarita fauna. As noted above, all of the species which inhabit Margarita Island also occur on _ the mainland of northeastern Venezuela. Only 13 of the taxa from Margarita have been recorded on Trinidad; the three exceptions are Leptonycteris curasoae, Pteronotus parnellii fuscus, and Glosso- phaga longirostris longirostris. The latter two species are represented on Trinidad by different geographic races; P. p. rubiginosus with its affinities apparently with the Amazon Basin fauna (Smith, 1972:76) and G. |. major apparently independently differentiated on Trinidad and Tobago. Compared with the chiropteran fauna of Surinam (Husson, 1962), only six taxa occurring on Margarita Island are presently known from that country also. Four, six, and four species found on Margarita also inhabit Aruba, Curacao, and Bonaire, respectively. Seven species are shared by Margarita and Grenada islands. From the foregoing account, it should be apparent that the chiropteran fauna of Margarita Island represents an attenuation of a fauna characteristic of northeastern South America (in- cluding Trinidad). The representation of most major taxonomic groups of bats on Margarita Island, but in markedly reduced numbers when compared to the adjacent Venezuelan fauna, may reflect (1) Saccopteryx, Micronycteris, in mist-nets ACADEMY OF SCIENCES VOLUME 73 the area of the island, (2) the proximity of the island to the source area, and (3) the ecological diversity of the island. These three factors have been discussed by Koopman (1958, 1968), Mac- Arthur (1965), MacArthur and Wilson (1963, 1967), and others. More recently, McNab (1971), MacArthur, Diamond, and Karr (1972), and Fleming, Hooper, and Wilson (1972) have approached the problems of island faunas from the viewpoint of the ecological role of the individual species which comprise the fauna. Unfortunately, such studies are greatly limited by the paucity of available information on the ecological roles of many species; this is especially true of bats. Inasmuch as the entire adaptive radiation of Neotropical bats seems to have revolved around diverse feeding habits, the availability of particular food sources and food items of particular sizes would appear to be primary parameters in the chiropteran niche. McNab (1971) also con- sidered these two factors to be most important in the niche partitioning of tropical bat faunas. A consistent classification of chiropteran food habits has not been readily available in the past and in fact, the food habits of many species simply are not known. Wilson (1973) made a noteworthy attempt at such a classification. In our analysis of the chiropteran fauna of Margarita Island and the adjacent mainland we have followed his method of categorizing the various species into trophic categories. These categories are (Fig. 3): carnivores (CARN.— feeding on tetrapods); piscivores (PISC.—feed- ing on fish); sanguinovores (SANG.—feeding on blood); foliage gleaners (FOL. GLE.—feeding on insects on foliage or on the ground); aerial insectivores (AER. INSECT.—feeding on flying insects); frugivores (FRUG.—feeding on fruit); and nectarivores (NECT.—feeding on flowers, nectar, and/or pollen). For comparative purposes, we have categorized the trophic habits of the species comprising the fauna of Margarita and those of 39 species actually captured by Smith on the adjacent Venezuelan mainland. We have adopted Wilson’s (1973) trophic values in most cases or only deviated slightly from his assessed value. In addition, our values are placed at the specific level rather than at the generic level. A combined, two-dimensional niche matrix for the bat fauna of Margarita Island and the Vene- zuelan mainland is presented in figure 3. Several interesting points can be seen in this diagram. 1974 BATS FROM MARGARITA ISLAND, VENEZUELA TABLE 6. Importance values for the seven trophic roles of the chiropteran faunas of nor America and the Antilles. The number in parentheses represents the numt of species in each of the updated faunas; see text for lior Carn. Pisc. Sang. Fol. Gle. Aer. Insect Pru Aruba (4) 0.0 0.0 0.0 2:5 50.0 7.5 40 Curacao (8) 0.0 Tao 0.0 2.5 55.0 15.0 0.0 Bonaire (6) 0.0 0.0 0.0 1.6 50.0 21.7 6.7 Margarita (16) 0.0 3.8 11.9 ths) 43.1 22.5 10.6 Venezuela (39)! 3.8 1.5 7.4 12.6 40.8 26.9 6.9 Surinam (61) Sei/ 1.0 372. 10.8 48.0 23.8 9,5 Trinidad (63) 3.5 0.9 3.0 11.3 48.9 26.8 5.6 Tobago (17) 2.3 355) 0.0 7.6 40.6 40.0 5.9 Grenada (12) 0.0 5.0 0.0 4.2 45.0 31.6 14.2 0.0 5.0 0.0 2.5 29.2 18.3 Dominica (12)* ‘ Based on actual captures. * Based on Jones and Phillips (1970). The 39 mainland species occupy 29 of the 49 cells of the matrix and the 16 bats from Margarita occupy 20 of these cells. At this time it is difficult to comment on the “unoccupied” cells in both faunas. Future studies no doubt will reveal these to be either nonexistent or to be occupied by other terrestrial vertebrates. With respect to bat faunas in general, we doubt that the six cells in the upper left-hand corner (small-sized carnivores, piscivores, and sanguinovores) exist for bat spe- cies. This is based on problems of body size versus prey size and various physiological param- eters. The unoccupied cells on the right-hand side of the matrix either reflect sampling biases or do not exist for bats in northeastern South America. With the exception of the sanguinovore cells, these unoccupied cells are indeed occupied by bats in other faunal regions. McNab (1971:354) notes that carnivorous species of bats, other than fishing species, are generally absent from island faunas. He cites species of the genus Phyllostomus as the only carnivorous species on Margarita and Tobago. Phyllostomus discolor is the species occurring on Margarita, and to our knowledge is not a carnivorous species (Fleming ef al., 1972:559; Goodwin and Greenhall, 1961:238). Therefore, the carnivorous cells of Margarita Island at this time appear to be unoccupied by bats. The apparent absence of carnivorous bats on islands is further illustrated in table 6. As can be seen in figure 3, the bulk of the bat faunas of Margarita and the adjacent main- land occupy the aerial insectivore-frugivore cells, 67.5 and 65.6 percent, respectively. In represen- tative chiropteran faunas from northern South 45.0 America and the Antilles (Table 6) these trophic roles are also predominant with the aerial insecti- vore complement comprising the larger of the two. Koopman (1958:434) points out that the eco- logical differences among the various islands off the northern coast of South America are rather extreme. Aruba, Curacao, and Bonaire, the most xeric of these islands, have a dispropor- tionately large aerial insectivore and nectarivore fauna compared to that of Trinidad, Tobago, and Grenada, which are the most mesic of the islands. Margarita Island, which has both xeric and mesic conditions, is intermediate, at least with regards to the nectarivorous trophic role. The foliage glean- ing trophic role appears to be reduced on all islands except Trinidad. There is a marked disharmony (MacArthur and Wilson, 1967:175— 78) in the bat faunas of Aruba, Curacao, Bonaire. Grenada, and Dominica, as compared with the mainland fauna, as illustrated by the absence of carnivores and sanguinovores (Table 6). The Margarita fauna is disharmonic with the mainland in lacking carnivorous species as is Tobago in lacking sanguinovorous species. It should be pointed out that Trinidad essentially represents an extension of the mainland chiropteran fauna with all trophic roles being represented. In conclusion, the bats of Margarita Island comprise a depauperate chiropteran fauna which is most closely allied to the adjacent Venezuelan bat fauna. Ecological diversity and of the island seem to be the most relevant factors influencing the composition of the Margarita fauna. The proximity to the source with regards to the islands off the the size area, at least north coast of Venezuela, does not appear to be a major factor for these respective faunas. We believe that the Margarita bat fauna is a result of repartitioning limiting reduced size of the or expansion of the mainland niches on the island. In examining increasing population density on species-poor islands MacArthur et al. (1972) note that species utilized more space (that is, wider range of altitude, or habitats, or vertical foraging strata, or any combination of these; wider range of foraging techniques; or broader utilization of food items). Although we do not have sufficient data relating to population densities of the chirop- teran species from Margarita Island, we suspect that they also utilize wider niche space which might account for the reduced bat fauna of this island. LITERATURE CITED Allen, G. M. 1902. The mammals of Margarita Island, Venezuela. Proc. Biol. Soc. Washington, 15:91-97. Davis. W. B. 1973. Geographic variation in the fishing bat, Noctilio leporinus. J. Mamm., 54: 862-874. Fleming, T. H., E. T. Hooper, and D. E. Wilson. 1972. Three Central American bat communi- ties: structure, reproductive cycles, and move- ment patterns. Ecology, 53:555—569. Genoways, H. H., R. J. Baker, and R. S. Loregnard. 1973. Two species of bats new to the fauna of Trinidad. Mammalia, 37:362—363. Goodwin, G. G. 1958. Bats of the genus Rhogeessa. Amer. Mus. Novit., 1923:1—17. Goodwin, G. G., and A. M. Greenhall. 1961. A review of the bats of Trinidad and Tobago. Bull. Amer. Mus. Nat. Hist., 122:187-302. 1962. Two new bats from Trinidad, with comments on the status of the genus Mesophylla. Amer. Mus. Novit., 2080: 1-18. 1964. New records of bats from Trinidad and comments on the status of Molossus trini- tatus Goodwin. Amer. Mus. Novit., 2195: 1-23. Hummelinck, P. W. 1940. A survey of the mam- mals, lizards, and mollusks. Studies on the Fauna of Curacao, Aruba, Bonaire and the Venezuelan Islands, 1:59—108. 8 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Husson, A. M. 1960. De Zoogdieren van de Neder- landse Antillen. Rijksmuseum van Naturalijke Historie, Leiden, viii + 170 pp. 1962. The bats of Surinam. handel., 58: 1-282. Zool. Ver- Jones, J. K., Jr., and C. J. Phillips. 1970. Comment on systematics and zoogeography of bats in the Lesser Antilles. Studies on the Fauna of Curacao and other Caribbean Islands, 32:131- 145. Jones, J. K., Jr., J. D. Smith, and R. W. Turner 1971. Noteworthy records of bats from Nica- ragua, with a checklist of the chiropteran fauna of the country. Occas. Papers Mus. Nat. Hist., Univ. Kansas, 2: 1-35. Koopman, K. F. 1958. Land bridges and ecology in bat distribution on islands off the northern coast of South America. Evolution, 12:429—439. 1968. Taxonomic and distributional notes on Lesser Antillean bats. Amer. Mus. Novit., 2333:1-13. LaVal, R. K. 1973. Systematics of the genus Rhogeessa (Chiroptera: Vespertilionidae). Occas. Papers Mus. Nat. Hist., Univ. Kansas, 19:1—47. MacArthur, R. H. 1965. Patterns of species di- versity. Biol. Rev., 40:510—533. MacArthur, R. H., J. M. Diamond, and J. R. Karr. 1972. Density compensation in island faunas. Ecology, 53:330-342. MacArthur, R. H., and E. O. Wilson. 1963. An equilibrium theory of insular zoogeography. Evo- lution, 17:373—387. 1967. The theory of island biogeography. Monogr. Pop. Biol., 1:xi + 1—203 pp. Marinkelle, C. J., and A. Cadena. 1972. Notes on bats new to the fauna of Colombia. Mammalia, 36:50-58. Matson, J. O., and F. P. Brown. 1974. Notes on some bats from a cave on Peninsula Paraguana, Venezuela. Bull. Southern California Acad. Sci., 13592 —93% McNab, B. K. 1971. The structure of tropical bat faunas. Ecology, 52:352-358. 1974 BATS FROM MARGARITA ISLAND, VENEZUELA Miller, G. S., Jr. 1897. Description of a new Power, D. M., and J. R, Tamsitt, 19 bat from Margarita Island, Venezuela, Proc. Phyllostomus discolor (Chiropter Biol. Soc. Washington, 11; 139. tidac). Canadian J, Zool., 51:46) _- 1913. Revision of the bats of the genus Sanborn, C. C. 1936. Records and m Glossophaga. Proc, U.S. Nat. Mus., 46:413— of Neotropical bats. Field Mus. Nat. H 429. Ser., 20:93—106. Musso Q., A. 1962. Lista de los mamiferos cono- ————, 1949, Mammals from the Rio | cidos de la Isla de Margarita. Mem. Soc, Gen, Peru. J. Mamm., 30:277-288 Nat. La Salle, 22: 163-180. Patten, D. R. 1971. A review of the large species of Artibeus (Chiroptera: Phyllostomatidae ) from western South America. Unpublished Ph.D. thesis, Texas A and M Univ., College Station, 175 pp. Phillips, C. J. 1971. The dentition of Glosso- phagine bats: development, morphological char- acteristics, variation, pathology, and evolution. Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 54: 1-138. Pine, R. H. 1972. The bats of the genus Carollia. Tech. Bull., Texas Agric. Exper. Stat., Texas A and M Univ., 8:1-125. Pirlot, P. 1965. Deux formes nouvelles de chirop- teres des genres Eumops et Leptonycteris. Le Naturaliste Canadien, 92:5—7. Pirlot, P., and J. R. Leon. 1965. Chiropteres de Vest du Venezuela. I. Région de Cumana et Ile de Margarita. Mammalia, 29:367—374. Smith, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Misc. Publ. Mus. Nat Hist., Univ. Kansas, 56:1—132. Thomas, O. 1911. The mammals of the tenth edition of Linnaeus; an attempt to fix the types of genera and the exact bases and localities of the species. Proc. Zool. Soc. London, pp. 120 158. 1928. The Godman-Thomas expedition to Peru. VII. The mammals of the Rio Ucayali. Ann. Mag. Nat. Hist., Ser. 10, 2:249-265. Valdez, R. 1970. Taxonomy and geographic varia- tion of bats of the genus Phyllostomus. Unpub- lished Ph.D. dissertation, Texas A and M Univ.. College Station. Wilson, D. E. 1973. Bat faunas: parison. Syst. Zool., 22:14—29. a trophi com- Accepted for publication February 14, 1974 MATERNITY HABITS OF MYOTIS LEIBIT IN SOUTH DAKOTA MERLIN D. TUTTLE! ABSTRACT: Badlands National Monument, South Dakota AND LAWRENCE R. HEANEY? Twelve active roosts containing 27 Myotis leibii were located in and near the Active and inactive roosts were characterized as to temperature inside and outside of the roosts, position and size of opening, slope facing, internal dimensions, and soil type. Two roosts each contained more than two individuals; all others held single individuals or a single female and her young. A possible instance of twinning was observed. Young bats were characterized as to weight, length of forearm, amount of pelage, and flying ability. Although Myotis leibii was first recognized more than 100 years ago, little is known concerning its summer roosting habits. Only one maternity colony has been reported and other observations of summer roosting have been limited. Koford and Koford (1948) reported a colony of at least 35 individuals, including juveniles, roosting under wallpaper in an abandoned wooden house in California. The only other summer colony re- corded was an estimated 12 individuals roosting behind a sliding shed door in Ontario (Hitchcock, 1955). Other bats of this species have been found roosting alone or in pairs under the fol- lowing circumstances: between boards leaning against the wall of a cottage (Stephens, 1945); in barns (Jones, 1964:84); under a loose strip of pine bark (Swenk, 1908:137; Jones, 1964:84); under a large flat rock at the edge of a quarry (Tuttle, 1964); under a stone on a hillside; in ero- sion crevices in the Badlands of South Dakota (Barbour and Davis, 1969:103): and in small pockets of eroded sandstone in the Badlands of Nebraska (Quay, 1948). Between 21 and 29 July 1972, we spent 30 man-hours searching for roosts of M. /eibii in and near the Badlands of South Dakota. Twelve of the more than 35 roosts (places containing guano) discovered were occupied by a total of 27 bats including seven lactating females, two postlactating females, one nonreproductive female, one adult male, four juvenile females, and seven juvenile males (Table 1). Five individuals escaped before sex or age could be determined. There was no evidence that the bats had altered the naturally formed roosts they occupied. Age of juveniles varied appreciably. The smallest weighed 1.6 gm and had a forearm length of 14.5 mm; the heaviest weighed 3.9 gm and had a forearm length of 31.0 mm. Only the largest 80 juvenile was volant and it could not maintain flight for more than 100 m. Six juveniles were nonvolant (forearms 14.5—24.5), and four were capable of slight forward progression (forearms 27.0-30.0) when released a few feet above ground level. Extremes for forearm lengths and weights for juveniles follow: females, 14.5 mm— 31.0 mm and 1.6 gm—3.9 gm; males, 17.5 mm—29.0 mm and 1.9 gm—3.8 gm. The smallest female had no body fur, and one male was only slightly furred; all others were moderately to completely furred. Only two roosts contained more than a single adult or an adult and its offspring. Roost number 3 held four lactating females nursing five non- volant juveniles; one female apparently was nurs- ing two juvenile males having the same weight and forearm length. Myotis austroriparius is the only member of this genus in the United States so far reported to produce twins (Barbour and Davis, 1969:61), but we interpret our findings as an indication of twinning in M. leibii. Roost 11 contained a postlactating female in addition to a lactating female and her single offspring. Adult males were absent from areas where females and their juvenile offspring were found; the single adult male, taken in roost number 1, was roughly 74 mi from the nearest other roost that was located. Roosts 1 and 2, at ca. 4 mi E Cottonwood, Jackson County, were in horizontal fractures in large, flat, siltstone boulders that measured 0.3 m <) ES G2 SECs: 0x 2. S56 ESE sens Bye), 0 xX 3. LOI <2: 3 > tua in cm Characteristics of roosts occupied by Myotis leibii. Maximum Inside Dimensions 1.5 9.0 14.0 cm e250 15.0 cm 2.0-3.5 & 15.0 31.0 cm DES EDS 5.0 cm 12555420:03 Allan Hancock Foundation, Univ. Southern Cal fornia, Los Angeles, California 90007. gica PO c AEs 96 BULLETIN SOUTHERN CALIFORNIA 120 105 120 105 Figure 1. by examination of gut contents. Intertidal species which were to be used in feeding studies were fixed immediately in the field. Specimens obtained by SCUBA diving were fixed as soon as possible, but in some cases up to six hours elapsed between collection and fixation. Following the termination of the cruise, the collections were taken to the Allan Hancock Foundation where final determinations were made by the second author. The crabs are housed in the Allan Hancock Foundation. Measurements given in this report refer to carapace length. References given with each species are to original descriptions; papers giving good illustrations and information on color and biology; and recent papers citing extensions of range. Our authority for the spelling of Spanish ACADEMY OF SCIENCES VOLUME 73 90 75 90 75 Collecting stations of Stanford Oceanographic Expedition 18. place-names is Index to Map of Hispanic America (“Millionth Map”) (U.S. Government Printing Office, 1945). SPECIES COLLECTED FAMILY COENOBITIDAE Coenobita compressus H. Milne-Edwards Cenobita compressa H. Milne-Edwards, 1837:241. Cenobita intermedia Streets, 1871:241. Cenobita panamensis Streets, 1871:241. Coenobita compressus: Boone, 1931:145, text-fig. 3; Glassell, 1937:242; Holthuis, 1954:16, text-figs. 4a,b; Bright, 1966:188, 189, text-figs. 4A-C; Haig, Hopkins, and Scanland, 1970:15, 27; Ball, 1972: 265 et seq. 1974 HERMIT CRABS FROM THE TROPICAL EASTERN PACIFIC TABLE 1. Station Data, Stanford Oceanographic Expedition 14 STATION LOCALITY DATE (196 POSITIO | Isla de Santa Clara, Golfo de Guayaquil, Ecuador 5~7 Apr x 10'S 2 Talara, Pert 8-9 Apr 4°34'S 3 Paita, Peru 10-13 Apr 5°05'S | 4 Salinas and vicinity, Ecuador 15-18 Apr 2°11’S—80759" 5 Manta, Ecuador 19-20 Apr 0°56'S—80 6 Punta Galera, Ecuador 22 Apr 0°50’N—80 7 Punta Sua, Ecuador 23 Apr 0°52’N—79 8 Atacames Reef, Ecuador 23 Apr 1°00’'N—79 9 Punta Barca, Bahia de Buenaventura, Colombia 26 Apr 3°50’N—77 10 Punta Mono, Isla Gorgona, Colombia 27 Apr 2°57’N—78 11 Straits between Islas Gorgona and Gorgonilla; Colombia 28 Apr 2°56’N—78 12 Bahia de Solano, Colombia 30 Apr—l May a. Punta San Francisco Solano 6°18’N—77° b. Punta Cotudo c. Punta Nabuga d. Muntas 6°16.4’N—77 6°22.8’N—77°2 13 Bahia de Cupica, Colombia 2 May a. Punta Cruces 6°39.3’N—77° b. Bahia Chicocoro 6°41’N—77 14 Balboa, Canal Zone and Panama City, Panama 5-6 May a. Islas Naos and Culebra, Canal Zone 8°54.8’N—79° b. Punta Paitilla, Panama City 8°58.1’N—79° 15 Isla Taboguilla, Bahia de Panama, Panama 7 May 8°48’N—79 16 Isla Montuosa, Panama 9 May 7°28’N—82° 17 Area of Islas Negritos and Cedro, Golfo de Nicoya, Costa Rica 11-12 May a. Isla Negrito-adentro 9°49’N—84° b. Isla Cedro 9°50.5’N—84° c. Puntarenas 9°59’N—84° 18 Bahia Brasilito, Costa Rica 13 May 10°25’N—85° 19 Punta Chiquirin and vicinity, Golfo de Fonseca, El Salvador 15-16 May a. La Union b. Punta Chiquirin c. Isla Chuchito 13°20’N—87° 13°18’N—87° 13°19.2’N—87° 20 Salina Cruz and vicinity, Oaxaca, México 20-22 May a. Bahia Ventosa 16°10.1’N—95 b. Salina Cruz harbor 16°9.5’N—95° c. Tartar Shoals 16°18.8’N—98° 21 Acapulco and vicinity, Guerrero, México 23-24 May 16°50’N—99° 22 Bahia Tenacatita, Jalisco, México 27 May 19°17’N—104 23 Bahia de la Magdalena, Baja California, México 2 Jun 24°24’N—112 Remarks: Specimens of Coenobita compressus collected and observed during Stanford Oceanographic Expedition 18 were the subject of a separate report (Ball, 1972). Distribution: Several Chilean records, Estrecho de Magallanes and northward, require confirmation. Paita, Peri, to Santa Rosalia, Golfo de California, Mexico; Bahia de la Magdalena on the outer Baja California peninsula; offshore islands including Gala- pagos, Isla del Coco, and Revillagigedos. FAMILY DIOGENIDAE Dardanus sinistripes (Stimpson) Pagurus sinistripes Stimpson, 1859:82. 30.6°W 24.5°W 31.9°W 31'W 31'W 14’°W 52’'W 52.2’'W S50'W 49'W SO’'W 47W 45.6°W "09"W 12.6°W 39'W 5W “SOW °04'W 7 a1 490 Dardanus sinistripes: Rathbun, 1910:556, 597. pl. fig. 2; Glassell, 1937:251; Haig, Hopkins. Scanland, 1970:16, 27. Dardanus imbricatus Rathbun, 1910:556, 597. pl fig. 3. Dardanus peruensis Balss. 1921:21. os BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Localities: Paita (sta. 3); Punta Galera (sta. 6); off Isla Gorgona (sta. 11); Punta Cotudo (sta. 12b); Isla Taboguilla (sta. 15); Bahia Brasilito (sta. 18); Salina Cruz (sta. 20b); Tartar Shoals (sta. 20c); El Morro and NE corner of La Roqueta, Bahia de Acapulco (sta. 21); La Manzanilla, Bahia Tenaca- Bahia de la Magdalena (sta. 23). Males 6.1 to 22.7 mm, females tita (sta. 22); Veasurements: 8.8 to 29.0 mm. Color description: Carapace varies from mottled salmon to tan. Eyestalks salmon, with a tuft of white setae just proximal to golden cornea; ventral side of stalk white, sometimes with two salmon-colored bands. white to light tan. Antennae uniform pale salmon. Chelipeds red with purplish tones and many purplish merus with a dark red band distally; smaller cheliped with coarse red and white hairs. Cutting edge of fingers white, with a narrow longitudinal line of red. Walking legs salmon or mottled salmon and white, with scattered purple tubercles; merus and carpus paler than the two distal segments. Second left walking leg with transverse ridges of purple crossing flattened surface of propodus and dactyl. Distribution: Bahia de Sechura, Pert, to Isla Tiburon, Golfo de California, México; outer Baja California peninsula as far north as Boca de Santo Domingo. Distribution for this species is subject to revision, however, since there is evidence that two closely related forms have been confused under the name D. sinistripes (Biffar and Provenzano, 1972: 799-800). Natural history: This species is widely distrib- uted both geographically and vertically. During Stanford Oceanographic Expedition 18 it was found in the littoral and to depths of 100 feet, usually on bottoms which were a mixture of sand and gravel. It may be locally very abundant (e.g., at sta. 18). The shells inhabited by this species are often enormous in relation to the size of the crab and it is sometimes difficult to imagine how the crab manages to move its shell. A variety of shells are occupied, although at depths greater than 50 feet there appears to be considerable uniformity in the type of shell occupied at a given locality. Commensal anemones were frequently present on the shell (stas. 15, 18, 20c, 21, 22). At Bahia de la Magdalena (sta. 23) a small speci- men had part of its shell coated with orange sponge, while the shells of three larger animals from the same locality bore a dense growth of a hydroid (Hydractinia sp.). This hydroid was in the area of the aperture on all three shells and entirely covered one of them. The zooids near the aperture are continuously moved around in the crab’s respiratory current when the crab is withdrawn; this Antennules tubercles; would presumably make the VOLUME 73 aperture a favorable place for the hydroid to grow. A large polynoid worm was living in one of the shells from sta. 23, and a porcelain crab, Porcellana paguriconviva Glassell, was living in the shell with a hermit from sta. 22. Petrochirus californiensis Bouvier Petrochirus californiensis Bouvier, 1895:6; Glassell, 1937:251; Steinbeck and Ricketts, 1941:454, pl. 12 fig. 1; Haig, Hopkins, and Scanland, 1970:25, 27; del Solar, Blancas, and Mayta, 1970:24. Petrochirus granulatus californiensis: Bott, 1955:53, pl. 5 figs. 7a,b. Localities: Isla Taboguilla (sta. 15); Isla Cedro (sta. 17b); Bahia Brasilito (sta. 18). Measurements: Males 23.8 and 29.5 mm, females 31.8 mm. Color description: Carapace pale, with a dark reticulate pattern. Eyestalks uniform reddish-brown with a hint of purple; distally a dark brown chevron, and a white band just proximal to the dark cornea. Antennules white with a longitudinal brown stripe laterally and mesially; brown at the base of the flagellar setae. Antennal flagella alternately banded brown and white, usually three to five brown segments alternating with three to five white segments. Chelipeds reddish-purple; cutting edge and tip of fingers white; mesial surface of both meri with two large maculations of deep purplish-brown at distal end, lateral surface with one such blotch at lower distal corner. Dactyl of walking legs dark reddish- brown, with abundant reddish-brown hairs on margins. Other segments of walking legs pale reddish-purple; merus with a large maculation of dark purplish-brown midway along dorsal margin. Distribution: Caleta La Cruz, Pert, to Punta Penasco, Golfo de California, México; north to Bahia de Santa Maria on the outer Baja California peninsula. Natural history: This species was found on bottoms of sand and gravel to a depth of 35 feet. It often carries a number of commensals within its shell. For example, one individual from sta. 15 with a carapace length of 31.8 mm, living in a shell of Vasum cestus (Broderip), shared it with two pairs of porcelain crabs, Porcellana cancri- socialis Glassell; a number of polychaetes; and several red polyclad flatworms, Emprosthopharynx opisthophorus Bock, ranging up to 20 mm in length. On the outside of the shell were barnacles, bryozoans, and algae. In the shell with a 23.8 mm Petrochirus from sta. 17b were two juvenile porcelain crabs, Porcellana paguriconviva Glassell, and a polyclad flatworm, Stylochus sp. A speci- men from sta. 18 bore several large commensal anemones on its shell. 1974 HERMIT CRABS FROM THE Aniculus elegans Stimpson Aniculus elegans Stimpson, 1859:83; Boone, 1931: 140, text-fig. 1; Walton, 1950:192; Haig, Hopkins, and Scanland, 1970: 16, 27. Aniculus longitarsis Streets, 1871:240. Localities: Salinas off Punta Mandinga (sta. 4); Manta (sta. 5); Atacames Reef (sta. 8); Punta Barca (sta. 9); Punta Cotudo (sta. 12b); Isla Taboguilla (sta. 15); Isla Naos (sta. 14a); Punta Paitilla (sta. 14b); Bahia Brasilito (sta. 18); Tartar Shoals (sta. 20c); Bahia de la Magdalena (sta, 23). Measurements: Males 19.9 to St.1) mm, non- ovigerous females 20.7 to 35.1 mm, ovigerous female 31.4 mm. Color description: Carapace shield with distinct grooves marking the various areas; protogastric and branchial areas deep red, others pink. Posterior carapace deep red with white splotches. Eyestalks uniform light tan; corneas dark brown. Eyescales reddish. Antennules uniform tan. Antennae uniform reddish-brown. Chelipeds mottled red and_ pink; transverse rings bordered with a fine red line and with a fringe of fine white hairs along their distal edge. Fingers with small, black, corneous spinules and tufts of coarse, dark red, white-tipped hairs. Walking legs mottled red and pink, dactyl red, propodus with a subdistal red band; transverse rings bordered with a fine red line and with long, red, white-tipped hairs. Distribution: Cabo de San Francisco, Ecuador, to Golfo de California, México (no precise locality in the Gulf has been specified, but the species occurs there at least as far north as Bahia de San Carlos); north to Bahia de la Magdalena on the outer Baja California peninsula. The known range is now ex- tended southward in Ecuador from Cabo de San Francisco to Punta Mandinga. Natural history: This species was collected by dredging in 13 fathoms (24 m) and by diving to depths of 35 feet, on a variety of substrates in- cluding sand and gravel, mixed rock and sand, rocky outcrops, and Pocillopora coral. Several times these animals were found clustered together in groups of two or three. The crab is frequently very small in proportion to the size of shell it occupies. Gut contents of three specimens from stas. 14a and 14b were examined and found to include unidentifiable organic matter and parts of a small crustacean. A 51.1 mm specimen from Isla Taboguilla had a pair of commensal porcelain crabs, Porcellana paguriconviva Glassell, and a 35.1 mm specimen from Bahia de la Magdalena harbored one crab of the same species. Trizopagurus magnificus (Bouvier) Clibanarius magnificus Bouvier, 1898:378. Clibanarius chetyrkini Boone, 1932:29, text-fig. 8. TROPICAL EASTERN PACIFIC Trizopagurus magnificu Forest, 19 figs. 2, 11, 18; Haig, Hopkins, and Scar 5 PA Localities: Manta (sta. 5); Punta Co 12b); Isla Montuosa (sta. 16); Isla Tabo 15); Isla Naos (sta. 14a); Isla Negrito (st Bahia Brasilito (sta. 18); El Morro, Bal Acapulco (sta. 21); Bahia de la Magdalena ( Measurements: Males 8.0 to 20.8 mm ovigerous females 6.3 to 15.2 mm, ovigerous femal: 6.4 to 17.1 mm. Color description: Shield brown with white spot posterior part of carapace salmon with white spot Eyestalks brown with white spots; cornea red. Proxi mal segment of antennules brown with white spot distal segment and flagellum uniform bright orange Antennae bright orange. Chelipeds brown with large white tubercles; cutting edge of fingers black. Walking legs brown with large spots varying from white to bright orange. Distribution: Isla de la Plata, Ecuador, to Golfo de California, México (apparently the southernmost part only); north to Bahia de la Magdalena on the outer Baja California peninsula; Archipiélago Galapagos. de Natural history: Trizopagurus was found mainly on rocky outcrops and on living heads of Pocillo- pora coral, from the intertidal zone to 30 feet. They seemed rather gregarious, with groups of seven or eight frequently observed. The crabs were sometimes so tightly wedged among the Pocillopora branches that it was very difficult to imagine how they were able to move. They oc- curred in a variety of shell types. At sta. 14a several Trizopagurus were observed feeding by scraping at the rocks with both cheli- peds. They appeared to be eating fine algae. Gut contents of 11 individuals from five localities included coarse sand and shell fragments, unidenti- fiable fine organic matter and small pieces of algae, two of which appeared to be Gelidiuin pusillum (sta. 12b) and Polysiphonia sp. (sta. 21). Clibanarius panamensis Stimpson Clibanarius panamensis Stimpson, 1859:84: Steinbeck and Ricketts, 1941:454, pl. 16 fig. 1: Holthuis. 1954:23, text-figs. 7. 8; Bott. 1955:53. pl. 5 figs. 6a,b; Haig, Hopkins. and Scanland. 1970:17. 27 Localities: Punta Barca (sta. 9): Muntas (sta. 12d): Isla Chuchito (sta. 19c). Males ovigerous females 11.5 to 11.9 mm. Color description: Carapace mottled and white. Eyestalks uniform light tan of green. Antennules olive drab: fl longitudinal stripe of bright orange at base of setae. 39.8 22.8 11.0 to 17.3 mm. ovigerous female Measurements: mm. non- olive drat 100 Antennae uniform olive drab. Chelipeds brown with longitudinal orange stripes; manus and fingers cov- ered with white tubercles. Ground color of walking legs dark brown, with stripes, varying in color from orange to white, running the entire length of each segment; outer surface with four or five light stripes on merus, four on carpus, propodus, and dactyl. On the dark light stripes are subequal in width. Distribution: sla de la Correa near Capon, Pert, to Santa Rosalia, Golfo de California, México; north to Bahia de Santa Maria on the outer Baja California peninsula. Natural history: This species was found inter- tidally on fine sediment in protected areas. At Punta Barca several crabs were observed on the bottom of an intertidal stream, moving their chelipeds rapidly back and forth over the sandy bottom while apparently feeding. Periodically a cloud of particles was shot out anteriorly. The gut of one of these animals was packed solidly with mud and fine organic matter cementing some larger particles of sand. A few small fragments propodus the and of algae were present. Clibanarius albidigitus Nobili Clibanarius albidigitus Nobili, 1901:24; Holthuis, 1954:25, text-fig. 9; del Solar, Blancas, and Mayta, 1970:23. Localities: Paita (sta. 3); Isla de Santa Clara (sta. 1); Punta Brava near Punta Mandinga (sta. 4); Manta and nearby Punta Mal Paso (sta. 5); Punta Galera (sta. 6); between Islas Gorgona and Gor- gonilla (sta. 11); Punta Mono (sta. 10); Punta Barca (sta. 9); Isla Montuosa (sta. 16); Isla Tabo- guilla (sta. 15); Punta Paitilla (sta. 14b); Isla Chuchito (sta. 19c). Measurements: Males 3.2 to 10.8 mm, _ non- ovigerous females 3.5 to 7.5 mm, ovigerous females 3.1 to 7.1 mm. Color description: Carapace mottled olive drab and tan. Eyestalks olive drab, cornea black. Antennular peduncles olive drab, flagellum orange. Antennae orange. Chelipeds olive drab with white tubercles. Walking legs olive drab with white tubercles; outer and inner sides of dactyls white. Holthuis (1954:27), describing the color of speci- mens from El Salvador, noted that they had a longitudinal white stripe on the propodus, carpus, and merus of the walking legs (fig. 9c) in addition to the characteristic broad, longitudinal white stripe on the dactyl. He added that in adults the white stripe was often lacking on the propodus, carpus, and merus (fig. 9b). In a series of more than 20 speci- mens from El Salvador collected during Stanford Oceanographic Expedition 18, the situation is exactly as described and illustrated by Holthuis, with most, although not all, of the crabs longitudinally striped BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 with white on all four segments. On the other hand, specimens collected further south, almost without exception, have only the very characteristic broad white stripe on the dactyl and no trace of it on any other segments of the walking legs. In a series of 56 specimens from Punta Paitilla a few very small individuals show some lack of dark pigment on the propodus. Distribution: Caleta La Cruz, Peri (del Solar, Blancas, and Mayta, 1970; other records listed by these authors are in error) to La Libertad, El Salvador. The known range is now extended southward in Pert from Caleta La Cruz to Paita. Natural history: This species appears to be highly adaptable, being found higher in the intertidal than any other of the marine hermits encountered, and existing over a considerable range of temperatures. It is typically found in rocky tidepools and seems to be especially abun- dant on rocky intertidal flats. A wide variety of mollusk shells is occupied at various localities. Calcinus obscurus Stimpson is usually found in the same environment, although its distribution is centered slightly lower in the intertidal; within the area of overlap the two species are usually found grouped together. At times when the Clibanarius are inactive they form piles consisting of up to a few hundred individuals under rocks or out in the open. Feeding is done by scraping at the surface of rocks with the chelipeds. Gut contents of 35 specimens from six localities were examined. In almost all cases the gut was packed with fine sediment and algal fragments (Bostrychia radicans being the only identifiable alga); diatom frustules and one copepod exoskeleton were also found. In the field this species was seen feeding on Enteromorpha and on Padina or something encrusting it. Clibanarius digueti Bouvier Clibanarius digueti Bouvier, 1898:379. Locality: Bahia de la Magdalena (sta. 23). Measurements: Male 10.0 mm, ovigerous female 8.5 mm. Color description: Carapace mottled tan. Eyestalks olive drab to dark green; cornea with white spots on a dark background. Antennules olive drab to dark green, with bright orange flagellum. Antennal peduncles red, flagellum orange. Chelipeds brown with abundant bluish-white tubercles; fingers orange. Walking legs brown with many white spots; dactyl reddish-orange distally, shading to brown at base of segment. Distribution: Throughout Golfo de California, Mexico. It is now recorded for the first time from the outer side of the Baja California peninsula. 1974 HERMIT CRABS FROM THE Habitat; This species was found in rocky pools in the lower intertidal. Calcinus obscurus Stimpson Calcinus obscurus Stimpson, 1859:83; Nobili, 1901: 26; Holthuis, 1954: 20, text-figs. 5, 6. Localities: Isla de Santa Clara (sta. 1); Salinas and vicinity (sta. 4); Punta Galera (sta. 6); between Islas Gorgona and Gorgonilla (sta. 11); Punta Mono (sta. 10); Punta Barca (sta. 9); Punta San Francisco Solano (sta. 12a); Punta Cruces (sta. 13a); Isla Montuosa (sta. 16); Isla Taboguilla (sta. 15); Isla Naos (sta. 14a); Punta Paitilla (sta. 14b); Isla Negrito (sta. 17a); Bahia Brasilito (sta. 18); Punta Chiquirin (sta. 19b). Measurements: Males to 18.0 mm, non-ovigerous females to 14.0 mm, ovigerous females 5.6 to 10.8 mm. Color description: Carapace shield olive green with punctae varying from light blue to white. Posterior carapace tan and white. Eyestalks olive green with a broad white ring just proximal to black cornea. Eyescales orange. Antennules olive green proximally; distal segment of peduncle and flagellum orange. Antennae uniform bright orange. Chelipeds olive drab; margins orange; cutting edge of fingers white. Walking legs vary from olive drab to brown, with whitish punctae; dactyl with a distal orange patch at base of nail and often with varying amounts of orange subproximally (in small specimens the dactyl generally has a proportionately greater area of orange than in large individuals). Distribution: Bahia de Santa Elena, Ecuador, to La Libertad, El Salvador. Natural history: This species is widely distrib- uted and often very abundant in the lower intertidal. Its intertidal distribution overlaps that of Clibanarius albidigitus, and the food and habitat requirements of the two species appear quite similar. Many types of mollusk shells are occupied by Calcinus obscurus. At several stations (especially 9 and 10) many large C. obscurus were out on the upper surface of rocks with the aperture of the shell facing upward and the crab withdrawn inside. Reese (1969:349, text-fig. 3A,B) reported similar be- havior for Calcinus laevimanus (Randall) and Clibanarius corallinus (H. Milne-Edwards) at Eniwetok Atoll, Marshall Islands. Feeding usually appears to involve scraping Most of cheliped fine algae and detritus from the rocks. the work is done by the small right while the much larger left cheliped is mainly as a support. These crabs were observed feeding on Padina or the material encrusting it (sta. 1), and on Gelidium pusillum (sta. 12a). used TROPICAL EASTERN PACIFIC 3) ral from { of the packed with fine sand and unidentifial matter with Identifiable Gut were examined contents Most put some recognizable algal fr material included Monostroima adoreanum (sta. 1), Gelidium sp. (sta what appeared to be fragments of small cru ceans (stas. | and 4). Calcinus californiensis Bouvier Calcinus californiensis Bouvier, 1898:380; Glassell 1937:252; Haig, Hopkins, and Scanland, 1970:16 Bille ?Calcinus californiensis: Chace, 1962:627, text-figs. Sh Gh Localities: NE corner of Isla Roqueta, SW of Bahia de Acapulco (sta. 21); La Manzanilla, Bahia de Tenacatita (sta. 22); Bahia de Ja Magdalena (Star 023) Measurements: Males 4.2 to 13.7. mm, non- ovigerous females to 11.4 mm, ovigerous female 4.2 Color description: Carapace shield olive drab with bluish punctae and orange margins. Posterior carapace mostly white. Eyestalks olive drab with a broad white ring just proximal to black cornea. white. Antennules olive drab proximally: distal seg- ment of peduncle and flagellum orange. reddish-orange. Chelipeds olive drab with punctae; margins reddish-orange; cutting fingers white. Walking legs bright reddish-orange: dactyl solid reddish-orange, without rings or patches of another color; merus of first walking legs with a large olive spot laterally. Distribution: México: Acapulco to Isla San José. Golfo de California, and to Bahia de la Magdalena, west coast of Baja California: Isla Clipperton. Habitat: This species was found on boulders interspersed with sand, from the lower intertidal to a depth of approximately 10 feet. mm. Eyescales Antennae bluish edge of Isocheles, sp. indet. Localities: Salinas at Punta Mandinga (sta. 4): near Puntarenas (sta. 17c). Measurements: Males 12.0 and 12.8 mm (sta. 4): male 8.2 mm, females 6.5 and 10.0 mm (sta. 17c). Color description: Carapace shield mottled brown and white. Eyestalks white with two longitudinal brown stripes, one dorsal and the other cornea black. Antennules white with two longitt brown stripes. one dorsal and the on the distal segment of the peduncle. flagella with a greenish band on each segment. of chelipeds mottled brown and whit with broad elongate areas of brown. c brown. Merus and carpus other of walkin 102 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES propodus brown dorsally and white ventrally, dactyl brown and white. The above notes were taken on specimens from Punta and markings of the individuals from Puntarenas are much the same, with the following exceptions: the longitudinal stripes on the eyestalks appear black instead of brown; the walking legs are tan with some green, and there is a broad white band at the distal end of the propodus and a little white at the proximal end of the dactyl. Habitat: Low intertidal, sand and = scattered rocks (sta. 4). At the water’s edge on a gently sloping sandy beach, low in the intertidal (sta. lsc) Remarks: Three species of /socheles are cur- rently recognized from the eastern Pacific, and still others are to be described by J. Forest of the Muséum National d'Histoire Naturelle (Paris), who is revising the genus (Forest and Saint Laurent, 1968:107). Until this revision is com- pleted the status of the “Te Vega” specimens must remain uncertain. Mandinga. The color Paguristes spp. Paguristes perrieri Bouvier, P. anahuacus Glassell, and five undescribed species of this genus were col- lected during Stanford Oceanographic Expedition 18. These will be treated in the second part of the present study. FAMILY PAGURIDAE Pagurus perlatus H. Milne-Edwards Pagurus perlatus H. Milne-Edwards, 1848:60; Haig, 1955:18, 21, text-figs. 3, 4; Haig, 1968:21; del Solar, 1970:46 [Pagarus]. Bernhardus obesocarpus Dana, 1852:445; Dana, 1855: pl. 27 figs. S5a-d. Locality: Paita Harbor (sta. 3). Measurements: Small male 5.5 mm. Color description: Carapace mottled reddish-brown and white. Eyestalks olive with fine yellow spots; cornea brown with gold flecks. Antennules with yellow spots on a transparent background; a large, dark green spot on distal part of terminal segment of peduncle, and a similar spot on flagellum. Antennal scale and peduncle with fine yellow spots on a tan background; flagellum irregularly ringed, usually with three brown segments alternating with a white segment. Proximal articles of chelipeds mottled brown and white; chela mostly white with some tan. Merus and carpus of walking legs mottled tan and white, changing to mostly white on propodus; dacty] white. Distribution: Puerto Corral, Chile, to department VOLUME 73 Peru. Unlike the other hermit crabs treated in this paper, Pagurus perlatus is a warm- temperate rather than tropical species. Habitat: This species was collected on a bottom of fine sand at a depth of approximately 10 feet. of Tumbes, Pagurus spp. (miamensis group) Forest and Saint Laurent (1968:116) designated as the ‘“miamensis group” a number of species of Pagurus with close affinities to P. miamensis Provenzano of the tropical West Atlantic. Seven species belonging to this group were collected during Stanford Oceano- graphic Expedition 18: Pagurus lepidus (Bouvier), P. benedicti (Bouvier), P. galapagensis (Boone), P. villosus Nicolet, and three undescribed species. We shall treat these forms in the third part of the present study. Pylopagurus varians (Benedict) Eupagurus varians Benedict, 1892:24. Pylopagurus varians: Glassell, 1937:253; 1954:141, 152, pl. 42 E-H. “Stag-horn”: Smith, 1966:30, 2 text-figs. Locality: Bahia de la Magdalena (sta. 23). Measurements: The single specimen, which had been kept alive in an aquarium, was eaten by another aquarium animal before it could be preserved and measured. Color description: Shield orange; posterior carapace with reddish-brown spots on an orange background. Eyestalks reddish-brown; cornea bright orange. Basal segments of antennules clear, distal segment of peduncle with alternating reddish-brown areas and dorsal white spots; flagellum reddish-brown. Antennal peduncle transparent except acicle, which has alter- nating transverse bands of white and reddish-brown; flagellum with three to five reddish-brown segments alternating with a white segment. Merus of large cheliped mottled red and white, carpus pink with a few deep red tubercles, chela reddish-brown to orange. Merus and chela of small cheliped mottled red and white, carpus with distinct transverse red and white bands. Walking legs mostly reddish-brown with a white area at distal end of each segment. Distribution: Islas Secas, Panama, to Bahia de Tepoca on the east side of Golfo de California, Mexico, and Arena Bank to Isla Angel de la Guarda on the west side of the Gulf. It is now recorded for the first time from the outer side of the Baja California peninsula, and also in somewhat shallower water than previously: the recorded bathymetric range is 6 to 100 fathoms (11 to 183 meters). Natural history: The specimen was taken on a mixed bottom of sand and rock at a depth of approximately 20 feet. The shell of this animal had been covered, and perhaps replaced, by a Walton, 1974 HERMIT CRABS FROM THE white hydrocoral which formed five antler-shaped branches. When turned completely upside down on these branches, the crab seemed to be unable to right itself. The hydrocoral had a number of amphipods associated with it. The association of Pylopagurus varians with a hydrocoral was briefly mentioned by Walton (1954), and also by Glassell (1937) who referred to it as a “bryozoan growth.” Smith (1966) discussed the association in more detail and illustrated the crab with its carcinoecium. His specimen from Golfo de California, living with a many-branched hydrocoral, was not mentioned by name but one of us: (J. Haig) has seen it and confirmed its identity. ACKNOWLEDGMENTS We wish to thank the participants in Stanford Oceanographic Expedition 18 who assisted in the collection of hermit crabs, and those persons who made identifications of other organisms: Judith Terry, mollusks; Stephen Prudhoe, polyclads; and Robert Waaland, algae. The expedition aboard “Te Vega” and the first author’s observations on the hermit crabs were made possible by NSF grants GB-6870 and GB-6871 to Stanford University. Earlier studies by the second author, which facilitated identifications and yielded other information included in this paper, were supported by NSF grants G-21956 and GB-2039 during 1962-1964. LITERATURE CITED Ball, E. E. 1972. Observations on the biology of the hermit crab, Coenobita compressus H. Milne Edwards (Decapoda; Anomura) on the west coast of the Americas. Rev. Biol. Trop., 20:265— 273. Balss, H. 1921. Results of Dr. E. MjGbergs Swedish scientific expeditions to Australia 1910-13. XXIX. Stomatopoda, Macrura, Paguridea und Galatheidea. K. Svensk. Vetensk. Handl., 61 (10) : 1-24. Benedict, J. E. 1892. Preliminary descriptions of thirty-seven new species of hermit crabs of the genus Eupagurus in the U.S. National Museum. Proc. U.S. Nat. Mus., 15:1—26. Biffar, T. A., and A. J. Provenzano, Jr. 1972. A reexamination of Dardanus venosus (H. Milne Edwards) and D. imperator (Miers), with a TROPICAL EASTERN PACIFIC description of a new species of Darda the western Atlantic (Crustacea, Diogenidac). Bull. Mar. Sci., 22:7 Boone, L. 1931. A collection of anom i macruran Crustacea from the Bay of Panar and the fresh waters of the Canal Zone. Bul Amer. Mus. Nat. Hist., 63:137—189. ———. 1932. The littoral crustacean fauna of the Galapagos Islands. Part I]. Anomura. Zoologica 14: 1-62. Bott, R. 1955. Dekapoden (Crustacea) aus EI Salvador. 2. Litorale Dekapoden, ausser Uca. Senckenbergiana Biol., 36:45—70. Bouvier, E.-L. 1895. décapodes recueillis en Basse-Californie par M. Diguet. Bull. Mus. Hist. Nat., Paris, 1:6—8. Sur une collection de crustacés 1898. Sur quelques crustacés anomoures et brachyures recueillis par M. Diguet en Basse- Californie. Bull. Mus. Hist. Nat., Paris, 4:371- 384. Bright, D. B. 1966. The land crabs of Costa Rica. Rev. Biol. Trop., 14: 183-203. Chace, F. A., Jr. 1962. pod crustaceans of Clipperton U.S. Nat. Mus., 113:605-635. The non-brachyuran deca- Island. Proc. Dana, J. D. 1852. Crustacea. United States Ex- ploring Expedition during the years 1838, 1839. 1840, 1841, 1842. . . Vol. 13(1). Philadelphia. viii + 685 pp. 1855. Crustacea, Atlas. United States Ex- ploring Expedition during the years 1838, 1839, 1840, 1841, 1842. Vol. 14. Philadelphia. 27 pp., 96 pls. del Solar, E. M. 1970. Crustaceos braquiuros (can- grejos), anomuros y estomatopodos de las zonas nerito-pelagica y litoral de Tumbes. Bol. Soc. Geogr. Lima, 89:40-48. 1970. 53 pp. del Solar, E. M., F. Blancas, and R. Mayta. Catalogo de crustaceos del Peru. Lima. Forest, J. 1952. Contributions 4 la revision des crustacés Paguridae. J. Le genre Trizopagurus. Mém. Mus. Hist. Nat., Paris, (A)5(1):1—40. Forest, J., and M. de Saint Laurent. 1968. Cam- pagne de la Calypso au large des cOtes atlantiques 104 de l'Amérique du Sud (1961-1962). 6. Crustacés décapodes: paguridés. Ann. Inst. Océanog., (n.s.) 45 (2):47-170. Glassell, S. A. 1937. The Templeton Crocker Ex- pedition. XI. Hermit crabs from the Gulf of California and the west coast of Lower California. Haipverd 955. Chile Expedition Anomura of Chile. 51(12):1-68. Reports of the Lund University 1948-49. 20. The Crustacea Lunds Univ. Arsskr., (2) 1968. A report on anomuran and brachy- uran crabs collected in Peru during cruise 12 of R/V Anton Bruun. Crustaceana, 15:19-30. Haig, J., T. S. Hopkins, and T. B. Scanland. 1970. The shallow water anomuran crab fauna of southwestern Baja California, Mexico. Trans. San Diego Soc. Nat. Hist., 16:13-31. Holthuis, L. B. 1954. On a collection of decapod Crustacea from the republic of El Salvador (Central America). Zool. Verhandel., Leiden, 23:1-43. Milne-Edwards, H. 1837. Histoire naturelle des crustacés, comprenant l’anatomie, la physiologie et la classification de ces animaux. Vol. 2. Paris. 531 pp. 1848. Note sur quelques nouvelles espéces du genre PAGURE. Ann Sci. Nat., (3) 10:59-64. Nobili, G. 1901. Viaggio del Dr. Enrico Festa nella repubblica dell’ Ecuador e regioni vicine. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 XXIII. Decapodi e stomatopodi. Boll. Mus. Zool. Anat. Comp. Univ. Torino, 16(415):1—58. Rathbun, M. J. Peru and the adjacent coast. Mus., 38:531-620. 1910. The stalk-eyed Crustacea of Proc. U.S. Nat. Reese, E. S. 1969. tidal hermit crabs. Behavioral adaptations of inter- Amer. Zool., 9:343—355. Smith, A. G. 1966. Stag-horns Pac. Disc., 19(5):30-31. and_ long-horns. Steinbeck, J., and E. F. Ricketts. 1941. Sea _ of Cortez; A leisurely journal of travel and research, with a scientific appendix comprising materials for a source book on the marine animals of the Panamic faunal province. Viking Press, x + 598 pp. Stimpson, W. 1859. Notes on North American Crustacea, no. 1. Ann. Lyc. Nat Hist., New York, 7:49-93. Streets, T. H. 1871. Catalogue of Crustacea from the Isthmus of Panama, collected by J. A. McNeil. Proc. Acad. Nat. Sci. Philadelphia, 23:238-243. Walton, B. C. 1950. Some new and rare Pacific pagurids. J. Washington Acad. Sci., 40:188—193. 1954. Anomura) two new species. 18:139-173. The genus Pylopagurus (Crustacea: in the Pacific with descriptions of Allan Hancock Pac. Exped., Accepted for publication September 15, 1973. SYSTEMATICS OF PTEROCLADIA MEDIA FROM CALIFORNIA JOAN -G. STEWART! Abstract: The finding of unilocular cystocarps on herbarium specimens of Gelidinm crinale var. luxurians from California requires that the plants be referred to the genu Pterocladia. Pterocladia media Dawson (1958) is expanded for this purpose. Gelidinm sinicola Gardner (1927) may possibly also belong here. Additional collections of this Plero cladia species are noted. The genus Pterocladia. J. Agardh (1851) was established for Gelidium lucidum (Turn.) Sond. Species in both genera are similar in vegetative morphology. Cystocarps in Pterocladia are uni- locular and in Gelidium, bilocular; a consequence of carposporophyte development from one surface of an axis, or from both. Studies of species re- ferred to these two genera (Fan, 1961; Feldmann and Hamel, 1936; Loomis, 1949; Okamura, 1934) have suggested characters by which non-cysto- carpic plants of one genus might be distinguished unequivocably from the other. Some distinctions have proved useful, but often there remain speci- mens of dubious or tentative assignment. Four assignments involving Gelidiuwm-Pterocladia col- lections from California are discussed here and an inclusive answer proposed. One such question arose during a study (Stewart, 1968) of the morphological variation relative to the ecology of Pterocladia in California. Speci- mens in certain collections were extremely narrow, and in addition, sparsely branched, compared with more regular pinnate branching and broader axes characteristic of P. capillacea. These specimens were set aside for further study and not included in the work on P. capillacea. Other collections of the distinctive entity have been made sub- sequently from several sites on the southern California coast. Pterocladia media Dawson (1958) was de- scribed from a single collection from La Jolla (Dawson, 15609, AHF 63142, 63143, two isotype sheets) and contained no fertile plants. He stated that these small plants differed in “size” from P. pyramidale (Gardner) Dawson (= P. capillacea). Pterocladia capillacea plants 10-20 cm in height are abundant intertidally but even much smaller specimens are regularly branched with the axes mostly as wide as in taller plants. Dawson (pers. comm.) commented that the comparative lack of branching and narrow axes were note- 105 worthy distinctions between the two species in San Diego. Attempts to locate additional material at the type locality were unsuccessful, and the species apparently was not known from other than the type collection. Gelidium crinale forma luxurians was described (Collins, 1906) from collections Mrs. Snyder from San Diego, having earlier distributed under this in Collins, Holden, and Setchell, Phycotheca Boreali-Americana No. 1138. Gardner (1927) referred some tetraspo- rangial plants from Santa Monica to this taxon and raised its status from “forma” to “variety.” Numerous California collections of small, narrow (thereby appearing terete), sparsely branched gelidiaceous algae have been identified as belong- ing to this variety. Sexual plants were not known from the Pacific coast. On several Dawson sheets in the Herbarium at Allan Hancock Foundation, there are comments indicating that he wondered whether they might better belong with Pterocladia on the basis of vegetative mor- phology. During a recent study of California Gelidium, three specimens attributed to G. crinale var. were found to have unilocular cystocarps (Fig. 1). This necessitates referring the collections of this variety, as known from California, to the genus Prterocladia. Another poorly defined taxon Gelidium sinicola Gardner (1927): described on the basis of a single non-fertile collection from San Fran- cisco, UC 276620. Since then the epithet has been applied inconsistently to several different small Gelidium spp. Measurement data (Table 1) made by been name luxurians is and general appearance of the type material of G. sinicola are similar to data concerning G. crinale Biology. Pr (i 1 Dept. Population and Environmental University of California, Irvine, California address: 1314 Ritter, Scripps Institution of Oceanog- raphy, P.O. 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Unilocular cystocarps on “Gelidiwm crinale var. luxurians” UC 262565. var. luxurians. Indeed, Gardner noted (p. 227, 1927), “This form of Gelidium crinale seems closely related to G. sinicola of this paper.” The similarity between vegetative specimens of Pterocladia media and Gelidium sinicola (Table 1) makes it highly probable that the two ought to be considered synonymous although final attribution must await the finding of sexual structures in plants resembling G. sinicola from central or northern California. Table 1 summarizes currently available infor- mation on the four entities outlined above. The only cystocarps observed were unilocular and occurred on plants previously identified as G. crinale var. luxurians. This taxon therefore must be transferred to the genus Pterocladia and the species with which the material can most appro- priately be placed is P. media. Other collections as discussed above of undetermined Pterocladia are now attributed to P. media. Pterocladia media Dawson Pterocladia media Dawson 1958: 68, pl. 21, fig. 3-4, pl. 24, fig. 11; Gelidium crinale forma luxurians Collins, in Collins, Holden, and Setchell, PBA 1138 (1903); Collins 1906: 111; Gardner 1927: 277, pl. 46, fig. 1, pl. 47, fig. 3 (as var. /uxurians). Redescription: Thalli 3—5 (7) cm high, axes growing in tufts, simple or branched, erect portions arising from a system of stoloniferous creeping parts attached by short peg-like haptera. Branching (when occurring) irregular and sparse, generally distichous, often secund or more regularly alternate or pinnate ter- minally. Lateral branchlets with or without slight constrictions at point of junction with proximal axis; all axes and branches slender, narrow, mostly SYSTEMATICS OF PTEROCLADIA MEDI of uniform dimensions from base to ay 100-200 pm thic to numerous, more often it broad, Rhizoidal inner mead relatively abundant seen throughout medull of several rows of pigmented cells, with similar size or with cells of inner twice as large as cells in the outer layer. Medul irregularly rOW rs | m shaped cells, fewer than cortical Tetrasporangia arranged in V-shaped rows apically or branches in some other sori plants, in without arrangement. Cystocarps develop and protrude or one surface of the apical region of a branch, typically with a single ostiole. Infrequent or rare, in sandy patches on rock sul strates, intertidal. Thalli are soft and lax, rather than cartilaginous, irregularly branched compared with Gelidium pusillum (Stackh.) Le Jolis which can be similar in height and habitat. Generally the latter is found in thicker clumps or dense mats. Pterocladia capillacea is more strikingly distichous and pinnate than P. media with precurrent stiff or and slender and main axes of larger dimensions than the ultimate branchlets and usually all parts are broader thus appearing more flattened than P. media. In conclusion, certain algae, non-fertile but Prero- cladia-like in vegetative considered as P. media. characters, can now be Southern California collec- tions of other Prerocladia-Gelidium species are some- times difficult to separate if cystocarps are absent. found on When Pterocladia-type cystocarps were some plants similar to Dawson’s P. media collection quite (but Dawson's identified as Gelidium crinale “Pterocladia” var. luxurians), use of for his entity was supported, and his description could then be expanded to include both formerly undetermined plants and plants incorrectly attributed to the genus Gelidium. Pterocladia media and P. capillacea occur intertidally in southern California in similar habitats but can generally be recognized one from the other by the degree of branching breadth. regularity and axis ACKNOWLEDGMENTS Support in the form of a Graduate Fellowship from the National Science Foundation is appreciated. I am grateful for the advice of P. S. Dixon on the presentation of this information. LITERATURE CITED Agardh, J. G. 1851. algarum. Lund.., Species genera et Vol. 2, Pt 2:337-504. Collins, F. S. 1906. the Phycotheca 8:104-113. New species. etc.. Boreali-Americana Rhodora, 108 1958. Notes on Pacific Coast marine Bull. So. California Acad. Sci., 57: Dawson, E. Y. algae VII. 65-80. Fan, K. 1961. Morphological studies of the Gelid- iales. Univ. California Publ. Bot., 32:315—368. Feldmann, J., and G. Hamel. 1936. Floridées de France. Gelidiales. Revue Algologique, 9:209— 264. Gardner, N. L. 1927. New species of Gelidium on the Pacific Coast of North America. Univ. California Publ. Bot., 13:273-281. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Loomis, N. H. 1949. New species of Gelidium and Pterocladia with notes on the structure of the thalli in these genera. Allan Hancock Found., Occas. Pap., (6): 1-8. Okamura, K. of Japan. 1934. On Gelidium and Pterocladia J. Fish. Inst. Tokyo, 29:41-67. Stewart, J. G. 1968. Morphological variation in Pterocladia pyramidale. J. Phycol., 4:76—84. Accepted for publication May 10, 1973. RELATIVE STEM WATER POTENTIALS OF THREE WOODY PERENNIALS IN A SOUTHERN OAK WOODLAND COMMUNITY JAMES P. SYVERTSEN! ABSTRACT: Dawn stem water potentials of Quercus agrifolia, Juglans california, and Heteromeles arbutifolia were measured with a pressure chamber during the drought of 1972. Individuals of a particular species growing at the top of a slope had lower water potentials than those at the canyon bottom. These differences became greater as the season progressed and soil drying occurred. During the summer months when evaporative demands were high, leaf water deficits were not overcome during the overnight equilibration period as soil-plant water potential gradients existed in the dawn hours. Juglans and Heteromeles on the south- facing slope were able to maintain higher diurnal stem water potential maxima than correspond- ing individuals on the north-facing slope. This apparent anomaly may result from the more xeric conditions on the south-facing slope inducing acclimation or providing selection pressure for drought resistance of these two species. Plant distributions depend on availability of water as much as any single environmental factor. The free energy of water in the soil and of that within plant cells is expressed by water potential (Slayter, 1967). Plant water potentials can be adequately estimated by measuring stem water potentials (SWP) in a pressure chamber (Scho- lander et al., 1965, Waring and Cleary, 1967). This instrument has been used in a variety of studies requiring data concerning plant water potentials (Hass and Dodd, 1972, Sucoff, 1972). It was my purpose in this study to compare the SWP of Quercus agrifolia Nee., Heteromeles arbutifolia M. Roem. and Juglans californica Wats. (Munz and Keck, 1959) growing at different slope positions and aspects in a Southern Cali- fornia Oak Woodland. This data could then be used to explain plant densities and distributions. Since the plant density on the north-facing slope was about six times greater than that on the south- facing slope, it was expected that individuals grow- ing at the top of the more xeric south-facing slope would have the lowest SWP. In the only other seasonal SWP study in an Oak Woodland, Griffin (1973), working in Cen- tral California, found that bottomland Quercus douglasti Hook. and Arn., QO. lobata Nee. and Q. agrifolia had higher water potentials than individuals growing in upland savannas. All three ‘ Dept. Biological Sciences, California State Univer- sity, Fullerton, California 92634, (Present address: Dept. Biology, New Mexico State University, Las Cruces, New Mexico 88003). i i] — 1974 STEM WATER POTENTIALS OF WOODY PERENNIALS Taste 1. Daily percent relative humidity minima Tas_e 3, Temperature data and precip: and temperature maxima on dates following the 12 — for the Puente Hills study area during August 1972 rainstorm in the Puente Hills study area. 1972 i ~ Minimum Maximum Precip, Femperature “¢ Precipitation Date % RV Temp, °C mm - Mean Mean I 12 74 37.9 8.64 Month max. min Mean Mu Min 13 60 26.7 = June 28.0 13.6 21.0 33.8 4 0.5) 14 42 26.7 = July 3255 lan 24-30 24035) e127 15 35 27.8 = Aug. Sil 5ienL6:5e) 24205 eA0iS) 6 22S 6S Sept. 28.3 14.6 21.3 32.2 10.0 6.5% Monthly mean minimum %RH = 36.4 species had higher water potentials following a wet season than after a dry season. The deciduous Q. douglasii on a north-facing slope had lower water potentials than those growing on the south- facing slopes. Maximum diurnal water potentials of the evergreen Q. agrifolia were as low as —27 bars in August of a dry year. Since this study was also conducted during a drought year, it was hoped that the Q. agrifolia data would be com- parable to Griffin’s and that additional insight to plant distributions could be gained from looking at the other two species. METHODS The study area was an east-west canyon on the Weaver Ranch in the Puente Hills of southeastern Los Angeles County, California. The vegetation on the opposing north-south-facing slopes was dominated by Q. agrifolia, J. californica, and H. arbutifolia. The understory was primarily com- posed of Sambucus mexicana Presl., Rhus inte- grifolia (Nutt) Benth. and Hook., R. ovata Wats., and R. diversiloba T. and G. (Munz and Keck, 1959). The canyon walls were approximately equal in height (100 m) and slope (50 percent) (Rowlands, 1972). I set up four sampling stations; one at the top and bottom of each of the two opposing slopes. At each sampling station, two to five healthy twigs were excised with a sharp razor blade from two comparable individuals each of Quercus, Taste 2. Total monthly precipitation in the Puente Hills study area from December, 1971 to August, 1972. Monthly Precip. (mm) Totals Dec 71 Feb 72 Apr 72 153.9 1.46 132. May 72 June 72 Aug 72 Del 0.51 8.64 Juglans, and Heteromeles, Quercus sampling meth- ods resemble those of Griffin (1973): twigs 8-15 cm long with at least three leaves. Heteromeles twigs were 10-15 cm long with 6-15 leaves; Juglans leaves were 15-25 cm long with 7-17 leaflets. All samples were collected in the pre- dawn or first light hours when the relative humid- ity was near its diurnal maximum and transpiration rates minimum. Samples from each species were collected normally twice a month from April through August, 1972. Preliminary tests which contrasted on-site mea- surements of SWP and those of stems transported to the laboratory in humidified polyethylene bags and measured 90 minutes significant differences. quently taken to the lab where measurements were made within 60 minutes after twig excision. later, indicated no All samples were subse- Sample materials were sealed into the pressure chamber and subjected to pressure increase rates of 0.5 bar sec—' as described by Waring and Cleary (1967) and Ritchie and Hinckley (1971). The balancing pressure was read at the first appearance of fluid on the reference surface of the twig or petiole. Relative humidity was recorded daily on a hygrothermograph located 10 km northeast of the site at California Polytechnical University, Pomona (Table 1). Relative humidity was also measured on sampling days at each station with a sling psychrometer. All other climatological data (Tables 2-3) were from the U.S. Dept. of Com- merce (1971-72); temperatures were from the Cal Poly Pomona Weather Station and the pre- cipitation information was from the Walnut Patrol] Station Weather Station approximately + north of the site. Both weather stations are located km RESULTS The data show that the individuals at the higher slope positions had signifi 110 2.8 S$ —-16 =-14 = 5.42 Ss i ©-10 = 78 E -6 = 25 -2 T2223 228 44 425 51 526 66 6-20 7-7 7-29 B15 DATE (month-day) Figure 1. Stem water potentials (SWP) in -bars of Quercus agrifolia growing on the north-facing slope. Open circles represent mean SWP of plants at the top of the slope and closed circles those at the canyon bottom: vertical lines represent standard deviations. (P <.05) lower SWP than those growing at the bottom slope positions (Figs. 1-6). These rela- tionships were not as apparent from January in the same drainage system as the study site but approximately 70 m lower in elevation. Differences in mean SWP (Figs. 1-6) were statistically analyzed using a randomized complete- block design (Steel and Torrie, 1960) analysis of variance in which the sampling dates were used as blocks. through April as they were during the summer months. Higher relative humidities and lower temperatures were observed throughout the study at the lower sampling stations. The few instances in which this slope-position difference was reversed were probably the result of sampling error. ' 1 Nw yo AS) (eo); - {) eee lt, oa Gi On B WATER POTENTIAL (-bars) o oo 5-26 6-6 620 77 729 8-15 DATE (month-day) 44 425 511 Figure 2. SWP of Quercus agrifolia growing on the south-facing slope. (See Fig. 1) BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 bars) WATER POTENTIAL ( 425 5-11 526 6-6 620 7-7 7-29 8-15 DATE (month-day) Figure 3. SWP of Juglans californica growing on the north-facing slope. (See Fig. 1) All SWP showed a significant (P < .U5) decrease from spring to summer (Figs. 1-6). July was the warmest, driest month of the summer as tempera- tures ranged from 12.7—40.5°C and there was no | precipitation (Table 3). Although the area re- | ceived 8.64 mm of precipitation the morning of | 12 August, it probably had no effect on soil | moisture in the root zone as the temperature | reached 32°C that afternoon and was followed |, by subsequent days of high temperatures (Table | 1). All SWP increased between 29 July and 15 | August. There is a difference (P < .07) between the SWP from the north and south-facing slopes. | Swnny @Ooon dey ron oe 1 ' ho ' an @ WATER POTENTIAL Chars) iS) —o— oO a —-O- 1 a 5-26 6-6 6-20 7-7 7-29 815 DATE (month-day) Figure 4. SWP. of Juglans californica growing on fil the south-facing slope. (See Fig. 1) 4-25 5-1] 1974 STEM WATER POTENTIALS OF WOODY PERENNIALS ~28 | 28 C 2% 4 26 -24 24 —-22 22 E fa 3-20 : 3-20 318 ’ | ° $ a 18 —- = -16 zs ry ao 5 : ! 5 a-12 ° | a -|2 cc c cc 43-10 | eo) = =z = -8 ; = 50) -6 -4 -2 5-26 6-6 620 7-7 7-29 8-15 DATE (month-day ) 4-25 5-11 Figure 5. SWP. of Heteromeles arbutifolia growing on the north-facing slope. (See Fig. 1) On most sampling days, Juglans and Heteromeles growing on the more xeric south-facing slope had higher SWP than that of the same species on the north-facing slope (Figs. 3-6). The lower SWP ‘of Juglans on the north-facing slope are more dramatic and are lower there than on the south- facing earlier in the spring (Figs. 3-4). Interspecific differences make SWP comparisons difficult unless these differences are accounted for (Boyer, 1967; Kaufmann, 1968). Ritchie and Hinckley (1971) reported that the magnitude of the error between stem and leaf water potentials and between SWP of different species is minimum at high water potentials. Since all data are near the diurnal maxima, comparisons can be made (cautiously) on a relative basis even though mea- surements were made on Quercus and Heteromeles |twigs and on Juglans leaves. These comparisons can be helpful in understanding distributional | differences. There is a significant (P <.05) difference in the SWP of the three species. Heteromeles gen- erally had the lowest SWP followed by Quercus and Juglans which generally had the highest values (Figs. 1-6). Distributionally, Quercus was most important (referring to importance value indices, Shimwell, 1971) on the north-facing slope and canyon bottom (presumably the more mesic sites) ; Juglans was most important on the south-facing slope on which Heteromeles was relatively rare (Rowlands, 1972). 5-26 6-6 6-20 7-7 7-29 8-15 DATE (month-day) 425 5-11 Figure 6. SWP. of Heteromeles arbutifolia growing on the south-facing slope. (See Fig. 1) DISCUSSION When soil water is not limiting, dawn soil and plant water potentials can be assumed to be equal even though leaf water deficits occur daily. When soil water becomes limiting or when high evap- orative demands cause extreme leaf water deficits, however, the overnight equilibration period may not be adequate to overcome internal plant water deficits (Slayter, 1967). In this case, dawn plant water potentials would be lower than soil water potentials. Soil water potentials were probably higher at the canyon botton due to down-slope water movement both as surface run-off and subterranean percolation. This would also result in a shallower water table at the canyon bottom. These facts along with lesser evaporative demands brought about by lower temperatures and higher relative humidities at the canyon bottom, explain the slope-position differences observed in all species. Since soil drying would occur faster on the slopes than at the base of the canyon. the general water potential decrease and the slope- position differences became more apparent as the dry season progressed. December rains may have wet soil profiles but precipitation in subsequent months was far below normal (Table 2). The rainstorm on 12 August which increased relative humidity minima on subsequent days (Table 1). decreased evaporative demands (and consequent leaf water deficits) and allowed all species to show an increased SWP between the 29 July and 15 112 August samples (Figs. 1-6) by enabling the soil and plant water potentials to approach equilibria. If one ignores the 29 July data, because the dawn values may reflect a soil-plant water potential gradient due to the previous days extreme leaf water deficits, the curves approximate more closely typical drying trends. ; The generally higher SWP observed for south- facing the values recorded on the north-facing slope are opposite from what one would expect. Individuals slope Juglans and Heteromeles relative to growing on the more xeric south-facing slope should exhibit lower SWP than those on the more slope. Soil water should be less and leaf water deficits greater on the south- slope due to greater evaporative demands. must be some mechanism that enables these mesic north-facing facing There plants to respond in this manner. Shropshire (1972) compared physiological re- sponses of H. arbutifolia that were grown in a mesic-conditioned environmental chamber with those grown in a garden under more xeric condi- tions. When both groups were subjected to rela- tively dry air, the plants grown under mesic conditions closed their stomates within 1.5 hours (stopping transpiration and COs exchange); the garden grown plants were able to maintain reduced gas exchange for 3.5 hours. She felt that the stomates of this species were in some way capable of acclimating to xeric environmental conditions which enabled them to continue gas exchange in dry air. Plants that had been acclimated to drier conditions were better able to maintain lower leaf water deficits or overcome these deficits than plants that were acclimated to mesic conditions. Lower water deficits should be reflected in the dawn SWP of plants grown in different habitats and can be used to explain my results. Since the 1971—72 season was one of the driest on record for Southern California (U.S. Dept. of Commerce, 1972), soil on both slopes probably had unusually low water potentials. Because the south-facing slope is characteristically more xeric, individuals on that slope would be acclimated to drier conditions than those on the opposing slope. This acclimation could be accomplished by any morphological or physiological modification (be it genetic or phenotypic) that would result in a reduction of water usage. This environmental pre-conditioning would have enabled the south- facing slope Heteromeles and Juglans to recover more quickly from daily leaf water deficits that occurred during the extreme drought. Heteromeles demonstrates a smaller north-south disparity than BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Juglans (Figs. 3-6). Although Heteromeles has deep tap roots (Hellmers et al., 1955), it probably is not rooted into the water table as is Juglans (T. L. Hanes, pers. comm.). The less extensive root systems of shrubs may negate any acclimation advantage since upper profiles were dry on both slopes. This also explains why Heteromeles had generally lower SWP than the two species of trees. It is possible that Juglans is more responsive to previous environmental selection. Griffin (1973) reported that QO. douglasii growing on the north-facing slopes had lower SWP than those on south-facing slopes. He attributed this disparity to increased root densities on the north-facing slopes which depleted soil water. The north-facing slope which I studied also was much more densely vegetated having approximately six times the density of the south-facing slope. Quercus shows no consistent north-south rela- tionship; soil water potential data would be helpful in interpreting these SWP values. Greater root densities should be found where there is more soil water and hence higher SWP values. It is possible that SWP measurements were made when the plants were transpiring due to an immediate response to the first light of dawn. Since the plants at the top of the slope are exposed to dawn light before the individuals at the canyon bottom, their stomates might open earlier and subsequent leaf water deficits would establish a water potential gradient earlier in the morning. Northeast-facing slope plants receive the earlier summer dawn light and hence establish water potential gradients sooner than do south-facing slope individuals. Interspecific water potential differences could be explained simply by dif- ferential metabolic water requirements. Further research needs to be done to determine the relative sequence and amount of light on a particular slope necessary to cause stomatal re- sponse in these species. Interspecific comparisons of transpiration rates, diurnal water potential curves and leaf water deficit recovery time should be made. It is possible that there is more water available to the south- facing slope plants due to substrate, root depth, and density differences and it would be interesting to re-examine this area in a normal rainfall season to see if these species would respond in the generally expected north-south slope manner. Seasonal soil water potential and water table information would be instructive in the further understanding of soil-plant relationships in these woody perennials. 1974 ACKNOWLEDGMENTS I would like to acknowledge T. L. without whose encouragement and technical advice this study would never have become a reality. Jack Burk was instrumental in manuscript preparation and in my understanding of the water potential data; Leo Song, Jr. helped with the climatological data and M. Lentner (Dept. of Experimental Statistics, New Mexico State University) did the statistical analysis. T would also like to thank all others, too numerous to mention, who read the manuscript and gave critical advice. Hanes, LITERATURE CITED Boyer, J. S. 1967. with a pressure 42:133-137. Leaf water potentials measured chamber. Plant Physiol., |Griffin, J. P. 1973. woodland oaks 54:152-159. Xylem sap tensions in three of Central California. Ecol., 1972. Water stress Ecol., 53:674—680. | Hass, R. H., and J. D. Dodd. patterns in honey mesquite. Hellmers, H., J. S. Horton, G. Juhren, and J. O’Keefe. 1955. Root systems of some chaparral plants in Southern California. Ecol., 36:667—678. Kaufmann, M. R. 1968. Evaluation of the pressure chamber technique for estimating plant water potential of forest tree species. For. Sci., 14: 369-373. Munz, P. A., and D. D. Keck. 1959. Flora. Univ. California Press, 1681 pp. A california Los Angeles, | Ritchie, G. A., and T. M. Hinckley. 1971. Evidence STEM WATER POTENTIALS OF WOODY PERENNIALS for error in pressure bomb estima xylem potentials, Ecol,, 52:534-536 Rowlands, P. G. 1972. Distribution and abundan southern ou lar Dept California State University, Fullerton of tree species in a ‘ (unpublished Master's of Biolo Californi thesis). Scholander, P. F., H. T. Hammel, E. D. and E. A. Hemmingsen. 1965. Sap pressure in vascular plants. Sci., 148:339~-346. sradstreet Shimwell, D. W. 1971. fication of vegetation. Seattle, 322 pp. The description and classi Univ. Washington Press, Shropshire, F. M. 1972. Convergent evolution of two chaparral shrubs under similar environments on different continents. Ph.D. dissertation, Univ. California, Los Angeles, 105 pp. Slayter, R. O. 1967. Plant water relationships. Academic Press, New York, 366 pp. Steel, R. G. D., and J. H. Torrie. 1960. Principles and procedures of statistics. McGraw-Hill Inc., 481 pp. Sucoff, E. 1972. Water potential in red pine: soil moisture, evapotranspiration, crown _ position. Ecol., 53:681—686. U.S. Dept. of Commerce. 1971, 1972. Climatologi- cal Data 75(13-14). 76(1-9). Waring, R. H., and B. D. Cleary. 1967. Plant moisture stress: an evaluation of the pressure bomb. Sci., 155:1248-1254. Accepted for publication October 10, 1973. RESEARCH NOTES THE FIRST REPORT OF THE GIANT POLYNOID POLYCHAETE HOLOLEPIDA MAGNA MOORE FROM CALIFORNIA Moore (1905) described a new genus and new species of a giant polynoid scaleworm, Hololepida magna, from a single specimen measuring 25 cm and possessing 120 segments. It was collected by the Albatross expeditions from Kasaan Bay, Prince of Wales Island, Alaska (corrected locality, Moore, 1908) from 173.7-198.5 m. Later Berkeley (1923) reported two specimens, length not specified, from 27.4—36.6 m depth near Nanaimo, British Columbia. This species has not been reported since that date. Recently specimens of what proved to be H. magna were turned over to me for identification by personnel from both the Los Angeles City and County Sanita- tion Districts. These specimens were collected by trawling in connection with monitoring programs. In discussion of this species with the late Olga Hartman, she showed me a_ specimen, previously unreported, dredged from Monterey Bay, California on 12 February 1931 by G. E. MacGinitie. The data on these new reports are summarized below. The lengths and number of segments given represent minimal figures since this species readily fragments. Monterey Bay, 1 specimen, length 11.6 cm, 83 segments; collected by G. E. MacGinitie, 12 February 1931. Santa Monica Bay, 2 specimens, length 22.9 and 21.6 cm, 94 and 96 segments, depth 54-8 m, collected by Los Angeles City Sanitation District on 31 May 1972. Palos Verdes Peninsula, 5 specimens, 1 measured 20.3 cm, 4 specimens fragmented, 89 segments, depths 59-137 m, collected by Los Angeles Sanitation District on 2 June 1972 (3), 14 June 1973 (1), and 5 December 1973 (1). Superficially this genus resembles a_ sigalionid more than a polynoid because of its large size, the large number of elytrae and the occurrence of elytrae on every posterior segment, but it can be readily distinguished from a sigalionid by the presence of dorsal cirri on segments lacking elytra. The possession of the nuchal hood (Fig. 1) distinguishes this genus from the other polynoid genera. Three species of the genus Hololepida are brown and may be separated as follows: H. magna: nuchal Prostomium deeply incised anteriorly; hood triangular; neuropodial bidentate setae strongly serrated with the two teeth of nearly equal length (Moore, 1905:541). Eastern Pacific Ocean. H. australis: Prostomium weakly incised anteriorly; nuchal hood rectangular; neuropodial bidentate setae strongly serrated (Monro, 1936:93). Falkland Islands. H. veleronis: Prostomium deeply incised anteriorly and with lateral lobes produced and rounded anteriorly; nuchal hood triangular; neuropodial bidentate setae weakly serrated (Hartman, 1939: 48). Gulf of California. All specimens of this genus were obtained by dredging and all were fragmented during the process of collection. Undoubtedly more specimens of H. magna will be encountered in the future especially as a result of the initiation of additional monitoring programs. The fact that the southern California specimens were collected in connection with monitor- ing domestic sewage discharges does not necessarily imply that this species is an indicator of polluted conditions. The author wishes to acknowledge thanks to Jan Stull, Los Angeles County Sanitation District and to Leland Hill, Los Angeles City Sanitation District for referring the material to me. Figure 1. Hololepida magna, anterior end, dorsal view. 114 1974 LITERATURE CITED Berkeley, FE. 1923. Polychaetous annelids from the Nanaimo District. 1. Syllidae to Sigalionidae. Contrib, Canadian Biol., Ottawa, 1:203—218. Hartman, O. 1939, Polychaetous annelids. Pt. 1. Aphroditidae to Pisionidae. Allan Hancock Pacific Exped., 7: 1-156. Monro, C. C. A. 1936. Polychaete worms. II. Discovery Reports, 12:59-198. Moore, J. P. 1905. New species of polychaetes from the North Pacific, chiefly from Alaskan waters. Proc. Acad. Nat. Sci. Philadelphia, 57:525-569. 1908. Some polychaetous annelids of the Northern Pacific Coast of North America. Proc. Acad. Nat. Sci. Philadelphia, 60:321—364. DonaLp J. RetsuH, Dept. Biology, California State University, Long Beach, California 90840. Accepted for publication March 19, 1974. NUTRITIVE VALUE OF THREE COMMON CHAPARRAL PLANTS FOR THE WOODRATS, NEOTOMA FUSCIPES AND N. LEPIDA The food values of different plants to woodrats have commonly been inferred from stomach contents, feces composition and nest caches (Horton and Wright, Ecology, 25:341-351, 1944; Linsdale and Tevis, The dusky-footed woodrat, 1951; Cameron, J. Mamm., 52:288—296, 1971). The actual nutritive value of a plant can only be determined by following a physio- logical index, such as body weight, during feeding trials. Only if there is a positive result can one say that a particular item is a physiologically available food (Chess and Chew, J. Mamm., 52:193-195, 1971). Three common chaparral plants were fed to Neotoma fuscipes (Dusky-footed woodrat), a charac- teristic chaparral rodent, and to N. lepida (Desert woodrat), typical of desert areas, but found in a great variety of vegetation types (Cameron, 1971). The N. fuscipes we used were trapped in the Santa Monica Mts. and San Gabriel Mts. in typical chapar- ral. The N. lepida were from near Mitchell Caverns, San Bernardino Co, California, from a desert com- munity dominated by Larrea tridentata, Opuntia spp.. RESEARCH NOTES 104 A fasciculatum Q wislizenii 104 kK = 100 DAYS S. mellifera %INITIAL WT ice) ice) ine) o Gs ia , H VET) Mee © @ oo any te | 2 Sit DAYS Figure 1. Body weight changes of woodrats fed on diets of a half maintenance ration of Purina Labo- ratory Chow and a surplus of the leaved stems of: (a) Adenostoma fasciculatum, (b) Quercus wislizenii. and (c) Salvia mellifera. Dashed line connects means for Neotoma fuscipes: solid line, N. lepida: broken line, control group of N. fuscipes, fed only half ration of chow. Vertical lines are the range of mean + 2 standard errors. In figure Ic the average values for N. lepida are not connected, in order to reduce confusion of lines. 116 and Yucca schidigera. The plant species used were fasciculatum shrub), Quercus wislizenii (evergreen oak, a tree), and Salvia mellifera (black sage, a deciduous shrub). another of evergreen 4denostoma (chamise, an evergreen Chamise and one species or oak are common dominants in chaparral of southern California, consequently they are potentially important food plants for woodrats. Linsdale and Tevis (1951) rated chamise and live oaks as Class 1 foods, food plants of N. fuscipes, on subjective grounds. Salvia mellifera is dominant only and most eaten in the coastal sage type of chaparral; we chose it because it has a strong odor from volatile oils, which might be deterrents to feeding. Linsdale and Tevis rated black sage as a Class 4 food, least eaten; black sage and other odoriferous were characterized as “not a well liked food” by N. fuscipes. The woodrats were kept at room temperature in 25 x 30 x 60 cm wire mesh cages with woodchips and shredded newspaper bedding and water as desired from a drinking bottle. During a preliminary period we determined the minimum amount of Purina Laboratory Chow that was necessary for each animal to maintain body weight. Weights averaged 182 g (108-238 g) for N. fuscipes and 102 g (79-143 g) for N. lepida. During a feeding trial each animal was given one half of its maintenance ration of chow plus an excess of leaved stems of one of the plants. Plants were freshly collected at the beginning of each trial (unless noted otherwise) and kept with their stems in water. A control group of three N. fuscipes was given only the half chow ration. Rats were weighed daily at the same time; weights were ex- pressed as a percentage of the initial weight. The woodrats ate leaves and bark of Q. wislizenii, but only leaves of the other two species. They used stems of all species to make “nests” in their cages. Body weight changes on the different diets are shown in figure 1. If the ranges of + 2 standard errors do not overlap, or overlap only slightly, One can assume statistical significance (P < 0.05). The standard errors are usually large due to small sample sizes (3—5 animals) and wide individual variation. The loss of weight of the controls is a baseline for interpretation of the nutritive value of the plant part of the diet. As shown in figure la and 1b, A. fasciculatum and Q. wislizenii were sufficiently nutritious to allow N. fuscipes to maintain normal weight (i.e. a weight not significantly different from initial weight). All N. lepida lost weight on these plants: their average weights were not significantly different several species BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 from the controls, which were starving on a half ra- The nutritive value of the oak seemed The first feedings tion of chow. to be affected by its freshness. were with stems picked 10 days before the first feeding. Although the stems were kept in water, the N. fuscipes fed on them dropped significantly below initial weight, although they were significantly above the controls. When fresh stems were given to both experimental groups beginning in the fourth day (fr in Fig. 1b) the weights of N. fuscipes returned to normal. Neotoma lepida continued to lose weight, but at a slower rate. When the control group was given fresh oak stems beginning in the sixth day (cf in Fig. 1b), they recovered their lost weight. The lesser value of the “old” oak stems causes us to wonder about the food value of stems found in woodrat nests. The continual loss of weight of N. lepida on the oak was surprising in view of Cameron’s (1971) report that N. lepida in Joshua Tree National Monument had a diet mainly of Quercus turbinella, another evergreen species. But, Q. wislizenii is a plant that is foreign to the experi- ence of the population of N. lepida that we sampled. The maintenance of weight of our N. fuscipes on chamise is not consistent with the unsupported statement of Linsdale and Tevis (1951:278) that “chamise foliage . . . will not sustain the rats.” Neither species of woodrat was able to maintain weight on S. mellifera, and they washed themselves excessively after beginning to eat the leaves. One N. fuscipes was unable to regain normal weight after the trial, even when given additional succulent foods (carrots and celery). The volatile substances may have had some effect on the functioning of the digestive tracts of the rats, as do oils of sage- brush for deer (Nagy et al., J. Wildlife Mgmt., 28:785-790, 1964). Learned or inherited behavior may be involved in the inability of our N. lepida to maintain weight on any of the three plants, which are all foreign to their environment. Lee (Univ. California Publ. Zool., 64:57-96, 1963) found differences in the way experienced and naive woodrats fed on Opuntia. Without further tests we are unwilling to conclude that there is a real difference in the nutritive value of chamise and oak for the two species of Neotoma. Ropert M. CuHEwWw and James C. WoopMan, Dept. Biological Sciences, Univ. Southern California, Los Angeles, California 90007. Accepted for publication March 18, 1974. INSTRUCTIONS FOR AUTHORS The BULLETIN is published three times each year (April, August, and November) and includes articles in English in any field of science. 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Printing charges accruing from excessive addi- tions to, or changes in, the proofs must be assumed by the author. Reprint orders are placed with the printer, not the Managing Editor. CONTENTS | Reproductive behavior of the gray whale Eschrichtius robustus, in Baja California. By William | EMS AIMARGS! sceys she, wieueed storage cists er sve cue aiientedeus (o\esaie re OsPeiyol a pereiatts oils ulsvalele geile tsetse Re ake 37) Bats of Margarita Island, Venezuela, with zoogeographic comments. By James Dale Smith and FURRY UG CNOWAYS ois cine cist) «etn sc eae evorsbs abe ioe ace sine vey ellie date toilet Nelle eh Set OCR e eS 64 ' Maternity habits of Myotis leibii in South Dakota. By Merlin D. Tuttle and Lawrence R. Heaney 80 | Influence of habitat structure on a population of voles. By George O. Batzli .............. 83 An unusual breeding aggregation of frogs, with notes on the ecology of Agalychnis spurrelli e (Anura: Hylidae). By Norman J. Scott, Jr. and Andrew Starrett ..........00eeseeeeee- Hermit crabs from the tropical eastern Pacific. I. Distribution, color, and natural history of some common shallow-water species. By Eldon E. Ball and Janet Haig .............. Systematics of Pterocladia media from California. By Joan G. Stewart .........0ceeeeaee Relative stem water potentials of three woody perennials in a southern oak woodland community. By ais WP sSVVeEriSen anesec ener ssc 0s ei otc | a ales eye teierioveterofels|e easiel eyeie atele Rach RECN RESEARCH NOTES The first report of the giant polynoid polychaete Hololepida magna Moore from California. By Donald J. Reish ...... AOC EID ME CIEL ACO eo Oe! OtorG Bid b)0 0-0\6.0-d.o Go Nutritive value of three common chaparral plants for the woodrats, Neotoma fuscipes and N. } lepida., By, Robert: M. Chew and James\G.'\Wocdman. 6... hase acer TS) ae NEW YORK BC 1ANICAL 3 x L. i ) ary =, CovER: California gray whale, Eschrichtius robustus, from off Islas Coronados, Baja California, Mexico. Photo by Stephen Leatherwood, courtesy Naval Undersea Center. Number 3 NOVEMBER ve Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 OFFICERS Jules Crane, Jr., President Donald Bright, First Vice President Wayland D. Hand, Second Vice President Richard E. Pieper, Secretary Donald R. Patten, Treasurer James Dale Smith, Technical and Managing Editor Roberta S. Greenwood, Index Editor BOARD OF DIRECTORS 1972-1974 1973-1975 1974-1976 Wayland D. Hand Donald Bright George Callison Donald R. Patten John J. Baird Kristen Fauchald Donald J. Reish Jules Crane, Jr. Takashi Hoshizaki Andrew Starrett Richard E. Pieper James Dale Smith Patrick H. Wells David P. Whistler Stuart L. 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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 « [FOR ACADEMY OF VOLUME 73 NOVEMBER 26, SICILEN CI 1974 TUBE DWELLING BEHAVIOR IN THE MARINE ANNELID PHRAGMATOPOMA CALIFORNICA (POLYCHAETA: (FEWKES ) SABELLARIIDAE ) PeTerR ALAN Roy! ABSTRACT: Abdominal tori position the body centrally in the tube, permitting lateral and dorsal body cilia to draw in water ventrally, pass it up between the parapodia, and expel it dorsally. Withdrawal involves contraction, peristalsis, and backward parapodial walking. Extensions of the parathoracic palae effect forward locomotion, and parathoracic neurosetae cause body twisting. In defecation abdominal neurosetae pass fecal strings anteriorly where a body retraction expels them ventrally from the tube. Members of the polychaete family Sabellariidae are suspension feeders that live in blind tubes of cemented sand grains and shell fragments. Ana- tomically and behaviorally they are adapted to a tube dwelling existence. Studies on the biology of the Sabellariidae have centered on morphology, embryology, larval behavior, and tube and reef building. The behavior of adult animals is still incompletely known. McIntosh (1922) included the observations of Arnold Watson on the natural history and anatomy of Tetreres (Pallasia) murata (Allen), Sabellaria spinulosa Leukart, and S. alveolata (L.). Wilson described the development and settling behavior of the larvae of S. alveolata (1929, 1968a, b, 1970a) and S. spinulosa (1929, 1970b). Wilson (1969) also described the natural history and defecation behavior of S. alveolata. Dales (1952) considered the development and structure of the anterior end of Phragmatopoma californica (Fewkes), and included observations on the ciliary tracks of the feeding tentacles and the food grooves leading to the mouth. Vovelle (1957a, b, 1958a, b, 1963, 1965) described the structure and construction of the tubes and the formation of colonies in S. alveolata. He noted that an adult worm removed from its tube was unable to build a new one. Kaestner (1967) briefly mentioned water currents within the tube, and noted they are responsible for irrigation and defecation. Kirtley’s (1968) account of P. lapidosa (Kinberg) described the processes of tube building, the ciliary and feeding currents, peristaltic pumping, locomotion, and defecation of this worm. While these accounts contribute much valuable information, the picture of the activities of a worm within its tube is not yet complete; the objective of the present paper is to fill this gap. Studies have centered on how the animal main- tains its position and posture within the tube, the manner in which withdrawal and forward loco- motion occur, the circulation of water within the tube by ciliary and other means, and the mecha- nism of elimination of fecal matter from the tube. METHODS Studies were conducted at the Hopkins Marine Station of Stanford University. Pacific Grove, California. Colonies of Phragmatopoma_ cali- fornica abound on the intertidal rocks between Mussel Point and Point Alones, adjacent to the marine station. Portions of colonies taken in the field were maintained in good condition for long as five months in tanks of running sea\ (14°-16° C) in the laboratory. 1 Hopkins Marine Station of Stanfor Pacific Grove, California 93950 Southeastern Massachusetts Universit mouth, Massachusetts 02747). 117 118 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 | la A Figure 1. Phragmatopoma californica in sand tubes constructed against glass. a. Worm in expanded position with the body twisted; dorsal surface exposed anteriorly and ventral surface posteriorly. b. Worm withdrawn into its tube, left side of anterior region. c. Worm withdrawn into its tube, anterior region, ventral side. A, anus; BO, building organ; C, cauda; CB, ciliary band; DC, dorsal cirrus; FG, food groove; M, mouth; NS, neurosetae; O, operculum; P, palae; PX, parathorax; PT, prostomal tentacle; T, torus; VC, ventral cirrus; VP, ventral plate. 1974 BEHAVIOR OF For studies of ciliary tracts on the body, speci- mens were first removed from their tubes and anesthetized with a solution of magnesium chloride isotonic with seawater (7.4 MeCh by weight). The worms were then replaced in sea- water and a suspension of colloidal graphite (Aguadag, Acheson Colloids Co.) added. For observations of living worms in tubes, a transparent tube preparation was developed. Worms taken in the field were brought into the laboratory and distal portions of the tube broken off until the worm was left with only enough tube to contain its fully contracted body. These worms were placed in petri dishes without lids, and the dishes submerged in an aquarium. Sand was sprinkled over the dishes until a layer 1-2 grains thick was present on the bottom. The worms extended partially from their tubes, and with this sand they constructed extensions of their original tubes against the bottom of the dish. These extensions were arches of sand built against the glass with the glass forming part of the tube wall. Additional sand was added daily as needed. If too much sand was provided the extension would be a complete cylinder of sand, at least in spots, obscuring subsequent observations. Within 2—3 days worms had constructed tube extensions 1-3 times their body length. At this time the original posterior portion of the tube was broken off and sand was sprinkled over the resulting posterior hole. This was shortly cemented in place by the worms, closing the posterior end of the tube and leaving the animal in a tube of sand built against the glass, allowing clear vision of the entire body (Fig. la). For observation and photography, the top of the petri dish was added, and the whole dish inverted under water. A suspension of carbon particles was added to facilitate the study of currents within the tube. Except in special instances worms were always completely immersed in water during observation, and the water was percent Was changed frequently to keep it cool. Drawings were made with a camera lucida. RESULTS Morphology of Phragmatopoma_ californica.— The body of P. californica is about 2—3 cm in length and as in other sabellariids is composed of four regions: head, thorax, abdomen, and cauda (Fig. 3a, b). The head (Dales, 1952) includes the operculum, opercular stalk, feeding tentacles. mouth, and building organ. The thickened muscu- PHRAGMA, POMA HEAD RY A ( rd » \ Wena THORAX ’ \\ar =} PS he \\\ yp ; {' s§ Gi ye \ My LEY vas n\ PY Se. 4 iB, Me ‘ a ~ N= Ly We \== 4 MEN 5A \—7 ABDOME SY es) g > Bye lem CAIN WSs. CAUDA Gg Ni BW: | LAN B iS 2a b a ¥ Figure 2. Phragmatopoma californica: entire animals, in expanded posture. tion of water around the body. a. right side. b. Ventral view. Arrows show major circula- Lateral view of lar thorax consists of five segments, the posterior four of which bear large dorsal cirri. The posterior three segments of the thorax are the parathoracic and these bear spear- shaped notopodial palae and capillary neurosetae. segments, The palae are smallest in the first parathoracic segment and largest on the third segment, while the reverse is true of the neurosetae. Ventrally the parathoracic segments form a muscular ventral plate. The abdomen 20-40 rather similar segments, all bearing dorsal cirri which, consists of as those of the thorax, possess blood vessels with a pulsating blood flow. The abdominal notopodia are uncinigerous tori. Each neuropodium consists of a ventrolateral lobe bearing capillary neuro- setae and lateral to it a broad fleshy lobe referred to as a “ventral cirrus” by McIntosh (1922). There is a gradual reduction in size of the dorsal cirri and tori from the first abdominal segment on back so that posteriorly the cirri are to papillae and the tori to elongate. stiff. paddle The groove shaped projections. abdomen is con ventrally, forming a that extends of the ventral plat ) the cauda. a ndrical The terminal region of the body lacking parapodia the posterior margin origin of the cauda. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 lcm Figure 3. Anterior body regions of Phragmatopoma californica, with feeding tentacles removed. Small arrows show the direction of ciliary beat; large arrows indicate the major flow of water over the body. a. Lateral view from right side. b. Dorsal view. view. See figure 1 for key to abbreviations. c. Ventral 1974 BEHAVIOR OF and external signs of segmentation, is reflected ventrally and anteriorly, It extends within the concavity of the abdomen for one-third to one-half the total length of the abdomen. The anus is at the tip of the cauda. Building of the tube extension and posture forward within the tube.—Tube building by worms. in glass dishes indicates that sand particles need not be in suspension to be used for construction. Any particles of sand contacting the head tentacles are passed by cilia to the mouth and then to the building organ where the grains are coated with cement. The head is then protruded and_ the grain manipulated into position and cemented at the mouth of the tube by the building organ. Tubes built against glass in the laboratory were thinner at their open ends and lacked the flared apertures normally present in colonies found in the field. However, tubes of solitary specimens of P. californica, which are built against surfaces of intertidal rocks around Hopkins Marine Station, also lack flared apertures and are otherwise similar to the tubes constructed in the laboratory. In natural colonies the head and thorax of an expanded worm lie within the flared tube opening. The opercular stalk is bent dorsally so that the feeding tentacles radiate outward beyond the tube mouth. Expanded worms in tubes con- structed against glass behaved similarly in all respects except that they protrude their opercula a little beyond the tube opening in order to fully extend the feeding tentacles (Fig. la). The worm occupies a central position within the tube along the tube axis leaving room for water to circulate on all sides. The body is braced in this position laterally by extension of the abdominal tori on each side so that their uncinigerous margins contact the tube wall. Ventrolaterally, the neuropodial “cirri” are braced against the tube wall and aid in the positioning of the body especially posteriorly where the notopodial tori are long and attenuated. Although the anteromedially directed abdominal neurosetae may contact the tube wall, they are not critical in maintaining normal position of the body in the tube. The thorax is braced in a central position by the posterolaterally directed parathoracic palae. The parathoracic neurosetae, directed antero- ventrally, and the neurosetae of the first abdominal segment, hold the thorax elevated above the floor of the tube. Worms are often observed to be twisted through 90° to 180° along the axis of their bodies (Fig. la). PHRAGMATOPOMA Withdrawal response and back Like rapidly otnet ed rar P ca withdray l " ea me tentacles into its tube Wh feeding tentacles are rapidly contr opercular stalk is bent ventrally to (Fig. 1b, c). Simultaneously the wort the anterior two-thirds of its body tt palae and tori of this region lose contact the tube wall. The posterior tori and perhay the neuropodial cirri maintain contact as an anchorage. Then the worm withdraws its anterior end into the tube by shortening the body If the stimulus is sufficiently strong or i continued, the animal backs further into the tube. This is accomplished in two ways. The first action is monotaxic and involves body peristalsis. The parathorax is expanded after completion of the initial withdrawal response. This increases its diameter and anchors the anterior body in the withdrawn position. Then the abdomen constricts, loses contact with the tube wall, and is elongated and extended poste- riorly where the tori gain a new posterior anchorage. Finally the thorax constricts and the body shortens, withdrawing the anterior portions. completing the cycle. The second method of back- ward than the involves parapodial walking. In walking back- locomotion is slower first and ward nearly the whole body loses lateral contact with the tube wall. The posterior abdominal tori maintain an anchorage. Backward movement of the body is initiated by walking motion of these posterior tori. Only the projecting of the last 6-10 abdominal involved, and the activity of successive segments appears relatively uncoordinated. During this type of backward movement the posterior abdomen is flexed dorsally. These two methods of backward locomotion often occur in sequence. with the peristaltic backing occurring directly after the initial withdrawal and the gliding walk deeper rear-to-front tori segments are following thereafter when the worm lies in its tube. Forward locomotion and twisting of the —Forward the tube is carried out by a mechanism best described as palea “walking.” The posterolaterally directed thoracic palae push against the tut thrust out of their setal sacs, ward force on the body. A worn the rear of the lateral forward by paleal thrusts locomotion in tube retracts its tori sing contact with the tube wall. and move VOLUME 73 OF SCIENCES SOUTHERN CALIFORNIA ACADEMY BULLETIN vc NS ‘7 ead TT cB lcm [-s) U 1974 BEHAVIOR OF shortened its body in an initial withdrawal re- sponse likewise re-extends anteriorly by paleal walking rather than by forceful forward muscular extension. This activity of setal extension appears nonmetameric, Body twisting (Fig. la), is accomplished by the action of the parathoracic neurosetae. These setal bundles are directed the mid-line ventrally, and when those of only one side are forcefully extended against the tube wall, a torque is exerted about the body axis in a direction Opposite to the thrust. Twisting often occurs in conjunction with forward locomotion. The neuro- setae can also twist the anterior body while the worm is in a stationary position, but not to the degree that occurs during forward movement. Ciliary currents within the tube.—The ciliary currents associated with feeding in Phragmato- poma have been observed by Dales (1952), but the currents within the tube were not de- scribed. Observations both on active worms within their tubes and on relaxed animals removed from their tubes show the major ciliary tracts responsible for water circulation within the tube to be as follows: (1) Discrete transverse bands of cilia are present along the entire dorsal surface of the abdomen, thorax, and opercular stalk (Figs. 3b and 4b). The beat of these cilia moves water anteriorly. (2) The dorsal toward cirri of the thorax and abdomen also bear transverse ciliated ridges on their medial surfaces. The bands are reduced posteriorly along with the reduction in size of the cirri. The dorsal cirri are normally directed anteromedially (Figs. la, b), hence their cilia move water anteriorly in the space between the dorsal body wall and the adjacent inner surface of the tube. (3) The lateral body wall between successive parapodia bears numerous longitudinally oriented bands of cilia. These bands are most extensive on the anterior abdominal region (Fig. 4a) and are reduced posteriorly as the notopodial tori become narrower and more elongate. No ciliary action was observed on the para- podia themselves. At the body, the opercular stalk also bears longitudinal anterior end of the PHRAGMATOVPOMA bands of cilia on its lateral move water toward the dorsum (Fi The ventral surfaces of the thor; abdomen, and the entire cauda are dé ciliary bands. The ventral surface of the oy cular stalk bears transverse bands of cilia wi beat anteriorly and function in the rejectio particles from the mouth (Fig. 4c) The bracing of the body in a central position along the axis of the tube by the tori provide space all around the body for water circulation Figure 2a shows the general circulation pattern When a worm is in an expanded condition with its head at the aperture of the tube, drawn into the tube past the ventral surfaces of the head and thorax. Some passed dorsally by the lateral cilia of the opercular stalk and the interpodial thoracic cilia (Fig. 3a) but most of it flows posteriorly along the channel beneath the abdomen. The beat of the interpodial cilia water is water is moves water dorsally between successive tori all along the abdomen (Fig. 4a). These segmentally separated streams join the main dorsal current which flows anteriorly and is powered by the cilia of the dorsum and the dorsal cirri. Figure 4b illustrates the interpodial and dorsal currents and the manner in which the interpodial ciliary bands merge with those of the dorsum. The dorsal exhalent passes out over the dorsal surface of the oper- cular stalk (Fig. 3b). Gametes have been observed emerging from the tube in the dorsal current during spawning. current An eddy exists in the water around the head of expanded worms. Figure 3a illustrates how the interaction of the dorsal exhalent the ventral inhalent current and the rejection current from the ventrum of the opercular stalk combine to produce this eddy. This phenomenon may facilitate the collection of particulate matter by the feeding tentacles of the head. Thoracic fornica exhibits a peristaltic pumping behavior when it is withdrawn in its tube. This behavior always occurs in withdrawn worms. whether they current, pumping.—Phragmatopoma _ cali- are only slightly withdrawn into the tube opening -com- or deeply placed in the tube. Pumping is a plished by a piston-like peristalsis confined the parathoracic region. Presumably this beh Figure 4. Somites of the anterior abdominal region of Phragmatopoma calijornica, sh a. Ventral views. t position of the cilial bands. b. Lateral view from the ri view with the cirri of the right side removed. See figure 1 for key to abbreviations 124 BULLETIN SOUTHERN CALIFORNIA serves to increase the rate of water exchange at the mouth of the tube when the worm is withdrawn. During pumping, the parathorax shortens and The thorax is pushed anteriorly on the forward stroke by the Then the parathorax constricts, losing its contact, and is retracted by a short longitudinal contrac- tion. A complete pumping stroke is repeated 40 to 60 times a minute. Defecation and fecal elimination—The cauda of P. californica does not extend forward more than one half the length of the abdomen of an expanded worm. Even in a contracted worm, broadens, contacting the tube wall. a single extension of parathoracic palae. the cauda has not been seen to extend anteriorly beyond the anterior end of the abdomen. The anus at the tip of the cauda lies in a mid- ventral position, facing into the ventral inhalent current. Feces are extruded from the anus in the form of continuous strings. The extrusion is accom- plished by peristaltic waves that begin at the anus and travel up the cauda to its origin at the abdomen. The abdominal neurosetae com- mence anteriorly directed stroking or rowing motions during defecation. These motions do not appear metameric. Nevertheless, the opposed tips of the neurosetae on the two sides of the body grasp the fecal string, and actually pull it out of the anus as it is extruded. The fecal matter is broken into 3 to 5 mm lengths during this process and is passed anteriorly to the posterior margin of the ventral plate. The animal then undergoes a withdrawal-like response which expels the feces ventrally from the tube. DISCUSSION Previous studies of sabellariid behavior have been carried out largely on other species, but have been limited as normal behavior of the worms could not be observed in situ. Never- theless, findings of earlier workers often agree with present californica. observations on Phragmatopoma Since there are close morphological similarities among the Sabellariidae, it is logical to assume that the behavior of P. californica may be considered generally representative for the family. The use of uncini or uncinigerous tori as a means of maintaining position within the tube is widespread within the Sedentaria. Phragmato- poma californica, and presumably other sabel- lariids, have developed this to a high degree. ACADEMY OF SCIENCES VOLUME 73 The extended tori are situated so that water circulation can take place freely around the body. Watson (in McIntosh, 1922) observed that the posterior tori of S. alveolata could... “be moved freely backward and forward, as well as retracted,” but he did not determine their locomotory function. Although there was no mention of the twisted posture so often seen in P. californica, Watson (in McIntosh, 1922) did deduce the functions of the parathoracic palae and neurosetae in S. spinulosa which could result in body rotation. Kirtley (1968) alludes to the locomotory function of the parathoracic palae of P. lapidosa, but included no details. He also assigned a loco- motory function to the abdominal neurosetae in that species which has not been observed in P. californica. Watson (in McIntosh, 1922) suggested that the parathoracic palae of S. spinulosa are involved in the shaping of the tube and Kirtley (1968) reported the same for P. lapidosa. The only observed uses of the thoracic palae by P. cali- fornica have been bracing the body, locomotion, and pumping. According to Kirtley (1968), P. lapidosa uses its opercular palae for grasping and manipulating sand grains during tube building. This behavior was not observed in P. californica during tube building in the petri dishes. Manipulation and positioning of the sand grains was accomplished by the building organ of P. californica in a manner similar to that shown by Vovelle (1965) for S. alveolata. Some previous workers have commented on the presence of cilia or on water currents about the bodies of sabellariids. Watson (in McIntosh, 1922) and Kaestner (1967) both noted the existence of a main exhalent dorsal current. Watson (in McIntosh, 1922) suggested that rowing motions of the abdominal neurosetae in S. spinulosa helped to create water currents, and that water was expelled from the tube anteriorly by undulatory movements of the body. Kirtley (1968) reported that in expanded speci- mens of P. lapidosa, currents in and around the tube were aided by “peristaltic motion of the body segments.” In P. californica circulation of water and expulsion of water from the tube were normally carried out by ciliary action; expulsion of water by body movement occurred only during pumping, withdrawal, and fecal elimination. Moreover, peristaltic thoracic pump- ing occurred only while the animal was withdrawn. 1974 BEHAVIOR OF Several workers have commented on the posi- tion of the cauda and the matter of defecation in different sabellariids. Watson (in McIntosh, 1922) suggested that the abdominal neurosetac hold the cauda in position. This is not the case lies freely neurosetae normally In S. alveolata in P. californica, where the cauda under the abdomen; the touch it only during defecation. Watson (in McIntosh, 1922) a young example the ejecta discharged from the gut in a thread-like form and worked forward by ciliary action and perhaps the movement of the bristles. The annelid then protruded its anterior region, and with a sudden jerk inward the refuse was dropped outside the tube.” Wilson (1969) indicated that defecation in S. alveolata takes place by the extension of the cauda forward so that the anus is protruded out of the tube mouth. This was never observed in P. californica. Kirtley (1968) stated that the abdominal neuro- setae of P. lapidosa were responsible for “removing metabolic wastes from the tube,” but does not report how this is done. Kaestner (1967) stated that the dorsal exhalent current carries feces out of the tube, which was never seen in P. californica. “also noticed in ACKNOWLEDGMENTS Special thanks are extended to Donald P. Abbott of the Hopkins Marine Station, who suggested the prob- lem and supervised the study with great patience. I would also like to thank John Reardon of Southeastern Massachusetts University, and Henry Lague of the Braintree, Massachusetts School Department, for making possible my initial visit to Hopkins Marine Station in 1968. LITERATURE CITED Dales, R. P. 1952. The development and structure of the anterior region of the body in the Sabel- lariidae, with special reference to Phragmatopoma Californica. Quart. J. Micro. Sci., 93:435—452. Kaestner, A. 1967. Invertebrate Zoology, vol. I. 2nd ed., (trans. H. W. Levi and L. R. Levi). Interscience, New York. 597 pp. Kirtley, D. W. 1968. The Reef Builders. Nat. Hist., 77:40—45. McIntosh, W. C. 1922. A Monograph on_ the British Marine Annelids. vol. IV, part 1. Hermellidae to Sabellidae. 1-250. Ray Soc., London, PHRAGMATOPOM Vovelle, J Remaraue ut tomique ct I'histocl ce fe laires intervenant dans l'édificatior Sabellaria alveolata (1..). Bull. So / de France, Paris, 263;202~20 1957b, substance cémentaire du tube de Sabellaria a lata (L.) Academic des Sciences, Paris Données histoc himique Annelide Polychete, Comptes Res ) i 244 2964-2966 1958a, tube de Sabellaria alveolata Remarquées sur Ja structure di) (L.) et les tions impliquees dans son édification forma Archive de Zoologie Expérimentale et Générale, 95:52 68. 1958b. Sur la sécrétion de composé phénoliques intervenant dans la stabilisation de la substance cémentaire du tube chez Sabellaria alveolata (L.). Comptes Rendus, Academic des Sciences, Paris, 246:472—474. 1963. cémentaire du ((U))- 99, Ultrastructure de la sécrétion tube chez Sabhellaria Proc. XVI Cong. Zool., Washington, p. alveolata 1965. Le tube de Sabellaria alveolata (L.) et son cément. Etude écologique, expérimentale, histologique, et histochimique. Archives de Zo- ologie Expérimentale et Générale, 106:1—187. Wilson, D. P. 1929. The larvae of the British Sabellarians. J. Mar. Biol. Assoc. U.K., 16: 221-269. 1968a. Some aspects of the development of eggs and larvae of Sabellaria alveolata (L.). J. Mar. Biol. Assoc. U.K., 48:367—386. 1968b. The settlement larvae of Sabellaria alveolata (L.). Assoc. U.K., 48:367-435. behavior of the J. Mar. Biol. 1969. The Honeycomb Worm. Sea Fron- tiers, 15:322-329. 1970a. Additional observations on | growth and settlement of Sabellaria aly Mar. Biol. Assoc. U.K., 50: 1-32. 1970b. The larvae of Sa and their settlkement behavi Assoc. U.K., 50:33-52. Accepted for publication Janua THE SYSTEMATICS AND DISTRIBUTION OF MARINE TUBIFICIDAE (ANNELIDA: OLIGOCHAETA) IN THE BAHIA DE SAN QUINTIN, BAJA CALIFORNIA, WITH DESCRIPTIONS OF FIVE NEW SPECIES Davip G. Cook! Apstracr: Marine Tubificidae (Annelida, Oligochaeta) from Bahia de San Quintin, Baja California, are described, and their distribution in the bay noted relative to the mean particle size of the sediments. Tubifex postcapillatus, n. sp. is characterized by its simple-pointed dorsal setae, and short hair setae which occur in postclitellar segments only. The setal pattern and details of the genital anatomy distinguish Thalassodrilus belli, n. sp. from other members of its genus. Both of these species are associated with the finest sediments which are located mainly at the north end of the bay. Limnodriloides monothecus, n. sp. with its single, mid-dorsal spermatheca unique to this genus, occurs mainly in silts and very fine sands (4 to 5 @). Limnodriloides verrucosus, n. sp. possesses a papillate body wall previously unknown in the genus, but reminiscent of the Peloscolex condition; it occurs in all sediment types in the bay (3 to 8 ¢) but is most abundant in the 5 to 6 # range. The dominant oligo- chaete, Limnodriloides barnardi, n. sp., closely resembles Limnodriloides winckelmanni Michaelsen, especially in its possession of elongate, grooved spermathecal setae enclosed in vacuolated sacs; the species are distinguished by details of their penial structures. Limno- driloides barnardi, n. sp. occurs in all sediment classes but its optimum range appears to be in 4 to 6 ¢ sediments. In a general report on the benthic exploration of Bahia de San Quintin, Baja California, in 1960— 1961, Barnard (1962) pointed out that unpolluted or unmodified bays and enclosures of the South- ern Pacific region of North America are both rare and poorly known biologically. Barnard (op. cit.) SEES the two primary objectives of the survey as . a search for basic information on an un- polluted enclosure as near southern California as possible, and the initiation of comparative investi- gations planned by the Institute of Marine Bio- Research on many such enclosures in the Eastern Pacific.” This paper reports on the systematics and distribution within Bahia de San Quintin of the tubificid oligochaetes. The recognition of Oligochaeta, mainly Tubi- ficidae, as regular components of the marine benthic community, has occurred only within the last decade. Previously oligochaetes were re- garded as more or less exclusively fresh-water or terrestrial with an occasional species “invading” the marine environment via estuaries and fresh- water sources in the littoral zone. Undoubtedly this may be true of some species, but the works of Brinkhurst (1963; 1966), Hrabé (1966: 1967: 1971a; 1971b), and Cook (1969; 1970a; 1971) have demonstrated that a significant number of truly marine Tubificidae exist, including some from the deep sea (Cook, 1970b). The collection of Tubificidae from Bahia de San Quintin, numbering 55 samples, each con- taining from 1 to about 500 worms, was re- ceived for examination from the Smithsonian In- stitution. Five species of Tubificidae have been recognized, namely, Tubifex postcapillatus n. sp., Thalassodrilus belli n. sp., Limnodriloides barnardi n. sp., Limnodriloides monothecus n. sp., Limnodriloides verrucosus n. sp. STUDY AREA AND METHODS Bahia de San Quintin is located on the Pacific side of Baja California with its northeastern tip at 30° 30’ N, 116° 00’ W. A full description of the study area, field, and laboratory methods, in- volved in the survey is available in Barnard (1962). Briefly, an area of six square miles in the eastern arm of the bay was sampled as uni- formly as possible at a density of 15 stations per square mile. A benthic sample was collected at each station (designated SQ 1 to 94) using a *National Museum of Natural Sciences, Ottawa, Ontario, Canada, K1A OMS8 (Present Address: Great Lakes Biolimnology Laboratory, Canada Centre for Inland Waters, Burlington, Ontario, Canada, L7R 4A6). 126 and q 1974 PACIVIC COAST modified Hayward orange-peel grab with a capacity of 650 sq cm (0,065 sq cm). Material from the grab was washed through a Tyler screen (12.6 mesh strands per cm; mesh diameter 0.495 mm), the animals preserved in 4 percent formalin and transported to the laboratory. Here they were rewashed through a screen (23.6 mesh strands per cm), sorted into taxonomic groups, stored in formalin for three months and finally transferred to 70 percent ethanol. The Oligochaeta had been lightly stained in eosin, dehydrated, and transferred to methyl salicylate by the late A. W. Bell, who had also made microscope slide preparations of one to four individuals from a number of samples; from this preliminary survey Bell had apparently recog- nized the presence of a Limnodriloides (some slides labelled as “L. winckelmanni”) and a spe- cies of “Peloscolex” or “Tubifex” (= T. post- capillatus). After Bell’s slides had been examined to assess the number and type of taxa involved, material in fluid was studied and identified using: a) temporary mounts in methyl salicylate; b) permanent whole mounts in Canada _ Balsam, stained in eosin; c) permanent mounts of dis- sected worms lightly stained in either eosin or Harris’ haematoxylin. In 25 samples the num- bers of worms were too large to permit identifica- tion of every individual, therefore subsamples of about 30 worms were taken by the following method which eliminates, or minimizes, any selec- tion bias for large individuals. Material was trans- ferred to a small dish where it was spread out uniformly (as estimated visually): 30 individuals were separated from one small area (size de- pendent on total sample size) and these identi- fied. A rough estimate of the total number of individuals (T) in these samples was arrived at by comparing the area occupied by 30 indi- viduals (As) with the total area of the sample dish (At); i.e., T = 30 (At) /As. SYSTEMATIC ACCOUNTS Tubifex postcapillatus, new species Figure 1 Holotype.—United States National Museum (USNM) catalogue number 45289. Bahia de San Quintin, Station number SQ 1. Depth, less than 2 m, in gray clay. Collected 22 April 1960. Paratypes—USNM 45290. Five individuals from SQ 9 (less than 2 m depth, in black sandy silt with small amounts of clay; collected 22 April 1960), and four individuals from SQ 5 (less than 2 m depth, TUBIFICIDATL in dark gr ollected pril 19 Museum of Natural Sciences, Ott ( catalogue number 3475, One ind holotype. Etymology.—"capillatu L, “lon hair’; hence species has hair setae posterior! Description.—Up to 58 mm long. Diameter 0.24 to 0.41 mm mm at segment XI, 0.33 mm tapering to a mir of 0.16 mm long as, or shorter than it is wide at the peristomiur junction. Clitellum weakly XI and % XII. Dorsal segments II each bundle contains 3 to 4, sometimes 5, bifid 75 to 85 w long, with the upper tooth slightly longer than the lower; from about segment XIII to terminal segment each bundle contains two or three smooth hair setae 110 to 125 yw long, and elongate, single-pointed crochets 80 to 95 u modified crochets and hair setae are similar in appear- ance but differ in the latter’s lack of a node, more slender form, and greater length. Ventral setae: teriorly each bundle contains three to five bifid setae 70 to 88 uw long, with the upper tooth slightly longer than the lower; posteriorly each bundle has two setae similar in form and length to anterior setae. No modified genital setae. Paired spermathecal and male pores are situated in the line of the ventral setae on segments X and XI respectively. Male genital system (all structures paired): y deferens 25 to 31 u diameter, about 1.25 mm long (in holotype), joins the atrium subapically and dorsally, opposite to the prostate gland. Atria hook—to commar-shaped, reflexed posteriorly, about 340 4» in total length, 54 to 100 « diameter; atria reach their maximum diameter distally to vas deferens prostate junctions; internal lining epithelium of this swelling more glandular than remaining atrial lining. Atria terminate in cuticularized penes 105 to 135 u long, 50 to 70 « diameter. Prostate gland relatively small, more or less enclosed by “hook” of atrium. Paired spermathecae with large ovoid to sacciform ampullae and barrel-shaped ducts about 135 uw long. 60 u diameter. Spermatozeugmata elongated ovoids, about 260 u long, 63 « maximum diameter. segment anteri rounded, a posteriorly. Prostomium developed on segment to X eae setac: in two or three long: an- as Remarks.—Tubifex postcapillatus differs from other members the genus in possessing hair setae and simple-pointed crochets only in of posterior segments. In other species, where they i tend occur, hair setae are always present in, and tend to be more numerous on, anterior red P. heterochaetus (Jaroshen! Peloscolex species [e. sen, 1926), P. swirencovi and P. intermedius Cook, 1969] but genus is characterized by the possession of a papillate or ridged body wall. at leas = t life cycle: the distinctions between 7 nd 128 Figure 1. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Tubifex postcapillatus, n. sp. A, Longitudinal view of genital segments; B, Distal end of atrium (optical section); C, Penis; D, Anterior dorsal seta; E, Anterior ventral seta; F, Posterior dorsal crochet; G, Part of hair seta (same bundle as F). vas deferens; 4, atrial ampulla; 5, 2, spermatozeugma; 3, 1, spermathecal duct; prostate gland; 6, penial sac; 7, atrial muscle layer; 8, glandular epithelium of distal atrium; 9, penial sheath. Peloscolex, which are complex and probably arti- ficial, are discussed in Pickavance and Cook (1971). Immature, semi-mature and mature indi- viduals of T. postcapillatus which were examined showed no trace of papillation, body wall ridges or accumulations of foreign particles, and is there- fore precluded from Peloscolex as this is presently defined. Thalassodrilus Brinkhurst, 1963 Definition—Hair setae __ absent. Penial setae present or absent. Body wall smooth. No gut divertic- ulae. Thick muscle layer associated with paired atria, at least terminally. Wide vasa deferentia, shorter than, or about as long as the atria, enter the latter apically. Each atrium with a small compact prostate gland. Spermathecae small or absent. Sperm in spermathecae in random masses; spermatozeugmata absent. Coelomocytes absent. Type species —Rhyacodrilus prostatus Kndollner, 1935. Remarks.—Thalassodrilus was established by Brinkhurst (1963) to accommodate R. prostatus. Hrabé (1967) rejected the genus and placed R. prostatus in Limnodriloides but later (Hrabe, 1971b) modified his opinion, redefined Thalasso- drilus, and included in it R. prostatus, Limno- driloides roseus Pierantoni, 1903, Limnodriloides pectinatus Pierantoni, 1903, and Limnodriloides gurwitschi Hrabé, 1971 (1971la). The modified definition of Thalassodrilus given above differs from Brinkhurst’s (1963) and Hrabé’s (1971b) conception of the genus by its specific reference to the heavy musculature associated with the male genitalia, and contains 7. prostatus (Knollner), T. gurwitschi (Hrabé) and T. belli, n. sp. The de- scriptions of L. roseus and L. pectinatus are in- 1974 adequate and in this author's opinion should remain as species inquirendae in Spiridion (Cook, 1969; Brinkhurst and Jamieson, 1971); they can- not be included in either Limnodriloides because they lack gut diverticulae, or Thalassodrilus be- cause they do not possess heavy atrial or pseudo- penial musculature. Some problems with Thalassodrilus do remain however. Thalassodrilus gurwitschi and T. belli are clearly closely related and, except for their lack of gut diverticulae and possession of relatively wide vasa deferentia, could be considered as spe- cies of Limnodriloides; both differ from 7. pro- status in their common possession of — thick muscular pseudopenes* and distinct thin-walled atria, rather than the thick, muscular, pear-shaped structures thought to be the atria in 7. prostatus. It is argued that the latter could be derived from a T. gurwitschi-like ancestor by the excessive or precocious development of the pseudopenial musculature which finally enveloped and_ in- corporated the original atrium both morpho- logically and functionally. The author feels that because of this possible mechanism together with the possession of relatively wide vasa deferentia and the other, mainly negative, characters included in the generic definition, the three species here included in Thalassodrilus form a convenient, and possibly monophyletic group. An additional ad- vantage of the arrangement is the removal of T. gurwitschi (and T. belli) from Limnodriloides which therefore remains the homogenous taxon defined in Cook (1969) (and see discussion under Limnodriloides, below). KEY TO SPECIES OF THALASSODRILUS la. Penial setae present. Atria pear-shaped with thick muscular walls _.. T. prostatus (Knollner) b. Penial setae absent. Atria elongate, thin-walled, with muscular pseudopenes terminally — 2 2a. Spermathecae present T. belli, n. sp. Spermathecae absent _...... T. gurwitschi (Hrabé) Thalassodrilus belli, new species Figure 2 Holotype-—USNM 45287. Bahia de San Quintin, Station SQ 3. Depth, less than 2 m, in gray clay. Collected 22 April 1960. Paratypes—USNM 45288. One individual from SQ 1 (in black mud); four individuals from SQ 2 (in dark gray clay); one individual from SQ 6 (very dark clay, some lighter clay); one individual from SQ 7 (black sandy silt); for all stations depth, PACIFIC COAST TUBIFICIDAL less than 2 m; One individual from SO dark gray silt. Collected 22 Etymology.—The species is of the late Dr. A. W. Bell part of this collection. Description.—55 to 63 collected 22 April 5. Depth, le 1960 1960 April named I who began to seu ments 13 mm long. 0.32 to 0.43 mm diameter anteriorly, 0.4 to 0.61 mm at segment XI, 0.4 mm tapering to 0.7 mm posteriorly. Prostomium shorter than it is at the peristomium junction. Clitellum segment XI and a small part of segment XII. Dorsal and ventral setae similar in number, size and shape: se i Wide restricted to tae three to five per bundle anteriorly, two per bundle posteriorly; setae bifid, 69 to 105 » long, with the upper tooth slightly thinner than, but as long as, the lower; setae in the middle region of the body tend to be shorter than others. Paired male and sperma- thecal pores open in the line of the ventral spermathecal pores open near intersegmental furrow IX/X. Pharyngeal glands in segments IV and V. agogen cells begin in segment VI. gut diverticulae. Male genital system (all structures paired): vas deferens, about 265 uw long, 20 to 30 uw diameter, joins atrium apically. Atrium about 335 uw long, with an apical swelling 47 to 54 u diameter, and a long duct 20 to 26 uw diameter which is con- voluted or folded proximally. Folded part is enclosed in a muscular sac, forming a protrusible pseudopenis. Prostate gland small, joined to the atrial swelling ventrally. Paired spermathecae relatively very small, pear-shaped, 70 to 110 uw diameter with very short, indistinctly separated ducts. Sperm in spermathecae in random masses. setac; Chlor- No discernable Remarks.—The general dimensions, the peculiar form of the male genitalia, the number and form of the setae, and the absence of gut diverticulae, are all characters similar in, or com- mon to, both T. belli and T. gurwitschi. The latter species, however, has no trace of spermathecae in the many mature and immature individuals studied by Hrabé (1971a; pers. comm.). A further dis- tinction between the two species is found in the setae; in T. gurwitschi the upper tooth of each seta is distinctly thinner at the base than the lower tooth (Hrabé, 1971a, Fig. 1 a—c) whereas in T. belli the basal width of the teeth are approxi- mately equal (Fig. 2d). Limnodriloides Pierantoni, 1903 Definition—Hair setae absent: somatic setae 2 bifid crochets. Penial setae absent. Body wall smooth or, rarely, with raised papillae. preclitellar region with a pair of diverticulae atria divisible into well-defined ampull regions, both with relatively thin muscle \ 130 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Figure 2. Thalassodrilus belli, n. sp. A, Longitudinal view of genital segments; B, Transverse section of male pore; C, Transverse section of spermatheca; D, Seta (segment VIII). 1, spermatheca; 2, vas deferens; 3, epithelium of clitellum; 7, circular muscle; 8, prostate gland; 4, atrial ampulla; 5, penial complex; 6, muscle sac surrounding penis; 9, lining epithelium of penial duct; 10, longitudinal muscle; 11, sperm mass. deferentia about as long as, or slightly longer than, the atrial ampullae (excluding the duct) which they join more or less apically. Compact prostate glands join atria on discrete, broad prostatic ducts. Penial structures variable in form but are rarely absent. Spermathecae relatively large. Spermatozeugmata absent, but in most species sperm in the spermathecae are organized into discrete, oriented bundles. Coelo- mocytes absent. Type species. antoni, 1903. Limnodriloides appendiculatus Pier- Remarks.——A_ short history of Limnodri- loides, a revised diagnosis of the genus, and rea- sons for removing S. roseus (Pierantoni) and S. pectinatus (Pierantoni) from it, are given in Cook (1969) (also vide supra under Thalasso- drilus). Briefly, the presence of the unique gut diverticulae found in the eighth or ninth segment is considered as the positive unifying somatic char- acter of Limnodriloides. Hrabe (1971b) appar- ently accepted this proposition but, on the basis of the structure of the intromittent organs, split the genus into two; he restricted Limnodriloides to those species possessing pseudopenes (containing L. appendiculatus, L. agnes, Hrabé, 1967, and L. winckelmanni Michaelsen, 1914) and established Bohadschia for the species with true penes (con- taining L. medioporus Cook, 1969, B. maslini- censis Hrabe, 1971, and B. pierantonii Hrabé, 1971). In this paper three new species are de- scribed; L. monothecus and L. verrucosus which both possess pseudopenes (i.e. Limnodriloides sensu Hrabé) and L. barnardi which has complex intromittent organs showing characteristics of both pseudopenes and true penes (see remarks following the description of this species). Be- cause of this intermediate type and because L. bar- nardi appears to be very close to L. winckelmanni, the distinction between Hrabé’s Limnodriloides and Bohadschia can no longer be considered valid, even at the subgeneric level, therefore Cook’s (1969) conception of Limnodriloides is retained in this account. Limnodriloides monothecus is analogous to P. monospermathecus (Knollner, 1935) in Phallo- drilus; in both cases the possession of a single 1974 dorsal spermatheca, rather than paired ventral ones, is considered to be significant at the species level only. However, the organization of the sperm in the spermatheca of L. monothecus, and other members of the genus, needs closer exam- ination. In L. appendiculatus, L. agnes, and L. medio- porus, the sperm is arranged in the spermathecae in oriented bundles (Hrabé, 1967; Cook, 1969); in L. winekelmanni and L. barnardi the sperm bundles are more regular in form than those of the three former species (Michaelsen, 1914; Hrabe, 1967; vide infra); both Michaelsen and Hrabe (op. cit.) refer to these structures as spermato- phores or spermatozeugmata in L. winckelmanni, and Hrabé (1967) is unable to decide whether the sperm bundles of L. agnes are spermatozeug- mata or not. Observation of the sperm bundles of L. medioporus and L. barnardi has shown that the sperm tails tend to be organized into parallel bundles, some of which appear to have coalesced or adhered closely to each other. The orientation of sperm is developed to a higher degree in L. monothecus; in this species the sperm bundles are very long and narrow with the sperm heads (darkly staining in haematoxylin) concentrated at one end of the bundle, and the sperm tails, arranged in very loose spirals, composing the remaining two-thirds of the bundle (Fig. 4b). The term “spermatozeugma” is used in the Tubi- ficidae to denote a sperm aggregation in which sperm heads occur around a central axis with the sperm tails, which are embedded in a cement- ing substance, radiating from this axis in a spiral manner (Stephenson, 1930; Brinkhurst and Jamie- son, 1971). The sperm bundles of Limnodriloides, including the highly organized bundles of L. monothecus cannot therefore be considered as spermatozeugmata. A character which has been added to the generic diagnosis of Limnodriloides is the nature of the body wall. This is necessary because of the dis- covery of L. verrucosus which, although clearly a Limnodriloides (form of the male genitalia and presence of gut diverticulae) has a papillate body wall superficially indistinguishable from that of many species of Peloscolex (see also remarks following the description of L. verrucosus). KEY TO SPECIES OF LIMNODRILOIDES la. One pair long, hollow-ended spermathecal setae, each contained in large muscle-coated sac, situ- ated on segment X. Atrial duct lined, at least in part, with glandular epithelium ——_ ; 2 PACIFIC COAST PUBIFICIDATL b. Modified spermathecal without glandular lining epitheli 2a. Intromittent organs small pseu surrounding ends of than setal shaft ity ipermathecal winckelmanni Mi b. Intromittent organs large penes (in part) er in voluminous penial sacs. Muscle surro ends of spermathecal setae about as thich shaft L. barnardi { 3a. Single, clongate, carrot-shaped spermatheca pre ent, opening mid-dorsally — L. b. Spermathecae paired, opening ventrally or laterally 4 4a. Body wall papillate, at least in postclitellar region L. verrucosus, n. sp. b. Body wall smooth 5 Sa. In posterior one per bundle Atrial duct at least as long as vas deferens L. agnes Hrabé. b. In posterior segments setae two per bundle. Atrial duct shorter than vas deferens 6 6a. Genital pores contained in median ventral folds in body wall, appearing externally as elongate median slits arranged transversely aie ee L. medioporus Cook. b. Genital pores paired, ventro-lateral 7 7a. Gut diverticulae in segment VIII : : _ L. appendiculatus Pierantoni. b. Gut diverticulae in segment IX 8 8a. In preclitellar segments setae two per bundle. Vasa deferentia 9 to 13 « wide. Terminal part of male ducts S-shaped : = eS L. maslinicensis (Hrabé). b. In preclitellar segments setae usually three per bundle. Vasa deferentia 16 uw wide. Terminal part of male ducts straight en Sere ss L. pierantonii (Hrabé). monothecus, 1 ventro segments setae Limnodriloides monothecus, new species we Figure Holotype-—USNM 45285. Bahia de San Quintin. station SQ 16. Depth, less than 2 m, in dark grey sandy silt. Collected 23 April 1960. Paratypes—USNM_ 45286. Six individuals. data as for holotype: one individual from SQ 37 (Depth, less than 2 m, in grey silt, collected 24 April 1960). NMC 3478. One individual from SQ 9 (Depth, less than 2 m, in black sandy silt with small amounts of clay, collected 22 April 1960). Etymolcgy.—*mono” = Gr. “one. single”: = Gr. and L. “case, receptacle”; hence “sing theca.” Description—Absut 46 segments. 9.5 mm ng 0.12 to 0.23 mm _ diameter .27 mm at segment XI. 0.23 O 3 mn posteriorly. Prostomium rounded. shorter than is wide at the peristomium jv Clitellun ery weakly developed or indiscern: Dors nd ven- ~ w& Ne Figure 3. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 56 VOLUME 73 Limnodriloides monothecus, n. sp. A, Longitudinal view of genital segments; B, Sperm bundle; C, Somatic seta; D, Proximal end of spermatheca; E, Penis sac (resting position); F, Penis sac (everted position). ; Spermathecal pore; 2, spermatheca; 3, vas deferens; 4, male pore; 5, prostate gland; 6, atrial ampulla; 7, atrial duct; 8, penis rectractor muscles; 9, penis sac epithelium. tral setae similar in number, size and shape: setae 3, sometimes two per bundle in segments II to IX, absent in X and XI, and two per bundle from seg- ment XII to terminal segment; setae bifid, 40 to 50 uw long, with upper tooth shorter and thinner than the lower. No modified genital setae. A single mid- dorsal spermathecal pore is situated near interseg- mental furrow IX/X. Male pores paired, ventro- lateral. Pharyngeal glands present in segments IV and V. A pair of diverticulae join the gut in the middle of segment IX. Male genital system (all structures paired): relatively thick vas deferens, 23 to 34 u diameter, 245 to 300 uw long, joins atrium more or less apically. Atrium consists of an ovoid ampulla, 90 to 130 uw long, 35 to 50 uw diameter, and an elongate, tubular duct 450 to 550 uw long, 13 to 27 u diameter, which opens to the exterior in an evaginable sac from the median side; sacs 90 to 105 uw long, 60 to 77 « maximum diameter when evaginated. Prostate gland joins atrium ventrally, near to the ampulla/ duct junction. Single median spermatheca elongate, carrot-shaped, 280 to 400 uw long, 55 to 85 mw maxi- mum diameter, tapering to about 20 u distally. Sperm in spermatheca oriented into long narrow bundles 200 to 260 uw long, 6.8 «~ maximum diameter, with the darkly staining sperm heads all occurring at one end of the bundle. Remarks.—The _ single, mid-dorsal theca found in L. monothecus, though unique in is also known in Phallodrilus monospermathecus. sperma- Limnodriloides, Limnodriloides verrucosus, new species Figure 4 Holotype—USNM 45283. Bahia de San Quintin, station SQ 41. Depth, less than 2 m, in gray very fine sandy silt. Collected 24 April 1960. 1974 Figure 4. Limnodriloides verrucosus, 0. transverse axis of body); E, Somatic seta. sp. A, Longitudinal view of genital segments: Prostomium and first two segments; C, Penis; D, Body wall of posterior segment (x — y 1, atrial ampulla; 4, prostate gland; 5, atrial duct; 6, circular muscle layer of atrial duct: PACIFIC COAST TUBIFICIDAI spermathecal ampulla; 2, vas deferens: atrial duct lining epithelium; 8, penis protractor muscle; 9, body wall papilla. Paratypes—USNM_ 45284. One individual from SQ 39 (in gray very fine sand; collected 23 April 1960); three individuals from SQ 46 (in gray silty very fine sand; collected 24 April 1960); two indi- viduals from SQ 49 (in gray very fine silty sand; collected 24 April 1960); two individuals from SQ 58 (in gray silty very fine sand; collected 24 April 1960). NMC 3476. One individual from SQ 54 (in gray silty fine sand; collected 24 April 1960). Depths (all paratypes), less than 2 m. Etymology.—‘verrucosus” = L. “warty.” Description—Up to 45 segments. 7.5 to 13 mm long. 0.12 to 0.18 mm diameter anteriorly, 0.18 to 0.25 mm at segment XI, 0.17 tapering to 0.08 mm posteriorly. Prostomium a little shorter than it is wide at the peristomium junction, with a small, thin- walled papilla anteriorly. Body wall covered with a sparse to very dense layer of raised papillae and small foreign particles, usually beginning just behind the clitellum in mature individuals, but occurring in anterior segments of immature specimens. Clitellum confined to segments XI and a small part of XII. Dorsal and ventral setae similar in number. size and shape: setae two per bundle in segments II to VII and one per bundle from VIII to the terminal seg- ment; setae absent on segments X and XI (no modi- fied genital setae); all setae bifid, 40 to 65 u long. with the upper tooth thinner than, but about as long as the lower. Male and spermathecal pores paired in the line of the ventral setae. Pharyngeal glands in segments IV and V. A pair of diverticulae join the gut ventro-laterally in the middle region of segment IX and extend anteriad to septum VIII/IX. Male genitalia (all structures paired): vas deferens, 7.3 to 10.5 uw diameter and about as long as, or slightly shorter than the atrium. join latter apically. Atrium thin, elongate, with thickly stalked prostate gland joining ventrally on a raised papilla. Atnum wit! drical ampulla about 100 u long. 13.7 eter, and a tubular duct 8 to 11.5 uw diar to about 21 u terminally: total length of 220 zp. Long of Muscles from swollen end of atria its axis atrium 134 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 wall, forming a protrusible pseudopenis. Paired Paratypes—USNM — 48731. Two — individuals, spermathecae with ovoid to elongate ampullae 97 data as for holotype. Two individuals from SQ 8 to 113 « long, 48 to 57 « wide, and discrete ducts (in black silt; collected 22 April 1960). One indi- 57 to 73 u long, 14 to 19 uw diameter. Sperm in vidual from each of SQ 19, 22, 30, 37, 46, SI, 85, spermathecae in random masses (no trace of oriented bundles). Remarks.—tThe | papillate wall of L. verrucosus is unique in Limnodriloides, but is morphologically indistinguishable from the con- dition found in many Peloscolex species. In the latter genus wide intraspecific variation of the nature of the body wall is known: Dahl (1960) found that Peloscolex benedeni (Udekem) shed its cuticle and papillae at maturity, and Brink- hurst (1964) states that the freshwater Peloscolex ferox (Eisen) attains the papillate condition from a nonpapillate juvenile and then periodically, though not strictly seasonally, sheds its papillae. A study of 197 individuals of L. verrucosus in different states of maturity (Types and USNM 45292) has revealed the following variation pat- tern in its body wall: papillae are developed on segments IV or V to the terminal segment, prior to, or immediately after, emerging from the cocoon; the papillae grow with the animal and may also develop on the first few anterior seg- ments; as the individual reaches sexual maturity there appears to be a progressive loss of papillae from the anterior end until the body wall is papillate only in the postclitellar region; after copulation and cocoon deposition, it appears that both the genitalia and the papillae regress, or the latter shed (as in Peloscolex). The general body form and dimensions, the papillate body wall, and the number and form of the setae of L. verrucosus are strikingly similar to those of Peloscolex gabriellae Marcus, 1950. The latter appears to be a widespread, amphi- american, marine species, or species complex, with very wide ecological tolerances (Cook, 1970b), and because its setal pattern is unique within Peloscolex, its presence is sometimes diag- nosed on the basis of immature individuals. How- ever, unless the gut diverticulum character, which is rarely checked in routine tubificid identifica- tions, is examined, immature P. gabriellae and L. verrucosus can be mistaken for each other. body Limnodriloides barnardi, new species Figure 5 Holotype-—USNM 48730. Bahia de San Quintin, station SQ 16. Depth, less than 2 m, in dark grey sandy silt. Collected 23 April 1960. 91: two individuals from SQ 15. NMC 3480. One individual from SQ 2. All paratypes collected April 1960 at a depth of less than 2 m. Additional (non-type) material—USNM_ 45291 and 45297. Etymology—Named_ for J. Laurens Barnard (Smithsonian Institution) who led the expedition to Bahia de San Quintin. Description—About 35 segments. 8 to 10 mm long. 0.12 to 0.20 mm in diameter anteriorly, 0.25 to 0.30 mm at clitellum, 0.10 to 0.16 mm posteriorly. Prostomium a little shorter than it is wide at the peristomium junction, with a small papilla anteriorly. Clitellum weakly developed on segments XI and XII. Dorsal and ventral setae similar in number, size and shape: somatic setae bifid, 40 to 60 » long, with subequal teeth; in preclitellar region each setal bundle contains 3, sometimes two or four setae; pos- teriorly each bundle contains two setae. Each ven- tral bundle of segment X contains one slender, straight or curved, elongate spermathecal seta 110 to 120 uw long, which has a hollow groove from the node to the distal end; proximal end of spermathecal setae, on which the setal protractor muscles are in- serted, are strongly hooked; distal half of sperma- thecal setae each enclosed in a_ pear-shaped sac, 65 to 70 uw long, 40 to 48 uw diameter, which consists of an inner layer of vacuolated cells and an outer muscle layer, and which connects with the body wall musculature. Ventral setae absent on segment XI. Paired male pores open in the line of the ventral setae on segment XI. Paired spermathecal pores open in the line of, or slightly lateral to the sperma- thecal setae (out of a total of 25 individuals ex- amined for the character, 18 had spermathecal pores located anterior to the spermathecal setae, five had pores posterior to the setae, and two were without spermathecal setae). Pharyngeal glands present in segments IV and V. Chloragogen cells begin in segment VI. Male genital system (all structures paired): conical male funnel situated ventrally on septum X/XI drains into the vas deferens, 9 to 12 mw diameter, about 180 uw long, which joins the atrium apically; atrium, whose long axis is directed anteriorly, consists of an ovoid to elongate ampulla, 100 to 110 uw long, 35 to 60 uw diameter, and an elongate duct, 95 to 135 u long, 25 to 35 « diameter, which terminates as the penis; a large, thickly-stalked prostate gland joins the atrial ampulla ventrally. Atrial ampulla consists of a thin outer muscle layer and an inner layer of thin lining cells. About the first 90 to 100 uw of the atrial duct (ad- jacent to the ampulla) is lined with thick, highly glandular cells. The penial complex consists of a more or less circular infolding of the body wall (the penial sac) 60 to 70 uw deep, 45 to 50 « maximum 1974 PACIFIC: COAST TUBIFICIDAT Figure 5. Limnodriloides barnardi, n. Spermathecal seta; C, Somatic seta. 1, muscular sac surrounding spermathecal seta; 5, male funnel; 6, vas deferens; 7, prostate glandular part of atrial duct. sp. diameter, and a truncated-cone-shaped penis, through which penetrates the terminal part of the atrial duct; the latter opens on the penis subterminally and laterally. Paired spermathecae with discrete ducts 40 to 55 uw long, 30 to 35 w maximum diameter, and elongate ampullae 200 to 220 u long, 70 to 120 u maximum diameter; latter tend to be constricted in the middle third of their length. Sperm in the sperma- thecae oriented into long narrow bundles, 95 to 135 uw long. Remarks.—Morphologically, L. barnardi is very closely related to L. winckelmanni. Their atria and atrial ducts with glandular lining cells are, on available evidence, indistinguishable, and_ their spermathecal setae contained within vacuolated sacs are unique in the genus; the number and form of their setae are also similar. A com- parison of the information available on L. winckel- manni, which is apparently derived from the type specimens alone (Michaelsen, 1914; Boldt, 1928; Hrabe, 1967; 1971b) with the description of L. barnardi, reveals that their intromittent organs differ (vide infra) and that the general dimen- sions of L. barnardi are smaller than those of L. winckelmanni (maximum body length, clitellum diameter and setal length of the two species are, respectively, 10 and 18 mm, 0.3 and 0.6 mm, 60 A, Longitudinal view of genital segments; B. sperm bundle; 2, spermathecal ampulla; 3, vacuolated 4, protractor muscles of spermathecal seta; gland; 8, penial sac; 9, atrial ampulla; 10, and 90 »). The dimensions are probably not im- portant characters as wide intra-specific variation in size is well known for many common tubificids and lumbriculids, but the different structures of their intromittent organs is more fundamental. Hrabé (1971b) showed that in L. winckelmanni small pseudopenial papillae the simple openings of the male ducts from the side. In L. barnardi, however, a large penial sac contains the terminal end of each atrial duct which opens sub- terminally on a truncated cone-shaped penis: em- ploying present definitions of the terms, it would appear that the proximal part of this organ can be termed a true penis while the part distal to the actual male orifice could be pseudopenis. cover considered as a ECOLOGICAL COMMENTS The eastern arm of Bahia de San Quintin (re- ferred to hereafter merely as “the bay is shallow body of water of more or less uniform salinity. At high tide the water depth over about 85 percent of the bay is less than two m: narrow. steep-sided channels to ei account for the remaining 15 (six 136 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Taste |. Station data and number of tubificids in each bay sample. A—station number; B—fraction of sample counted and identified (S indicates subsamples of 30 worms); C—total number of tubificids, including immature specimens (numbers in brackets are estimated); D—mean particle diameter in 4»; 1—Tubifex post- capillatus; 2—Thalassodrilus belli; 3—L. monothecus; 4—L. verrucosus; 5—L. barnardi. A. ay, C. Da i, 2. Be 4. 5. 1 Ss (130) 7 4 7 = 13 3 2 all iI 6 _ 8 — D} 1 3 all 9 7 = 4 mu - 2 4 all 1 7 I = = 2 ~ 5 Ss (300) 6 3 | = 2 5 6 S ((Z/S))) 9 - 2 - 13 = 7 all 44 30 3 = 12 10 8 Ss (200) 13 = = 5 = 13 9 Ss (100) 16 4 2 1 4 5 10 S (200) 38 = = 2 = 22 11 Ss (150) 21 = = 3 us 17 12 all 23 98 = 1 = 8 13 10% 4 26 = = = = 2} 14 Ss (250) 56 = =~ 3 2 17 15 20% 33 71 = ze, 1 = 16 16 all 34 32 = = 8 1 16 17 Ss (150) 22 = = 2 = 21 18 all 11 61 = = 1 = 1 19 Ss (300) 49 = - 5 3 16 20 all 26 105 - = 3 4 11 22 all 1 174 = = = = 23 Ss (400) 36 = = 5 = 23 24 all 7 13 = = = 1 1 25 all 2 143 = = = = 1 26 S (300) 19 11 = = 5 2 30 Moe = 6 19 = = = 4 31 10% 21 52 i 2 ES = 4 32 all 16 33 = = I = 3 35 all 47 58 = = 3 18 16 36 Ss (80) 51 = a 5 = 21 37 Ss (250) 31 = = 3 6 9 38 all 58 43 = = 1 26 21 39 Sand 15% (100) 54 = = = 23 q] 40 all 3 242 = = 1 = 1 4] S (150) 33 = = 5 11 11 42 Ss (100) ? = = = 17 5 43 S (500) 53 13 = = 1 11 45 all 6 36 = = = 1 2 46 all 11 51 = = = 10 1 47 Ss (275) 79 = = 6 = 21 49 Ss (225) ? = = = 28 = SO all 37 ? - = 37 = 51 Ss (75) 63 = = 2 4 16 53 all 1 78 = — = a 1 54 Ss (175) 9 a = ze 25 5 58 all 30 46 = “ = 23 7 60 S (75) 54 = = a 28 2 64 Ss (250) 48 = = 2: 14 13 74 all 4 103 es a Bs ” 4 75 all 4 142 - = = 1 2 76 Ss (75) 66 = 9 2 17 79 all 1 55 = = = 1 A 85 all 11 88 = 2 ws = 8 88 all 18 137 = ss zs 1 15 1974 1962) and at low tide much of the bay floor is exposed or only sparsely covered by shallow pools. The depth and salinity of the water, per se, are not considered to be significant factors in’ con- trolling the distribution of the Tubificidae in this study although the amount of exposure of the habi- tat at low tide, the degree of evaporation, and the water retention characteristics of the sediment, probably are. Such data are not available, but related sediment parameters, the mean particle size and sorting coefficient, are known for most of the bay stations. A textural analysis of the bottom sediments demonstrated that a continuous range of conditions exist in the bay, from coarse, well-sorted sands, to ‘poorly-sorted clays which dominate the north end of the bay; a complex mosaic of intermediate sediment types occur in the central regions, and sands predominate in the bay’s south end (Gorsline and Stewart, 1962). The latter authors also showed that, in general, the organic content of the sediment increased with decreasing particle size, hence it is probable that the amount of food available to oligochaetes is inversely proportional to this parameter. There- fore, the mean particle size of the sediment is thought to be a direct or indirect measurement of some factors determining tubificid distributions in this study, and will be considered in more detail below. The numbers of each tubificid species identi- fied from each San Quintin station (each sample = 0.065 m*) and other relevant data are sum- marized in Table 1; for purposes of brevity, and to facilitate comparisons with other studies, mean sediment particle sizes are discussed in terms of units (¢@ = logy particle diameter in mm, hence diameters of 4-8 » = 8 6, 8-16 n=7 g, 16 —- 31 p= 6 ¢, 31-62 p=5 g, 62-125 p= 4 4, 125 — 250 » = 3 g). From this information, and the locations of the benthic stations (Fig. 6) the distribution and density of each species can be inferred. Abstracting and comparing pertinent data from table 1, it is evident that the three species of Limnodriloides bear a definite relationship to very fine sand and silt (4 to 6 ¢ optimum) in terms of both their frequency of occurrence and population size. The dominant tubificid, L. bar- nardi, occurred at 48 stations comprising all sedi- ment classes, but was most frequently found in 4 and 5 ¢@ sediments (21 percent and 40 percent of its stations respectively); maximum population density of L. barnardi occurred in the 4 to 6 4 range. L. monothecus occurred at 21 stations PACIFIC COAST TUBIFICIDAE of GAM OUINTIM BAHIA BENTHIC STATIONS Figure 6. Locations of benthic sampling stations in Bahia de San Quintin; redrawn from Barnard, 1962. of which 52 percent had 5 ¢ sediments and 24 percent were 4 ¢. However. the population den- sity of this species was remarkably constant over its whole sediment range of 4 to 7 ¢ (about 280/m2). Of the 28 stations occupied by L. verrucosus, half had sediments in the 5 ¢ class. with the remainder distributed fairly evenly over the whole sediment range of 3 to 8 ¢: however. the abundance of L. verrucosus was highest in 5 to 6 ¢ and 8 ¢ substrates which may indicate that while it is capable of exploiting the finest sediments, it is partially excluded from them in this situation by competition with other deposit feeders or by the effects of predation. The two remaining tubificids, Tubijex post- capillatus and Thalassodrilus belli, appear to be associated with the finest sediments which are located mainly at the north end of Tubifex belli was present in only eigl but half of these were in the § ¢ class 138 the maximum abundance for this species (460 m2) was also recorded. Only six stations were occupied by 7. postcapillatus but three of these had 8 g sediments. strikingly more abundant in the 5 and 6 ¢ classes (about 3300 and 1700 worms/m* respectively, compared with about 250/m? in 8 ¢ sediments); except for the 8 @ class, these density figures were calculated from single samples whose num- bers were, in turn, derived from subsamples, there- fore large errors in them are possible. If the figures do reflect real trends, however, it is possible that T. postcapillatus has a wide sediment tolerance but is competitively restricted by T. belli or other detritivores in fine substrates (hence its low densities) and by Limnodriloides species in intermediate sediments; the latter is supported by the fact that at the 5 and 6 ¢ sediment stations where 7. postcapillatus was abundant, L. mono- thecus was absent while L. and L. barnardi were at relatively low densities. However, the species was verrucosus DISCUSSION That the distribution of freely burrowing benthic animals which feed upon their surrounding me- dium should be determined, in part at least, by sediment characteristics, is hardly surprising; at the extremes, a well-sorted, coarse-sand environ- ment could be compared to rocky ground, while fine silts and clays are comparable with a rich and fertile loam. In Bahia de San Quintin, avail- able habitats are restricted to the productive end of the scale and hence discrete communities of oligochaetes, such as those demonstrated in Cape Cod Bay, Massachusetts (Cook, 1971) cannot be delineated. However, even in the comparatively restricted sediment range of the bay, it is possible to recognize that 1) the finest silts and clay-silt are usually inhabited by T. postcapillatus and T. belli and 2) intermediate silts and fine-sands are usually exploited by L. barnardi, L. monothecus, and L. verrucosus. Although complicated by the differing availability of organic matter in the different substrate types, these trends support the conclusions of Bulow (1957), Brinkhurst and Kennedy (1962), Lasserre (1967), and Cook (1971) who all demonstrated that zones of differ- ing sediments support different oligochaete faunas in the marine environment; similarly Reish (1963) found that many polychaetes in Bahia de San Quintin were associated with definite sediment types. West coast marine oligochaetes are very poorly BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 known; the studies by Brinkhurst and Simmons (1968) reporting on the oligochaetes, mainly Tubificidae, of the San Francisco Bay system, Tynen’s (1969) description of some littoral Enchytraeidae from Vancouver Island, and the present paper, summarize existing knowledge of the class in the northeastern Pacific. Brinkhurst and Simmons (1968) found Peloscolex gabriellae to be the dominant oligochaete in San Pablo Bay, and south San Francisco Bay where Peloscolex apectinatus and Peloscolex nerthoides also oc- curred in significant numbers; these authors did not relate substrate particle size with species dis- tributions as it was felt that the study area was uni- formly available to the constituent species and that other, chemical or biological, determining factors dominated. This is undoubtedly true in some situa- tions, and especially so in the case of polluted waters (as in San Francisco Bay). As well as in- creasing nutrient concentrations, many sources of organic pollution can be expected to modify bottom deposits in the direction of lowering mean particle size and hence to decrease the diversity of niches available to free-burrowing benthic organisms. Thus these results, and those of authors who report definite correlations between sediment and species distributions, are not necessarily con- tradictory. For example, organic enrichment or increased siltation of Bahia de San Quintin could be expected to eliminate some of the tubificids described in this paper, and from the data pre- sented here it is possible that L. barnardi and L. monothecus could be the sensitive species; the modified distribution of the remaining three species of Tubificidae may, under these circum- stances, be very difficult to relate to any sedi- ment parameter. Indeed these speculations lead to the con- clusion that failure to detect a clear relationship between species distributions and physical sedi- mentary parameters may, if a diversity of sediment types are known to exist, indicate environmental deterioration. A growing preoccupation of aquatic biologists is the concern to detect, assess and monitor sources of environmental contamination and approaches have varied from a search for indi- cator species to sophisticated treatments of whole systems. The success of any biological investiga- tion, depends taxonomy; the point is stressed because this study however, basically on accurate has demonstrated that confusion between the two marine tubificids Limnodriloides verrucosus and Peloscolex gabriellae could arise if identifications 1974 are based on superficial characteristics (papilla- tion and setal pattern). The importance of this distinction further lies in the fact that P. gabriellae is resistant to high levels of organic pollution while the tolerance characteristics of L. verrucosus are unknown. ACKNOWLEDGMENTS My thanks to Marian H. Pettibone (USNM)_ for bringing this collection to my attention, and to J. Laurens Barnard (USNM) for helpful suggestions and advice. The study was supported by a _ Post- doctoral Fellowship sponsored by the National Re- search Council of Canada, and the National Mu- seum of Natural Sciences, Ottawa, whose assistance is greatly appreciated. LITERATURE CITED Barnard, J. L. 1962. Benthic marine exploration of Bahia de San Quintin, Baja California, 1960-61. Pac. Nat., 3:251-274. Boldt, W. 1928. Mitteilung uber Oligochaeten der Familie Tubificiden. Zool. Anz., 75:145—151. Brinkhurst, R. O. 1963. Notes on the brackish- water and marine species of Tubificidae (An- nelida, Oligochaeta). J. Mar. Biol. Assoc. U. K., 43:709-715. 1964. Observations on the biology of lake- dwelling Tubificidae. Arch. Hydrobiol., 60: 385-418. 1966. of the marine Tubificidae (Annelida, chaeta). Biol. Bull., 130:297—303. A contribution to the systematics Oligo- Brinkhurst, R. O., and B. G. M. Jamieson. 1971. Aquatic Oligochaeta of the World. Oliver and Boyd, Edinburgh, 1971: 860 pp. Brinkhurst, R. O., and C. R. Kennedy. 1962. Some aquatic Oligochaeta from the Isle of Man with special reference to the Silver Burn Estuary. Arch. Hydrobiol., 58 :367—376. Brinkhurst, R. O., and M. L. Simmons. 1968. The aquatic Oligochaeta of the San Francisco Bay system. California Fish Game, 54: 180-194. Bulow, T. 1957. Systematisch-autokologische Stu- dien an eulitoralen Oligochaeten der Kimbrischen Halbinsel. Kieler Meerforsch., 13:69-116. PACIFIC COAST TUBIFICIDAT Cook, D. G. 1969 The Tul Oligochaeta) of Cape Cod Ba revision of the Phallad) 1902, Picrant Spiridion Knollner Biol. Bull genera Limnodriloides 1935 1970a. Peloscolex dukei, n aculeatus, n. sp. (Oligochaeta, Tubificidac Atlantic, the latter the north-west being fron abyssal depths. Trans. Amer. Micro. S« 492-497, a 1970b. Bathyal and abyssal Tubificidae (Annelida, Oligochaeta) from the Gay Head Bermuda transect, with descriptions of new gen era and species. Deep-Sea Res., 17:973-981. 1971. The Tubificidae (Annelida, chaeta) of Cape Cod Bay II: ecology systematics, with the description of Phallodrilus parviatriatus nov. sp. Biol. Bull., 141:203-221 Oligo- and Dahl, I. O. 1960. The oligochaete fauna of 3 Danish brackish water areas. Medd. Dan. Fisk. Havunders., 2(26) :3—20. Gorsline, D. S., and R. A. Stewart. 1962. Benthic marine exploration of Bahia de San Quintin, Baja California, 1960-61. Marine quater- nary geology. Pac. Nat., 3:283-319. and Hrabé, S. 1966. New or insufficiently known spe- cies of the family Tubificidae. Spisy Piirodoved Fak. Univ. Brne, 470:57-77. 1967. Two new species of the family Tubi- ficidae from the Black Sea, with remarks about various species of the subfamily Tubificinae. Spisy Prirodoved Fak. Univ. Brne. 485:331-—356. 1971a. A note on the Oligochaeta of the Black Sea. Vést. Cs. Spol. Zool., 35:32-34. 1971b. On new marine Tubificidae of the Adriatic Sea. Scripta Fac. Sci. Nat. Ujep Brunensis, Biol., 3:215—226. Jaroshenko. M. F. 1948. Oligochaeta Dneprobug- skogo Limana. Nauch. Zap. Moldavskaya Nauch Issled. Baza Akad. Nauk SSR. 1:57-72. Kndéllner, F. H. 1935. Okologische atische Untersuchungen iiber litorale und marine Oligochiten der Kieler Bucht. Zool. Jb. (Syst 66:425-512. und System- Lasserre. P. 1967. Oligochétes m de France ITI. Roscoff, Penpoull. 140 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 de Concarneau: systématique, ecologie. Cah. Pierantoni, U. 1903. Altri nuovi oligocheti del Biol. Mar., 8:273-293. Golfo di Napoli (Limnodriloides n. gen.)—IL nota sui Tubificidae. Boll. Soc. Natur. Napoli, Marcus, E. du B.-R. 1950. A = marine tubificid 17: 185-192. from Brazil. Commun. Zool. Mus. Hist. Natur. Montevideo, 3(59): 1-6. Reish, D. J. 1963. A quantitative study of the benthic polychaetous annelids of Bahia de San Michaelsen, W. 1914. Beitrage zur Kenntnis der Quintin, Baja California. Pac. Nat., 3:399-436. Land-und — Siisswasserfauna Deutsch-Sitidwest- afrikas: Oligochaete a rg, pp. 139-182. : : atrikas:s OligochactajHambnre,spe yl] 7aas Stephenson, J. 1930. The Oligochaeta. Clarendon ; Press, Oxford. 978 pp. ———. 1926. Oligochaeten aus dem Ryck bei Greifswald und von benachbarten Meeresgebieten. : 2 Mitt. Zool. St. Inst. Hamburg, 42:21—29. Tynen, M. J. 1969. New Enchytraeidae (Oligo- chaeta) from the east coast of Vancouver Island. Pickavance, J. R., and D. G. Cook. 1971. Tubifex Can. J. Zool., 47:387-393. newfei n. sp. (Oligochaeta, Tubificidae) with a preliminary reappraisal of the genus. Can. J. Zool., 49:249-254. Accepted for publication February 12, 1974. A NEW SPECIES OF FOSSIL NUTTALLIA (MOLLUSCA: BIVALVIA) FROM THE PLIOCENE OF SONOMA COUNTY, CALIFORNIA Barry RotTH! AND RAJ GURUSWAMI-NAIDU? Apstract: WNuttallia jamesi, new species, is described from strata assigned Pliocene age in the Sebastopol quadrangle, central Sonoma County, California. The species is similar to N. nuttallii (Conrad), late Miocene to Recent of California and Baja California, and to N. olivacea (Jay), Miocene to Recent of Japan, but differs from both in having an acuminate posterior end. The genus Nuttallia is restricted, in both Recent and fossil occurrence, to the north Pacific basin. It represents a temperately adapted branch of the predominantly tropical Sanguinolariinae. Several years of field work by one of us (Guru- FAMILY PSAMMOBITDAE FLEMING, 1828 swami-Naidu) at an exposure of soft sandstone SUBFAMILY SANGUINOLARIINAE GRANT AND GALE, in central Sonoma County, California, mapped as 1931 Merced Formation by Travis (1952), have re- sulted in a collection of more than forty marine NSTC EN spccies mostly, moO) aa mu Nuttallia Dall, 1898: 58 [proposed as “section” of ous bird, fish, and mammal remains. Among the gayeuinolaria Lamarck, 1799]; Dall, 1900; Coan, mollusks represented is a new species of the 1973. tellinacean bivalve genus WNuttallia, described herein. An account of the associated fauna and paleoecology will appear in a report now in Genus Nuttallia Dall, 1898 ‘Dept. Geology, California Academy of Sciences, ; San Francisco, California 94118. preparation. “638 Hunter Lane, Santa Rosa, California 95404. 1974 Type species, by original designation, Sanguinolaria nuttallii Conrad, 1837; late Miocene to Recent, Cali- fornia and Baja California, Dall (1900) characterized Nuttallia as follows: “Shell large, suborbicular, inequivalve, more or less twisted, the right valve slightly flatter, the posterior cardinal in the left valve obsolete; the pallial sinus narrower in front and somewhat de- tached from the pallial line.” He gave this con- trasting description of Sanguinolaria, sensu stricto: “Shell moderately large, thin, equivalve, short, rose-colored or white, with short, inconspicuous nymphs, two bifid cardinal teeth in each valve; pallial sinus deep, widest in front, confluent with the pallial line below,. the epidermis thin, dehis- cent” (Dall, 1900). Living species of Nuttallia have a persistent, dark, generally glossy perios- tracum and a strong cardinal ligament borne on a projecting nymph. Fossil specimens with valves paired are relatively common. Coan’s (1973) review of Recent northwest American Psammo- biidae included a discussion of Nuttallia nuttallti (Conrad, 1837), type species of the genus and the only species known to be living in eastern Pacific waters. Kira (1953) reviewed the Recent Japanese species; Habe and Ito (1965) provide a later summary and synonymy. Recent authors are almost evenly divided on whether to call Nuttallia a genus or a subgenus. It is often ranked as a subgenus of Sanguinolaria Lamarck, 1799 (type species, Solen sanguino- lentus Gmelin, 1791), and occasionally as a subgenus of Soletellina Blainville, 1824 (type species, Soletellina radiata Blainville, = Solen diphos Linnaeus, 1771). (Soletellina is a junior synonym of Hiatula Modeer, 1793, since Solen diphos Linnaeus is the type species of the latter genus by subsequent designation [Stoliczka, 1870: 114]. A later designation [Winckworth, 1935] of Mya truncata Linnaeus, 1758, as type species of Hiatula is invalid.) As originally proposed by Grant and Gale (1931) the family Sanguinolari- idae contained the genera Gari Schumacher, 1817, Tagelus Gray, 1847, and Sanguinolaria (including “section” Nuttallia), and was thus practically coextensive with Coan’s (1973) usage of Psam- mobiidae Fleming, 1828. Keen (1969) reduced Sanguinolariidae to subfamily rank, including only the genus Sanguinolaria and under it seven sub- genera, Nuttallia among them. In both morphol- ogy and distribution, Nuttallia seems sufficiently distinct from Sanguinolaria to receive generic rank, with its affinity expressed at the subfamilial level. Typical Sanguinolaria species are tropical in NEW PLIOCENE BIVALVE FROM ¢ {LILFORNIA distribution and occur in the ¢ Pacific, and Panamic (eastern Paciti In the Panamic province, the northern for species of Sanguinolaria, sensu strict Gulf of California. That for S. (Psammot San Ignacio Baja California (1 1971), approximately 26° 45’ the 19° ¢€ 1950: fig. 2). Species of Nuttallia now occur in warm-temperate to cool waters in the north Pacili Lagoon, north latitude February surface isotherm (Durham Ocean, and fossil occurrences are restricted to the margins of the north Pacific basin. Recent N. nut tallii ranges from Bodega Bay, California (38° 20° N), to Magdalena Bay, Baja California (24° 40° N) (Coan, 1973). Its range includes the southern portion of the Oregonian molluscan province, all of the Californian province, and the northern part of the Surian province, in the Valentine (1966). Valentine's Surian province is essentially equivalent to the portion of the west coast of Baja California regarded by Addicott (1966) as an area of provincial overlap, with definitions of Californian forms predominating in outer-coast biotopes and Panamic forms dominant in pro- tected biotopes. By neither criterion is Nuttallia nuttallii a truly Panamic species, as one might infer from Coan’s (1973: table 3) distributional tabulation. Kuroda and Habe (1952) cite the western Pacific species Nuttallia petri (Bartsch, 1929) as ranging from 43°-46° N, Nuttallia olivacea (Jay, 1857) from 30°-43° N, and Nut- tallia “nuttallii(?)” [= N. ezonis Kuroda and Habe in Habe, 1955] from 39°—51° N. Both the western and eastern Pacific species which range into warm water—N. nuttallii and N. are also found considerably farther north, in cool- temperate marine climates. Phyletic separation of Nuttallia from Sanguino- laria dates from the Paleocene. The oldest known Nuttallia, N. townsendensis (Clark, 1925) from the late Eocene Quimper sandstone of Washing- ton, occurs in a fauna of distinctly tropical aspect (Durham, 1950). The similar N. Kamada, 1962, from the upper Oligocene or lower Miocene Asagai Formation of Honshu, occurs in a molluscan assemblage suggestive of warm-tem- perate conditions moderate depths. increasing adaptation to cooler conditions. olivacea— uchigoensis and deposition offshore in Later Tertiary species shov 7 species Nuttallia jamesi. Figures 1—3, 5— Diagnosis: A small, thin-shelled species of allial sinus, with acuminate posterior end. narrow pa 142 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Figures 1-3, 5. Nuttallia jamesi, new species; external and internal views of holotype (CAS Department of Geology No. 54330). Length 53.4 mm. Figure 4. Nuttallia nuttallii (Conrad); internal view of right valve with ligament attached; Recent, southern California (CAS Department of Geology No. 54335). Length 62.2 mm. Figure 6. Nuttallia jamesi, new species; dorsal view of paratype (CAS Department of Geology No. 54331). Length 46.8 mm. Figure 7. Nuttallia jamesi, new species; detail of hinge of left valve of paratype (CAS Department of Geology No. 54332), nymph partly broken. Figure 8. Nuttallia olivacea (Jay); internal view of right valve; Recent, Iwai Mura, Chiba Prefecture, Japan (CAS Department of Geology No. 54334). Length 54.0 mm. and hinge line sinuous in dorsal view, curving to the left posteriorly, to the right anteriorly. Description: Shell small for the genus, thin; in- equivalve, left valve more inflated than right valve; beaks situated at midline; anterior end of shell broad, evenly rounded, posterior end acuminate; surface of valves smooth except for incremental lines of varying strength. Hinge line shallowly sinuous in dorsal view, curving to the left posteriorly, to the right anteriorly. Right valve most inflated at about 14 of distance from beak to anterior end; compressed along both dorsal margins and along postero-ventral margin; with a radial angulation slightly below posterior dorsal mar- gin; beak small, pointed; hinge plate narrow, with two diverging, oblique cardinal teeth, the posterior tooth larger and bifid; anterior dorsal margin con- vex; posterior dorsal margin concave, with a narrow escutcheon proximally, in which rises an elongate, 1974 broad-topped nymph; pallial sinus narrow, deep, ex- tending about 4% of the distance anterior to midline, recurving before joining pallial line at an acute angle: muscle scars large; internal margin smooth. Left valve most inflated directly below beak; slightly compressed along posterior dorsal margin; with a blunt radial angulation running from beak to extreme posterior point of valve, and a shallow radial sulcus below this; beak elevated, small but prominent; hinge plate narrow, somewhat excavated, with a projecting, bifid anterior cardinal tooth (broken on holotype) and a small, nearly horizontal, laminar posterior cardinal tooth separated from the anterior tooth by a hollow; no lateral teeth present; posterior dorsal mar- gin with a narrow escutcheon, in which rises a nymph, pairing with that of the right valve; pallial sinus and muscle scars like those of the right valve; internal margin smooth. Dimensions: Holotype, length 53.4 mm; height 37.0 mm; diameter (both valves) 14.8 mm. Measured paratypes: (a) length 49.2 mm; height 36.6 mm; diameter 13.5 mm; (b) length 46.8 mm; height 33.9 mm; diameter 12.5 mm; (c) length 44.0 mm; height 32.9 mm; diameter 11.4 mm. Type material: Holotype, right and left valves, No. 54330, Department of Geology, California Academy of Sciences. Paratypes, Nos. 54331-54333, same institution. Paratypes have been deposited in the following institutions: Department of Paleobiology, U. S. National Museum of Natural History; Museum of Paleontology, University of California, Berkeley; Department of Paleontology, Los Angeles County Museum of Natural History; Department of Geology, Stanford University; Institute of Geology and Paleon- tology, Tohoku University, Sendai, Japan. Type locality: California Academy of Sciences De- partment of Geology locality No. 54164, roadcut in grayish-white, poorly indurated and_ highly fossil- iferous sandstone on north side of River Road, 0.2 miles north of town of Trenton and 0.3 miles east of intersection of River Road and Trenton-Healds- burg Road, United States Geological Survey Sebasto- pol, California, quadrangle, 15 minute series (topo- graphic), 1954 edition, Sonoma County, California (approximately 38° 29’ N, 122° 51’ W). On the geologic map of the Sebastopol quadrangle pre- pared by Travis (1952: pl. 1), the locality is at the fault contact between the Pliocene Merced Formation and “Jurassic” serpentine. The species is named for Mr. James Nikas, student of west American paleontology, who first suggested that the taxon was undescribed. DISCUSSION Comparisons: The only Nuttallia previously recorded from Pliocene strata in western North America is N. nuttallii (Conrad, 1837). It has been reported in the Jacalitos, Etchegoin, and San Joaquin Formations on the west side of the San NEW PLIOCENE BIVALVE FROM CALIFORNIA Joaquin Valley (Nomland, 1917 Stewart and Richards, 1940 [1941]). It most occurrence, Recent or fossil, is in the Pliocene Falor Formation (University ( fornia Museum of Paleontology invertel locality A-4234) of the Blue Lake quadran Humboldt County, California. These mens were reported by Manning and Ogle (1950) two speci as Sanguinolaria nuttallii, and typical of the species. Nuttallia nuttallii has a geo logic range of late Miocene (Neroly) (Weaver, 1949) to Recent. Nuttallia nuttallii (Fig. 4) is more inequivalve than N. jamesi, with the right valve nearly flat appear to be and the left valve strongly convex. The posterior end of N. jamesi is acutely pointed; that of N. nuttallii is broad, ending in an obtuse posterior angle. In dorsal view, the hinge line of N. nuttallii is less conspicuously sinuous than that of N. jamesi. The beaks of Nuttallia nuttallii are more anterior and the cardinal nymph is proportionally shorter and higher than on like-sized specimens of N. jamesi. The nymph of large specimens of N. nuttallii is low and elongate. In contrast to the relatively narrow pallial sinus of N. jamesi, the sinus of N. nuttallii is broad and trapezoidal and, although variable, is generally deeper, extending about half the distance anterior to the midline. The material at hand indicates that N. jamesi is a smaller species than N. nuttallii, which commonly reaches 90 mm and may reach 150 mm in length. The nominal form Sanguinolaria orcutti Dall, 1921, described from the Pleistocene of San Quintin Bay, Baja California, and considered by Grant and Gale (1931) and Coan (1973) to be synonymous with N. nuttallii, is also larger than N. jamesi, with a broad, obtuse posterior end and trapezoidal pallial sinus. Nuttallia toulai (Hertlein and Jordan, 1927) from the Miocene of Baja Cali- fornia, is another large, suborbicular form, with- out the acuminate posterior end of Nuttallia jJamesi. Nuttallia olivacea (Jay, 1857) (Fig. 8), a common Recent species in Japan, also recorded as early as the Miocene, is in some respects the spe- cies most similar to N. jamesi. Both species are of similar size, with pallial sinuses of the same gen- eral shape. Nuttallia however. has a bluntly angular posterior end, hinge line strai in dorsal view, a solid hinge plate, and both dorsal margins convex, while N. jamesi has a sharp poste- olivacea. rior angle, hinge line sinuous in dorsal view. narrow hinge plate, and a concave posterior dorsal margin. 144 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Other Asian Pliocene and Quaternary species Nuttallia Kuroda and 1955, is morphologically very are less similar. ezonis Habe, close to N. Habe in under which name it was reported by The “Sanguinolaria (Nuttallia) nuttalli’” from upper Pliocene strata on the western .coast of Kam- chatka (Slodkewitsch, 1938) is probably referable to N. ezonis. Nuttallia commoda (Yokoyama, 1925), described from the Pliocene of Japan, is a large, subquadrate, equivalve species with con- centric ridges crossing the shell. The Recent northwestern Pacific Nuttallia petri (Bartsch, 1929) is regarded by Habe and Ito (1965) as a synonym of N. Nuttallia (Slodkewitsch, 1936), an upper Pliocene species from the western coast of Kamchatka, also has low concentric ridges and, if not identical with N. commoda, is probably closely related to it. Age: Travis (1952) considered the Merced Formation in the Sebastopol Quadrangle to be “probably upper Pliocene in age,” based on both faunal criteria and local superposition with the middle to upper Pliocene Petaluma Formation. This age assignment of the Petaluma formation is based on remains of the horse, Neohipparion gidleyi Merriam, 1915, a species of Hemphillian age (Stirton, 1952). Berggren’s (1971: table 52:40) Cenozoic radiometric time-scale places the Hemphillian stage in the interval 3.5-10 mil- lion years before present, late Miocene to mid- Pliocene in terms of the European series-epoch classification. Fossils from Wilson Grove, 2 miles north of Trenton, were regarded by Dickerson (1922) as equivalent to the fauna of the type Merced For- mation on the northern San Francisco Peninsula. Recent field work has shown several distinct faunal assemblages, indicating different environ- mental and depositional conditions, within the Sonoma County Merced Formation. The strati- graphic relationships of these and other published localities to the type locality of Nuttallia jamesi are obscured by lack of exposures and local fault- ing. A radiometric age of six million years has been obtained from tuff interbedded in Merced Formation sandstones in the Sebastopol quadrangle (G. H. Curtis, personal communication). We have not determined the stratigraphic relationship of this tuff to the type locality of N. jamesi. Approximately one-third of the molluscan taxa at CAS locality 54164 are extinct. Protothaca staleyi (Gabb, 1866), Nucella trancosana (Arnold, 1908), (Dall, nuttallit, earlier workers. record of commoda. ochotica Nassarius (Caesia) grammatus VOLUME 73 1917), Nassarius (Demondia) californianus (Con- rad, 1856), Ophiodermella graciosana (Arnold, 1907), and Megasurcula remondii (Gabb, 1866) are among the extinct species widespread in strata traditionally classified as Pliocene in the west American provincial sequence. The presence of Nassarius (Caesia) grammatus and the absence of N. (C.) (Martin, 1914) would seem to exclude an uppermost Pliocene or early Pleistocene age for this fauna (cf. Addicott, 1965: B6-B8, fig. 1). This line of evidence places CAS locality 54164 stratigraphically below all or part of the Merced section at Bolinas, Marin County, California, the type locality of Nassarius morani- anus. The presence in the fauna of an undescribed echinoid of the genus Scutellaster may afford more precise placement within the west coast faunal sequence (J. W. Durham, pers. comm.). We conclude that the assemblage is Pliocene in the traditional terminology of west coast paleontolo- gists, but possibly should be called late Miocene in reference to the European series-epoch classi- fication. moranianus Paleoecology: The molluscan assemblage at the Trenton locality suggests deposition on a silty sand bottom in 15 to 40 meters depth, with several elements transported downslope from rocky and sandy intertidal and shallow subtidal areas probably less than a mile distant. As will be discussed in a forthcoming paper, several features of the fauna support analogy with present-day conditions at the southern end of Monterey Bay, California. Paired, intact valves (seven pairs in the type lot) of the rather thin-shelled Nuttallia jamesi might be interpreted as suggesting burial in place, rather than transport into the assemblage. Much of the fossil material from locality 54164, however, consists of indeterminable pelecypod fragments; thus the type lot represents an un- known fraction of the total Nuttallia component of the faunule. Recent Nuttallia nuttallii usually lives inter- tidally in protected bays, particularly in entrance channels washed by fast-moving tides, buried in (Fitch, It is less common on the open coast, The greatest depth from which the species is reported is 10 meters (Coan, 1973). The Japanese species, Nuttallia olivacea, has been reported from sandy and muddy bottoms, intertidally to 10 m (Kuroda, Habe, and Oyama, 1971). Nuttallia ezonis was cited by Habe (1964) as occurring on fine sand loose sand or mixed sand and gravel 1953%)2 where it occurs in sand below wave-base. 1974 bottoms from low tide level to “several” meters depth. ACKNOWLEDGMENTS The authors are indebted to J. Wyatt Durham, Peter U. Rodda, and Eugene V. Coan for advice on various aspects of this study. Warren O. Addicott loaned specimens for comparison. A. Myra Keen advised us concerning a nomenclatural detail. G. H. Curtis kindly allowed citation of a radiometric date. The professional assistance of Clara Gross, Assistant Li- brarian, California Academy of Sciences, is especially appreciated, LITERATURE CITED Addicott, W. O. 1965. Some western American Cenozoic gastropods of the genus Nassarius. U.S. Geol. Sury. Prof. Pap., 503-B:BI—B24. 1966. Late Pleistocene marine paleo- ecology and zoogeography in central California. U.S. Geol. Surv. Prof. Pap., 523-C:C1l-C21. Berggren, W. A. 1971. Tertiary boundaries and correlations. Pp. 693-809 in Micropalaeontology of the Oceans. (B. M. Funnell and W. R. Riedel. eds.), Cambridge Univ. Press. Clark, B. L. 1925. Pelecypoda from the marine Oligocene of western North America. Univ. California Publ. Bull. Dept. Geol. Sci. 15: 69-136. Coan, E. V. Psammobiidae. 1973. The northwest American The Veliger, 16:40—57. Dall, W. H. 1898. Synopsis of the Recent and Tertiary Psammobiidae of North America. Proc. Acad. Nat. Sci. Philadelphia, 50:57—62. 1900. Contributions to the Tertiary fauna of Florida with especial reference to the Silex beds of Tampa and the Pliocene beds of the Caloosahatchie River including in many cases a complete revision of the generic groups treated and their American Tertiary species [Teleodes- macea: Solen to Diplodonta|. Trans. Wagner Free Inst. Sci., 3:949-1218. Dickerson, R. E. 1922. Tertiary and Quaternary history of the Petaluma, Point Reyes and Santa Rosa quadrangles. Proc. California Acad. Sci. ser. 4, 7:527-601. NEW PLIOCENE BIVALVE FROM CALIFORNIA Durham, J. W. 1950, Cenozoic 1 of the Pacific Coast gull, Geol 61:1243~1264, Fitch; J. E. 1953 Common marine bj California Fish Gar | 102. California. Bull, 90:1 Dept. Grant, U. S., IV, and H. R. Gale. 1931 of the marine Pliocene and Pleistocene Moll Catalogue of California and adjacent regions. Mem. San Diego Soc. Nat. Hist., 1:1—1084. Habe, T. 1955. Fauna of Akkeshi Bay, XXI. Pelecypoda and Scaphopoda. Publ. Akkeshi Mar. Biol. Sta., 4:1—34. 1964. Shells of the western Pacific in color, vol. Il. Hoikusha Publ. Co., 233 pp. Habe, T., and K. Ito. 1965. colour. Vol. I. The northern Pacific. Publ. Co., 176 pp. Shells of the world in Hoikusha Kamada, Y. 1962. Tertiary marine Mollusca from the Joban coal-field, Japan. Paleontol. Soc. Japan, Spec. Paper, 8:1—187. Keen, A. M. 1969. Superfamily Tellinacea de Blainville, 1814. Pp. N613—N642 in Treatise on invertebrate paleontology, part N, [eae (R. C. Moore, ed.), Geol. Soc. America, 952 pp. vols. 1971. Sea shells of tropical west America: marine mollusks from Baja California to Peru. ed. 2. Stanford Univ. Press, xiv + 1064 pp. Kira, T. 1953. On the Japanese species of the genus Nuttallia (Pelecypoda). Venus, 17 144-151. Kuroda, T., and T. Habe. 1952. Check list and bibliography of the Recent marine Mollusca of Japan. Leo W. Stach, publ., 210 pp. Kuroda, T., T. Habe, and K. Oyama. The sea shells of Sagami Bay collected by His Majesty the Emperor of Japan. Maruzen Co., 741 pp. {in Japanese] + 489 pp. [in English]. 1971. Nomland, J. O. 1917. The Etchegoin Pliocene o middle California. Univ. California Publ. Bull. Dept. Geol., 10:191-254. Manning, G. A., and B. A. Ogle. 1950. Geology of the Blue Lake quadrangle, California l fornia Div. Mines Bull. 148:1—36. 146 Slodkewitsch, W. S. 1938. Tertiary Pelecypoda from the Far East, Pt. 2. U.S.S.R. Acad. Sci. Paleontol. Inst., Paleontol. of U.S.S.R., 10 (3): 1-275. Stirton, R. A. 1952. Are Petaluma horse teeth reliable in correlation? Bull. Amer. Assoc. Petrol. Geol., 36:2011—2025. Stoliczka, F. 1870-1871. Cretaceous fauna of Vol. III. The Pelecypoda, with a review of all known genera of this class, fossil and Recent. Mem. Geol. Soc. India, Palaeontol. Indica, ser. 6, 3:1-537 [pp. 1-222 issued- 1870; 223-537, 1871]. 2 southern India. Travis, R. B. 1952. Geology of the Sebastopol BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 quadrangle, California. California Div. Mines Bull. 162:1-33. Valentine, J. W. 1966. Numerical analysis of marine molluscan ranges on the extratropical northeastern Pacific shelf. Limno. Oceanog., 11:198-211. Weaver, C. E. 1949. Geology of the coast ranges immediately north of the San Francisco Bay region, California. Mem. Geol. Soc. America, 35:1-242. Winckworth, R. 1935. Notes on nomenclature. Proc. Malacol. Soc. London, 21:321-324. Accepted for publication January 29, 1974. TWO NEW SPECIES OF LAND SNAILS FROM THE PINALENO MOUNTAINS, ARIZONA W. O. Grecc! AND W. B. MILLER? ABSTRACT: in southeastern Arizona. The Pinaleno Mountains, sometimes called the Graham Mountains, lie in line with the Chiricahua system in southeastern Arizona and are separated from the northern end of the Dos Cabezas Moun- tains by a wide mesa. The Pinalenos have long been neglected by malacologists. A brief account is given (Pilsbry and Ferriss, Proc. Acad. Nat. Sci. Philadelphia, for 1918: 282-334, 1919) of a collecting trip made by Ferriss, 14 October 1913, “Besides Sonorella and Oreohelix, Vitrina alaskana was abundant and two young Vallonias were found.” Pilsbry and Ferriss described Sonorella grahamensis from this material but there was no further mention of the Oreohelix. On 21 April 1954, M. L. Walton and one of Two new species of land snails are described from the Pinaleno Mountains Leaving Safford on US highway 666, we traveled south seven miles, then turned west on the “Swift Trail,” now a well-maintained mountain road. We continued about 20 miles which took us to near 9000 feet altitude. Stopping now and then to look for snails, we found Sonorella and Oreo- helix and in the same microhabitat, Microphysula ingersolli (Pilsbry and Ferriss), Retinella indentata paucilirata (Morelet), Zoni- toides (Say), (Pilsbry and Ferriss), meridionalis meridionalis Discus Striatura Deroceras sp., arboreus 1 Dept. Invertebrate Zoology, Natural History Mu- seum of Los Angeles County, California 90007. 2 Dept. Biological Sciences, University of Arizona, us (Gregg) visited the Pinaleno Mountains. Tucson, Arizona 85721. Figure 1. Sonorella imitator, new species. Genitalia drawing made from projection of stained whole mount: ag, albumin gland; ec, epiphallic caecum; ep, epiphallus; go, genital orifice; hd, hermaphroditic duct; ot, ovotestes; ov, oviduct; pe, penis; pr, penial retractor; ps, penial sheath; pt, prostate; sd, spermathecal duct; sp, spermatheca; ta, talon; ut, uterus; va, vagina; vd, vas deferens; ve, verge. TWO NEW LAND SNAILS FROM ARIZONA 5.0mm 148 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 cronkhitei. (Newcomb), Punctum californicum Pilsbry, and Vertigo gouldi arizonensis Pilsbry. These snails were found under isolated rocks; no definite rockslides were present. The Oreohelix from this locality immediately appeared to be different from any described species. The Sono- rella specimens appeared to be S. grahamensis Pilsbry and Ferriss but subsequent dissection re- vealed two distinct species, S$. grahamensis and an undescribed species, both sympatric in the same microhabitat. The new Oreohelix and the new Sonorella are described below. All measure- ments are given in millimeters. FAMILY HELMINTHOGLYPTIDAE Sonorella imitator, new species Figures 1 and 2 This name was cited as a nomen nudum by Bequaert and Miller, The Mollusks of the Arid Southwest, p. 71973" Diagnosis——A_ species of moderate size, finely striate body whorl, characterized anatomically by a very long verge and a short oviduct. Description of Holotype—Shell heliciform, de- pressed conic, whorls four and one half, increasing gradually to the last whorl which is moderately ex- panded and descends behind the aperture so that the last one eighth of the body whorl lies at an angle of about 25 degrees with the transverse axis of the shell. Whorls convex, flattened above, rounded on the base. Aperture rounded-oval and lying at an angle of about 45 degrees with the vertical axis. Peristome very slightly expanded, not appreciably thickened, and with a thin parietal callus; the columellar end of the peristome slightly reflected over the margin of the umbilicus. Width of the umbilicus contained about seven times in the greater diameter of the shell. Sur- face glossy, color light sayal brown, lighter on the base and on each side of the peripheral chestnut brown band. Embryonic shell consists of one and one half whorl, apex smooth, followed by indistinct radial wrinkling which continues over the remainder of the embryonic portion of the shell and as far as the first half of the penultimate whorl where it is replaced by delicate growth lines. Early whorls have a minutely granulose appearance with here and there suggestions of papillae which have been nearly effaced by erosion. There are numerous impressed spiral lines on the upper surface of the body whorl and on the last half of the base of the body whorl. Figure 2. Sonorella imitator, new species, three views. Figure 3. Oreohelix grahamensis, new species. Genitalia drawing made from projection of stained whole mount. See figure | for key to abbreviations. 1974 TWO NEW LAND SNAILS FROM ARIZONA 150 Height of holotype 12.0, diameter 21.8, umbilicus 3.0. Average of 17 paratypes: Height 12.2, diameter 21.9, umbilicus 3.0. Genitalia—The penis (Fig. 1) is about equal in length to the diameter of the shell and contains a large verge nearly as long as the penis. The verge gradually widens to a blunt distal end. Penis sheath short, penial retractor muscle inserted at the junction of the penis and epiphallus, epiphallus about half the length of the penis and bears at its proximal end a very short epiphallic caecum which is adnate to the adjacent portion of the vas deferens, oviduct very short. Measurements of genital structures: penis 20.8, penis sheath 5.4, verge 18.1, epiphallus 9.3, epiphallic caecum 1.3, penial retractor 7.4, vagina 12.5, oviduct 2.8, spermathecal duct 15.5, spermatheca 2.9 < 2.9, atrium 1.4. Type Locality—South slope of Mt. Graham, Pinaleno Mountains, Graham Co., Arizona; along “Swift Trail” highway, on north side, at a point 20.7 road miles from its beginning at US highway 666; elevation ca. 9000 feet, collectors M. L. Walton and W. O. Gregg, 21 April 1954. Type Material—Holotype, Natural History Mu- seum of Los Angeles County No. 1154. Paratypes: California Acad. Sci., Dept. Geol. No. 12948, San Diego Nat. Hist. Museum, invertebrate type coll. 2545, and private collections of S. S. Berry 31923, W. O. Gregg 8755, W. B. Miller 5357, and M. L. Walton 9279. Remarks.—On four paratypes which apparently represent senile specimens the peristome is slightly thickened and the parietal callus is moderately thickened. These snails were associ- ated with Sonorella grahamensis Pilsbry, from which they are indistinguishable on casual exam- ination. A more careful examination reveals that they fall into two separate size groups, S. gra- hamensis with an average diameter of 19.2 (14 specimens measured), S. imitator 21.9 (average 17 paratypes). The latter has a more expanded body whorl which is more flattened above. The penial length of Sonorella imitator is approxi- mately equal to the width of the shell; it has a long, robust, blunt-tipped verge occupying nearly the entire length of the penis cavity, whereas S. grahamensis has a penis about half as long as the diameter of the shell with a shorter verge which tapers to an acute tip. The oviduct is short in S. imitator whereas that of S. grahamensis exceeds the length of the verge. The genital anatomy of S. imitator closely resembles that of S. ambigua Pilsbry and Ferriss and S. ambigua verdensis Pilsbry, but the shell sculptures are considerably different. Accordingly, the authors consider the similarity of genital anatomy to be simply a case of convergent evolution. BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Sonorella imitator has also been found on Heliograph Peak in the Pinaleno Mountains (Bequaert and Miller, The Mollusks of the Arid Southwest, p. 117, 1973) where it is relatively common, in association with S. grahamensis, at the 10,000 foot level. At lower elevations, how- ever, such as along Wet Creek at the 6300 foot level where S. grahamensis is abundant, §. imitator is absent. It is apparently restricted to the high elevations of the Pinalenos, ca 9000 feet and above. FAMILY OREOHELICIDAE Oreohelix grahamensis, new species Figures 3 and 4 This name was cited as a nomen nudum for a sub- species of Oreohelix concentrata (Dall) by Bequaert and Miller, The Mollusks of the Arid Southwest, p. LEM OTSS Diagnosis ——A_ moderate-sized species of the O. yavapai-group, with closely set spirals on the body whorl, rather prominent lines of growth, and a prominent cornuate process at the proximal end of the penis. Description of Holotype——Shell of moderate size, low-conic, widely umbilicate. Whorls five and one half, gradually increasing in size, convex above and below the periphery which is rounded on the last three fourths of the body whorl, a slight keel per- sisting in the first quarter of the body whorl. The last quarter of the body whorl descends slightly below the strongly keeled periphery of the penultimate whorl. Peristome subcircular, simple, with a thin parietal callus. Aperture lying at an angle of about 45 degrees with the vertical axis of the shell. All whorls visible in the umbilicus which is contained about four times in the greater diameter of the shell. Embryonic shell consists of two and one eighth whorls and is about 4 mm in diameter. First whorl smooth (eroded); traces of growth lines appear on the second whorl at somewhat irregular intervals and con- tinue over the remainder of the shell. Fine spirals appear on the third whorl and continue nearly to the aperture. These spirals are closely set and regular on portions of the shell whereas at intervals they are interrupted by lines of growth; are less closely spaced over the rounded base of the body whorl and are clearly visible in the umbilicus. The coiling is somewhat irregular and in places the suture descends beneath the acute keel of the earlier whorl. On portions of the shell there are traces of a peripheral cord. Color light snuff brown above, much lighter on the base, with a chestnut band about 1 mm wide above the periphery and another band of the same color 1974 Figure 4. Oreohelix grahamensis, new species, three views. and somewhat wider just below the periphery. Height 10.8, diameter 20.0, umbilicus 5.0. On uneroded specimens the spirals appear on the last half of the second whorl and the radial lines appear on the first whorl. On some specimens the peripheral cord is continuous though variable. TWO NEW LAND SNAILS FROM ARIZONA Genitalia,—Internally ribbed port (Fig. 3) is more than half the total bulbously expanded; epiphallus spindle-sh half the length of the penis. In stained ar specimens, the distal end of the epiphallu protrude the giving opposite side of this verge-like structure, the proxin end of the penis bears a into the upper end of pen the appearance of a short verge. On 1} prominent cornuate pre jection. These characters of penis and epiphallu constant in all five anatomies examined. Albumin gland rather small; “talon” well-developed, black Measurements of genital structures: penis 21.4, in ternally ribbed portion 12.0, epiphallus 9.1, penial retractor 8.8, vagina 6.6, oviduct 5.6, spermathecal duct 21.5, spermatheca 1.1 135% Type Locality—South slope of Mt. Graham, Pinaleno Mountains, Graham Co., Arizona: along “Swift Trail” highway, on north side, at a point 20.7 road miles from its beginning at US highway 666; elevation ca. 9000 feet, collectors M. L. W. O. Gregg, 21 April 1954. Type Material_—Holotype, Natural History Mu- seum of Los Angeles County No. 1155. Paratypes: California Acad. Sci., Dept. Geol. No. 12947, Field Museum Nat. Hist. 109587, San Diego Nat. Hist. Museum, invertebrate type coll. 2546, and private collections of S. S. Berry 31922, W. O. Gregg 7038, W. B. Miller 4831, M. L. Walton 6844. Walton and Remarks.—The characteristics of the penis place O. grahamensis in the Oreohelix yavapai- group of Oreohelix sensu stricto (Pilsbry, Acad. Nat. Sci. Philadelphia, Monograph 3, vol. 1(1): 573, 1939). Oreohelix closely related to the O. concentrata-complex. From typical O. concentrata (Dall), O. graham- ensis is distinguished by its wider umbilicus, more elevated spire, spiral striation and uniformity of coloration. Bequaert and Miller (The Mollusks of the Arid Southwest, p. 127, 1973) considered that the pop- ulations of Oreohelix from the Pinaleno Moun- tains were only subspecifically distinct from Oreohelix concentrata populations in the Hua- chuca Mountains. Careful comparison of anatom- ical and shell features as well as consideration of prolonged and extensive geographical isolation have convinced the authors that the heterogeneity which has developed between the populations of these two mountain ranges is most probably sufficient to cause reproductive isola- tion. Accordingly, Oreohelix grahamensis is con- sidered to be a distinct species. grahamensis is most genetic Accepted for publication January 29. 1974. OBSERVATIONS ON THE LITHODID CRABS OF PERU, WITH DESCRIPTION OF TWO NEW SPECIES JANET Haic ! Apstract: Eight species of Lithodidae (Crustacea, Decapoda, Anomura) are reported from deep water off Peru. Two are described as new and a third is illustrated for the first time. The other five species, which were briefly cited from Peruvian waters (del Solar, 1972), are discussed in more detail. Attention is drawn to morphological changes that take place with growth in family Lithodidae. lithodids is discussed. The Lithodidae are a family of anomuran crabs, some of them of large size, which inhabit the littoral zone and depths to at least 2400 m. The family comprises more than 50 species, including such economically important forms as the Alaska king crab, Paralithodes camtschaticus (Tilesius), in the north Pacific, and the centolla, Lithodes antarcticus Jacquinot, off southern Chile and Argentina. Until recently no lithodids were known from Peruvian waters. In recent years the Instituto del Mar del Pert (IMARPE) has been conducting investigations in deep water, particularly on crustaceans of poten- tial economic importance. Since late 1970 E. M. del Solar, scientific advisor for IMARPE, has collected material of eight species of Lithodidae aboard “SNP-1,” “Wiracocha,” and the Japanese trawler “Challua japic.”” Samples of these, which were sent to me for identification by del Solar, form the basis of this report. Two of the species were previously undescribed. A third was described by Benedict (1895) from a specimen collected at an unspecified locality by the U.S. Fisheries vessel “Albatross”; it is illus- trated for the first time in this paper. The other five species were described by Faxon (1893) from material collected by the “Albatross” in the eastern Pacific between 07°31/30” N and 05°26’ 20” N; they were listed, with the first Peruvian records, by del Solar (1972). The presence of Lithodidae in Perit had been recognized earlier on the basis of unidentified fragments taken from the stomach of “cachalotes”’ (sperm whales). A ninth Peruvian species is represented by legs and part of the carapace of one such specimen, which is housed in a whaling museum at Tierra Colorada to the south of Paita; del Solar believes it to be a Lithodes (del Solar, 1972: 6, and pers. comm.). I have seen a photo- graph of this crab and agree that it bears a general Distribution of the tropical component of the west American resemblance to Lithodes. However, examination of the rostrum (if present) and the abdomen might prove it to belong to Neolithodes. Neo- lithodes diomedeae (Benedict) is reported from Chile and from México and is therefore to be expected in Peruvian waters. Measurements given in this report, except where there is a statement to the contrary, refer to the length of the carapace. All measurements were taken to 0.5 mm using a dial caliper. Specimens, including holotypes of the two new species, are in the collections of the Allan Hancock Foundation (A. H. F.). This paper is contribution No. 351 from the Allan Hancock Foundation. Lithodes Latreille, 1806 Lithodes Latreille, 1806: 39. Type species by mono- typy: Cancer maja Linnaeus. Bouvier, 1896: 10, 20. Carapace spiny; rostrum generally prominent and armed with spines. Antennal acicle (scaphocerite ) either well developed, rudimentary, or lacking. Walk- ing legs considerably longer than greatest width of carapace. Second (basal) abdominal somite with median plate fused to lateral plates, and usually with laterals fused to marginal plates; on abdominal somites 3—5, median plate replaced by a membrana- ceous area covered with calcified platelets; lateral plates of somites 3—5 distinct and paired; marginal plates of somites 3—5 distinct and variable in number, present on both sides in males, on right side only in females. Lithodes panamensis Faxon, 1893 Lithodes panamensis Faxon, 1893: 166; Faxon, 1895: 50, pl. 10 figs. 1, 1 a-c; Bouvier, 1896: 24; del Solar, 1972: 5, 14. 1 Allan Hancock Foundation, Univ. Southern California, Los Angeles, California 90007. 152 1974 Diagnosis: Carapace covered with low, warty pro tuberances; two pairs of spines on. gastric region, two spines on each branchial region, a pair on care diac and a pair on intestinal region; lateral spines consisting of outer orbital, anterolateral, hepatic, and three branchials, Gastric and branchial regions very convex, defined by deep depressions. Rostrum strongly inclined upward, terminating in a median and two lateral spines; ventral process long but not visible in dorsal view. Antennal acicle long, rudi- mentary, or absent. Abdominal plates coyered with low tubercles; basal (second) somite with a distinct suture on each side separating marginal plate from fused median and lateral plates. Previous records: 07°31'30" N, 79°14’ W, 458 fms [837 m] (Faxon, 1893, 1895). 03°48’ S, 81°22’ W, 680 m; 07°59’ S, 80°22’ W, 760-800 m; latitude of Pisco [13°44’ SI, from stomach of a sperm whale; 17°34’ S, 71°55’ W, 850 m (del Solar, 1972). Material examined: Ovigerous female; off Peru, 07°59’ S, 80°22’ W; 760-800 m, hard bottom; November 1971; E. M. del Solar on trawler “Challua japic.” Measurements: Length of carapace excluding ros- trum, 100 mm; maximum width of carapace excluding lateral spines, 108 mm; length of right, third walking leg: merus, 78 mm; carpus, 44 mm; propodus, 70.5 mm; dactyl, 45.5 mm. Remarks: My specimen agrees well with the description and illustration of the holotype, a female 79 mm long excluding the rostrum. A few minor differences were noted. The three terminal rostral spines are more elongate than the rostral spines of the type; the proximal portion of the rostrum is also longer and advanced beyond the eyestalks. A pair of small spines on the cardiac region of the carapace was not mentioned by Faxon, although his illustration, perhaps errone- ously, shows a median spine in that area. The lateral hepatic spines are much larger than they appear in the illustration of the holotype. On each antennal peduncle there is a rudimentary, conical acicle, instead of the long, slender one occurring on the right peduncle in the type. A specimen collected by del Solar at 17°34’ S, has a carapace 190 mm long and measures 970 mm between the tips of the extended legs (del Solar, 1972: 14). Lithodes wiracocha, new species Figure 1 Lithodes n. sp.: del Solar, 1972: 14. Description: Carapace a little longer than broad: surface and all margins densely covered with small, sharp spinules, some of the marginal ones tending to NEW LITHODID CRABS FROM PERU become enlarged. Gastric region bears short, stout spines, and separated fr region by a decp, transverse depression, ¢ region well defined, with a pair of small Branchial regions separated from cardiac region deep, oblique grooves; cach branchial region with well developed spine opposite sulcus separating tric and cardiac regions. Intestinal region of small spines. Outer long, slender spine; a shorter anterolateral angle; a long, strong marginal hepatic spine; three lateral branchial spines, the most anterior one largest others not much larger than a few enlarged, laterally situated surface spinules. with a pair orbital angle marked by a one at Dorsal portion of rostrum inclined upward, with two terminal spines and a pair of median spines; proximal half densely covered with minute spinules to base of median spines; ventral process rather short and not visible in dorsal view, covered proximally with minute spinules. Eyestalks short, somewhat constricted at middle, and unarmed, with cornea situated laterally and ventrally. Second segment of antennal peduncle with several spines on its outer side, the most distal one large and elongate and reaching end of penulti- mate (fourth) segment; acicle rudimentary, conical. Chelipeds subequal in length, but right somewhat stouter than left; small, thornlike spines, disposed in lateral about longitudinal rows, on merus, carpus, and chela; spines somewhat enlarged on ventral surface of ischium. Walking legs long and slender: ischium, merus, carpus, and propodus covered on all sides by longitudinal rows of thorny spines, these a_ little larger dorsally and ventrally; dactyl with several rows of spines reaching almost to tip. Abdominal plates, and platelets of somites 3—5, densely covered with sharp spinules. Basal abdominal somite with a median pair of small spines; a narrow, spinule-free line on each side in position of suture which separates marginal from fused median and lateral plates in a few species of Lithodes, but in this case no actual suture visible. Lateral plates of left side edged with numerous long, sharp spines. many themselves bearing two or three spinules along each side; marginal plates of right side drawn out into still longer compound spines. Previous records: 03°48’ S, 81°22’ W, 680 m: 07° 59’ S, 80°22’ W, 800 m (del Solar, 1972). Material examined: Holotype, ovigerous female. A.H.F. 712; 12 mi. SW Banco de Mancora, Peru: 620 m, mud bottom; 15 March 1971: E. M. del Solar on trawler “Wiracocha.” Measurements: Length of carapace excluding ros- trum, 103.5 mm; rostrum, 16 mm; maximum width of carapace excluding lateral spines. 97 mm: 2 f right, third walking leg: merus. 78 mm: carpus. 48.5 mm; propodus. 79 mm; dactyl, 47 mm. Etymology: From Tici Wiracocha, the creator god of Inca mythology. and his namesake. the trawler “Wiracocha” aboard which the holotype ] Was cOol- 154 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 75 a ‘ Figure 1. Lithodes wiracocha, new species, holotype. A, animal in dorsal view; B, anterior part of carapace, dorsal view; C, same, lateral view; D, right antennal peduncle. Scale A = 30 mm; B= 15 mm; C= 10 mm; D = 5 mm. 1974 lected, ‘To be treated as a noun in apposition to the generic name. Remarks: This species is distinguished from all other described Lithodes by the dense spinulation of the carapace and walking legs. As in L. murrayi Henderson (a southern hemisphere spe- cies) and L. tropicalis A. Milne-Edwards, the } rostrum is inclined upward and bifurcate at the tip. Lithodes panamensis Faxon also has an up- | turned rostrum, which however is trifurcate at | the tip; Faxon’s species is further distinguished from L. wiracocha by the pronounced convexity of the gastric and branchial regions of the cara- pace. The dorsal portion of the rostrum of L. antareticus Jacquinot, which inhabits the southern part of the South American continent, is shorter than the ventral process and the latter is clearly visible in dorsal view. Several stalked barnacles are attached to the chelipeds and walking legs of the holotype; these have not yet been identified. Del Solar (1972: 15) reported a specimen with a span of 1.07 m between the tips of the extended legs. An equivalent measurement on the holotype is only 0.58 m. Paralomis White, 1856 Paralomis White, 1856: 134. Type species by mono- typy: Lithodes granulosa Jacquinot. Bouvier, 1895: 185; Bouvier, 1896: 12, 21. Leptolithodes Benedict, 1895: 484. Type species not designated. Pristopus Benedict, 1895: 486. Type species not designated. Carapace with spines, tubercles, or other orna- mentation according to the species; rostrum short and generally armed with spines. Antennal acicle (scapho- cerite) well developed, usually triangular and armed with several spines on margins. Second (basal) abdominal somite fused into a single plate; median plate of somites 3—5 entire; lateral plates of somites 3-5 distinct and paired; in males, marginal plates usu- ally fused to laterals, at least in part, on somite 3 and distinct on somites 4 and 5; in females, marginal plates of somites 3—5 present on right side only. Paralomis longipes Faxon, 1893 Paralomis longipes Faxon, 1893: 165; Faxon, 1895: pl. 9; Bouvier, 1896: 25; del Solar, 1972: 5, 14. Leptolithodes longipes: Faxon, 1895: 48. Diagnosis: Carapace and abdomen thickly covered with small, blunt tubercles, each encircled by a ring of short, stiff setae; a median spiniform tubercle on anterior part of gastric region; anterolateral and outer orbital spines sometimes enlarged, other laterals (hepatic and three or four branchials) short. Gastric, NEW LITHODID CRABS FROM PERU cardiac, and branchial regions well d tuberant; grooves defining sides of car becoming indistinct posteriorly and not to posterior margin of carapace. Rostrum term in a median inferior and two lateral dorsally a superor pir median spinule at base of Jatter 4 and one or two lateral spinules on cach side; vents with a Antennal spines. Walking legs very long, prismatic, tudinal rows of strong, thornlike spines. Previous records: 05°26'20" N, 86°55’ W, 770 fm [1410 m] (Faxon, 1893, 1895). 07°59’ S, 80°22’ W, 760-800 m; 16°29’ S, 73°33’ W, 1300 m (del Solar, 1972). Material examined: Male; off Pera, 07°59’ S, 80°22’ W; 760-800 m, hard bottom; November 1971; E. M. del Solar on trawler “Challua japic.” Measurements; Length of carapace excluding ros- trum, 106 mm; rostrum, 14 mm; maximum width of carapace, 117 mm; length of right, third walking leg: merus, 101 mm; carpus, 56 mm; propodus, 99 mm; dactyl, 76.5 mm. surface protuberance bearing one or tw spinules, acicle with several long, slender with longi Remarks: Faxon’s description and _ illustration were based on a male 84 mm long including the rostrum. My specimen differs in only a few details. The outer orbital and anterolateral spines are no larger than the other lateral spines, whereas they are enlarged in the illustrated type. In the Peruvian specimen there are two distinct spinules on each side of the rostrum proximal to the termi- nal pair of spines. The acicle of the left antenna has six spines and that of the right antenna eight; Faxon described and figured only five spines on an acicle of the male type. Numerous stalked barnacles are attached to the chelipeds and walking legs. Paralomis aspera Faxon, 1893 Paralomis aspera Faxon, 1893: 164; Faxon, 1 pl. 8; Bouvier, 1896: 26; del Solar, 1972: 5, 14. Leptolithodes asper: Faxon, 1895: 47. Diagnosis: Carapace and abdomen thickly covered with papillae or tubercles, each encircled by a ring of stiff setae; no surface spines: a sharp outer orbital and anterolateral spine, and four or five lateral branchial spines. Gastric, cardiac. and branchial regions well defined and prominent. Rostrum short. indistinctly tripartite, multispinose, lower part armed with as many as five spines. Antennal acicle with several spines on each margin and one on both uppe and lower side near base. Walking legs of moderate length, stout, densely spinose. Previous records: 07°06’15” N. 80°34’ W. 695 fms [1270 m] (Faxon, 1893, 1895). 03°48’ S, 81°20’ W. 560 m (del Solar, 1972). Material examined: Male (juvenile); off Peru: 1971; E. M. del Solar. 156 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Figure 2. Paralomis papillata (Benedict). A, animal in dorsal view; B, anterior part of cara- pace; C, rostrum, lateral view; D, right antennal peduncle; E, tubercles of carapace. Scale = 30mm; B= 15 mm; C=6mm;D=4mm; E=ca.7 mm. 1974 Measurements: Length of carapace including ros trum, 75 mm; maximum width of carapace, 75 mm; length of right, third walking leg: merus, 30 mm; carpus, 20 mm; propodus, 27 mm; dactyl, 24 mm. Remarks: The above diagnosis was derived from the description and illustration of the holo- type, a female 122 mm long including the ros- trum. The much smaller Peruvian specimen shows striking differences in the surface orna- mentation. Each papilla of the carapace and ab- domen is raised on a stalk, the whole forming a blunt-tipped (papilliferous) spine; at the base of the papilla is the ring of setae described and figured by Faxon. Toward the carapace margins, and on the chelipeds and walking legs, these spines become more elongate and in many, particularly } those of the walking legs, the terminal papilla is | tipped with a minute, sharp, corneous spinule. | Comparison of a good series of specimens of | different sizes would be of interest. Increase of size of individuals is probably accompanied by a gradual reduction of the surface armature from elongate, papilliferous spines bearing setae at the base of the papilla to low, papilliferous tubercles on the carapace and abdomen, these still retaining a ring of setae, and to spines of the usual kind on carapace margins and pereiopods. The rostrum in the Peruvian specimen is multi- spinose as in the type, but shorter and more sym- metrical. The eyes have two strong terminal spines, one dorsal and one mesial, extending well beyond the cornea, and there are about twelve smaller spines on the dorsal surface of the eye- stalk. All these spines have a ring of setae below the papillose tip. Faxon (1893: 165) mentioned that the type was “infested with a huge Peltogaster 36 mm _ in breadth.” My specimen also bears a large abdom- inal sacculinid, in this case 34 mm across. Paralomis papillata (Benedict, 1895) Figure 2 Leptolithodes papillatus Benedict, 1895: 485. Paralomis papillata: Bouvier, 1896: 25. Diagnosis: Carapace and abdomen thickly covered } with small, papilliform tubercles, each bearing stiff | setae over summit; no surface spines; outer orbital | spine well developed, lateral margins otherwise with small tubercles only. Branchial regions more pro- tuberant than gastric and much more so than cardiac; grooves defining sides of cardiac well defined, meet- ing at posterior end of this region and continuing as a single deep groove to posterior margin of carapace. Rostrum terminating in a median inferior and two lateral superior spines, latter pair not reaching end NEW LITHODID CRABS FROM PERU of eyes; ventral surface with an unarmed o 1 spinulate protuberance, Antennal acicle long, slender spines, Walking leg er prismatic, with longitudinal rows of tuber short, stout spines. Previous records: “Off Lower California, or haps south of that region” (Benedict, 1895) Material examined: Male; off Perd, 06°31’ S 81°O1’ W; 712-744 m, mud and hard sand; 17 May 1971; E. M. del Solar on R/V “SNP-1.” Measurements: Length of carapace excluding ros trum, 100 mm; rostrum, 12 mm; maximum width of carapace, 112 mm; length of right, third walking leg merus, 64 mm; carpus, 40.5 mm; propodus, 63 mm; dactyl, 5S mm. Del Solar’s collection includes closely allied but distinct species, both of which agree in most respects with the original descrip- tion of Paralomis papillata. Since no illustration of that species was ever published, it was necessary to compare material of both Peruvian forms with the holotype. Photographs of the type, provided by Henry B. Roberts, confirm the identity of the above specimen with Benedict’s species. The type (USNM 18536), which was collected by the “Albatross,” is the ecdysal cast of a large male. It is accompanied by the following informa- tion: “No label but with spns. from off lower Cal.” According to Mr. Roberts the length of the carapace with rostrum is 118 mm, length of the rostrum 10 mm, and breadth of the carapace 130 mm. My specimen agrees very closely with the type. The carapace is not quite so broad in pro- portion to its length. The ventral side of the rostral protuberance is minutely denticulate, while in the type this area is practically smooth. Both acicles are similar in structure to the left acicle of the holotype; but in the type specimen the right acicle has a strong terminal spine, and three lateral and two mesial spines which are con- siderably shorter. The Peruvian specimen has a few stalked bar- nacles attached to the walking legs: these are neither so large nor so numerous as those found on the A.H.F. specimens of Paralomis longipes and Lithodes wiracocha. Remarks: two Paralomis inca, new species Figures 3 and 4 Description (Adults): Carapace a little broader than long. covered with tubercles of different sizes each bearing a cluster of very short. stiff setae over summit. Gastric region moderately convex, some of 158 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Figure 3. Paralomis inca, new species, holotype. A, animal in dorsal view; B, anterior part of carapace; C, rostrum, lateral view; D, right antennal peduncle; E, tubercles of carapace. Scale A = 27 mm; B = 15 mm; Cand D = 4 mm; E=ca. 5 mm. 1974 its tubercles small and rounded, others larger, conical, with papilliform or pointed tip (latter type almost spiniform in largest individuals, definitely so in small- est adult). Cardiac region separated from gastric .by a deep, rectangular depression; with several conical tubercles and smaller, rounded ones. Branchial re- gions more protuberant than gastric and much more so than cardiac, separated from latter region by dis- tinct grooves joining at its posterior end and continu- ing as a single deep groove to posterior margin of carapace; branchials with both large and small tuber- cles as on other regions, and also with scattered swell- ings each bearing several small tubercles, a few of latter tending to become spiniform. A number of large, low, pustulous tubercles grouped on posterior part of carapace near midline. Outer orbital angle marked by a sharp spine; lateral margins otherwise with large and small tubercles, some of the larger ones spiniform. Rostrum terminating in a median inferior and two lateral superior spines, latter pair reaching or surpass- ing end of eyes; dorsally a median tubercle just proxi- mal to superior pair of spines, and a pointed tubercle on either side at base; underside of inferior spine with a large, distinctly spinulate protuberance. Eyestalks armed with a few spinules dorsally, larg- est One terminal and extending beyond cornea. Second segment of antennal peduncle with three or four strong spines laterally, distal one reaching onto terminal segment of peduncle. Acicle with two very strong spines, both appearing to be terminal and of about equal length; also a strong lateral spine and One or two lateral spinules, two strong mesial spines, and two or three very small dorsal spinules. Chelipeds subequal in length, but right slightly stouter than left. All segments with strong, thorn- like spines, each bearing short setae at tip. Walking legs long, prismatic, with longitudinal rows of strong, thornlike spines and short, pointed tubercles, these setiferous as elsewhere on body. Abdomen covered with small tubercles, conical or papilliform and setiferous at summit. (Juvenile): Carapace covered with long, slender spines, each with a tuft of long, flexible setae at and near tip; tip rounded and with a minute corneous spinule. A number of longer, stouter spines occurring as indicated on figure 4A; these sharp-tipped with rudi- mentary setae. Median area of posterior part of cara- pace with pustulous tubercles of different sizes; most of these raised on short, stout stalks and thus almost mushroomshaped. Spinulation of chelipeds and walk- ing legs as in adults, but spines proportionately longer and more slender; terminal setae in a long tuft. Ab- domen densely covered with long, slender spines, each tipped with a tuft of long, flexible setae. Material examined: Holotype, ovigerous female, A.H.F. 718; off Pert, 06°31.5’ S, 81°01.5’ W; 712- 744 m, mud and hard sand; 17 May 1971; E. M. del Solar on R/V “SNP-1.” Paratype, male (juvenile); 12 mi SW Banco de NEW LITHODID CRABS FROM PERU Figure 4. type. A, 30 mm: Paralomis inca, new species, juvenile para- carapace; B, carapace spines. Scale A = Bes ¢cas.5) mm: Mancora, Perti; 620 m, mud bottom; 15 March 1971: E. M. del Solar on trawler “Wiracocha.” Paratypes, male and ovigerous female; off Peru. 07°59’ S, 80°22’ W: 760-800 m, hard bottom; Novem- ber 1971; E. M. del Solar on trawler “Challua japic.” Measurements in mm: The measurements for the (holotype), a female, and two males, respectively. are as follows: length of carapace excluding rostrum, (108.0), 93.5, 80.0, 69.0; rostrum, (13.0), 10.0, 10.5. ; maximum width of carapace, (123.0), 109.0. 93.0, 69.0; length of right, third walking leg — merus. (53.0), 50.5, 47.0, 38.0: carpus, (37.0), 35.0. 31.0. 21.5; propodus, (50.0), 48.0, 44.0, 28.5: dactyl. (43.0), 41.5, 40.0, 26.5. Etymology: As a noun, “Inca” can refer either t re) e an emperor or chief of Perti in the days before t Spanish Conquest, or to a member of the dominan tribe: as an adjective, of or pertaining to the Incas or their empire. Here it is to be considered to be a noun in apposition to the generic name. Remarks: Paralomis inca and the closely allied P. papillata share a number of characters, includ- 160 BULLETIN SOUTHERN CALIFORNIA ing the swollen branchial regions defined by grooves which enclose the sunken cardiac region and extend as a single deep groove to the poste- rior margin of the carapace; the lack of well developed spines on the lateral margins, aside from the outer orbital spine; armature of the carapace consisting of papilliform tubercles with setae over the summit (not forming a ring below the summit as in P. longipes and P. aspera); and walking legs long and prismatic, with a row of spines on each longitudinal ridge. This combination of characters distinguishes both species from all other members of the genus. The tubercles of the carapace of P. inca are much less uniform in size and shape than they are in P. papillata; the lateral superior spines of the rostrum are longer, and the ventral protuberance of the median spine is more strongly spinulate; the walking legs are proportionately shorter, and their spines are longer, slenderer and more numerous. The 69 mm juvenile male is an ecdysal cast. Many of the spines of the carapace are bent near the tip or in the distal half, with the bent spines oriented in no particular direction; this condition may be due to the soft state of the integument. The dorsal portion of the rostrum is missing. The rather startling difference in appearance between juvenile and adults is duplicated in P. aspera Faxon, as I have noted earlier, and prob- ably in other species of Paralomis as well. Dis- tinctions between species of this genus are based in large part on the form of the surface orna- mentation, but little consideration has been given to changes in this ornamentation that may take place with growth. Del Solar (1972: 14) listed “Paralomis sp.” from 06°31’ S, 81°01’ W in 712-744 m. This record might refer to either the new species or conical or P. papillata, which were collected together at that locality. Lopholithodes Brandt, 1848 Lopholithodes Brandt, 1848 [July]: 174. Type species by monotypy: Lopholithodes mandtii Brandt. Echinocerus White, 1848 [November]: 47. Type species by monotypy: Echinocerus (Lithodes) cibarius White [= Lopholithodes mandtii). Ctenorhinus Gibbons, 1855: 48. Type species by monotypy: Ctenorhinus setimanus Gibbons [= Lopholithodes mandtii]. Echidnocerus: Bouvier, 1896: 12, 21. Carapace tuberculate, broader than long, anterior part of branchial margin extended laterally to cover base of walking legs; rostrum short, armed with spini- ACADEMY OF SCIENCES VOLUME 73 form tubercles. Antennal acicle (scaphocerite) tri- angular, with row of spines on margins. Carpus of chelipeds with a prominent lobe on inner margin, sometimes covering mouthparts; chelipeds and walk- ing legs fitting neatly together when folded, and sometimes folding under carapace. Second (basal) abdominal somite fused into a single plate; median plate of somites 3—5 entire; lateral plates of somites 3-5 distinct and paired; in males, marginal plates distinct or fused to laterals on somite 3, distinct on somites 4 and 5; in females, marginal plates of so- mites 3-5 present on right side only. Lopholithodes diomedeae (Faxon, 1893) Echinocerus diomedeae Faxon, 1893: 164; Faxon, 1895: pl. 7 figs. 3, 3 a, b. Paralomis diomedeae: Faxon, 1895: 46. Echidnocerus diomedeae: Bouvier, 1896: 27. Lopholithodes diomedeae: del Solar, 1972: 5, 14. Diagnosis: Carapace tuberculate; anterolateral mar- gins irregularly toothed. Rostrum short, with three spines or sharp tubercles, one median inferior and two paired superior. Antennal acicle with four or five spines on each margin. Carpus of chelipeds with inner lobe toothed on margin; outer margin with an unarmed crest. Walking legs longer than chelipeds and longer than greatest width of carapace; on first pair, anterior margin of carpus with an entire crest, of propodus with crest along proximal half and two or three teeth distally; on second and third pairs, carpus and propodus toothed along anterior margin. Previous records: 07°31'30" N, 79°14’ W, 458 fms [837 mm]; 07°21’°N; 79°35’) WS" fimss[93'5) mi] (Faxon, 1893, 1895). 03°48’ S, 81°22’ W, 680 m; 10°01’ S, 79°10’ W, 830 m (del Solar, 1972). Material examined: Male; off Peri, 10°01’ S, 79° 05’ W; 830 m, hard mud bottom; 19 May 1971; E. M. del Solar on R/V “SNP-1.” Measurements: Length of carapace excluding rostrum, 101 mm; rostrum, 9 mm; maximum width of carapace, 128 mm; length of right, third walking leg: merus, 56 mm; carpus, 38 mm; propodus, 39 mm; dactyl, 47 mm. Remarks: Faxon described this species from two female specimens. My male differs from the description and illustration of the larger, 64 mm type in having the various articles of the chelipeds and walking legs more elongate. The genus Lopholithodes contains two other species, L. mandtii Brandt and L. foraminatus (Stimpson), both from the temperate north Pa- cific. In those species the walking legs are about as long as the chelipeds and shorter than the greatest width of the carapace, and all the pereio- pods are modified to fold under the carapace, the inner marginal crest of the carpus of the chelipeds covering the mouthparts and the whole forming a boxlike structure. In Lopholithodes diomedeae 1974 the legs are long and do not fold beneath the carapace, and the chelipeds are clongate enough so that the carpal crest cannot cover the mouth- parts. In the Peruvian specimen, the first male to be reported, the marginal plates of the third abdominal somite are almost completely fused with the lateral plates. In this character L. dio- medeae is closer to L. foraminatus than to L. mandtii, in which the marginal plates are distinct on the third abdominal segment of males. In the original description, Faxon (1893) placed this species in Echinocerus and compared it with E. foraminatus Stimpson; later (1895) he changed his mind as to its affinities and allied it with Paralomis granulosa (Jacquinot). Bouvier (1896) retained it in Echinocerus (with spelling Echidnocerus) alongside E. cibarius, E. setimanus (both = mandtii) and E. foraminatus. Lopho- lithodes has since been shown to have priority over Echinocerus, so the correct name for Faxon’s species is Lopholithodes diomedeae. It was re- ported for the first time in this combination by del Solar (1972), for whom I had provided an identification. Glyptolithodes Faxon, 1895 Glyptolithodes Faxon, 1895: 42. Type species by monotypy: Riinolithodes cristatipes Faxon. Carapace with a large, conical prominence on gastric region, one on each posterolateral margin, and two on posterior margin; a prominent, crescentic, rounded ridge on each branchial region, enclosing cardiac region in a deep fossa; rostrum conical, with a short, laterally compressed ventral process. An- tennal acicle (scaphocerite) triangular, margins spined. Walking legs flattened, nearly spineless, margins lobed or dentate. Second (basal) abdominal somite fused into a single plate; median plate of somites 3—5 entire: lateral plates of somites 3—5 distinct and paired; in males, marginal plates partially fused to laterals on somite 3 and distinct on somites 4 and 5; in females, marginal plates of segments 3—5 present on right side only. Glyptolithodes cristatipes (Faxon, 1893) Figure 5 Rhinolithodes cristatipes Faxon, 1893: 163; Faxon, 1895: pl. 7 figs. 2, 2 a-c; Bouvier, 1896: 27. Glyptolithodes cristatipes: Faxon, 1895: 43; del Solar, 1972555 13: Rhinolithodes (Glyptolithodes) monde, 1967: 3, pl. 1. Diagnosis: Carapace broader than long, covered with small granules; outer orbital and anterolateral angles with a small, blunt tooth; anterolateral (an- cristatipes: | Baha- NEW LITHODID CRABS FROM PERU terior branchial) margin with a few con or teeth; prominences on carapace a generic diagnosis; cardiac fossa Rostrum open ™ short, conical, unarmed dorsall process denticulate anteriorly, not Antennal acicle with a terminal two to four spinules on each margin peds flattened, outer visible in de view. pine and Carpus of che margin cristate and sometime cut into two or three lobes, inner expanded and cut dentiform with several sharp teeth. Walking legs greatest width of carapace in adults; anterior margin of merus, carpus, and propodus cristate, crest or cut into two to several of those articles dentate. Abdomen tuberculate. Previous records: 07°09'45"” N, 80°50’ W, 322 fm: [590 m] (Faxon, 1893, 1895). S of Banco de Man- cora, Pert, 400 m; 03°51’ S, 81°18’ W, 800 m; 07° 42’ S, 80°26’ W, 693 m (del Solar, 1972). Off Iquique, Chile, depth unrecorded; 25°11’ S, 70°31’ W, 245-266 m (Bahamonde, 1967). Material examined: Female (juvenile); S of Banco de Mancora, Pert, 03°51.3’ S, 81°18.2’ W; 795-800 m, mud bottom; 11 January 1971; E. M. del Solar on R/V “SNP-1.” Female; off Peri, 06°31.5’ S, 81°01.5’ W; 712-744 m, mud and hard sand; 17 May 1971; E. M. del Solar on R/V “SNP-1.” Male; off Puerto Chicama, Pert, 07°42’ S, 80°26’ W; 693 m, rocky bottom; 2 March 1971; E. M. del Solar on R/V “SNP-1.” Measurements: Length of male carapace excluding rostrum, 79.5 mm; rostrum, 6 mm; maximum width of carapace, 98 mm; length of right, third walking leg: merus, 44.5 mm; carpus, 28 mm; propodus, 38 mm: dactyl, 37.5 mm. Length of female including ros- trum, 89.5 mm. Length of juvenile including rostrum, 27 mm. margin with resi into lobes or armed longer than entire lobes; posterior margin Remarks: The original description of this species was based on a juvenile male specimen. The juvenile collected by del Solar is larger than the 16.5 mm holotype, but agrees closely with it. Adults, on the other hand, show a number of dif- ferences, some of which were pointed out by Bahamonde (1967), who reported six large speci- mens from off Iquique and Taltal, Chile. The carapace is broader than long instead of about as long as broad; the setae which decorate its lateral prominences in juveniles are absent in adults. There are three or four spines, instead of two. on each margin of the antennal acicle. The walking legs are much longer than the carapace width, and the various articles of these legs are propor- tionately more elongate than they are in juveniles. On the illustrated specimen (Fig. 3A) the artist has depicted an abnormally small (probably re- generating) third walking leg. The third walking 162 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Figure 5. legs are normally about the same size as the other two pairs. The abdomen of the 89.5 mm female bears a sacculinid 45 mm across. In his original description of this species, Faxon (1893) referred it provisionally to Rhinolithodes Brandt. Later he examined material of R. wos- nessenskii Brandt and concluded that the two species are generically distinct. Characters by which he distinguished Glyptolithodes from Rhinolithodes (Faxon, 1895: 43) include the form of the rostrum and legs and of the cardiac region, which is elevated and spherical in R. wosnessenskii, the type species of Rhinolithodes. Bouvier (1896), who evidently did not know of Faxon’s 1895 work, failed to mention Glypto- lithodes and retained G. cristatipes in Rhino- lithodes, a genus characterized by having the marginal plates of abdominal segments 3—5 com- Glyptolithodes cristatipes (Faxon). carapace in lateral view; C, abdomen of male; D A, male in dorsal view; B, same specimen, D, abdomen of female. Scale A= 30 mm. pletely fused with the corresponding lateral plates (Bouvier, 1895: 187; 1896: 21). This is true of both sexes, as I have confirmed by examination of a male and a female R. wosnessenskii in the col- lections of the Allan Hancock Foundation. The very different structure of the abdomen in G. cristatipes (Fig. 5 C, D) confirms Faxon’s opin- ion that his species belongs to a separate genus. REMARKS ON DISTRIBUTION Bouvier (1896) showed that the Lithodidae, a family adapted to cold and temperate waters, originated in the North Pacific and underwent tropical submergence during its migration south- ward along the Pacific coast of the Americas. On that coast it re-emerged at about 42° S, from which latitude southward it is represented by 1974 Lithodes antarcticus Jacquinot and Paralomis granulosa (Jacquinot) in the littoral and to depths of about 150 and 100 m, respectively. The deep- water, tropical component of this fauna first became known when (1893) described Lithodes panamensis, Paralomis longipes, P. asper, Lopholithodes diomedeae, and Glyptolithodes cristatipes from between 07°31/30” N and 05°26’ 20” N, in 590 to 1410 m. Shortly afterwards, Benedict (1895) described Paralomis papillata from a specimen collected at an unrecorded depth and an uncertain locality, probably off México. Rathbun’s (1910) pioneer compilation of the decapod Crutacea of Pert’ included no Litho- didae, because at that time practically no explora- tion of the deep sea had been made in Peruvian waters. This situation continued to exist until very recently, when the investigations of IMARPE (del Solar, 1972) yielded all five of Faxon’s species as well as Paralomis papillata (Benedict) and the two new species described in the present report. Glyptolithodes cristatipes was recorded from Chile at 25°11’ S by Bahamonde (1967), who noted that fragments of two undetermined species, a Paralomis and a Leptolithodes |[=Para- lomis], have been taken from stomachs of sperm whales captured in Chilean waters. Lithodes pana- mensis and Glyptolithodes cristatipes are known to occur off northern México (unpublished rec- ords). Thus, although much deep-water explora- tion of the Pacific American coast remains to be done, a general picture of the “tropical com- ponent” of the Lithodidae is becoming clearer: it consists of at least eight benthonic species and, considered as a unit, reaches its northern limit in tropical México but penetrates the temperate Chilean waters to the south. Faxon (1895: 51) reported small, indetermin- able juveniles of “two more species of Lithodes” from the collections of the “Albatross” off trop- ical western America. One of these, taken be- tween 16°33’ N and 0°04’ S in 1207 to 1847 m, belongs to Neolithodes A. Milne-Edwards and Bouvier, 1894. Neolithodes diomedeae (Bene- dict) was described off southern Chile from 42° 36’ S and 45°35’ S in 2454 and 1920 m, respec- tively, and has since been reported from 23°39’ N in 1382 m (Parker, 1964: 163). Until more is learned about the genus in the eastern Pacific, it is impossible to say whether Faxon’s specimens belong to N. diomedeae or to another, as yet undiscovered form. As I complete the final draft of this report, E. M. del Solar informs me (pers. comm.) that he has found a juvenile Neolithodes from Peruvian waters, and that he intends to Faxon NEW LITHODID CRABS FROM PERU investigate its and that of Neolithodes taken in Pera from the tol sperm whale (see introduction) status A deep-water species of Lithodes and Paralomis occur in the northeast Pacific and least as far south as southern California; they ar not well known from the southern part of their range, which may overlap to some extent the range of the tropical group of lithodids. Faxon’s second undetermined Lithodes (Faxon, 51, pl. 10 fig. 2), a juvenile collected at 21°19’ N, is perhaps a very small specimen of L. couesi Benedict, whose reported range is Bering Sea to San Diego, Cali- fornia. ACKNOWLEDGMENTS [ am pleased to acknowledge the help I received from several sources, without which this study could never have been completed. Enrique M. del Solar not only furnished the specimens on which it was based, but provided information on additional Peruvian material as well as constant encouragement. Henry B. Roberts provided photographs and information on the holotype of Paralomis papillata (Benedict). A sub- vention from the Sociedad Nacional de Pesqueria of Lima, Pert, made possible the illustrations, which are the work of Jerry J. Battagliotti. LITERATURE CITED Bahamonde, N. 1967. Rhinolithodes (Glyptolitho- des) cristatipes Faxon frente a la costa chilena (Crustacea, Decapoda, Anomura, Lithodidae). Not. Mens. Mus. Nac. Hist. Nat., Santiago, 136: 3-7. Benedict, J. E. 1895. Descriptions of new genera and species of crabs of the family Lithodidae. with notes on the young of Lithodes camtschati- cus and Lithodes brevipes. Proc. U.S. Nat. Mus., 17:479-488. Bouvier, E.-L. 1895. Recherches sur les affinités des Lithodes & des Lomiis avec les Pagurideés. Ann. Sci. Nat., Zool., (7) 18:157-213. 1896. Sur la classitication des Litt nés et sur leur distribution dans les oceé Ann. Sci. Nat., Zool., (8) 1:1-46. Brandt, J. F. 1848. Die Gattung Lithodes Latr nebst vier neuen ihr verwandten von Wosnes- senski entdeckten, als Typen einer besonderm Unterabtheilung (Tribus Lithodea) der Ed- 164 BULLETIN SOUTHERN CALIFORNIA wards’schen Anomuren. Imp. Sci. St. Petersbourg, Bull. Phys.-Math. Acad. 7:171-176. del Solar, E. M. 1972. Addenda al catalogo de Crustaceos del Peru. Inf. Inst. Mar Peru, 38: 1-21. Faxon, W. 1893. Reports on the dredging opera- tions off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California . .. by the U.S. Fish Commission steamer “Albatross,” during 1891. ... WI. Preliminary descriptions of new species of Crustacea. Bull. Mus. Comp. Zool., Harvard Univ., 24:149-220. 1895. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands . . . by the U.S. Fish Commission steamer “Albatross,” during 1891. ... XV. The stalk-eyed Crustacea. Mem. Mus. Comp. Zool., Harvard Univ., 18:1- 292. Gibbons, W. P. 1855. [Description of a new genus and species of crab.] Proc. California Acad. Sci., 1:48—49. Latreille, P. A. 1806. Genera crustaceorum et insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisque plurimis ACADEMY OF SCIENCES VOLUME 73 explicata. Vol. 1. + 302 pp. Paris and Strasbourg. xviii Milne-Edwards, A., and E. L. Bouvier. 1894. Troi- siéme campagne du yacht l’Hirondelle, 1887. Neolithodes, genre nouveau de la sous-famille des Lithodinés. Bull. Soc. Zool. France, 19:120— 122. Parker, R. H. 1964. Zoogeography and ecology of some macro-invertebrates, particularly mollusks, in the Gulf of California and the continental slope off Mexico. Vidensk. Medd. Dansk Naturh. Foren., 126:1-178. Rathbun, M. J., 1910. The stalk-eyed Crustacea of Perti and the adjacent coast. Proc. U.S. Nat. Mus., 38:53 1-620. White, A. 1848. Description of Echinocerus ciba- rius, a new species and subgenus of Crustacea. Proc. Zool. Soc. London, 16:47—49. 1856. Some remarks on Crustacea of the genus Lithodes, with a brief description of a species apparently hitherto unrecorded. Proc. Zool. Soc. London, 24:132-135. Accepted for publication March 29, 1973. RANGE EXTENSIONS FOR SOME CALIFORNIA MARINE ISOPOD CRUSTACEANS ERNEST W. IVERSON! ABSTRACT: New range extensions are given for 22 species of California marine isopods. Brief ecological data and intermediate localities are Excirolana kincaidi are added to the California fauna. included. Dynamenella dianae and New localities are given for four species previously known only from the type localities. Numerous collections of marine isopods have been made by the author along the California coast in the past several years. Examination of this and other material has resulted in 22 new range ex- tensions and numerous intermediate localities being recognized. New records were also en- countered while curating a portion of the marine isopod collection in the Department of Inverte- brate Zoology, California Academy of Sciences (CAS), the Hopkins Marine Station collection (HMS), Pacific Grove, California (recently trans- ferred to the CAS collection), and from materials sent to the author for identification. Several new records were obtained from a collection of iso- 1 Dept. Invertebrate Zoology, California Academy of Sciences, Golden Gate Park, San Francisco, Cali- fornia 94118. 1974 pods dredged by the R/V Amigo and the R/V Falcon during a benthic survey of the northern sector of Monterey Bay, California. The survey was conducted by the Moss Landing Marine Laboratory (MLML), Elkhorn slough, California, under contract to the Association of Monterey Bay Area Governments and later under the Sea Grant Program. The sublittoral isopod fauna ob- tained showed a remarkable resemblance to the southern California fauna reported by Menzies and Barnard (1959). Seven species were collected by the survey, two of which, Austrosignum sp. and Tecticeps convexus have not been reported from southern California. One species Bathycopea dal- tonae has previously been reported from off San Francisco, California (Loyola e Silva, 1971). The museum records reported herein may represent temporary non-reproducing populations resulting from introductions by man or from un- usual environmental conditions such as the oc- casional intrusion of warm water into central California from the south. Early records of introduced species are reported to provide possible base dates from which times of introduction and subsequent dispersal along the coast may be judged. Two species, Dynamenella dianae and Exciro- lana kincaidi are here added to the California fauna, while four species are reported for the first time outside their type localities. The specimens upon which the present range extensions are based have been deposited in the collections of the Department of Invertebrate Zoology, California Academy of Sciences or the Moss Landing Marine Laboratory (MLML benthic survey material only). SUBORDER ANTHURIDEA FAMILY ANTHURIDAE Haliophasma geminata Menzies and Barnard Previous distribution: Southern California coastal shelves and slopes, 9.2 to 512.4 m; Santa Catalina Island, 73.2 to 122.6 m; Santa Rosa Island, 14.6 m (Menzies and Barnard, 1959:19), and San Quintin Bay, Baja California, Mexico (Menzies, 1962:339). New northern record: Station 1159, SW Rio Del Mar, Monterey Bay, California, 36° 57.1’ N. Lat., 121° 56.2’ W. Long.; 3 May 1972; MLML benthic survey; dredged in 15 m over grey sand. Collected from other parts of Monterey Bay at depths ranging from 15 to 65 m by the MLML sur- vey. Bottom sediments varied from grey sand _ (shallowest stations) to mud (deepest stations). RANGE EXTENSIONS OF CALIFORNIA MARINE ISOPODS 165 SUBORDER ASELLOTA FAMILY JANIRIDAL lais californica (Richardson ) Previous distribution: Australia, New Zealand, Singa pore, and California from Bodega Bay, Tomales Bay, Bolinas Lagoon, and San Francisco Bay (Rotramel, 1972:193-194), and Newport Bay (Anonymous, 1965:203). New northern record: Near Samoa, Humboldt Bay, California; 29 April 1972; E. Iverson and R. Tal- madge; commensal on the sphaeromid isopod Sphaeroma quoyana (= S. Richardson, 1904). pentodon laniropsis epilittoralis Menzies distribution: Dillon’ Beach, Marin Co., California, and Pacific Grove, Monterey Co., Cali- fornia (Menzies, 1952:150—151). New southern record: 3.2 miles N Arroyo Laguna Creek, San Luis Obispo Co., California; 5 February 1972; E. Iverson; several specimens from green fila- mentous algae in the high intertidal. Previous FAMILY MUNNIDAE Munna halei Menzies Previous distribution: Tomales Point, Marin Co., California (Menzies, 1952:134). New southern record: El Capitan Beach, San Luis Obispo Co., California; 28 December 1971; E. Iver- son; numerous specimens from among the spines of the purple sea urchin, Strongylocentrotus purpuratus, taken among rocks and debris in the mid-tide zone. SUBORDER FLABELLIFERA FAMILY CIROLANIDAE Excirolana kincaidi (Hatch) Previous distribution: San Juan Island, Washington (George and Stromberg, 1968:252) to Coos Bay, Oregon (Hatch, 1947:208). New southern record: Torrance Beach, Los Angeles Co., California; 13 September 1972; E. Iver- son; screened from fine wet sand just below the maxi- mum reach of the waves near the high tide mark. The related isopod E. linguifrons (Richardson, 1899) was also collected from the same habitat. Further records: Intermediate localities in Cali- fornia include Drakes Beach (20 July 1969. G. L. Rotramel), and mouth of Bolinas Lagoon. Marin Co. (19 December 1972, John Chapman; Aquatic Park Marina, San Francisco Bay (25 May 1972. D. Behrens); and Princeton Harbor, Half Moon Bay. San Mateo Co. (5 July 1971, E. Iverson and D. Behrens). 166 FAMILY CYMOTHOIDAE Lironeca californica (Schioedte and Meinert) Previous distribution: Alaska (Hatch, 1947:211) to San Pedro, Los Angeles Co., California (Richardson, 1905a:260). New southern record: Entrada Point, San Quintin Bay, Baja California, Mexico; 23 March 1949; R. R. Harry ef al.; parasitic on the gills of the Dwarf Surf- perch, Micrometrus minimus. Remarks: Lironeca californica previously has been reported parasitizing three other species of fish. This is the second member of the family Embiotocidae to be infected. FAMILY SPHAEROMIDAE Dynamenella dianae (Menzies) Previous distribution: San Quintin Bay, Baja Cali- fornia, Mexico; Mayaquez Bay, Puerto Rico, and Eniwetok Atoll, Marshall Islands (Glynn, 1970:20). New northern record: Ventura Yacht Harbor, Ventura Co., California; 20 September 1972; E. Iverson; from fouling on floats and under rocks used as shoring. Further records: Additional new localities in south- ern California include Marina Del Rey Harbor (16 September 1972, E. Iverson) and Redondo Beach, Los Angeles Co. (14 September 1972, E. Iverson); Oceanside Harbor (19 September 1972, E. Iverson), and San Diego Bay, San Diego Co. (19 September 1972, E. Iverson). Exosphaeroma inornata Dow Previous distribution: Palos Verdes, Los Angeles Co., California (Dow, 1958:93). New northern record: Bune Point, Humboldt Bay, California, 29 April 1972; E. Iverson; from algae covered rocks. Further records: A eurytopic species from many additional localities in California between Horse- shoe Cove, Bodega Bay to near Arroyo Laguna creek, San Luis Obispo Co. Exosphaeroma octoncum (Richardson) Previous distribution: California (Richardson, 1899:836). New northern record: Tomales Head, Marin Co., California; 1 July 1969; D. Beach and B. Roth; one specimen from the CAS collection. Monterey Bay, Gnorimosphaeroma lutea (Menzies) Previous distribution: Popoy Island, Alaska (Menzies, 1954:14) to Oso Flaco Lake, San Luis Obispo Co., California (Eriksen, 1968:5). New southern record: El Capitan Beach, San Luis BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 73 Obispo Co., California; 28 December 1971; E. Iver- son; one specimen near the mouth of a small stream, under marine algae cast ashore. Gnorimosphaeroma noblei Menzies Previous distribution: Upper region of Tomales Bay, California (Menzies, 1954:21—22), and Urup Island, Kurile Islands, Sea of Okhotsk (Kussakin, 1972:156). New northeastern Pacific northern and southern records: Near Samoa, Humboldt Bay, California; 29 April 1972; E. Iverson and R. Talmage; numerous specimens from the mud overhang of small channels crossing a Salicornia flat and under marine algae at the edge of the flat; and Palos Verdes, Los Angeles Co., California; 13 September 1972; E. Iverson; under rocks in tide pools of the middle tide zone on the semi-protected outer coast. Sphaeroma quoyana Milne-Edwards Previous distribution: Australia, Tasmania, New Zealand, and in northeastern Pacific bays: in Cali- fornia: Bodega Bay, Tomales Bay, Bolinas Lagoon, San Francisco Bay (Rotramel, 1972:193-194), San Pedro, Los Angeles Harbor (Johnson and Snook, 1927:287), Newport Bay (Menzies, 1962:340), San Diego Bay (Johnson and Snook, 1927:287), and in Mexico: San Quintin Bay, Baja California (Menzies, 1962:340). New northern record: Near Samoa, Humboldt Bay, California; 29 April 1972; E. Iverson and R. Tal- madge; numerous specimens burrowing in the mud banks of the small channels running through the Salicornia flat. One dried lot of S. quoyana was found in the HMS collection from Humboldt Bay (1931, G. MacGinitie collector). The commensal lais californica is present in both the 1931 and 1972 collections. Further records: Morro Bay, San Luis Obispo Co., California (24 September 1972, J. Carlton). Remarks: Sphaeroma quoyana (as S. pentodon Richardson, 1904) has been reported twice from Alaska, first, under questionable circumstances, by Atwood and Johnson (1924:26) and again by John- son and Snook (1927:288). These records probably resulted from Richardson including the original de- scription of S. pentodon in the reports of the Harri- man Alaska Expedition (1904:214-215). Her de- scription was based on ten specimens collected by Ritter and party at Sausalito, San Francisco Bay. Al- though several authors have studied the fauna of the upper northeastern Pacific, additional records are not available. Thus the Alaskan records are probably not valid. Tecticeps convexus Richardson Previous distribution: Monterey Bay, California (Richardson, 1899:838). New northern and southern record: 0.5 miles SW 1974 Crescent City Whistle Buoy, California; 13 August 1940; collector unknown; 24 specimens trawled from 31 m; and near Point Conception, Santa Barbara Co., California; 14-17 July 1916; Carl L. Hubbs; both lots are from the CAS collection. Further records: Additional intermediate localities in California include: 1.5 miles SW off the Mad River (9 August 1940; collector unknown); Horse- shoe Cove, Bodega Head (7 August 1971); J. Carl- ton); off Doran Beach, Bodega Bay (29 March 1967, A. Kuris and J. Ackeman), and Drakes Bay, Marin Co., (2 March 1936; L. Hertein). SUBORDER, GNATHIIDEA FAMILY GNATHIIDAE Gnathia crenulatifrons Mond Previous distribution: Southern California coastal shelves and slopes, Santa Catalina Island and basin at depths from 9.2 to 1259 m (Menzies and Barnard, 1959:29). New northern record: Station 1152, south of Point Santa Cruz, Monterey Bay, California; 36° 54.8’ N. Lat., 122° 1.0’ W. Long.; 20 August 1971; MLML benthic survey; dredged in 37 m over grey sand. SUBORDER VALVIFERA FAMILY ARCTURIDAE Idarcturus allelomorohus Menzies and Barnard Previous distribution: Point Conception to Laguna Beach, California, and Cortes Bank, SW San Clemente Island, California, at depths from 12.8 to 91.5 m (Menzies and Barnard, 1959:23). New northern record: Station 1153, SW Soquel Point, Monterey Bay, California; 36° 56.7’ N. Lat., 121° 59.2’ W. Long.; 21 August 1971; MLML ben- thic survey: dredged in 35 m over grey sand. FAMILY IDOTEIDAE Edotea sublittoralis Menzies and Barnard Previous distribution: Southern California coastal shelves from Point Conception to the Mexican border at depths from 13.7 to 64 m (Menzies and Barnard, 1959:22). New northern record: West of Daly City, San Francisco Co., California; 9 September 1972: John Chapman; near a sewer outfall at a depth of 9.8 m. Further records: Several specimens were collected in the northern sector of Monterey Bay, California by the MLML benthic survey in 12 to 35 m over a sandy substrate. RANGE EXTENSIONS OF CALIFORNIA MARINE ISOPODS 167 Idotea (Idotea) rufescens (Fee) Previous distribution: Gabriola Pass, Departure Bay Vancouver Island (Fee, 1926:18) to Marin Co., Cali fornia (Menzies, 1950:170), New southern record: South of Carmel, Monterey Co., California; 8 January 1967; D. Wabbler: one specimen from the CAS collection. Remarks: This specimen, an ovigerous female, agrees with the diagnosis given by Menzies (1950: 170) except for the apex of the frontal process, which is notched. Similar variation is known from one other eastern Pacific idoteid, /. aculeata. (Pentidotea) Idotea (Pentidotea) montereyensis (Maloney) Previous distribution: Seabech, Kitsap Co., Washing- ton (Hatch, 1947:219) to Estero Bay, San Luis Obispo Co., California (Menzies, 1950:187). New northern and southern records: Boundary Bay, British Columbia; July 1916; F. W. Weymouth: one specimen from the CAS collection; and Pismo Beach, San Luis Obispo Co., California; 23 October 1970; E. Iverson; one specimen from drifting brown algae in the surf. Idotea (Pentidotea) resecata Stimpson Previous distribution: Karta Bay, southeast Alaska (Richardson, 1905b:216) to Los Coronados Islands, SW San Diego, California (Miller, 1968:20). New southern record: At sea in the mouth of the Gulf of California between 23° 12’ N. Lat., 106° 29° W. Long., and 23° 3’ N. Lat., 109° 31’ W. Long.; 3 August 1932; Crocker Galapagos Expedition: one specimen from the CAS collection. Remarks: In California, [. (P.) resecata is com- monly found in the intertidal zone among rocks and algae; subtidally it is found on the eelgrass Zostera and in the Macrocystis canopy offshore. It is not known if the present specimen was associated with drifting algae. Idotea (Pentidotea) schmitti (Menzies) Previous distribution: Bering Sea to Monterey Bay. California (Menzies, 1950:171). New southern record: Punta Banda, Baja del Norte, Mexico; 30 December 1971: D. Conners: one large male from the CAS collection. Further records: Additional intermediate localities in California include Point Sal (1915, collector un- known) and near Point Conception. Santa Bar- bara Co. (14-15 July 1916, Carl L. Hubbs). Both lots are from the CAS collection. Idotea (Pentidotea) stenops (Benedict) Previous distribution: Coos Bay, Oregon (Hatch, 1947:217) to San Simeon Bay. San Luis Obispo Co.. California (Miller, 1968:21). 168 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES New southern record: Near Point Conception, Santa Barbara Co., California; 2—3 June and 14-17 July 1916; Carl L. Hubbs; two specimens from the CAS collection. Synidotea magnifica Menzies. and Barnard Previous distribution: Southern California coastal shelves: Point Conception to Oceanside at depths from 54.9 to 91.5 m (Menzies and Barnard, 1959:26). New northern record: 6.5 miles, 159° true from Point San Luis Light, San Luis Obispo Bay, Cali- fornia, 35° 3.4’ N. Lat., 120° 43.0’ W. Long.; 12 September 1938; Crocker-Stanford Expedition; dredged in 36.6 m over fine green mud. One speci- men from the CAS collection. After the acceptance of this paper, I received a copy of the Moss Landing Marine Laboratories benthic survey report (Hodgson, A. T. and J. W. Nybakken, 1973. A Quantitative Survey of the Ben- thic Infauna of Northern Monterey Bay, California. Moss Landing Marine Laboratories Technical Publi- cation 73-8, 241 pp.). The range extensions for Haliophasma geminata, Gnathia crenulatifrons, and Idarcturus allelomorohus are duplicated here. ACKNOWLEDGMENTS I am very grateful to Peter N. Slattery (MLML), James T. Carlton (CAS), David W. Behrens, and John Chapman (California State University, San Francisco) for making their material available. Robert R. Talmadge, of Eureka, California, kindly showed me several rich collecting areas in Humboldt Bay and at Trinidad Head, California. James T. Carlton offered many helpful suggestions on the manuscript. LITERATURE CITED Anonymous. 1965. An oceanographic and_bio- logical survey of the southern California main- land shelf. California State Water Quality Con- trol Board, Publication no. 27, xiii + 232 pp. Atwood, W. C., and A. A. Johnson. 1924. Marine structures their deterioration and preservation. Nat. Res. Counc., Washington, D.C., 534 pp. Dow, T. G. 1958. Description of a new isopod from California, Exosphaeroma inornata. Bull. So. California Acad. Sci., 57:93-97. Eriksen, C. H. 1968. Aspects of the limno-ecology of Corophium spinicorne Stimpson (Amphipoda) and Gnorimosphaeroma_ oregonensis (Dana) (Isopoda). Crustaceana, 14:1-12. Fee, A. R. 1926. The Isopoda of Departure Bay and vicinity, with descriptions of new species, VOLUME 73 variations, and colour notes. Contrib. Can. Biol. Fish., n.s., 3:15—46. George, R. Y., and Jarl-Ove Stromberg. 1968. Some new species and new records of marine isopods from San Juan Archipelago, Washington, U.S.A. Crustaceana, 14:225—254. Glynn, P. W. 1970. A systematic study of the Sphaeromatidae (Crustacea: Isopoda) of Isla Margarita, Venezuela, with descriptions of three new species. Memoria de la Sociedad de Ciencias Naturales la Salle, 30(85):48 pp. Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions. Univ. Washington Pub. Biol., 10:155—274. Johnson, M. E., and H. J. Snook. animals of the Pacific coast. New York, 659 pp. 1927. Seashore Macmillan Co., Kussakin, O. G. 1972. Isopoda from the coastal zone of the Kurile Islands. I. Janiridae and Jaeropsidae from Urup Island. Crustaceana, sup- plement 3: 155-165. Loyola e Silva, J. 1971. Sobre os géneros Ancinus Milne Edwards, 1840 e Bathycopea Tattersall, 1909, da colegao U.S. Nat. Mus. (Isopoda, Crustacea). Arquivos do Museu Nacional, 14: 209-233, 7 figs. Menzies, R. J. 1950. The taxonomy, ecology, and distribution of northern California isopods of the genus /dothea with the description of a new species. Wasmann J. Biol. 8:155—-195. 1952. Some marine asellote isopods from northern California, with descriptions of nine new species. Proc. U.S. Nat. Mus., 102:117— 159. 1954. A review of the systematics and ecology of the genus “Exosphaeroma,” with the description of a new genus and new species, and a new subspecies (Crustacea, Isopoda, Sphaeromidae). Amer. Mus. Novit., no. 1683, 23 pp. 1962. The marine isopod fauna of Bahia — de San Quintin, Baja California, Mexico. Pac. Nat., 3:337-348. Menzies, R. J., and J. L. Barnard. 1959. Marine Isopoda on coastal shelf bottoms of southern California: Systematics and ecology. Pac. Nat., 1:3-—35. 1974 Miller, M. A. 1968. Isopoda and Tanaidacea from buoys in coastal waters of the continental United States, Hawaii, and the Bahamas (Crustacea). Proc. U.S. Nat. Mus., 125:53 pp. Richardson, H. R. 1899. Key to the isopods of the Pacific coast of North America, with de- scriptions of twenty-two new species. Proc. U.S. Nat. Mus., 21:815—869. 1904. Isopod crustaceans of the north- west coast of North America. Pp. 211-230 in Harriman Alaska Expedition, Vol. 10, Crus- taceans, Doubleday, Page and Co., New York, 337 pp. ERRATUM Syvertsen, J. P. 1974. Relative stem water | potentials of three woody perennials in a southern joak woodland community. Bull. So. California | Acad. Sci., 73(2): 108-113. Page 109 (bottom second column) should read as follows: Relative humidity was recorded daily on a hygrothermograph located 10 km northeast of the site at California Polytechnical University, Pomona (Table 1). Relative humidity was also measured on sampling days at each station with a sling jpsychrometer. All other climatological data (Tables 2-3) were from the U.S. Dept. of Com- merce (1971-72); temperatures were from the ‘Cal Poly Pomona Weather Station and the pre- RANGE EXTENSIONS OF CALIFORNIA MARINE ISOPODS 169 ——. 1905a. Isopods from the Alaska Salmon Investigation, Bull. U.S. Bur. Fish. 1904, 24 209-221. ———. 1905b. A monograph on the isopods of North America. Bull. U.S. Nat. Mus., no. 54: 727 pp. Rotramel, G. L. 1972. lais californica and Sphaeroma quoyanum, two symbiotic isopods introduced to California (Isopoda, Janiridae and Sphaeromatidae). Crustaceana, suppl. 3:193- 197. Accepted for publication October 24, 1973. cipitation information was from the Walnut Patrol Station Weather Station approximately 4 km north of the site. Both weather stations are located in the same drainage system as the study site but approximately 70 m lower in elevation. Differences in mean SWP (Figs. 1-6) Statistically analyzed using a randomized complete- block design (Steel and Torrie, 1960) analysis of variance in which the sampling dates were used as blocks. were RESULTS The data show that the individuals growing at the higher slope positions had _ significantly Lines in italic moved from page 110 (first column). RESEARCH NOTES A NEW SPECIES OF COMATACARUS (ACARINA: TROMBICULIDAE) FROM CENTRAL UNITED STATES Studies of North American chiggers of the subfamily Leeuwenhoekiinae have revealed a new species of Comatacarus Ewing, a genus recently revived by Reed (1973). Examples of this species were reported previously as Leeuwenhoekia (Comatacarus) ameri- cana (Ewing) in studies of the larvae (Loomis, 1956, and Finley, 1958) and nymphal stage (Crossley, 1960). The type series is in the University of Kansas chigger collection currently housed at California State University, Long Beach. However, the holotype will be sent, on loan, to the Rocky Mountain Laboratory (RML and USNM). 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. Comatacarus pusillus, new species Figure 1 Types.—Holotype (KU 6171, in RML) plus 4 para- types (KU 6172-6175) from 24 km N, 19 km W St. Francis, Cheyenne Co, Kansas, from Peromyscus maniculatus (RL 521101-14) taken 1.XI.1952 by R. B. Loomis; 4 paratypes (KU 6167-6170) same data as holotype except host Reithrodontomys mega- lotis (RL 521101-15). Diagnosis.—Palpal setal formula B/B/BBB + 7B; chela with tricuspid cap; galeala branched; 2 genualae I, genuala II and III; tibiala III; subterminala and parasubterminala I; femur I with incomplete suture; sensilla nude; PW/SD = 1.4. Description of holotype (all measurements in mi- crons with means and extremes of the nine types in parentheses ).—Larva: red in life measuring 400 x 270, partially engorged. Idiosoma: Dorsal body setae arranged 2-44-12- 14-14-14-8-6-4 total 118: measurements; humerals 57; anterior dorsal 66-39. Ventral setae 94; measure- ments; 2 sternal setae 42, 48; preanal setae 27-30, 44; postanal setae 36-42; total body setae 220. Gnathosoma: palpal setal formula B/B/BBB + 7B; chela 44, with a tricuspid cap; trifurcate palpal claw; galeala branched. Scutum: With few and scattered puncta anterior margin biconcave, posterior margin smoothly rounded; AM _ seta with short accessory branch on_ basal third; AL bases anterior to AM bases; SB slightly anterior to PL bases; PW/SD = 1.4; sensilla nude. Scutal measurements: AW 72 (73, 72-75); PW 87 (87, 81-89); SB 30 (27, 21-30); ASB 39 (35, 30- 39); PSB 27 (27, 27-30); AP 39 (38, 36-40); AM 48 (47, 45-50); AL 51 (53, 48-60); PL 45 (48, 45-55); S 84 (84, 78-87); nase 16 x 10. Legs: 6-6-6 segments, terminating in pair of claws and clawlike empodium, without onychotriches. Leg I: 360 (350-400); coxa with 2B (branched setae); trochanter with 1B; femur with 1B proximal, incomplete suture; and 5B distal; genu with 4B, 2 genualae, microgenuala; tibia with 8B, 2 tibialae, microtibiala; tarsus 117 30 with 40B, tarsala 18 (18-21), microtarsala, parasubterminala, subtermi- nala, pretarsala. Leg II: 312 (312-355); coxa with 1B; trochanter with 1B; femur with 6B; genu with 4B, genuala, microgenuala; tibia with 6B, 2 tibialae; tarsus 93 30 with 25B, tarsala 21 (21-24), microtarsala, pretarsala. Figure 1. Larva of Comatacarus pusillus, new species, A, scutum and eyes; B, dorsal aspect of gnathosoma: C, ventral aspect of palpal tibia and tarsus; D, selected setae (H, humeral; AD, anterior dorsal; PD, posterior dorsal; S, sternal; V, ventral); E, leg I distal three segments showing nude setae with measurements in microns and bases of branched setae: F, leg II as above; G, leg III as above. 170 1974 Leg IIT: 378 (354-400); coxa with IB; trochanter with 1B; femur with 5B; genu with 4B, genuala; tibia with 5B, tibiala; tarsus 114 % 24 with I8B. Taxonomic Remarks. —Comatacarus — pusillus is most closely similar to C. americanus Ewing (1942). Examination of the lectotype of C. americanus (USNM 1416) revealed that it has a nude lateral palpotibial seta, shorter tarsal segments (1, 60; TI, 55; Ill, 65) and an femur I. Both C. americanus and C. pusillus have 2 genualae I whereas the other three North American species, C. sfewarti (Gould), C. dolosa (Gould) and C.. inconspicuus Reed, have only one genuala I. As noted by Vercam- men-Grandjean et al. (1973) C, stewarti also appears to have a divided femur’ I. Reed (1973) restored Comatacarus to generic status, and we agree with its removal from Leeuwwenhockia, as confirmed by examination of the holotype of the type-species, L. verduni (OQudemans). Reed (1973) reported that all species of Comata- carus had an accessory branch on the AM seta. Not only was this character present in C. pusillus, it also was seen in three North American species of | Odontacarus; O. chiapanensis (Hoffmann), O. hirsutus | (Ewing) and O. villosus Goff and Loomis, all mem- | bers of the subgenus Tarsalacarus Vercammen-Grand- | jean. Crossley (1960) described a nymph, under the name Leeuwenhoekia americana, reared from a larva | taken with the type series of C. pusillus. | Specimens examined (16).—Comatacarus pusillus (10): KANSAS. Cheyenne Co, 24 km N, 19 km W | St. Francis, 1.X1.1952, Peromyscus maniculatus (5, holotype + 4 paratypes), Reithrodontomys megalotis (4 paratypes), COLORADO. Boulder Co, 21 km S Estes Park, 8.VIII.1947, Neotoma cinerea orolestes (1, KU 6130). Comatacarus americanus (6): OREGON. Port- land, 20.V.1936, western mole (USNM 1416, lecto- type + 5). undivided ACKNOWLEDGMENTS We are indebted to Dr. James M. Brennan of the Rocky Mountain Laboratory, Hamilton, Montana for advice and the loan of specimens, and to Janice Fisher for the illustrations. LITERATURE CITED Crossley, D. A. 1960. Comparative external mor- phology and taxonomy of nymphs of the Trombiculidae (Acarina). Univ. Kansas Sci. Bull., 40:135-321. Ewing, H. E. 1942. Remarks on the taxonomy | | | | of some American chiggers (Trombiculinae), RESEARCH NOTES 171 including the descriptions of new and species, J. Parasit., 28:485—493, genera Finley, R. B., Jr. 1958. The wood rats of Colorado distribution and ecology. Uniy. Kansas Publ., Mus. Nat. Hist., 10:213—552. Loomis, R. B. 1956. The chigger mites of Kansas (Acarina; Trombiculidae). Univ. Kansas Sci. Bull., 37: 1195-1443. Reed, J. T. 1973. Comatacarus, formerly subgenus of Leeuwenhoekia, restored to generic status, with description of a new species (Acarina: Trombiculidae). J. Med. Ent., 10:315—317. Vercammen-Grandjean, P. H., R. L. Langston, and J. R. Audy. 1973. Tentative nepophylogeny of trombiculids. Folia Parasitologica (Praha), 20:49-66. M. Lee GorF and RICHARD B. Loomis, Dept. Biology, California State University, Long Beach, California 90840. Accepted for publication March 13, 1974. BUZZ POLLINATION OF CASSIA QUIEDONDILLA (LEGUMINOSAE) BY BEES OF THE GENERA CENTRIS AND MELIPONA Van der Pijl (1939), Michener (1962). Linsley (1958. 1962a, b), and Wille (1963) described an effective mode of pollen collection (now termed buzz pollina- tion) that is used by female oligolectic bees while foraging for pollen on flowers having anthers with terminal pores. We now know that hollow. tubular anthers with apical pores are found in at least 400 genera in many plant families (Harris, 1905). Our knowledge of “buzz” (vibratile) pollination is still very incomplete. The present study is a preliminary note on the pollination ecology of an additional Cassia species. Field observations on bee visitations to Cassia quiedondilla Micheli, were conducted on 23 January 1974 on the highway midway between Tixtla and Chilapa, in the State of Guerrero, Mexico. Visitations by pollinating bees were observed for two hours, from 1100 to 1300. Cassia quiedondilla is a large (to 3 m.) caesalpina- ceous legume with large yellow flowers (Fig. 1) (Standley, 1927). The flowers are composed of two large curved petals and three highly reduced petals. The larger two petals function in combination as a = 172 BULLETIN SOUTHERN CALIFORNIA 10mm Figure 1. Cassia quiedondilla (Caesalpinaceae) The flower has four short “fodder” stamens, three short vestigial stamens and three long “pollinating” stamens. Centris anthracina Snelling and Melipona fasciata guerreroensis Schwarz vibrate their indirect flight muscles to release the pollen from the long curved pollinating anthers. landing platform and resonating chamber for the bees. There are three large (10 mm.) stamens and four smaller (6 mm.) stamens functioning as “fodder” stamens (Faegri and van der Pijl, 1966; Meeuse, 1961). In addition, there is a third group of three very small (2.5 mm.) rudimentary stamens. The stamens are arcuate and heteranthic. The flowers are a uniform orange-yellow with prominent dark veins, especially in the two large curved petals. There are no markings on the flowers which might function as nectar guides. Pressed voucher specimens of the flowers were photographed under ultraviolet light, using a Sony videorecorder T. V. camera fitted with a U. V. transmitting lens and filter. The flowers proved to be entirely ultra-violet absorptive making the flowers yellow not “bees purple,” (Jones and Buchmann, 1974). The three categories of vibratile pollinators de- scribed by Michener (1962), are: 1) the buzzing bees, 2) the biting bees, and 3) the gleaning bees. The two species of bees, Centris anthracina Snelling and Melipona fasciata guerreroensis Schwarz, which were observed collecting pollen from the flowers of C. quiedondilla used only the buzzing technique. Neither biting nor gleaning bees were observed on the flowers, although the time spent watching the pollina- tion interaction was too short. While working the flowers for the abundant pollen, ACADEMY OF SCIENCES VOLUME 73: both species of bees emitted loud intermittent buzzing sounds which could be heard up to five meters away) from the group of Cassia plants. The large black Centris anthracina is an extremely rapid flier, spending only a few seconds on an individual flower. It pro- duces much louder intermittent buzzes than the smaller Melipona fasciata guerreroensis. The behavior of both Centris and Melipona is essentially the same on the flowers, although the. heavier Centris causes the flower to hang in a pendant fashion. The lighter Melipona (approximately the size of Apis mellifera L.) does not cause this inversion of the flowers. Only female bees were observed in the Cassia flowers, since there is no nectar produced to attract male bees. Both bees land on the modified lower petal and move up until the head is positioned at the base of the banner petal. Curling their bodies tightly over the long pollinating anthers, the bees vibrate their indirect flight muscles (wings are held closed over the back) in one to three extremely short bursts of high (higher than normal flight sounds) frequency sounds. This vibration of the flower results. in loosening of the pollen and it shoots out of the terminal anther pores in the form of a small cloud. Most of the small light pollen is lost in this manner, but much of the pollen lands on the ventral part of the bee’s thorax and abdomen. After receiving pollen’ on its venter, the bee backs out of the flower and in mid-air combs the pollen into the scopa. Previous studies have indicated that some Solanum species produce an unusually high percentage of in- viable (judged by staining with cotton-blue) pollen grains (Linsley and Cazier, 1963). Linsley and Caziet1 reported testing pollen loads from Solanum elaegni- folium pollinators and found that 64 percent of the pollen grains were inviable. They suggested that the high proportion of inviable pollen grains was due to. some inherent genetic factor and not the result of high external temperatures as reported by Stow (1927). When mixed populations of S. rostratum and S. elaegni- folium were tested, Linsley and Cazier (1963, 1970) reported that the percentage of inviable grains was only 15.6 percent of the total. In Solanum douglassii Dunal, Jones, and Buchmann (unpublished) found that only 5 percent of the pollen produced was inviable. Cassia quiedondilla produces an extremely high percentage of inviable pollen grains. Tests made using a cotton-blue with lactophenol stain, revealed that inviable pollen averaged 92.6 percent of that pollen produced by the three anther types. The nor- mal pollinating anthers produced 95 percent inviable pollen (549 grains counted), while the “fodder” anthers tested revealed that 96 percent were inviable (500 grains counted) and the rudimentary anthers produced 87 percent inviable pollen (500 grains counted). As reported by various writers, Cassia produces two sizes of pollen grains. Linsley and Cazier (1972), have reported that Cassia bauhinoides 1974 produced 79 percent of the small-grained type (2,565 grains counted). The large grains were about 80 microns in diameter, whereas the small grains ‘were only 27 microns in diameter. The large grains in C. quiedondilla are 32.5 microns in diameter and the smaller grains are 20 microns in diameter, ‘These abundant small but viable pollen grains have been interpreted by Linsley and Cazier (1972) as a food “bonus” for pollinating bees. The ratio of small pollen grains in C. quiedondilla was found to be 43.4 percent of the 1,949 grains counted. Undoubtedly, these pollen grains act as an extra food reward for pollinat- ing bees. In addition, the presence of 43.4 percent small grains in the Cassia anthers may function as billiard balls and serve to loosen more of the pollen from the anthers. When the anthers are vibrated, the smaller grains would serve to knock more of the pollen out into the pollen cloud that is characteristic of buzz pollination in species of Solanum and Cassia. ACKNOWLEDGMENTS I would like to thank P. A. Adams, T. L. Hanes, B. H. Brattstrom, J. D. Smith, R. Snelling, M. Meden, and especially C. E. Jones, for comments on_ this manuscript. The pollinator determinations were made by Roy Snelling and the bees are on deposit in the collection of the Los Angeles County Museum of Natural History. Voucher specimens of C. quiedon- dilla used in this study are on deposit in the Faye MacFadden Herbarium, California State University, Fullerton, numbers 016358 to 016361. Special thanks goes to Paula Kay McKenzie for illustrating the Cassia flower in figure 1, and to David A. Young for providing transportation to the site. LITERATURE CITED Faegri, K., and L. van der Pijl. 1966. The prin- ciples of pollination ecology. Pergamon Press, London, 248 pp. Harris, J. A. 1905. The dehiscence of anthers by apical pores. Missouri Bot. Gardens, 16th Ann. Rept.: 187-257. RESEARCH NOTES 173 Jones, C. E., and S. L. Buchmann. 1974. Ultra violet floral patterns as functional orientation cues in hymenopterous pollination system Anim. Behav., 22:481-485. Linsley, E. G. 1958. The ecology of solitary bees Hilgardia, 27(19):543—S85. 1962a. The colletid Priloglossa arizonensis Timberlake, a matinal pollinator of Solanum. Pan-Pacific Ent., 38(2):75—82. 1962b. Ethological adaptations of solitary bees for the pollination of desert plants. Sartryck ur Meddelande nr 7 Sveriges Froodlareforbund, 189-197. Linsley, E. G., and M. A. Cazier. 1963. Further observations on bees which take pollen from plants of the genus Solanum. Pan-Pacific Ent., 39(1):1-18. 1970. Some competitive relationships among matinal and late afternoon foraging activities of caupolicanine bees in southeastern Arizona (Hymenoptera, Colletidae). J. Kansas Ent. Soc., 43(3) :251—261. 1972. Diurnal and seasonal behavior pat- among adults of Protaxaea terns gloriosa (Hymenoptera, Oxaeidae). Amer. Mus. Novit., 2509:1—25. Meeuse, B. J. D. 1961. The story of pollination. Ronald Press, New York, 243 pp. Michener, C. D. 1962. An interesting method of pollen collecting by bees from flowers with tubular anthers. Rev. Biol. Trop., 10(2):167— 17/5 Pil, L. van der 1939. Over de meeldraden van enkele Melastomataceae. De Trop. Nat., 28: 169-172. Standley, P. C. 1927. Trees and shrubs of Mexico. Contrib. Nat. Herbarium, 400-411. Stow, I. 1927. A _ cytological study on pollen sterility in Solanum tuberosum. Japanese J. Bot., 3(3):217-237. Wille, A. 1963. Behavioral adaptations of bees for pollen collecting from Cassia flowers. Rev. Biol. Trop., 11(2):205-210. STEPHEN L. BUCHMANN, Dept. Biology, California State University, Fullerton, California 92634. Accepted for publication May 24, 1974. EDITORIAL: CRITICAL REVIEW OF MANUSCRIPTS As indicated in the Instructions for Authors (inside back cover), all manuscripts submitted to SCAS BULLETIN are reviewed by referees who critically read for scientific content, originality, and clarity of presentation, and who assist the editor in making judgments as to acceptability for publication. The reviewers also assist authors by offering constructive suggestions which may lead to improvement of the text or illustrations of manuscripts. By contribution of their time and professional expertise the referees measurably enhance the quality of SCAS BULLETIN. In behalf of our readers, authors, and myself, I thank the following reviewers for valuable editorial assistance in recent months: Phillip A. 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All changes in galley proof attributable to the author (misspellings, inconsistent abbreviations, deviations from style, ete.) will be charged to the author. Reprint orders are placed with the printer, not the Managing Editor. CONTENTS Tube dwelling behavior in the marine annelid Phragmatopoma californica (Fewkes) (Polychaeta: Sabellariidae) (By RetersAlanwRoyininstenveehe= -Varlcisienernenel eroledlelcichelstoiclee teh tenn mer enee ne 117 The systematics and distribution of marine Tubificidae (Annelida: Oligochaeta) in the Bahia de San Quintin, Baja California, with descriptions of five new species. By David G. Cook .... 126 A new species of fossil Nuttallia (Mollusca: Bivalvia) from the Pliocene of Sonoma County, California. By Barry Roth and Raj Guruswami-Naidu..........0.. eee e eee e eee e nee 140 Two new species of land snails from the Pinaleno mountains, Arizona. By W. O. Gregg and W. BaMVALlers. vii trarvercupereitechaliaiaretevene oleve (eneisn caeralouctotiaiausyabsneneterte te eiiclstrene re oT HRT eR ReMekeRe 146 Observations on the lithodid crabs of Pert, with description of two new species. By Janet Haig 152 Range extensions for some California marine isopod crustaceans. By Ernest W. Iverson ...... 164 ) 2 Te: Na) Von a een etn Mn eee RE it RE SA ie ee eee Eee RUA DRS bith G o'o.g 000 0'0 0 169 RESEARCH NOTES A new species of Comatacarus (Acarina: Trombiculidae) from central United States. By M. Lee GojffrandvRichard BMLOOMIS Sanececis oad eee 170 Buzz pollination of Cassia quiedondilla (Leguminosae) by bees of the genera Centris and Melip- Ona YByuStephen vl BUCHMAN. woes oleieliscisiotae ce oe oe yO Riieiee reiote 171 Editorialsii@niticalireview of Manuscripts}