ISSN 0038-3872 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES BOLLETIN Volume 89 Number 3 BCAS-A89(3) 97-148 (1990) DECEMBER 1990 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 © Southern California Academy of Sciences, 1990 OFFICERS Camm C. Swift, President June Lindstedt Siva, Vice-President Hans M. Bozler, Secretary Takashi Hoshizaki, Treasurer Jon E. Keeley, Technical Editor Gretchen Sibley, Managing Editor BOARD OF DIRECTORS 1988-1990 1989-1991 1990-1992 Sarah B. George Takashi Hoshizaki Jack W. Anderson Margaret C. Jefferson George T. Jefferson Hans M. Bozler Susanne Lawrenz-Miller David L. Soltz Theodore J. Crovello John D. Soule Camm C. Swift Peter L. Haaker Gloria J. Takahashi Robert G. Zahary June L. Siva Membership is open to scholars in the fields of natural and social sciences, and to any person interested in the advancement of science. 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Date of this issue 6 December 1990 THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 97-108 Southern California Academy of Sciences, 1990 Prairie Dog Food Preference and the Photosynthetic Pathway-Selective Herbivory Hypothesis Harrington Wells, Waynelle Mason, and James R. Stewart Faculty of Biological Science, University of Tulsa, Tulsa, Oklahoma 74104 Abstract. —Black-tail prairie dogs (Cynomys ludovicianus) at the Tulsa Zoological Park were offered diets consisting of C, and C, plant species under conditions of controlled food species abundance and availability, in order to test whether the photosynthetic pathway-selective herbivory hypothesis is predictive of foraging behavior. The C, species tested were Setaria italica, Zea mays, Eragrostis curvula, and Salsola kali. The C, species tested were Arachis hypogea, Daucus carota, Helianthus annus, and Lactuca sativa. Four experiments were performed. I. Each food was presented separately to test whether C. ludovicianus would eat each species. II. All four C, species were presented simultaneously to test for preferences within the C, plant food group. III. All four C, species were presented simulta- neously to test for preferences within the C, food group. IV. Finally, all four C, and all four C, species were presented simultaneously to test for preferences between C, and C, food groups. Data demonstrated that prairie dogs have sig- nificant food preferences within both the C, and C, species groups tested. However, predictions of the photosynthetic pathway-selective herbivory hypothesis were not met. No statistical preference for either C, or C, species existed. The black-tail prairie dog, Cynomys ludovicianus, inhabits shortgrass prairies from eastern Montana through southwest North Dakota south to western Texas, New Mexico, and southeastern Arizona. The North American prairie supported approximately five billion black-tail prairie dogs during the 1 9th century (Coppock et al. 1983), at which time C. /udovicianus formed one of the region’s largest mammal populations. Prairie dog populations have now been greatly reduced in the wild. Presently, C. /udovicianus occupies less than 10% of its former range (Hoogland 1981), and existing populations are less than 2% of what they were a century ago (Coppock et al. 1983). Herbivory, pasture disturbance, and potentially explosive population growth have made the black-tail prairie dog incompatible with both agriculture and ranching. The observed population decline is largely the result of intentional eradication by man. Means of extermination have been developed that are in- expensive and effective (i.e., suffocation by gasoline; Kansas ranchers, personal communication). Relict populations in a few reserves (e.g., Wind Cave National Park, Devils Tower National Monument, and a reserve at Lubbock, Texas), and colonies maintained in Zoological parks, therefore, figure heavily in survival of the species. Determining black-tail prairie dog food preference is, thus, one im- portant component in their preservation and management. C. ludovicianus is strictly herbivorous. Natural diet of the black-tail prairie dog oi 98 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES includes seeds, shoots, and roots (Fagerstone and Williams 1982). Potential her- bivore forage in temperate North America includes both C, and C, plants. The photosynthetic pathway-selective herbivory hypothesis predicts that herbivores should tend to feed on C, rather than on C, plants in temperate climates when a choice exists, given that both types are palatable (Caswell et al. 1973). The selective herbivory hypothesis is based upon the assumption that C, plants, statistically, are a nutritionally poorer food resource than C; plants in temperate regions. Evidence that a general nutritional food difference for herbivores exists between C, and C, plants in temperate climate comes from two types of study. First, data from plant composition experiments depict a trend in which nitrogen and phos- phorus contents tend to be greater in C, plants than C, plants in temperate regions (National Research Council 1958; Wilson and Haydock 1971; Caswell et al. 1973). Other plant composition studies show a trend in which the ratio of digestible to non-digestible carbohydrates tends to be greater in temperate climate C, plants than C, plants (National Research Council 1958; Wilson and Ford 1971; Wilson and Haydock 1971; Minson 1971), and that herbivores select species greater in digestible carbohydrates (Spalinger et al. 1988). These nutritional differences may be related to carbon assimilation efficiencies of C, and C, species in temperate regions (Pearcy and Pjorkum 1983; Tolbert and Zelitch 1983), since transport and phloem unloading of nutrients (both carbon and nitrogen) from leaves is linearly dependent on the rate of carbon assimilation (Giaquinta 1980; Thorne 1985). Correspondingly, fruit and seed nutritive stores have been shown to be affected by the rate of leaf carbon assimilation during development of reproductive structures (Pate 1984). The second type of data supporting the hypothesis that a general nutritional difference exists between C, and C, plants in temperate regions comes from studies of herbivore survival and fecundity responses under laboratory conditions. These experiments generally use domesticated plants as food for native herbivore species in order to minimize palatability complications. The data show a trend of in- creased survival and/or fecundity when the diet consists of C, rather than C, plants (Smith et al. 1952; Wilbur 1954; Barnes 1955; Mulkern et al. 1962; Putnam 1962; Pickford 1962, 1963; Coupe and Schultz 1968; Delvi and Pandian 1971). Existence of a reward (nutritional) difference does not automatically mean that the difference can be perceived by a foraging animal (Wenner 1971; Wells and Wells 1983). Data from both field and laboratory herbivore food preference stud- ies, however, lend credibility to the proposal that nutritional differences between C, and C, plants actually affect herbivore foraging. Herbivores from temperate regions, in taxa ranging from insects to mammals, tend to select C; over C, plants as food when a choice exists in both lab preference tests and in observations of natural diets (Scharff 1954; Jantz 1962; Thompson 1965; Shade and Wilson 1967; Pruess 1969; Hansen and Ueckert 1970; Ueckert and Hansen 1971; Evans and Tisdale 1972; Flinders and Hansen 1972). Furthermore, the photosynthetic pathway-selective herbivory hypothesis ap- pears to be a robust model in two respects. First, the selective herbivory theory is not phylum specific. Second, the feeding preference of specific herbivores ap- pears to be the result of evolution to selectively forage on C, plants in general, rather than only selection to feed on specific C,; plants with which a particular herbivore has locally evolved. Eurasian herbivores given North American C, and PRAIRIE DOG FOOD PREFERENCE 99 C, plant species tend to select C, over C, species, as do North American herbivores given Eurasian C, and C, plant species (Scharff 1954; Thompson 1965; Jantz 1962; Pruess 1969; Evans and Tisdale 1972). This is not surprising since the ability of generalist foragers to detect reward quality as well as quantity differences is central to optimal foraging theory and would be predicted evolutionarily (Schoe- ner 1971; Cody 1974; Pulliam 1975; Pyke et al. 1977; Krebs 1978; Belovsky 1986). In fact, animal selection of food based upon ability to determine qualitative nutrient differences has been observed in a wide range of taxa (Moss et al. 1972; Pulliam 1975; Rozin 1976; Gross-Custard 1977; Wells and Wells 1984, 1986; Belovsky 1986; Provenza et al. 1987). This study was designed to test whether the photosynthetic pathway-selective herbivory hypothesis is predictive of black-tail prairie dog food preference under controlled conditions of plant species availability and abundance. Methods The study was conducted at the Tulsa Zoological Park, Tulsa, Oklahoma. Ex- periments I, II, and III were conducted from March through May, while exper- iment IV was performed from May to October, 1988. The colony of prairie dogs consisted of seven animals in a natural outdoor exhibit, without vegetation. Food for the prairie dogs prior to this study was Purina Monkey Chow. Prairie dog food preferences were tested using four C, plant species and four C, plant species. Domestic plant species were used to minimize unpalatability problems that can result from secondary plant compounds (Owen-Smith et al. 1983), following the design of prior experiments using other organisms (Scharff 1954; Jantz 1962; Pruess 1969). Each food used in a particular experimental trial was weighed prior to being offered to the animals, and a time limit was set for feeding the animals. The time limit was 30 minutes per trial in experiments I, II, and III and 60 minutes per trial in experiment IV. After that interval had elapsed, any remaining food was removed and weighed. Plant species were selected for this study on the basis of year-round availability and general palatability to herbivores. None are found naturally in the prairie dog range; thus specific co-evolved relationships between animals and plants should not confound the study. The C; species tested were Lactuca sativa (lettuce leaves; 56 g/trial = 8 g/animal-trial), Arachis hypogea (unshelled peanuts; 28 g/trial = 4 g/animal-trial), Daucus carota (carrot roots; 56 g/trial = 8 g/animal-trial), and Helianthus annus (unshelled sunflower seed; 14 g/trial = 2 g/animal-trial). The C, species tested were Setaria italica (millet seed; 56 g/trial = 8 g/animal-trial), Zea mays (dry corn seed removed from cob; 56 g/trial = 8 g/animal-trial), Era- grostis curvula (lovegrass seed; 28 g/trial = 4 g/animal-trial), and Salsola kali (Russian thistle seed; 56 g/trial = 8 g/animal-trial). Amounts of each food species were chosen such that 100% could be consumed by the prairie dogs in a trial, and so that volumes were approximately equal. Thus, abundance of a food would not affect prairie dog food preferences. Food was placed in ceramic bowls when given to test animals. Each plant species was placed in a separate bowl. Experiments were run at normal feeding periods. Only one experiment (food offering) was performed per day. There were four parts to this study. I. One plant species presented per trial. —First, prairie dogs were tested to see if they would eat each species of plant chosen for the experiment. Each of the 100 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES four C, plant species and each of the four C, plant species of food were presented to the test animals five times. Only one plant species was presented at a time, only one species per day, and the order in which plant species were presented was random. The animals were presented food for 30 minutes during each trial. II. C; plant species presented simultaneously.—The second part of this study was to determine prairie dog food preference among the species of C, plants. Animals were presented samples of the four C, species simultaneously, once each day, for ten consecutive days. Preference was determined by ranking the C; species, based on percentage of each plant species eaten. Animals were presented food for 30 minutes during each experimental trial. III. C, plant species presented simultaneously. —The third experiment was iden- tical in design to the second, with the exception of plant species used. The four C, species were offered simultaneously to the animals on ten consecutive days, once each day. Preference was determined by ranking within the C, species, based on percentage of each plant species eaten. Animals were presented food for 30 minutes during each experimental trial. IV. C; and C, plant species presented simultaneously. —Finally, the animals were offered, simultaneously, all four C, and all four C, plants chosen for this study. This part of the study was also conducted for a total of ten days, with feeding limited to once each day. Preference was determined by ranking species offered, based on percentage eaten. Animals were presented food for 60 minutes during each experimental trial. The Kruskal-Wallis one-way analysis of variance by ranks was used in exper- iments II, III, and IV to test whether the responses to food species within an experiment were different (Siegel 1956; Leach 1979). The null hypothesis in each experiment predicted that no difference in response to different species of foods existed. Data from experiment IV were partitioned into two groups: C, species and C, species. If no preference by prairie dogs exists overall between C, and C, plants, then the rank sums of the two groups should be approximately equal. The null hypothesis that there is no preference for either group was tested using the Mann- Whitney U statistic, of which the Kruskal-Wallis statistic is a direct extension for more than two groups (Leach 1979). Multiple comparison tests were used to supply detail about the differences detected by the ANOVA. Two statistics were used. The first test, the REGWQ multiple-stage statistic, controls maximum experimentwise error rate (a = 0.05 overall) under any complete or partial null hypothesis (Ryan 1959, 1960; Einot and Gabrial 1975; Welsch 1977). The second statistic was Fisher’s unprotected LSD test in which comparisonwise error rate is controlled (a = 0.05 each com- parison), but not experimentwise error rate (Miller 1981; SAS/STAT 1988). Finally, food preference was compared to food composition. Published data on forage composition exist for all species used except E. curvula and S. kali seeds (Henry and Morrison 1922, 1959; USDA Handbook 8-11 1980). Composition of E. curvula seed and S. kali seed were determined by laboratory analysis, fol- lowing the methods established by the Association of Official Analytical Chemists (Williams 1984). Whole seeds of these two species were weighed, and then dried at 60°C to a constant mass. Dried seeds of each species were assigned to one of three groups. Neutral lipids were extracted from ground seeds of the first group PRAIRIE DOG FOOD PREFERENCE 101 of each species using a Goldfisch lipid extractor (Labconco Model 35001) for eight hours with petroleum ether as a solvent. The second sample of seeds from each species was ground and analyzed for total nitrogen using an ammonia se- lective electrode and an Orion microprocessor ionalyzer (Model 901) following Kjeldahl digestion. Total protein was estimated by multiplying total nitrogen by a factor of 6.25 following Williams (1984). The third group of seeds from each species was ashed at 550°C for 24 hours in a Thermolyne ashing furnace. Car- bohydrate was estimated by subtraction of the combined estimates of protein and lipid from ash free dry seed mass. Kendall rank correlations (Siegel 1956) were performed to examine the relation of forage composition to food preference. Results I. One plant species presented per trial.—The prairie dogs ate 100% of the single species of plant presented each day, regardless of the plant species or whether the food was a C, or C, plant. Therefore, C. /udovicianus will eat all plant species used in this set of experiments. Furthermore, the animals in this study can regularly consume the quantity of any one of the food species used in experiments II, III, and IV. II. C; plant species presented simultaneously. — Black-tail prairie dogs showed a distinct food preference among the C, group of plants (Table 1). Based on rank sums, peanut appears to be most favored, and lettuce least favored. III. C, plant species presented simultaneously. —Food preference by black-tail prairie dogs exists within the C, group of plants (Table 2). Again based on rank sums, thistle appears to be preferred, and corn least favored. Furthermore, the similar values of the Kruskal-Wallis statistic (H) in parts II and III suggest that approximately equal disparity in preference exists within the C, group as within the C, plants. IV. C; and C, plant species presented simultaneously. —Food preference exists, as predicted from parts II and III, within the combined group of C, and C, plants (Table 3). However, a general preference either for C, or for C, plants does not appear to exist (Table 3). Both multiple pairwise comparison test results (Figs. 1 and 2) show A. hypogea (C;) being the preferred food, followed by H. annus (C;), S. italica (C,), and S. kali (C,). Fisher’s LSD test separates A. hypogea into a unique most preferred group (Fig. 2), whereas the REGWQ test combines the first three taxa into a single group, and places S. kali into the less preferred food groupings (Fig. 1). Zea mays (C,), E. curvula (C,), and L. sativa (C,) are placed into groupings in both analyses representing foods that tend to be avoided (Figs. 1 and 2). Salsola kali (C,) and D. carota (C;) in both Fisher’s LSD and the REGWQ methods are intermediate in preference (Figs. 1 and 2). Thus, the multiple pairwise comparisons do not separate the C, and C, plants into distinct groups, or even segregate C, seeds from C, seeds, based on daily quantities eaten. The observed feeding preference of the black-tail prairie dog is partially ex- plained by food composition. The correlation of C. ludovicianus food preference to forage percentage lipid content is 7 = 0.50 (significance level: P = 0.054), and to percentage protein content is 7 = 0.61 (P = 0.031); based on percentage wet mass (dry mass results were 7 = 0.47 and 0.31 respectively). The two preferred foods correspond to forage with highest lipid and protein composition. However, 102 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1. Results of prairie dog food preference when only C, plant species were presented simul- taneously. Data are expressed as percent by weight of food consumed. Significant food preferences existed (H = 20.57, df = 3, P < 0.001). Percent food consumed each day Day A. hypogea HA. annus D. carota L. sativa 1 100 29 39 66 Dy 100 64 18 29 3 79 43 02 43 4 86 00 00 39 5 100 75 63 47 6 88 88 03 13 7 100 50 81 50 8 100 63 47 59 9 81 75 06 06 10 100 75 100 13 Mean % 93.4 56.2 35.9 36.5 Rank sum 343 200 140 137 food preference differences in feed with lower protein and lipid content are not explained by comparing either percentage protein, lipid, or carbohydrate content (Table 4). The reason S. italica (C,) seed is a preferred food, while Z. mays (C,) and E. curvula (C,) are avoided, is not known. Seed size does not appear to be the deciding factor since Z. mays, H. annus, and A. hypogea all have large seed, while S. kali, E. curvula, and S. italica all have small seed. Discussion Herbivore foraging behavior under natural conditions can be dependent on several factors related to forage. The cumulative effects of these factors, in turn, determine optimal foraging in a particulr habitat. Forage availability, handling Table 2. Results of prairie dog food preference when only C, plant species were presented simul- taneously. Data are expressed as percent by weight of food consumed. Significant food preferences existed (H = 19.09, df = 3, P < 0.001). Percent food consumed each day Day S. kali S. italica E. curvula Z. mays 1 38 14 04 11 2 66 80 39 18 3 52 23 04 09 4 Si, 09 00 09 5 78 13 13 00 6 100 78 31 34 7 75 06 06 03 8 81 44 38 16 9 72 03 13 00 10 100 50 38 47 Mean % 71.9 32.0 18.6 14.7 Rank sum 336.5 204.5 148.5 130.5 PRAIRIE DOG FOOD PREFERENCE 103 Table 3. Results of prairie dog food preference when C, and C, plant species were presented simultaneously. Data are expressed as percent by weight of food consumed. Significant food preferences existed (H = 40.71, df = 7, P < 0.001), but not between the groups C, and C, plants (U = 717.5, n, =n, = 40, P= 0.156). Percent food consumed each day C, Cy A. H. D. L. S. Ss: Z. E. Day hypogea annus carota sativa italica kali mays curvula 1 81 75 00 06 50 41 13 13 D 100 100 25 06 47 56 16 31 3 100 75 56 16 84 91 19 75 4 81 75 47 00 94 63 06 19 5 88 75 DD 16 72 31 34 23 6 100 63 44 19 41 53 38 38 7 81 63 94 28 56 88 84 38 8 100 63 69 25 100 59 63 44 9 100 63 25 13 59 44 50 19 10 100 63 100 06 100 97 91 56 Mean % 93.1 71.5 48.2 13.5 70.3 62.3 41.4 35.6 Species rank sum 697.5 539.5 362.5 105.0 519.5 456.0 306.0 254.0 Group rank sum C; C, 1704.5 1535.5 time, and secondary plant compounds, as well as nutritional value, can be im- portant in determining foraging behavior (Eloff 1983; Owen-Smith et al. 1983; Garnett et al. 1985; Jonsdottir-Vivaas and Saether 1987; Provenza et al. 1987; Spalinger et al. 1988). Foraging behaviors of specific species appear in some instances to have evolved primarily in response to only a single factor. Presumably, some environments present conditions in which a single factor is of dispropor- tionate selective importance. Innate behavior resulting from selection in these instances can result in non-optimal foraging in alternative environments (Horn et al. 1986; Provenza et al. 1987). Other herbivore species seem to respond to all of these factors, and to generally forage as predicted by optimal foraging theory REGWQ MEAN FOOD FOOD CARBON GROUPING RANK TYPE SPECIES PATHWAY 69.75 seed Arachis hypoqgea C3 E 53.95 seed Helianthus annus C3 51.95 seed Setarta stalica C4 45.60 seed Salsola kal: C4 36.25 root Daucus carota C3 30.60 seed Zea mays C4 i" 25.40 seed Eragostis curvula C4 10.50 leaf Lactuca sativa C3 Fig. 1. REGWQ multiple-stage pairwise comparison test. Maximum experimentwise error rate under any complete or partial null hypothesis is controlled (a = 0.05), but not comparisonwise error rate. Neither C, forage in general nor only C, seeds are separated from C, forage (seeds). 104 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES LSD MEAN FOOD FOOD CARBON GROUPING RANK TYPE SPECIES PATHWAY F 69.75 seed Arachis hypogea C3 53.95 seed Helianthus annus C3 51.95 seed Setarta staltca C4 E 45.60 seed Salsola kal: C4 36.25 root Daucus carota C3 30.60 seed Zea mays CG, 25.40 seed Eragostis curvula Or - 10.50 leaf Lactuca sativa C3 Fig. 2. Fisher’s LSD multiple pairwise comparison test. Comparisonwise error rate (a = 0.05), but not experimentwise error rate is controlled. Neither C, forage in general nor only C, seeds are separated from C, forage (seeds). (Belovsky 1984a, b). Non-optimal foraging of some herbivores regardless of the environment has also been observed, and appears to be explainable by overriding selective forces such as competition, social learning, and predation risk avoidance behavior (Belovsky 1981; Meserve 1981; Grubb and Greenwald 1982; Holmes 1984; Carey and Moore 1986; Peacock and Jenkins 1988). Prairie dog diet in natural populations, determined by gut contents, has been reported to be extremely diversified both in plant taxa and parts of plants eaten. However, the authors, by limiting analysis to the grass portion of the diet, con- cluded that prairie dogs did not avoid C, plants (Fagerstone and Williams 1982). Rather, the data subset showed an actual preference for some C, plants. Two C, grass species, in particular, Bouteloua gracillis and Aristida longiseta, were found to predominate in gut contents. However, prairie dogs tended to switch from C, to C, plants seasonally. C. /udovicianus showed a preference for C; plants in spring. During the summer, fall, and winter months C, plants were preferred (Fagerstone et al. 1981; Fagerstone and Williams 1982). The parts of a plant eaten also varied Table 4. Food composition: percent wet mass/percent dry mass. Species are presented in the order they appear in Table 3. Carbo- Species H,0 Ash Protein Lipid hydrate C; A. hypogea (seed) 5.3/5.6 2.3/2.4 30.5/32.2 47.7/50.4 14.2/15.0 H. annus (seed) 4.5/4.7 3.8/4.0 27.2/28.3 41.4/43.1 23.5/24.6 D. carota (root) 88.3/754.7 1.2/10.3 1.2/10.3 0.2/1.7 9.1/77.7 L. sativa** (leaf) 95.7/2175.0 0.5/11.4 1.0/22.7 0.2/4.5 2.7/61.4 C, S. italica (seed) 10.2/11.3 5.6/6.2 10.7/11.9 4.7/5.2 68.8/76.6 S. kali* (seed) 3.8/4.0 5.8/6.0 20.6/21.4 36.0/37.4 33.8/35.1 Z. mays (seed) 9.3/10.3 1.8/2.0 11.5/12.7 7.9/8.7 69.5/76.6 E. curvula* (seed) 7.7/8.3 3.4/3.7 19.7/21.3 4.9/5.3 64.3/69.7 From: Henry and Morrison (1922, 1959). * Laboratory analysis by authors (N = 5 samples). ** USDA Handbook 8-11, 1980. PRAIRIE DOG FOOD PREFERENCE 105 seasonally (Fagerstone et al. 1981). A large percentage of the prairie dog’s diet was leaves from grasses in spring, the proportion of seeds in the diet increased throughout summer, and roots were a large portion of their diet in winter. Changes in gut contents observed could reflect differences in availability rather than chang- ing tastes or nutritional needs, although the relationship of prairie dog diet to available forage has been reported to be weak (Uresk 1984). Social influences involved in risk avoidance behavior is another reported factor that could lead to the changing diets recorded in prairie dog populations (Devenport 1989). Prairie dogs can unquestionably differentiate between plant food species, and preferentially select certain species as food both in field and experimental situa- tions. The results of the present study, however, suggest that earlier field study results, depicting a trend to feed on C, plants, may have resulted from changing availability, social learning, and/or risk avoidance behavior, rather than preference by the black-tail prairie dog. According to the photosynthetic pathway-selective herbivory hypothesis, prairie dogs should forage selectively on C, plants. Ac- cording to the field studies cited above, they prefer C, plants, at least seasonally. According to the present study, food preferences are unrelated to carbon cycle. Acknowledgments Special thanks to Dr. Jeffrey H. Black for presenting our research proposal for consideration to the Tulsa Zoological Park, and for experimental design sugges- tions. We are grateful to the Tulsa Zoo Directors, Curators, and Staff for allowing, and cooperating with, our research. Finally, we thank Drs. Steven Goldsmith and Patrick H. Wells for critically reading this manuscript, and Dr. Peyton Cook for advice on statistical analyses. Literature Cited Barnes, O. L. 1955. Effects of food plants on the lesser migratory grasshopper. J. Econ. Entomol., 48:119-124. Belovsky, G. E. 1981. Food plant selection by a generalist herbivore: the moose. Ecology, 62:1020— 1030. . 1984a. Summer diet optimization by beaver. Am. Midl. Nat., 111:209-222. . 1984b. Herbivore optimal foraging. A comparative test of three models. Am. Nat., 124:97- 115. . 1986. Generalist herbivore foraging and its role in competitive interactions. Am. Zool., 26: 51-69. Carey, H. V., and P. Moore. 1986. Foraging and predation risk in yellow-bellied marmots. Amer. Midland. Nat., 116:267—-275. Caswell, H., F. C. Reed, S. N. Stephenson, and P. A. Werner. 1973. Photosynthetic pathways and selective herbivory: a hypothesis. Amer. Nat., 107:465—480. Cody, M. L. 1974. Optimization in ecology. Science 85:1156—-1164. Coppock, D. L., J. K. Detling, J. E. Ellis, and M. I. Dyer. 1983. Plant-herbivore interactions in a North American mixed-grass prairie: 1. Effects of black-tailed prairie dogs (Cynomys ludovi- cianus) on intraseasonal aboveground plant biomass and nutrient dynamics and plant species diversity. Oecologia, 56:19. Coupe, T. R., and J. T. Schultz. 1968. The influence of controlled environments and grass hosts on the life cycle of Endria inimica (Homoptera: Cicadellidae). Ann. Entomol. Soc. Amer., 61:74— 77. Devenport, J. A. 1989. Social influences on foraging in black-tailed prairie dogs. J. Mammal., 70: 166-168. Delvi, M. R., and T. J. Pandian. 1971. Ecophysiological studies on the utilization of food in the paddy field grasshopper Osya velox. Oecologia, 8:267-275. 106 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Einot, I.,and K. R. Gabrial. 1975. A study of the powers of several methods of multiple comparisons. J. Amer. Stat. Ass., 70:351. Eloff, A. K. 1983. Fiber digestion in the hyrax. S. Afr. J. Anim. Sci., 13:28-30. Evans, G. C., and E. W. Tisdale. 1972. Ecological characteristics of Aristida longiseta and Agropyron spicatum in west-central Idaho. Ecology, 53:137—142. Fagerstone, K. A., and O. Williams. 1982. Use of C, and C, plants by black-tailed prairie dogs. J. Mamm., 63:328-331. , H. P. Tietjen, and O. Williams. 1981. Seasonal variation in the diet of black-tailed prairie dogs. J. Mamm., 62:820-824. Flinders, J. T., and R. N. Hansen. 1972. Diets and feeding habits of jackrabbits within a short grass ecosystem. IBP Grassland Biome. Colo. State Univ. Range Sci. Dept. Sci. Ser. No. 12. Garnett, S. T., I. R. Price, and F. J. Scott. 1985. The diet of the green turtle, Chelonia mydas (L.), in the Torres Strait. Aust. Wildl. Res., 12:103-112. Giaquinta, R. T. 1980. Transport of sucrose and oligosaccharides. Jn Biochemistry of plants, Vol. 3. (J. Press, ed.), Academnic Press, N.Y. Gross-Custard, J. D. 1977. Predator responses and prey mortality in the redshank Tringa totanus (L.) and a preferred prey Corophium volutator (Pallas). J. Anim. Ecol., 46:21-36. Grubb, T. C., and L. Greenwald. 1982. Sparrows and a brushpile: foraging responses to different combinations of predation risk and energy cost. Anim. Behav., 30:637-640. Hansen, R. M., and D. N. Ueckert. 1970. Dietary similarity of some primary consumers. Ecology, 51:640-648. Henry, W. A., and F. B. Morrison. 1922. Feed and feeding, 18th ed. Henry Morrison and Company, Madison. , and 1959. Feed and feeding, 22nd Ed. Henry Morrison and Company, Madison. Holmes, W. G. 1984. Predation risk and foraging behavior of the hoary marmont in Alaska. Behav. Ecol. Sociobiol., 15:293-301. Hoogland, J. L. 1981. The evolution of coloniality in white-tailed and black-tailed prairie dogs (Sciuridae: Cynomys leucurus and C. ludovicianus). Ecology, 62(1):252—272. Horn, M. H., M. A. Neighbors, and S. N. Murray. 1986. Herbivore responses to a seasonally fluctuating food supply: growth potential of two temperate intertidal fishes based on the protein and energy assimilation from their macroalgal diets. J. Exp. Mar. Biol. Ecol., 103:217—239. Jantz,O.K. 1962. Food plants of Melanoplus femurrubrum femurrubrum (DeGeer) in the bluestem grass region of Kansas. Unpubl. M.S. Thesis. Kansas State Univ., Manhattan. Jonsdottir-Vivaas, H., and B. E. Saether. 1987. Interactions between a generalist herbivore, the moose Alces alces, and its food resources: an experimental study of winter foraging behavior in relation to browse availability. J. Anim. Ecol., 56:509-520. Krebs, J.R. 1978. Optimal foraging: decision rules for predators. Jn Behavioral ecology. (J. R. Krebs and N. B. Davies, eds.), Sinauer Associates, Sunderland, Mass. Leach, C. 1979. Introduction to statistics. John Wiley & Sons, N.Y. Meserve, P. L. 1981. Resource partitioning in a Chilean semi-arid small mammal community. J. Anim. Ecol., 50:745-757. Miller, R. G. 1981. Simultaneous statistical inference. Springer-Verlag, N.Y. Minson, D. J. 1971. Influence of lignin and silicon on a summative system for assessing the organic matter digestibility of Panicum. Australian J. Agr. Res., 22:589-598. Moss, R., G. R. Miller, and S. E. Allen. 1972. Selection of heather by captive red grouse in relation to age of the plant. J. Appl. Ecol., 9:771-781. Mulkern, G. B., J. F. Anderson, and M. A. Brusven. 1962. Biology and ecology of North Dakota grasshoppers. I. Food habits and preferences of grasshoppers associated with alfalfa fields. North Dakota Agr. Exp. Sta. Res. Rep. No. 7. ———., K. P. Pruess, H. Knutson, A. F. Hagen, J. B. Campbell, and J. D. Lambley. 1969. Food habits and preferences of grassland grasshoppers of the north central great plains. North Dakota Agr. Exp. Sta. Bull. No. 481. National Research Council. 1958. Composition of cereal grain and forages. Washington, D.C. Owen-Smith, N., S. M. Cooper, and P. A. Novellie. 1983. Aspects of the feeding ecology of a browsing ruminant: the kudu. S. Afr. J. Anim. Sci., 13:35-38. Pate, J.S. 1984. The carbon and nitrogen nutrition of fruit and seed—case studies of selected grain legumes. Jn Seed physiology. (D. R. Murray, ed.), Academic Press, N.Y. PRAIRIE DOG FOOD PREFERENCE 107 Peacock, M. M., and S. H. Jenkins. 1988. Development of food preferences: social learning by Belding’s ground squirrels Spermophilis beldingi. Behav. Ecol. Sociobiol., 22:393-399. Pearcy, R. W., and O. Pjorkum. 1983. Physiological effects. In CO, and plants. (E. R. Lemon, ed.), AAAS Selected Symposium 84. Westview Press, Inc., Boulder. Pickford, R. 1962. Development, survival, and reproduction of Melanoplus bilituratus reared on various food plants. Can. Entomol., 94:859-869. 1963. Wheat crops and native prairie in relation to the nutritional ecology of Camnula pellucida (Scudder) (Orthoptera: Acrididae) in Saskatchewan. Can. Entomol., 95:767-—770. Pruess, K. P. 1969. Food preference as a factor in the distribution and abundance of Phoetaliotes nebrascensis. Ann. Entomol. Soc. Amer., 62:323-327. Provenza, F. D., J. T. Flinders, and E. D. McArthur. 1987. Domestic herbivore foraging tactics and landscape pattern. Gen. Tech. Rep., Internt. Res. Stn., 137-140. Pulliam, H. R. 1975. Diet optimization and nutrient constraints. Amer. Natur., 109:765-768. Putnam, L. G. 1962. Experiments with some native and introduced plants as foods for Camnula pellucida (Scudder) (Orthoptera: Acrididae) in western Canada. Can. J. Plant Sci., 42:589-595. Pyke, G. H., H. R. Pulliam, and E. L. Charnov. 1977. Optimal foraging: a selective review of theory and tests. Quart. Rev. Biol., 52:137-154. Rozin, P. 1976. The selection of foods by rats, humans and other animals. Jn Advances in the study of behavior, Vol. 6. (R. A. Hinde, C. Beer, and E. Shaw, eds.), Academic Press, N.Y. Ryan, T. A. 1959. Multiple comparisons in psychological research. Psychol. Bull., 56:26—-47. . 1960. Significance test for multiple comparison of proportions, variances, and other statistics. Psychol. Bull., 57:318—-328. SAS/STAT. 1988. Guide for personal computers version 6 edition. SAS Institute, Inc., Cary, N.C. Scharff, D. K. 1954. The role of food plants and weather in the ecology of Melanoplus mexicanus. J. Econ. Entomol., 47:485—489. Schoener, T. W. 1971. Theory of feeding strategies. Ann. Rev. Ecol. Syst., 2:369-404. Shade, R. G., and M. C. Wilson. 1967. Leaf-vein spacing as a factor affecting larval feeding behavior of the cereal leaf beetle Oulema melanopa (Coleoptera: Chrysomelidae). Ann. Entomol. Soc. Amer., 60:494—496. Siegel, S. 1956. Nonparametric statistics for the behavioral sciences. McGraw-Hill Book Company, Inc., N.Y. Smith, D. S., R. H. Hanford, and W. Chefurka. 1952. Some effects of various food plants on Melanoplus mexicanus mexicanus (Sauss.) (Orthoptera: Acrididae). Can. Entomol., 84:113- 117. Spalinger, D. E., T. A. Hanley, and C. T. Robbins. 1988. Analysis of the functional response in foraging in the black-tail deer. Ecology, 69:1166-1175. Thompson, D. Q. 1965. Food preferences of the meadow vole (Microtus pennsylvanicus) in relation to habitat affinities. Amer. Mid. Nat., 74:76-86. Thorne, J. H. 1985. Phloem unloading of C and N assimilates in developing seeds. 7n Annual review of plant physiology 36. (W. R. Briggs, ed.), Annual Reviews, Inc., Palo Alto. Tolbert, N. E., and I. Zelitch. 1983. Carbon metabolism. Jn CO, and plants. (E. R. Lemon, ed.), AAAS Selected Symposium 84. Westview Press, Inc., Boulder. Ueckert, D. N., and R. M. Hansen. 1971. Dietary overlap of grasshoppers on sandhill rangeland in northeastern Colorado. Oecologia, 8:276-295. Uresk, D. W. 1984. Black-tailed prairie dog food habits and forage relationships in western South Dakota. J. Range Manage., 37:325-329. USDA Handbook 8-11. 1980. Composition of foods. United States Department of Agriculture, Agriculture Handbook 8-11, Washington, D.C. Wells, H., and P. H. Wells. 1983. Honey bee foraging ecology: optimal diet, minimal uncertainty, or individual constancy. J. Anim. Ecol., 52:829-836. , and 1986. Optimal diet, minimal uncertainty and individual constancy foraging of honey bees, Apis mellifera. J. Anim. Ecol., 55:881-891. Wells, P. H. and H. Wells. 1984. Can honey bees change foraging patterns? Ecol. Entomol., 9:367— 473. Welsch, R. E. 1977. Stepwise multiple comparison procedures. J. Amer. Stat. Ass., 72:359. Wenner, A. M. 1971. The bee language controversy: an experience in science. Educational Programs Improvement Corporation, Boulder, Colo. 108 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Wilbur, D. W. 1954. Host plants of the leafhopper Endria inimica (Say) (Homoptera: Cicadellidae). Kansas Acad. Sci. Trans., 57:139-146. Williams, S. 1984. Official methods of analysis of the association of Official Analytical Chemists, 14 ed. The Association of Analytical Chemists, Arlington. Wilson, J. R., and C. W. Ford. 1971. Temperature influences on the growth, digestibility, and carbohydrate composition of two tropical grasses Panicum maximum var. trichoglume and Setaria sphacelata, and two cultivars of the temperate grass Lolium perenne. Australian J. Agr. Res., 22:563-571. , and K. P. Haydock. 1971. The comparative response of tropical and temperate grasses to varying levels of nitrogen and phosphorous nutrition. Australian J. Agr. Res., 22:573-587. Accepted for publication 29 November 1989. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 109-114 © Southern California Academy of Sciences, 1990 Revision of Two Dorvilleid Species From the Pacific Coast of North America (Annelida: Polychaeta) James A. Blake and Brigitte Hilbig Science Applications International Corporation, 89 Water Street, Woods Hole, Massachusetts 02543 Abstract.—Two species of Protodorvillea described from California and Wash- ington have been reexamined and determined to belong to the genus Pettiboneia. Structures originally described as branchiae have been determined to bear internal aciculae and are actually dorsal cirri. The numerous rows of accessory denticles originally noted as specific characters are generic level characters in Pettiboneia. The two species, P. pugettensis and P. dibranchiata, are redescribed and compared with related species of Pettiboneia. In 1978, Armstrong and Jumars described two new species of Protodorvillea from the Santa Catalina Basin (California) and Puget Sound (Washington): P. dibranchiata and P. pugettensis. In the following year, Blake (1979) reported on the discovery of Pettiboneia sanmatiensis Orensanz, 1973 in Tomales Bay, Cal- ifornia; this genus and species was heretofore known only from Argentina. Upon examination of the two papers it became apparent to us that both of Armstrong and Jumars’ Protodorvillea species were very similar to Pettiboneia in the com- position of the jaw apparatus. Pettiboneia is an unusual genus in having maxillae with numerous rows of denticles and no basal plates. We recently reexamined the type material of Armstrong and Jumars and have confirmed that both species do indeed belong to the genus Pettiboneia. In order to clarify the position of these two species, we present complete redescriptions and comparisons with related species that have been described in subsequent years (Campoy and San Martin 1980; Westheide and von Nordheim 1985; Wolf 1987; Hilbig and Ruff 1990). A key to all known species of Pettiboneia is presented in a companion paper (Hilbig and Ruff 1990). Type specimens examined as part of this study are now deposited in the Los Angeles County Museum of Natural History and include the former type desig- nations of the Allan Hancock Foundation (AHF). Two non-type specimens are deposited in the Royal British Columbia Museum (RBCM). Pettiboneia pugettensis (Armstrong & Jumars, 1978), new comb. Figure 1 Protodorvillea pugettensis Armstrong & Jumars, 1978:133-135, fig. 1. Material examined. — Puget Sound, Washington, intertidal, holotype (AHF Poly 1259) and 11 paratypes (AHF Poly 1260-63).—British Columbia, Chemainus Bay, subtidal, 11-12 m, sand-gravel, 30 September 1974, coll. K. D. Hobson, 2 specimens (BCPM). 109 110 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 1. Pettiboneia pugettensis (Paratypes, AHF Poly 1261-3): A, anterior end, dorsal view; B, parapodium, setiger 8 (arrow denotes notoacicula); C, uppermost compound falciger; D, lowermost compound falciger; E, furcate seta; F, capillary seta; G, maxillae; H, mandibles. A-G from paratypes, H from Anderson and Jumars (1978). Description. —Holotype 4.6 mm long, 0.3 mm wide for 48 setigers. Prostomium bluntly triangular, wider than long, recessed into achaetous peristomial ring; one pair faint eyes located at bases of antennae; antennae small, simple, clavate; palps well-developed, about twice as long as antennae, biarticulate (Fig. 1A). Peristomi- um with two achaetous rings, fused dorsally. Parapodia uniramous in setigers 1—2, subbiramous in setigers 3—10(—-14), uni- ramous in remaining setigers. Notopodia elongate, glandular, with fine notoacicula (Fig. 1B); some notopodia with very distinct vessel loops; neuropodia long, pointed apically and longer than notopodia when fully extended, truncate and shorter than REDESCRIPTION OF DORVILLEID POLYCHAETES 111 neuropodia when retracted. Ventral cirri short, inconspicuous (Fig. 1B), absent in posterior parapodia. Supraacicular fascicle with 1—2 subdistally serrated capillaries (Fig. 1F) and 1—- 2 furcate setae with slender, tapering, subequal tines and 3 rows of fine serrations at base of shorter tine (Fig. 1E). Subacicular fascicle with 2-3 compound falcigers (Fig. 1C, D); blade of uppermost seta elongate, tapering to hook-shaped tip, with very fine, long serrations along edge; often appearing spinigerous (Fig. 1C). Blades of lower compounds smooth, much shorter and wider (Fig. 1D); shafts of all compound setae with 1—2 subdistal rows of oblique serrations. Inferiormost com- pound seta emerging from tip of retractable setal lobe (Fig. 1B); in far posterior setigers occasionally accompanied by very fine, smooth capillary. Pygidium rounded, bearing 4 short clavate anal cirri. Maxillae composed of two main rows of free denticles on each side (Fig. 1G) and additional rows in apical part of pharynx; maxillary carriers and basal plates absent. Basal teeth of main rows elongate plates with main fang and coarsely serrated cutting edge; apical teeth of main rows rounded plates covered with surficial spines, with finely serrated cutting edge. Denticles of additional rows large, delicate, rounded plates with surficial spines. Mandibles curved, anteriorly flared, with scalloped cutting edge; posterior handle short and narrow (Fig. 1H). Remarks. — The original description of P. pugettensis contains some misinter- pretations of morphological characters. The structure addressed as “‘branchia”’ is clearly identifiable as a notopodium due to the presence of an acicula. The no- topodia may, however, function as gills because large vessel loops are usually present. The “‘cirrus-like protuberance”’ at the ventral neuropodial edge is not a consistent structure and is probably an artifact of preservation. The maxillae do not include maxillary carriers and basal plates or any fusion product; the four main rows are free from one another and contain only free denticles. The basal structure depicted in Armstrong and Jumars (1978: Fig. 11) is most likely a ligament-like structure connecting the jaws to the pharyngeal muscles. Pettiboneia pugettensis is readily distinguished from its congeners by the oc- currence of notopodia from setiger 3 rather than 2. The species is most closely related to P. sanmatiensis Orensanz from Argentina and California as redescribed by Blake (1979). Two specimens from British Columbia identified as P. sanma- tiensis by Blake (1979) have been reexamined. Both have the notopodia beginning on setiger 3 rather than setiger 2 and are therefore referred to P. pugettensis. In P. pugettensis the peristomial rings are fused dorsally instead of being separate, and the blades of the short-bladed compound setae are smooth instead of serrated. Pettiboneia dibranchiata (Armstrong & Jumars, 1978), new comb. Figure 2 Protodorvillea dibranchiata Armstrong & Jumars, 1978:135—137, fig. 2. Material examined. —Santa Catalina Basin, California, bathyal: holotype (AHF Poly 1264) and paratype (AHF Poly 1265). Description. —Holotype 5.8 mm long, 0.4 mm wide for 47 setigers. Prostomium bluntly triangular, wider than long, with two clavate to indistinctly biarticulate antennae and two biarticulate palps (Fig. 2A); antennae as long as greatest prosto- 112 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES a a cf, a *. if weaee l, i h 1, Sa My, =, u € aN x “5/ Cs gps ay Fig. 2. Pettiboneia dibranchiata (Paratype, AHF Poly 1265): A, anterior end, dorsal view; B, parapodium from anterior setiger (arrow denotes notoacicula); C, bifid tip of compound falciger; D, furcate seta; E, maxillae; F, mandibles. mial width, palps slightly longer; eyes absent. Peristomium with two rings, the first one much shorter and narrower than the second (Fig. 2A). Parapodia uniramous in setiger 1, subbiramous from setiger 2 through 13-17 and uniramous in remaining setigers. Notopodia very long, extending well beyond neuropodia, with distinct vessel loops and heavy ventral ciliation. Notopodial acicula very thin, visible only at notopodial base (Fig. 2B). Neuropodia blunt or REDESCRIPTION OF DORVILLEID POLYCHAETES 113 apically pointed if fully extended; ventral cirri short, digitiform, present from setiger 2 (Fig. 2B). Dorsal branchiae present from setiger 1 through setiger 13-— 17, decreasing on subsequent parapodia to an inconspicuous protuberance of the dorsal neuropodial margin. Supraacicular setae comprising 1-2 finely serrated capillaries with long, hairlike tips and | furcate seta with slender, unequal tines; short tine with 3 rows of basal serrations and narrow wing along inner side (Fig. 2D). Subacicular fascicle con- sisting of 3-4 compound setae with bifid, finely serrated blades (Fig. 2C); blade longest on seta closest to acicula; inferiormost seta emerging from tip of setal lobe when parapodium fully extended. Pygidium rounded, with two clavate dorsal anal cirri and two shorter ventral ones. Maxillae consisting of two main rows of denticles and approximately 12 ad- ditional rows on each side (Fig. 2E); maxillary carriers and basal plates absent. Denticles of row I large, delicate, rounded plates covered with surficial spines; proximal denticles of row II small, rectangular plates with main fang and coarsely serrated cutting edge; distal denticles of row II rounded, with surficial spines. Denticles of additional rows rounded, with surficial spines. Mandibles weakly sclerotized, with elongate, scoop-shaped, scalloped cutting edge and short, narrow handle (Fig. 2F). Remarks.—Some misinterpretations in the original description by Armstrong and Jumars (1978) are corrected in this redescription. The structure originally named “proximal branchia”’ contains a fine acicula and can therefore be identified as a notopodium. It may function as an additional gill, however, because of its very distinct vessel loop and ciliation. The “distal branchia”’ is a true gill. Wolf (1987) described a similar arrangement for P. duofurca; he found that the gills of P. duofurca are not present in all specimens. This might prove to be true for P. dibranchiata if additional material were available for examination. None of the Protodorvillea-like structures of the maxillary apparatus, such as fused maxillary carriers and basal plates, are present in the jaws of the paratype; instead, the main denticle rows are free from each other and consist only of free denticles. The two branchiate Pettiboneia species differ most obviously in the following characters: (1) notopodia are present from setiger 2 through 13-17 in P. dibran- chiata and from setiger 2 through 8-9 in P. duofurca; (2) gills are present from setiger 1 in P. dibranchiata and from setiger 3 in P. duofurca; (3) the furcate setae have unequal tines in P. dibranchiata and subequal tines in P. duofurca (adults); (4) the dorsal anal cirri are slightly longer than the ventral ones in P. dibranchiata and about 6—7 times longer in P. duofurca; (5) the mandibles are scoop-shaped in P. dibranchiata and flat, slightly curved in P. duofurca. The number of additional denticle rows seems to differ greatly between the two species; however, it should not be used as a diagnostic specific character because the apparent number of denticle rows may differ substantially even among specimens of the same species due to differences in the way the pharyngeal wall is spread on the slide (see Hilbig and Ruff 1990). Acknowledgments The original loan of the types of these species was by Dr. Jerry D. Kudenov, formerly of the Allan Hancock Foundation. A recent re-examination of the types 114 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES was based on a loan provided by Ms. Leslie Harris of the Los Angeles County Museum of Natural History. We are grateful to Dr. J. F. Grassle and the Biology Department of the Woods Hole Oceanographic Institution for providing working space and facilities in 1989. Literature Cited Armstrong, J. W., and P. A. Jumars. 1978. Branchiate Dorvilleidae (Polychaeta) from the North Pacific. Bull. So. Calif. Acad. Sci., 77:133-138. Blake, J. A. 1979. A redescription of Pettiboneia sanmatiensis Orensanz (Polychaeta: Dorvilleidae) and a revised key of the Dorvilleidae. Bull. So. Calif. Acad. Sci., 78:136—140. Campoy, A., and G. San Martin. 1980. Pettiboneia urciensis sp. n.: un Dorvilleidae (Polychétes: Errantes) de la Méditerranée. Cah. Biol. Mar., 21:201-207. Hilbig, B., and R. E. Ruff. 1990. Remarks on the genus Pettiboneia (Polychaeta: Dorvilleidae) with descriptions of two new species. Bull. So. Calif. Acad. Sci., 89:115-123. Orensanz, J. M. 1973. Los Annelidos Poliquetos de la Provincia Biogeografica Argentina. III. Dor- villeidae. Physis, Sec. A, 32(85):325-342. Westheide, W., and H. von Nordheim. 1985. Interstitial Dorvilleidae (Annelida, Polychaeta) from Europe, Australia and New Zealand. Zool. Scr., 14:183-199. Wolf, P.S. 1987. Two new species of Pettiboneia (Polychaeta: Dorvilleidae) primarily from the Gulf of Mexico. Proc. Biol. Soc. Wash., 100:28-34. Accepted for publication 30 May 1990. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 115-123 © Southern California Academy of Sciences, 1990 Remarks on the Genus Pettiboneia (Polychaeta: Dorvilleidae) with Descriptions of Two New Species Brigitte Hilbig! and R. Eugene Ruff 1 Science Applications International Corporation, 89 Water Street, Woods Hole, Massachusetts 02543 2 Ruff Systematics, 41 Chapel Street, Suite 101, Needham, Massachusetts 02192 Abstract. —Two new species of the genus Pettiboneia Orensanz, 1973, are described from the North Pacific and North Atlantic Oceans. Comments on within-species variability, generic relationships, and species distributions are presented, and a key to the species of Pettiboneia is provided. The genus Pettiboneia Orensanz, 1973, was originally described from seven specimens found in shallow water in the Gulf of San Matias, Argentina. Members of the genus are very small, typically averaging only about 5 mm in length, and are easily overlooked. However, within the last decade an additional four species have been discovered, all in intertidal to shelf depths, and in tropical or subtropical waters. Blake and Hilbig (this issue) are transferring two additional species orig- inally described as Protodorvillea to the genus. In this paper two new species from Alaska and from the western North Atlantic are described. These species are from deeper and more northern waters than previously reported, raising the number of species of Pettiboneia to nine. The Alaskan specimens were collected as part of a monitoring study conducted in two southeast Alaska fjords between 1979-1983. The Atlantic specimens were collected on the continental slope and rise between Cape Cod, Massachusetts, and Cape Lookout, North Carolina, between 1982-1986. The holotype and some paratypes are deposited in the U.S. National Museum of Natural History (USNM). Additional paratypes are deposited in the British Museum of Natural History (BMNH) and in the Zoological Museum of the Uni- versity of Hamburg (ZMH). Some specimens have been retained by the authors or have been transferred to the Smithsonian Institution. Pettiboneia brevipalpa n. sp. Figure | Pettiboneia sp.—VTN Consolidated, Inc., 1980. Material examined. —Boca de Quadra, Alaska: Sta. 200, 55°18.1'N, 130°30.6’W, 150 m, Apr 1980, 3 paratypes (USNM 130085); Jul 1980, 1 paratype (USNM 130086). Sta. 201, 55°18.3’N, 130°30.9’W, 145 m, Apr 1980, 1 paratype (USNM 130087). Sta. 277, 55°18.1'N, 130°30.6’W, 150 m, Sep 1983, 1 paratype (USNM 130088). Sta. 400, 55°16.7'N, 130°31.9’W, 150 m, Apr 1980, 3 specimens. Sta. 401, 55°16.7'N, 130°32.1’W, 140 m, Apr 1980, 1 paratype (USNM 130089); Jul 115 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 116 Fig. 1. Pettiboneia brevipalpa: A, anterior end, dorsal view; B, parapodium; C, capillary supraacicu- lar seta; D, furcate supraacicular seta; E, subacicular seta; F, maxillae; G, mandibles. 1980, 2 paratypes (ZMH P-20324). Sta. 500, 55°15.1’N, 130°33.1'W, 210 m, Apr 1980, 2 specimens; Jul 1980, 3 specimens. Sta. 501, 55°15.0’N, 130°32.8'W, 195 m, Jul 1980, 1 specimen. Sta. 502, 55°14.9'N, 130°32.6’W, 195 m, Jul 1980, 1 specimen. Sta. 600, 55°12.4’N, 130°35.8’W, 280 m, Jul 1980, 1 specimen. Sta. 601, 55°12.6’N, 130°36.1’'W, 275 m, Jul 1980, 3 specimens. Sta. 602, 55°12.3’N, REMARKS ON THE GENUS PETTIBONEIA 117 130°35.5’W, 275 m, Jul 1980, 3 paratypes (BMNH ZB 1990.29-31). Sta. 700, 55°10.2'N, 130°39.3'W, 280 m, Apr 1980, holotype (USNM 130084); Jul 1980, 5 paratypes (USNM 130090). Sta. 800, 55°05.9'N, 130°43.5'’W, 330 m, Apr 1980, 3 paratypes (USNM 130091). Sta. 801, 55°06.1’N, 130°43.9’'W, 330 m, Apr 1980, 1 specimen. Sta. 802, 55°05.7'N, 130°43.3'W, 380 m, Jul 1980, 1 paratype (ZMH P-20325).—Smeaton Bay, Alaska. Sta. 010, 55°18.7’'N, 130°41.4’W, 241 m, Oct 1980, 1 paratype (BMNH ZB 1990.32). Description. — Holotype complete, 2.4 mm long, 0.1 mm wide for 35 setigers. Other complete specimens up to 4.5 mm long for 66 setigers. Body slender, fragile, unpigmented in alcohol. Prostomium pear-shaped, about as long as wide (Fig. 1A); antennae clavate, about as long as greatest prostomial width; biarticulate palps half antennal length, inserted ventrolaterally behind antennae, with short, inconspicuous palpophores; eyes absent; narrow band of cilia encircling prosto- mium between antennae and palps; large yellowish-brown nuchal organs on pos- terior margins of prostomium. Two subequal asetigerous peristomial rings, shorter than setigerous segments; complete ciliary bands on both rings and on anterior setigers. Cirriform notopodia with embedded acicula present from setiger 2 through setigers 8-11; absent posteriorly; as long as or slightly longer than neuropodial acicular lobe. Neuropodia with conical acicular lobe and inferior retractable setal lobe supported by inferiormost seta (Fig. 1B); ventral cirri short, cirriform. Supraacicular setae include 1-3 serrated capillary setae (Fig. 1C) and 1 (occa- sionally 2) furcate seta with unequal tines and 1-2 rows of fine serrations below shorter tine; longer tine with delicate wing on inner side (Fig. 1D). Subacicular fascicle with 2-4 compound setae having long to short unidentate, finely serrated blades; shafts bifid with coarse subdistal serrations (Fig. 1E). Far posterior setigers occasionally with simple pointed inferior setae. Pygidium rounded, longer than preceding setigers, with 2 pairs of clavate sub- terminal cirri. Maxillae with two main rows of free denticles and about six additional rows on each side; maxillary carriers and basal plates absent (Fig. 1F). Basal teeth of main rows smooth, delicate, rounded plates with serrated cutting edge; middle and upper teeth rounded plates covered with surficial spines. Teeth of additional rows including smooth, anteriorly serrated plates proximally and spinose plates distally. Mandibles elongate, slightly curved, anteriorly flared, smooth and weakly incised (Fig. 1G). Remarks. — Pettiboneia brevipalpa is similar to P. sanmatiensis in the number of notopodia, the shape of the prostomium, the number of maxillary rows, and the size and shape of the pygidial cirri. It differs from the latter species in lacking eyes and in having long rather than short notopodia. In addition, P. brevipalpa differs from all other known species in possessing palps that are much shorter than the antennae, and palpophores shorter than the palpostyles. Of the 42 specimens examined, eight were gravid females and ten appeared to be developing or mature males. There are up to 8 round or elongated eggs per setiger averaging 40 um in length and 27 wm in width. The eggs are first present between setigers 14—21, and the sperm appear between setigers 15-19. The gametes are associated with the bases of the parapodia and continue for most of the length of the worm. 118 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Etymology. —The specific name refers to the unique short palps which distin- guish this species from its congeners. Distribution.—Known only from Boca de Quadra and Smeaton Bay in the Alaskan panhandle, 140-380 m, in fine silts. This is the northernmost record for the genus. Pettiboneia bathyalis n. sp. Figure 2 Pettiboneia sp. 1.— Blake et al. 1987; Maciolek et al. 1987a, b. Material examined.—U.S. Atlantic Continental Slope and Rise Program, off New England, Sta. N2, 40°57.2'N, 66°13.9’W, 2100 m, Nov 1984, 1 specimen; Apr 1986, 1 specimen. Sta. N8, 40°10.3'’N, 67°37.4'W, 2180 m, Nov 1984, 1 paratype (USNM 130093); Apr 1986, 1 paratype (ZMH P-20326). Sta. N13, 39°48.4'N, 70°54.3’W, 1250 m, Nov 1985, 1 specimen. Sta. N14, 39°41.0'N, 70°54.3'W, 2105 m, Nov 1984, 1 specimen.— Off Delaware and New Jersey: Sta. M1, 38°36.0’N, 72°53.0'W, 2195 m, Aug 1984, 1 specimen; Dec 1984, 2 paratypes (USNM 130094); Aug 1985, 1 paratype (USNM 130095). Sta. M2, 38°35.8’N, 72°53.7'W, 2020 m, Aug 1984, 2 specimens; Aug 1985, 1 specimen; Nov 1985, holotype (USNM 130092), 2 paratypes (USNM 130096). Sta. M3, 38°36.8'N, 72°51.4'W, 2055 m, May 1984, 1 specimen; Dec 1984, 1 specimen. Sta. M4, 38°44.5'N, 72°33.0’W, 2100 m, May 1984, 1 specimen; Dec 1984, 1 specimen. Sta. M5, 38°50.5’N, 72°33.0’W, 2065 m, Nov 1985, 1 specimen. Sta. M6, 39°05.5'N, 72°03.0'W, 2090 m, Nov 1984, 1 specimen. Sta. M7, 38°27.4'N, 73°03.4'’W, 2100 m, Dec 1984, 1 specimen; Aug 1985, 2 paratypes (BMNH ZB 1990.33-34), 1 specimen. Sta. M9, 38°17.3'N, 73°14.5'W, 2105 m, May 1984, 4 paratypes (ZMH P-20327); Aug 1984, 2 specimens; Nov 1984, 5 paratypes (USNM 130097), 2 specimens; May 1985, 4 specimens; Aug 1985, 3 paratypes (USNM 130098); Nov 1985, 1 specimen. Sta. M10, 37°51.8’N, 73°19.8’W, 2095 m, May 1984, 1 spec- imen; Aug 1985, 1 specimen. Sta. M11, 38°40.2’N, 72°42.2’W, 1515 m, Aug 1984, 1 specimen; Dec 1984, 1 specimen. Sta. M12, 38°29.3'N, 72°42.2'W, 2505 m, Dec 1984, 1 specimen. Sta. M13, 37°53.3’N, 73°45.1'W, 1613 m, Nov 1984, 3 specimens; May 1985, 3 specimens; Nov 1985, 1 specimen. — Off North Carolina: Sta. S3, 34°14.8'N, 75°40.1’W, 1500 m, Mar 1984, 1 specimen. Sta. S4, 34°11.4’N, 75°38.8'W, 2000 m, May 1984, 1 specimen; Sep 1985, 2 paratypes (USNM 130099); Nov 1985, 6 paratypes (BMNH ZB 1990.35—40). Sta. S6, 34°49.5'N, 75°13.4'’W, 2004 m, May 1984, 5 specimens; July 1984, 3 specimens. Sta. S10, 35°26.3'N, 74°41.4'W, 2003 m, Nov 1985, 4 specimens. Sta. $12, 33°00.3’N, 76°07.4’W, 1996 m, Nov 1985, 1 paratype (USNM 130100). Sta. S14, 32°23.6'’N, 77°01.1'W, 805 m, Nov 1985, 1 specimen. Description. —Holotype complete, 5.5 mm long, 0.3 mm wide for 65 setigers. Other complete specimens to 6.0 mm long with up to 70 setigers. Body stout, dorsoventrally compressed, unpigmented in alcohol. Prostomium broadly rounded anteriorly, slightly wider than long, with two ciliary bands straddling antennae (Fig. 2A); antennae short, smooth, slightly cla- vate, half as long as prostomial width; palps longer, biarticulate, with elongate palpostyles; eyes absent. First peristomial ring short, about as wide as prostomium, often telescoped beneath the second, wider, longer ring; each ring with ciliary band close to posterior margin. REMARKS ON THE GENUS PETTIBONEIA 119 Fig. 2. Pettiboneia bathyalis: A, anterior end, dorsal view, ciliation not shown; B, parapodium; C, capillary supraacicular seta; D, geniculate supraacicular seta, setiger 1; E, geniculate supraacicular seta, setiger 3; F, furcate supraacicular seta, setiger 5; G, subacicular seta, middle position; H, subacicular seta, inferiormost position; I, maxillae; J, mandibles. Cirriform aciculate notopodia present from setiger 2 through setigers 7-9. Neu- ropodia with acicular lobe and retractable inferior setal lobe; filiform ventral cirri inserted subdistally, extending beyond acicular lobe (Fig. 2B). Supraacicular fascicles with 1-3 capillary setae (Fig. 2C) and 1-2 bifid, serrated geniculate setae in anteriormost parapodia (Fig. 2D); geniculate setae progressively 120 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES changing within anterior 5 setigers to furcate setae having unequal truncate tines (Fig. 2E, F); shaft coarsely serrated below shorter tine. Subacicular fascicles with 3-4 compound setae having finely serrated falcigerous blades; serrations longer than blade width; inferior blades shortest; shaft with a few coarse subdistal teeth (Fig. 2G, H). Posterior setigers occasionally with simple cultriform seta in ven- tralmost position. Pre-pygidial setigers short. Pygidium wider than long, with 2 pairs of clavate subterminal cirri; dorsal pair longest. Maxillae consisting of two main rows of free denticles and numerous additional denticles arranged in 6—8 rows on each side; maxillary carriers and basal plates absent (Fig. 21). Basal teeth of main rows smooth, rounded plates with finely serrated cutting edge; middle teeth smooth, rectangular, with proximal main fang and coarsely serrated cutting edge; anterior teeth oval plates with surficial spines, proximal main fang and coarsely serrated cutting edge; main fang lacking in last 1-4 denticles. Denticles of additional rows large, delicate, rectangular to oval plates with surficial spines and finely serrated cutting edge, arranged in several imbricated distal rows and one proximal row on each side. Mandibles L-shaped with short, sclerotized handle and long, delicate, scoop-shaped cutting edge with 10-12 rounded teeth in one row and 2-3 teeth in additional rows (Fig. 2J). Remarks. — Pettiboneia bathyalis appears to be close to P. urciensis in prostomial shape and number of notopodia. It differs from the latter species in lacking eyes and in having elongated palpostyles. P. bathyalis differs from its congeners in possessing geniculate rather than furcate setae in the anteriormost setigers. Of the 83 specimens examined, 8 were gravid females and 19 were males. The eggs are irregularly polygonal in shape, averaging | 16 wm in the longest dimension. There are about 8 eggs per segment after setiger 10 packed across the ventrum and extending into the parapodia. Sperm occur from setigers 13-20, continuing to near the end of the body. Etymology. —The species name refers to the bathyal regions where it was col- lected. Distribution. —P. bathyalis is distributed from Cape Cod, Massachusetts, to Cape Lookout, North Carolina, 800—2500 m, in muddy sands. This is the first deep-sea species of Pettiboneia discovered, and the first record for the western North Atlantic. Discussion The diagnostic characters delineating the genus Pettiboneia were provided by Orensanz (1973), and the structure of the jaw apparatus was elucidated by Blake (1979). The two new species conform well to the genus concept as reviewed by Wolf (1987) with the exception that in P. brevipalpa the biarticulate palps are distinctly shorter than the antennae. The generic diagnosis should therefore be emended to encompass this feature. Abundant material permitted a detailed assessment of within-species character variability. Many features, such as the shape of the prostomium, the length of the palps and antennae, and the length of the notopodia relative to the neuropodia, were found to be relatively constant within each species. The ciliary bands also appeared to be invariable, although these features were not always discernible. Examination of the jaw apparatus of at least ten specimens of each species REMARKS ON THE GENUS PETTIBONEIA 121 revealed very different appearances of the maxillary arrangement. The apparent number of maxillary rows varied between 8 and 14, and in most cases these rows were only discernible in the anterior part of the pharynx. The differences seemed to be a function of the maceration of the very thick pharyngeal muscle rather than reflecting a real morphological variability. The arrangement of maxillary plates as a diagnostic character on the species level should therefore be used with caution until the true morphology of the jaw apparatus can be revealed. Examination of dissected and dorsally opened pharynges with SEM may be an appropriate tech- nique. Anterior notopodia begin on setiger 2 and generally extend through setiger 11 in P. brevipalpa and through setiger 9 in P. bathyalis. However, in a number of specimens the posteriormost | to 3 notopodia were not observed (Table 1). Since these structures are supported by an internal acicula and do not tend to be de- ciduous, the variability in the total number of notopodia appears to be a real feature rather than an artifact. All of the neuropodia possess an inferior setal lobe supported by the ventralmost seta, a feature that has not been reported in the descriptions of other species within the genus. The length of this lobe was often variable in adjacent parapodia, and at times it was inconspicuous. This feature appears to reflect the degree of extension of the ventral seta, and the length is therefore an artifact of preservation. The inferior setal lobe is supported by a simple pointed seta rather than a compound falciger in the posterior setigers of about half of the examined speci- mens of P. brevipalpa and about a quarter of the P. bathyalis material. The setiger of its first occurrence varies greatly, and the seta is not always present in all consecutive setigers. Similar simple inferior setae have also been noted in other dorvilleid genera such as Ophryotrocha (Hilbig and Blake in press). The retractable setal lobe is also present in many of the species within that genus, although it has often been overlooked in the past (Hilbig and Blake in press). These two features appear to be widespread among dorvilleids and may represent plesiomorphic characters. The change from serrated bidentate geniculate setae to short-tined furcate setae noted in the anterior region of Pettiboneia bathyalis is also known from Dorvillea rudolphi and an undescribed Atlantic deep-sea species of Meiodorvillea reported in Blake et al. 1987, and Maciolek et al. 1987a, b, indicating a close relationship between these genera. The relationship between Pettiboneia and Meiodorvillea is further confirmed by the absence of maxillary carriers and basal plates and the presence of surficial spines on the anteriormost maxillary plates of Meiodorvillea muinuta. With recent discoveries, it appears that the genus Pettiboneia is widely distrib- uted throughout the world’s oceans (Fig. 3). A total of nine species are now known from tropical habitats to boreal waters, and from the shallow subtidal to depths exceeding 2500 m. Key to the known species of Pettiboneia NAMEBENES: PrESEN Oe tere HOUe tee BRE IGP AIAN eS ON Seah ooh he Nene Lule rat Rene ae I NDE VES AWSOME toc rg seers oy oc. Gass hls oes oth oun RaMOHOU ee Sask GREE 5 2a. Notopodia beginning on setiger 3; peristomial rings fused dorsally .... Bee Rete cose cog tog Ie ech ae P. pugettensis (Armstrong & Jumars, 1978) 122 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 2b. Notopodia beginning on setiger 2; peristomial rings distinct and not fused LORS ALLY yer ce arse ee ag AB ete eek a Mel rein ah AEE) ied beh i 3 3a. Notopodia shorter than neuropodia; furcate setae with subequal tines LMA Tere eet RS Dieter? Ci Rohe Lahn Su SR AN P. sanmatiensis Orensanz, 1973 3b. Notopodia as long or longer than neuropodia; furcate setae with unequal TCS Fee ave ates Bard © ee ee eae ee ed ethey aiaivert ely, ieee 4 4a. Prostomium pear-shaped; eyes large, situated behind antennae ....... PR ea iy tadiy yt iRth rh S52 oi ORM poe dR Ren ace sm ears PCN P. blakei Wolf, 1987 4b. Prostomium rounded; eyes small, situated in front of antennae ...... BEPC Ri doen eile iotaste is ei hapbin ean P. urciensis Campoy & San Martin, 1980 5a. Prostomium pear-shaped; palps shorter than antennae . P. brevipalpa n. sp. 5b. Prostomium rounded; palps longer than antennae ................. 6 6a. Notopodia longer than neuropodia; anterior neuropodia with superior branchiae: 2k Saver eee Pe ee 7 6b. Notopodia subequal to neuropodia; anterior neuropodia without supe- TOL branchiacyss was ie Ws RRR eee ed hye eed nyt a ee 8 7a. Branchiae beginning on setiger 1; notopodia extending through setigers 13-17; all anal cirri short ... P. dibranchiata (Armstrong & Jumars, 1978) 7b. Branchiae beginning on setiger 3; notopodia extending through setigers §—9:dorsalianal cirri long. 4a ee eee P. duofurca Wolf, 1987 8a. Notopodia extending through setiger 9; furcate setae in median setigers Serrated awathaunequal tineSus3) ype eras ee eee P. bathyalis n. sp. 8b. Notopodia extending through setiger 19; furcate setae in median setigers smooth: with:subequall tines! 9 3. 4e5.. ss ee eee as ch ortega, tie: expe P. australiensis Westheide & von Nordheim, 1985 P. brevipalpa P. sanmatiensis Ul P. dibranchiata P. sanmatiensis U Q Fig. 3. Worldwide distribution of species of Pettiboneia. REMARKS ON THE GENUS PETTIBONEIA 123 Table 1. Distribution of notopodia in the anterior setigers of P. bathyalis and P. brevipalpa. Number of specimens Last setiger bearing notopodia P. bathyalis P. brevipalpa 7 6 — 8 9 4 9 39 12 10 — 7 11 — 17 Acknowledgments We wish to thank Dr. James A. Blake for critically reviewing the manuscript. The material from Boca de Quadra and Smeaton Bay, Alaska, was collected by the benthic group at VIN Oregon, Inc., under contract to the United States Borax & Chemical Corporation. The Atlantic material was collected by Battelle Me- morial Institute under Contract No. 14-12-0001-30064 from the U.S. Department of the Interior, Minerals Management Service. Literature Cited Armstrong, J. W., and P. A. Jumars. 1978. Branchiate Dorvilleidae (Polychaeta) from the North Pacific. Bull. So. Calif. Acad. Sci., 77(3):133-138. Blake, J. A. 1979. A redescription of Pettiboneia sanmatiensis Orensanz (Polychaeta: Dorvilleidae) and a revised key to the Dorvilleidae. Bull. So. Calif. Acad. Sci., 78(2):136-140. , and B. Hilbig. 1990. Revision of two dorvilleid species from the Pacific coast of North America (Annelida: Polychaeta). Bull. So. Calif. Acad. Sci. (This issue). , B. Hecker, J. F. Grassle, B. Brown, M. Wade, P. D. Boehm, E. Baptiste, B. Hilbig, N. Maciolek, R. Petrecca, R. E. Ruff, V. Starczak, and L. Watling. 1987a. Study of biological processes on the U.S. South Atlantic slope and rise. Phase 2. Final Report prepared for the U.S. Department of the Interior, Minerals Management Service, Washington, D.C., under Contract No. 14-12- 0001-30064. 11 + 414 pp. and 13 appendices. NTIS PB87-214359. Campoy, A.,andG. San Martin. 1980. Pettiboneia urciensis sp. n.: un nouveau Dorvilleidae (Polyché- tes: Errantes) de la Méditerranée. Cah. Biol. Mar., 21:201—207. Hilbig, B., and J. A. Blake.. Accepted. Dorvilleidae (Annelida: Polychaeta) from the U.S. Atlantic slope and rise. Description of two new genera and 14 new species, with a generic revision of Ophryotrocha. Zoologica Scripta. Maciolek, N., J. F. Grassle, B. Hecker, P. D. Boehm, B. Brown, B. Dade, W. G. Steinhauer, E. Baptiste, R. E. Ruff, and R. Petrecca. 1987a. Study of biological processes on the U.S. Mid-Atlantic slope and rise. Final Report prepared for the U.S. Department of the Interior, Minerals Man- agement Service, Washington, D.C., under Contract No. 14-12-0001-30064. 1 + 314 pp. and 13 appendices. NTIS PB88 183090. , J. F. Grassle, B. Hecker, B. Brown, J. A. Blake, P. D. Boehm, R. Petrecca, S. Duffy, E. Baptiste, and R. E. Ruff. 1987b. Study of biological processes on the U.S. North Atlantic slope and rise. Final Report prepared for the U.S. Department of the Interior, Minerals Management Service, Washington, D.C., under Contract No. 14-12-0001-30064. 358 pp. and 12 appendices. NTIS PB88 196514/AS. Orensanz, J. M. 1973. Los annelidos poliquetos de la provincia biogeografica Argentina. III. Dor- villeidae. Physis, Sec. A, 32(85):325-342. VTN Consolidated, Inc. 1980. Boca de Quadra Baseline Report: Coastal and Marine Biology Pro- gram, Quartz Hill Molybdenum Project, Southeast Alaska. Report prepared for United States Borax & Chemical Corporation and Pacific Coast Molybdenum Company, December, 1980. Westheide, W., and H. von Nordheim. 1985. Interstitial Dorvilleidae (Annelida, Polychaeta) from Europe, Australia and New Zealand. Zool. Scr., 14(3):183-199. Wolf, P.S. 1987. Two new species of Pettiboneia (Polychaeta: Dorvilleidae) primarily from the Gulf of Mexico. Proc. Biol. Soc. Wash., 100(1):28-34. Accepted for publication 30 May 1990. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 124-129 © Southern California Academy of Sciences, 1990 A New Species of Marine Amphipod (Gammaridea: Ampeliscidae) from the Sublittoral of Southern California James D. Roney Biology Laboratory, Hyperion Treatment Plant, 12000 Vista del Mar, Playa del Rey, California 90293 Abstract.—A new species of ampeliscid amphipod, Ampelisca brachycladus, is described from southern California. The species is distinctive in having the inner ramus of uropod 1 approximately one-half the length of the outer ramus. Ampelisca brachycladus occurs along the southern California bight in shallow water (10-50 m) on a variety of substrates. The genus Ampelisca Kroyer 1842 is at present represented by 34 species on the continental shelf of the northeastern Pacific and 214 worldwide. This genus is exclusively marine and estuarine (Enequist 1949) and found on sublittoral sand and mud bottoms. Approximately 80% are sublittoral and 20% are bathyal to abyssal (Barnard 1969). The genus Ampelisca has been examined in many major studies along the eastern Pacific coast (Holmes 1908; Barnard 1954, 1960, 1967, 1971; and Dickinson 1982) and will not be reviewed in this paper. Only those diagnostic characters in combination that distinguish the new species from all known species of Ampelisca are illustrated. All illustrations and descrip- tions are based on type material. Ampelisca brachycladus n. sp. Figures 1-3 Description. —Female holotype, 7.3 mm. Head (Fig. 1) as long as first three pereonites combined; lower front margin of head oblique; oblique margin slightly concave; frontal margin of head not incised. Two corneal lenses on each side; lower cuticular lens positioned at lateral angle of head. Antenna | short, slender, not reaching end of peduncle of antenna 2; peduncular article 1 one-half length of article 2; article 3 short, flagellum subequal to peduncle. Antenna 2 of moderate length; article 3 short; article 4 longer than article 5; flagellum bearing sparse medium setae. Upper lip with median apical notch. Mandibular palp (Fig. 2A) 3- articulate; first article borne on medial elevated process bearing patch of minute setules, and armed with 2 lateral setae. (Note: stippled area on medial surface of first article of Fig. 2A indicating cuticular fold not intersegmental arthrodial mem- brane.) Left lacinia with 4—5 cusps; spine row with 8 large spines; molar toothed and ridged. Lower lip with mandibular lobe weak or lacking; inner lobe well developed. Maxilla 1 (Fig. 2B) outer lobe with 10 denticulared spines; palp with 4 strong apical spines. Maxilla 2 (Fig. 2C) inner plate setose. Maxillipedal palp (Fig. 2D) 4-articulate; outer lobe with 10 spines (medial group of spines spatulate and terminal ones long and slender); inner lobe setose and with a single apical spatulate spine. Pereonal segments dorsally smooth. Coxae 1-4 longer than broad; 124 NEW SPECIES OF GAMMARID AMPHIPOD 125 Fig. 1. Ampelisca brachycladus n. sp., female paratype, Santa Monica Bay, California. Scale: 1.0 mm. lower posterior corners untoothed. Coxa 1 expanded distally. Coxae 1-2 with a single row of plumose setae along entire ventral margin. Coxa 3 with sparse plumose setae on posterior ventral margin. Coxa 4 with sparse short setae along ventral margin; upper one-third strongly excavate posteriorly. Gnathopods | and 2 simple; both pereopods with pectinate setae. Gnathopod 1 moderately setose along posterior margin of articles 5 and 6, and setose on distal half of anterior margin of articles 5 and 6. Gnathopod 2 articles 5 and 6 moderately setose along posterior margin. Pereopods 3 and 4 similar to each other; pereopod 3 article 4 with setae only on anterior distal margin; pereopod 4 article 4 with setae along the entire anterior margin. Pereopods 5 and 6 similar (see Fig. 3A); article 6 with two spines on posterior margin, with setal tuft on distal end; article 5 with three sets of spines on posterior edge, with setal tuft at distal end; dactyl short, hooklike, and reversed. Pereopod 7 (Fig. 3B) lower lobe of article 2 reaching joint between articles 4 and 5; lower posterior edge of lobe oblique; article 4 Z << Zz —g —PLAZ —<— Fig. 2. Ampelisca brachycladus n. sp.: A, right mandible; B, first maxilla; C, second maxilla; D, maxilliped; symbols: LFT MD B, C; 0.2 mm for A, D. proximal article. Scales: 0.1 mm for left mandible; PROX ART NEW SPECIES OF GAMMARID AMPHIPOD 127 Fig. 3. Ampelisca brachycladus n. sp.: A, pereopod 6; B, pereopod 7; C, urosome; D, telson. Scales: 0.5 mm for A; 0.3 mm for B, C; 0.6 mm for D. 128 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES posterior lobe reaching more than halfway along hind margin of article 5; article 4 posterior lobe setose; article 5 distal anterior edge notched; article 6 longer than article 5; article 7 equal in length to article 5. All pleonal segments dorsally smooth. Epimera 2 and 3 posterodistal corners subquadrate. Urosomite 1 (Fig. 3C) dorsally a pointed hood. Urosomites 2 and 3 dorsally smooth. Uropod 1 reaching end of uropod 2; inner ramus one-half length of outer ramus; peduncle short and stout. Uropod 2 outer ramus lacking subapical spine. Uropod 3 rami lanceolate. Telson (Fig. 3D) deeply cleft, with terminal spine and seta on each lobe; each lobe with single spine halfway along median edge, and two spines inserted on outer margin 4th distance from proximal end. Relationship. —Ampelisca brachycladus n. sp. is most closely related to Am- pelisca agassizi (Judd 1896) (see Dickinson 1982) in having subquadrate epimera 2 and 3 and in having the urosome compressed lengthwise. However, they are distinguishable by article 5 of pereopod 7 of the new species being anterodistally notched; their telsons are also quite dissimilar. Ampelisca brachycladus is distin- guished from all known congeners by the short inner ramus of uropod 1. Material examined. — All specimens deposited in the Los Angeles County Mu- seum of Natural History. LACMNH No. 85-199.1, Aliso Beach, California (33°28'06"N, 117°44'06’W), Jan 1985, 35 m, 1 holotype female (7.3 mm); LACMNH No. 88-116.1, Santa Monica Bay, California (33°58'47’N, 118°30'27"W), July 1985, 27 m, 8 paratypes; LACMNH No. 85-200.1, Santa Monica Bay, California (33°52'34’N, 118°26'02”W), July 1985, 26 m, 3 paratypes. Distributional ecology.—Geographic range—Santa Monica Bay, California to San Diego, California. Bathymetric range—sublittoral 10 m to 50 m depth. Sed- iment preference— mixed bottom areas of silt and sand. Etymology.—The specific epithet, brachycladus, is a combination of the Greek words brachys (=short) and klados (=branch), alluding to the short inner ramus on uropod |. It is a masculine noun in apposition to the generic name. Acknowledgments I thank Charles A. Phillips, Hyperion Treatment Plant, for his assistance in obtaining and reviewing obscure literature. The illustrations are by Gregory B. Deets, Hyperion Treatment Plant. Finally, I wish to thank Dr. J. Laurens Barnard, Smithsonian Institution, NMNH, and Dr. Masahiro Dojiri, Hyperion Treatment Plant, for critically reviewing this manuscript. This paper was partially funded by grant number 89-1 from the Southern California Association of Marine In- vertebrate Taxonomists. Contribution Number 1 of the Southern California Association of Marine In- vertebrate Taxonomists. Literature Cited Barnard, J. L. 1954. Amphipoda of the family Ampeliscidae collected in the eastern Pacific Ocean by the Velero III and Velero IV. Allan Hancock Pac. Exped. 18, 137 pp. . 1960. New bathyal and sublittoral ampeliscid amphipods from California, with an illustrated key to Ampelisca. Pac. Nat., 1(16):1-36. . 1967. New species and records of Pacific Ampeliscidae (Crustacea: Amphipoda). Proc. U.S. Nat. Mus., 121:1—20. . 1969. The families and genera of marine gammaridean Amphipoda. U.S. Nat. Mus. Bull. 271, 535 pp. NEW SPECIES OF GAMMARID AMPHIPOD 129 . 1971. Gammaridean Amphipoda from a deep sea-transect off Oregon. Smithson. Cont. Zool. 61, 86 pp. Dickinson, J. J. 1982. Studies on the Amphipod crustaceans of the northeastern Pacific region. 1. Family Ampeliscidae, Genus Ampelisca. Natl. Mus. Can., Pub. Biol. Oceano., 10, 1-40. Enequist, P. 1949. Studies on the soft-bottom amphipods of Skagerak. Zool. Bidr. Upps., 28:297— 492. Holmes, S. J. 1908. The Amphipoda collected by the U.S. Bureau of Fisheries steamer “Albatross” off the west coast of North America, in 1903 and 1904, with descriptions of a new family and several new genera and species. Proc. U.S. Natl. Mus., 35:489-543. Judd, S. D. 1896. Description of three species of sand fleas (Amphipods) collected at Newport, Rhode Island. Proc. U.S. Natl. Mus., 18:593-603. Kroyer, H. 1842. Une nordiske slaegter og arter af amfipodernes orden, Lenhorende til familien Gammarina. Naturh. Tid Sskr., 4:141-166. Accepted for publication 26 September 1989. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 130-136 © Southern California Academy of Sciences, 1990 Two Medusae New to the Coast of California: Carybdea marsupialis (Linnaeus, 1758), a Cubomedusa and Phyllorhiza punctata von Lendenfeld, 1884, a Rhizostome Scyphomedusa Ronald J. Larson! and A. Charles Arneson? 'Harbor Branch Oceanographic Institution, 5600 Old Dixie Highway, Fort Pierce, Florida 34946 ?Scripps Institution of Oceanography, La Jolla, California 92093 Abstract.—The cubomedusa Carybdea marsupialis and the rhizostome scypho- medusa Phyllorhiza punctata are reported from waters of California and the east- ern Pacific for the first time. Previously, C. marsupialis was only known from the Atlantic Ocean. It apparently is native to this area but was confused with Carybdea rastoni which is known from Hawaii and the central and western Pacific. P. punctata was recently found in San Diego Bay, previously it was known in the Pacific from Hawaii and the tropical Indo-Pacific. Its recent appearance in Cal- ifornia was probably due to transport of polyps on the hulls of ships. A synopsis of cubomedusae and scyphomedusae from the waters of California is presented; a total of 19 species are known. Although the marine fauna of California is relatively well studied some inver- tebrate groups are in need of further work. The scyphomedusae are one such group. Nearly one-half of the 18 known species from California were reported after 1960 (Table 1). Most of these new records are for deep water species. Alvarino (1967, 1976) added two previously unreported coronates, Atorella vanhoeffeni Bigelow, 1909 and Nausithoe rubra Vanhoffen, 1902, to the fauna of California and Russell (1967) and Smith (1982) found two previously unreported mesope- lagic semaeostomes, Deepstaria enigmatica Russell, 1967 and Poralia rufescens Vanhoffen, 1902, which were collected using submersibles. Some neritic species have also been recently found. Two new stauromedusae, benthic scyphozoans which generally live subtidally attached to algae, were recently described from California. Larson (1988) described Kyopoda lamberti, which represents a new family, and Larson and Fautin (1989) described a new species belonging to the genus Manania. Here we report on the recent collection of two medusae, Carybdea marsupialis (Linnaeus, 1758), and a cubomedusan Phyllorhiza punctata von Len- denfeld, 1884, previously was not known from the eastern Pacific. Most recently, a darkly pigmented, undescribed species of Chrysaora, up to 25 cm diameter, has appeared in waters of southern California (J. Martin, pers. comm., 1989). Cubomedusae can be common in warm marine waters. Two species occur in inshore waters of the east coast of the U.S. Tamoya haplonema Miller 1859 has been collected along the north coast of the Gulf of Mexico (Phillips and Burke 1970), along the Georgia coast (Kraeuter and Setzler 1975), and as far north as Long Island (Mayer, 1910). Chiropsalmus quadrumanus (Miller, 1859) is known 130 NEW MEDUSAE FROM CALIFORNIA 131 Table 1. Checklist of Cubomedusae and Scyphomedusae known from the coast of California. Taxon References* Class Cubozoa Carybdea marsupialis (Linnaeus, 1758) 6, 11, 13, 17, 18 Class Scyphozoa Order Stauromedusae Suborder Eleutherocarpidae Haliclystus octoradiatus (Lamarck, 1816) 7 Haliclystus “‘californiensis” 7 “‘Stenoscyphopsis vermiformis”’ 7 Kyopoda lamberti Larson, 1988 1 Suborder Cleistocarpida Manania gwilliami Larson and Fautin, 1989 16 Order Coronatae Atolla wyvillei Haeckel, 1880 4,9, 12 Atorella vanhoeffeni Bigelow, 1909 12 Nausithoe rubra Vonh6ffen, 1902 9 Periphylla periphylla (Péron & Lesueur, 1809) 4,9, 12 Order Semaeostomeae Aurelia aurita (Linnaeus, 1758) 1 Chrysaora fuscescens Brandt, 1835 I, 2 3, VF Chrysaora sp. 17 Deepstaria enigmatica Russell, 1967 10 Pelagia colorata Russell, 1964 WY Phacellophora camtschtica Brandt, 1835 1,4, 17 Poralia rufescens Vanh6ffen, 1902 14 Order Rhizostomeae Phyllorhiza punctata von Lendenfeld, 1884 18 Stomolophus meleagris L. Agassiz, 1862 5,6 * References: 1 = Agassiz (1862), 2 = Fewkes (1889), 3 = Kishinouye (1899), 4 = Bigelow (1913), 5 = Bigelow (1914), 6 = Stiasny (1922), 7 = Gwilliam (1956), 8 = Russell (1964), 9 = Alvarino (1967), 10 = Russell (1967), 11 = Gladfelter (1973), 12 = Alvarino (1976), 13 = Satterlie (1979), 14 = Smith (1982), 15 = Larson (1988), 16 = Larson and Fautin (1989), 17 = Larson unpubl., 18 = this report. from the northern Gulf of Mexico (Guest 1959; Phillips and Burke 1970), and along the east coast of the U.S. from Florida (Larson, unpublished observations), Georgia (Kraeuter and Setzler 1975), and North Carolina (Mayer 1910). Several other species, i.e., Carybdea alata Reynaud, 1830 and Carybdea mar- supialis, although common in the Caribbean (Mayer 1910; Bigelow 1938), have not been reported in neritic waters of the U.S. east coast. Surprisingly, no cubomedusae are known from the coast of the tropical eastern Pacific even though one species, Carybdea rastoni Haacke, 1886, has been reported from California. The first report of this species from the west coast of the U.S. was based on material collected at La Jolla (Stiasny 1922). Additionally, this species has been collected at Santa Barbara (Gladfelter 1973; Satterlie 1979). Rhizostomes are also mainly subtropical/tropical, reaching their greatest di- versity in the Indo-Malayan region (Kramp 1970). In the tropical Indo-Pacific these medusae are so abundant that they are harvested for their collagenous bell (Omori 1978, 1981). Only a single species, Stomolophus meleagris L. Agassiz, 1862, was previously known from California (Bigelow 1914; Stiasny 1922). 132 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 1. A. Side view of preserved Carybdea marsupialis, 40 mm bell height, collected off Point Moremesa, Santa Barbara, California, 10 August 1985. B. Side view of live Phyllorhiza punctata (oral arm appendages not shown), 50 mm bell diameter, collected in Mayaguez Harbor, Puerto Rico, August 1975. Results Carybdea marsupialis (Linnaeus, 1758) Material examined: Thirteen specimens. One specimen 25 mm bell height; collected off Scripps pier, La Jolla, 7 October 1961; collector: R. Cutter. Five mature specimens, 36-40 mm bell height; collected off Point Moremesa, Santa Barbara; 10 August 1985; 5 m depth; collectors: S. Anderson and J. McCullach. Seven mature specimens, 28-35 mm bell height collected off Point Moremesa, Santa Barbara; 11 November 1984; collectors: S. Anderson and J. McCullach. For comparison, we examined C. rastoni specimens collected at the type locality, the Gulf of St. Vincent, Australia. In addition, C. marsupialis specimens from the Bahamas were studied. Discussion: The California material fits the diagnosis of C. marsupialis as de- scribed by Bigelow (1938) in his revision of the family Carybdeidae. The inter- radial phacellae are single stalked. These specimens most closely resemble the Mediterranean morph which is larger and has more obvious exumbrellar sculp- turing than medusae from the western Atlantic (Mayer 1910; Bigelow 1938) (Fig. 1). Carybdea marsupialis is distinguished from the closely related C. rastoni, in NEW MEDUSAE FROM CALIFORNIA 133 size and arrangement of the gastric cirri. In the larger C. marsupialis, reaching 4 cm bell height, the phacellae (dendritic gastric cirri present at each of the 4 interradial stomach corners), originate from a single trunk, or rarely from two adjacent trunks (Bigelow 1938, text-figs. 3-5). Whereas, C. rastoni is usually less than 3 cm bell height and has gastric cirri that originate from multiple trunks arranged in four interradial rows in the stomach corners (Haacke 1886, fig. 4b; Bigelow 1909, plate 10, fig. 7). Carybdea marsupialis was previously known only from the tropical/subtropical Atlantic and Mediterranean. Although, two species of Carybdea may occur along the coast of southern California, we believe that there is only one. We have examined material from the same locations along the coast of California where C. rastoni was previously reported and all specimens were C. marsupialis. It is possible that Stiasny (1922) made an incorrect determination. The first author examined Stiasny’s specimen which is in the Zoologiske Museum, Copenhagen. The specimen was a C. mar- supialis. Evidently, Stiasny had not examined the phacellae closely because the bell apex had not been dissected open to expose them. Stiasny probably identified the specimen as C. rastoni after only a cursory examination because that species was then the only known cubomedusan from the eastern Pacific since it was known from Hawaii. Although, Gladfelter (1973) and Satterlie (1979) examined living Carybdea material there is no indication on what they based their determinations. Based on the material available to us, there is only one species of Carybdea, namely C. marsupialis, from the California coast. It is enigmatic that C. marsupialis has not been reported from other areas of the eastern Pacific because in the Atlantic it is widely distributed and relatively common. Possibly the California population represents a recent introduction from the Atlantic as polyps attached to a ship hull or as medusae in bilge water. Yet, this seems unlikely because the polyps are small and delicate (Studebaker 1972) and probably could not have survived the trip. The medusae are also sensitive to water quality. | More likely, C. marsupialis is a resident species of the eastern Pacific, having occurred in the contiguous waters of the Caribbean and eastern Pacific prior to the emergence of the Isthmus of Panama about 3 million years ago (Woodring 1966). Why this species was not previously reported in the eastern Pacific is unknown, but it may be that C. marsupialis is not as common there as it is in the Caribbean, and the medusan fauna of the tropical/subtropical eastern Pacific has not been as well studied as that of the Caribbean. The distribution of C. marsupialis is apparently unique among medusae, being widespread in the subtropical/tropical Atlantic but restricted in the Pacific to the eastern region. The fact that the specimens from the Pacific more closely resemble the Mediterranean morph is puzzling but may be due to ecological factors, e.g. lower temperatures and/or higher prey concentrations. Carybdea marsupialis is common near Santa Barbara where it occurs from September to November (Gladfelter 1973; Satterlie 1979; S. Anderson and J. McCullugh, pers. comm. 1986). It is mostly seen in the water column a few meters above the bottom in about 10 m of water. Gladfelter (1973) reports that it feeds on mysids. Satterlie (1979) has described aspects of its neurophysiology. 134 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Phyllorhiza punctata von Lendenfeld, 1884 Material examined: One specimen (7 cm bell diameter); collected in Mission Bay (30 July 1981; surface). Numerous live specimens have also been observed by one of us (A. Arneson) in San Diego Bay. Discussion: This species is readily identified when alive by its bluish exumbrella with white spots (Fig. 1) and by its long oral-arm clubs with blue and white tips. The taxonomy of P. punctata is highly confused, mainly owing to poorly pre- served or damaged material. As suggested by Cutress (1971), P. punctata is prob- ably synonymous with a number of Mastigias species (i.e., M. albipunctata Stiasny, 1920; M. andersoni Stiasny, 1926; and M. scintillae Soares Moreira, 1961) and probably also with Cotylorhiza pacifica Mayer, 1915 and Cotylorhizoides pacifica Light, 1921. The life history of P. punctata has been elucidated by Cutress (1971). Phyllorhiza punctata was previously known in the Pacific, only from the Indo- Pacific (Mayer 1910). But recently, it was found in Hawaii (Devany and Eldredge 1977) as well as in the western tropical Atlantic, i.e., Brazil (Soares Moreira 1961), Puerto Rico (Cutress 1971), and Jamaica (Larson unpublished). Its recent ap- pearance in these areas was probably due to transport of polyps on the hulls of ships, as previously suggested by Cutress (in Doty, 1961). This hypothesis is supported by observations that P. punctata medusae are mostly confined to har- bors used by ocean-going ships. The San Diego Bay and adjacent Mission Bay populations may have originated from polyps carried from Honolulu Harbor by naval vessels. It is unlikely that medusae were the dispersal stage since they are very active and therefore have high food demands that would probably not be met in oligotrophic oceanic waters between Hawaii and California. Similarly, scyphistomal introduction has been hypothesized for the recent ap- pearance of another rhizostome, Anomalorhiza shawi Light, 1921, in Hawaii. Cooke (1984) postulated that this species was transported to Hawaii from the Philippines as part of the ship-borne fouling community. Apparently, P. punctata has become well established in San Diego Bay, judging by the seasonal presence of large numbers of medusae. Synopsis of California Scyphomedusae One species of cubomedusa and 18 species of scyphomedusae are now known from California (Table 1). Since about half of these species have been reported only in the last 20 years, it is likely that additional species may be discovered when further collecting is done in offshore waters and when material from existing collections is examined. Acknowledgments Thanks are due to S. Anderson and J. McCullach for collecting cubomedusan material, and to L. Marsh, K. Petersen, and G. Snyder for loan of specimens. A. Alvarino and anonymous reviewers provided helpful comments on the manu- script. This is contribution no. 739 of Harbor Branch Oceanographic Institution. Literature Cited Agassiz, A. 1862. Pp. 1-380 in L. Agassiz, 1862. Contributions to the natural history of the United States of America, Vol. 4. Little Brown, Boston. NEW MEDUSAE FROM CALIFORNIA 135 Alvarifio, A. 1967. Bathymetric distribution of Chaetognatha, Siphonophorae, Medusae, and Cte- nophorae off San Diego, California. Pac. Sci., 21:474—485. . 1976. Los indicadores planctonicos: distribuciOn batimétrica de alguanas medusas. Pp. 161- 190 in Memorias del II Simposio Latinoamerican Sobre Oceanographia Biologica. Universidad de Oriente, Venezuela. Bigelow, H. B. 1909. The Medusae. Rep. Sci. Res. Eastern Trop. Pac. Exped. U.S. Fish. Comm. Steamer Albatross 1904-1905. Mem. Mus. Comp. Zool. Harv. Coll., 37:1-243. . 1913. Medusae and Siphonophorae collected by the U.S. Fisheries Steamer Albatross in the north-west Pacific, 1906. Proc. U.S. Nat. Mus., 44:1-119. 1914. Note on the medusan genus Stomolophus from San Diego. Univ. Calif. Publ. Zool., 13:239-241. 1938. Plankton of the Bermuda Oceanographic Expeditions. VIII. Medusae taken during the years 1929 and 1930. Zoologica, N.Y., 23:99-189. Cooke, W. J. 1984. New scyphozoan records for Hawaii: Anomalorhiza shawi Light, 1921, and Thysanostoma loriferum (Ehrenberg, 1835); with notes on several other rhizostomes. Proc. Biol. Soc. Wash., 97:583-588. Cutress, C. E. 1971. Phyllorhiza punctata in the tropical Atlantic. Proc. Assoc. Island Marine Lab. Caribbean, 9:14. Devany, D. M., and L. G. Eldredge. 1977. Reef and shore fauna of Hawaii. Section 1: Protozoa through Ctenophora. Bishop Mus. Spec. Publ., 64:1-277. Doty, M. S. 1961. Acanthophora, a possible invader of the marine flora of Hawaii. Pac. Sci., 15: 547-552. Fewkes, J. W. 1889. New Invertebrates from the coast of California. Bull. Essex Inst., 21:99-146. Gladfelter, W. B. 1973. A comparative analysis of the locomotory systems of medusoid Cnidaria. Helgo. wiss. Meeresunters., 25:228-272. Guest, W. 1959. The occurrence of the jellyfish Chiropsalmus quadrumanus in Matagorda Bay, Texas. Bull. Mar. Sci. Gulf. Carib., 9(1):79-83. Gwilliam, G. F. 1956. Studies on West Coast Stauromedusae. Ph.D. Thesis, Univ. of California, Berkeley, 186 pp. Haacke, W. 1886. Die Scyphomedusen des St. Vincent Golfes. Jen. Z. Naturw., 20:589-638. Kishinouye, K. 1899. A new medusa from the California coast. Zool. Anz., 22:44—-45. Kramp, P. L. 1961. Synopsis of the medusae of the world. J. Mar. Biol. Ass. U.K., 40:1-469. . 1970. Zoogeographical studies on Rhizostomeae (Scyphozoa). Vidensk. Meddr. dansk naturh. Foren., 133:7-30. Kraeuter, J. N., and E. M. Setzler. 1975. The seasonal cycle of Scyphozoa and Cubozoa in Georgia estuaries. Bull. Mar. Sci., 25(1):66—74. Larson, R. J. 1988. Kyopoda lamberti gen. nov., sp. nov., an atypical Stauromedusa (Scyphozoa, Cnidaria) from the eastern Pacific, representing a new family. Can. J. Zool., 66:2301—2303. , and D. G. Fautin. 1989. Stauromedusae of the genus Manania (=Thaumatoscyphus) (Cni- daria, Scyphozoa) in the northeast Pacific, including descriptions of new species Manania gwilliami and Manania handi. Can. J. Zool., 67:1543-1549. Mayer, A. G. 1910. Medusae of the world. III. The Scyphomedusae. Carnegie Inst. Wash. Publ., 109:499-735. Omori, M. 1978. Zooplankton fisheries of the world: A review. Mar. Biol., 48:199-205. . 1981. Edible jellyfish (Scyphomedusae: Rhizostomeae) in the far east waters: A brief review of the biology and fishery. Bull. Plank. Soc. Japan, 28:1-11. Phillips, P. J., and W. D. Burke. 1970. The occurrence of sea wasps (Cubomedusae) in Mississippi Sound and the northern Gulf of Mexico. Bull. Mar. Sci., 20(4):853-859. Russell, F. S. 1964. On scyphomedusae of the genus Pelagia. J. Mar. Biol. Ass. U.K., 44:133-136. 1967. Ona remarkable new scyphomedusan. J. Mar. Biol. Ass. U.K., 47:469-473. Satterlie, R. A. 1979. Central control of swimming in the cubomedusan jellyfish Carybdea rastonii. J. Comp. Physiol., 133:357-367. Smith, K. L. 1982. Zooplankton of a bathyl benthic boundary layer: In situ rates of oxygen con- sumption and ammonium excretion. Limnol. Oceanogr., 27:461-471. Soares Moreira, M.G. B. 1961. Sobre Mastigias scintillae sp. nov. (Scyphomedusae, Rhizostomeae) das costas do Brasil. Bol. Inst. Oceanogr. S. Paulo, 11:5-29. Stiasny, G. 1922. Die Scyphomedusen-Sammlung von Dr. Th. Mortensen nebst anderen Medusen 136 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES aus dem Zoologischen Museum der Universitat in Kopenhagen. Pap. from Dr. Th. Mortensen’s Pac. Exped. 1914-1916. XIII., 73:513-555. Studebaker, J. P. 1972. Development of the cubomedusa, Carybdea marsupialis. M.S. Thesis, Univ. of Puerto Rico, Mayaguez, 60 pp. Woodring, W. P. 1966. The Panama land bridge as a sea barrier. Proc. Amer. Phil. Soc., 110:425- 433. Accepted for publication 26 July 1989. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 137-138 © Southern California Academy of Sciences, 1990 Research Notes The Tropical Colonial Stony Coral Tubastrea coccinea at Cabo San Lucas, Mexico Three coralla of the tropical stony coral Tubastrea coccinea Lesson, 1829 (see Wells 1983 for list of junior synonyms) were collected on July 17, 1989 at 4 m depth from the north side of a granite islet at Cabo San Lucas, Baja California Sur, Mexico. The live specimens were brilliant orange in color. The two largest coralla are 10 cm (Fig. 1) and 15 cm in diameters and each has approximately 100 corallites. The smallest corallum is 3 cm in diameter and has 15 corallites. Fig. 1. Colonial stony coral Tubastrea coccinea. 137 138 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES The coralla of Pocillopora spp. reported by Wilson (1988) occur on the south side of the same islet. This report extends the known range of Tubastrea coccinea from Bahia Pulmo in the Gulf of California (Brusca and Thompson 1977) approximately 80 km SSW to the border of the Pacific Ocean. The previous known range in the eastern Pacific Ocean extended from Bahia Pulmo north into the Gulf of California and south along the mainland shores and islands of Mexico, Costa Rica, Panama, Ecuador, and the Galapagos Islands; in the western Pacific Ocean, 7. coccinea has been recorded from the Hawaiian Islands west to Murray Island, Australia (Brusca 1980; Durham 1947; Durham and Barnard 1952; Horst 1922; Squires 1959; Wells 1983). Squires’ (1959, table 3) indication of a western coast of Baja California occurrence apparently is an error because there are no recorded localities for the species in that region. The specimens were deposited in the Invertebrate Zoology Section, Natural History Museum of Los Angeles County. Permission to collect kindly was obtained through M. en C. Jorge Garcia P. of the Departamento de Biologia Marina, Universidad Autonoma de Baja California Sur, through the courtesy of M. en C. Oscar Arizpe C. of the same department. I am indebted to Srs. Paulino Perez G. and Hector Reyes B. of La Paz for assistance in collecting. Literature Cited Brusca, R.C. 1980. Common intertidal invertebrates of the Gulf of California. University of Arizona Press, Tucson, 513 pp. , and D. A. Thompson. 1977. Pulmo reef: the only “coral reef’ in the Gulf of California. Ciencas Marinas, 1(3):37-53. Durham, J. W. 1947. Corals from the Gulf of California and the north Pacific coast of America. Geol. Soc. Amer., Mem., 20:1-68. , and J. L. Barnard. 1952. Stony corals of the eastern Pacific collected by the Velero III and Velero IV. Allan Hancock Pac. Exped., 16(1):1-110. Horst, C. J. van der. 1922. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, vol. 7, no. 9, Madreporaria: Agaricidae. Trans. Linnean Soc. London, ser. 2, 18:417-429. Squires, D. F. 1959. Results of the Puritan-American Museum of Natural History expedition to western Mexico. 7. Corals and coral reefs in the Gulf of California. Bull. Amer. Mus. Nat. Hist., 118(7):370-431. Wells, J. W. 1983. Annotated list of the scleractinian corals of the Galapagos. Pp. 213-295, in Corals and coral reefs of the Galapagos Islands. (P. W. Glynn and G. M. Wellington, eds.), University of California Press, Berkeley, 330 pp. Wilson, E. C. 1988. The hermatypic coral Pocillopora at Cabo San Lucas, Mexico. Bull. Southern California Acad. Sci. 87(2):79-83. Accepted for publication 12 September 1989. Edward C. Wilson, Curator, Section of Invertebrate Paleontology, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 139-142 © Southern California Academy of Sciences, 1990 Harbor Porpoises Utilize Tidally-induced Internal Waves Little information exists to associate small cetaceans with physiographical phe- nomena known to affect oceanic prey distribution. We present data which suggest that harbor porpoises, Phocoena phocoena, feed in surface slicks generated by tidally-induced internal waves. Internal waves are evident at the water surface as parallel streaks of flat water (“‘slicks”’) surrounded by rippled water in coastal areas and harbors when the wind is light (Ewing 1950; LaFond 1959). Internal waves and the resultant surface slicks are common features in many ocean basins (Ewing 1950; LaFond 1959; Cairns 1967; Hendrickson 1973; Lepley et al. 1977; Shea and Broenkow 1982; Chereskin 1983). The waves often result in areas of increased ievels of biological activity due to their tendency to concentrate small organisms (Zelids and Jillett 1982; Kingsford and Choat 1986; Shanks and Wright 1987) and to transport pelagic larval invertebrates and fish (Norris 1966; Shanks 1983, 1988). Fish that feed on plankton concentrated in the internal waves are also aggregated in these areas (Norris 1966; Kingsford and Choat 1986). In addition, cetaceans exhibit afhlia- tions with surface slicks and these animals may feed within the waves as well. Silber (1990) noted that Gulf of California harbor porpoises, Phocoena sinus, tended to be sighted more often in slicks caused by internal waves than in sur- rounding waters. However, there is little quantitative documentation of cetaceans associating with slicks of internal wave origin, nor are there data which suggest that cetaceans feed within the subsurface waves. In offshore areas, slicks may be caused by different factors, such as windrowing, fronts, and eddies, where waters of differing physical properties converge (Bowman and Esaias 1978). Smith et al. (1986) found that ribbons of increased biomass, represented by intermediate levels of the food web, along fronts or convergent zones were exploited by foraging cetaceans. Researchers in the western North Atlantic reported that sei, Balaenoptera borealis, and right whales, Eubalaena glacialis, followed offshore slicks as a source of concentrated planktonic prey (Watkins and Schevill 1976, 1979, 1982). Feeding humpback, Megaptera no- vaeangliae, and fin whales, Balaenoptera physalus, were also associated with slicks in areas where concentrated schools of fish prey were correlated with dense plank- ton patches (Watkins and Schevill 1979). Other pelagic cetaceans are known to utilize surface slicks, including pygmy killer whales, Feresa attenuata (Pryor et al. 1965), rough-toothed dolphins, Steno bredanensis, and false killer whales, Pseudorca crassidens (K. S. Norris, Long Marine Lab, 100 Shaffer Drive, Santa Cruz, California 95060, pers. comm.). During recent field studies on the behavior of harbor porpoises in Monterey Bay, California, we observed that porpoises surfaced most frequently in or near surface slicks. To quantify the association between surface slicks and behavioral activity, a subsequent study was conducted in which harbor porpoise behavior was monitored from a 67.5 meter bluff at Sunset Beach State Park overlooking 139 140 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Monterey Bay, from 10 September to 1 November 1988, during sea states of Beaufort 2 (no whitecaps, windspeed 7-11 km/hour). The study area consisted of water less than 20 m deep with a sandy substrate. At one-minute intervals, the most recent behavioral state of a focal group (milling or traveling) observed within the previous minute was noted, as was the nature of the water surface (slick or rippled) within which the activity occurred. The first porpoise(s) observed within 1.5 km of the study site was selected as the focal group. If a porpoise group traversed between slick and interslick areas, which occurred rarely, the group was categorized as having surfaced in the water type that it appeared in during its ultimate surfacing. Each surfacing sequence by a porpoise group was treated as an independent event, because we could not be sure whether or not the group under observation had been previously sampled. The minimum observation pe- riod for a focal group was set at three minutes for analysis. The width of the slicks was not measured, because the location and dimensions of the slicks and interslick areas were dynamic and constantly shifting. Traveling was defined as unidirectional movement for at least three surfacings. Milling was defined as nondirectional movement for three consecutive surfacings, generally consisting of criss-crossing or circling movement. Although milling be- havior may represent a wide variety of activities in cetaceans and the exact function of milling could not be established in our work, foraging activity in harbor porpoises apparently occurs during milling (Goetz 1983; Watson and Gas- kin 1983; Sekiguchi 1987). We saw fish jumping at the water surface near milling harbor porpoises on three occasions, which corroborates, but does not confirm a connection between milling and feeding. A total of 532 behavioral and associated water surface states was collected on 96 porpoise groups during a total of 19.4 hours on 15 days. Harbor porpoises were generally common within 3 km of shore, but we focused our attention on those animals closest to shore (<1.5 km from the observation site), because they were easiest to observe. We used seven power binoculars to spot porpoises. While in slicks (N = 222), harbor porpoises spent significantly more time milling (N = 139) than traveling (N = 83) (Chi-square = 13.6, df = 1, P < 0.001). No significant difference was found between the amount of time spent milling (N = 142) or traveling (N = 168) within rippled (non-slick) water (N = 310) (Chi-square = 2.02, P > 0.10). When the two habitats (slick versus non-slck) were compared, a significantly greater tendency for milling while in slicks was found (Chi-square = 24.2, df = 1, P < 0.001). Most harbor porpoise prey items are relatively small gregarious species (Jones 1981), that may be drawn to internal waves to feed on aggregated larvae or zooplankton. In an analysis of harbor porpoise stomach contents from Monterey Bay, Sekiguchi (1987) reported that market squid (Loligo opalescens), the northern anchovy (Engraulis mordax), the spotted cusk eel (Chilara taylori), rockfishes (Sebastes spp.), the plainfin midshipman (Porichthys notatus), the jack mackerel (Trachurus symmetricus), and the shiner surfperch (Cymatogaster aggregata) were the numerically dominant prey. Jones (1981) reported that two-thirds of all fish found in stomachs of harbor porpoises from north-central California live in open ocean or are inshore schooling species. The Pacific hake (Merluccius productus), the Pacific tomcod (Microgadus proximus), rockfishes, and northern anchovies accounted for 97% of all otoliths in harbor porpoise stomachs sampled. Inver- RESEARCH NOTES 141 tebrates, mostly Loligo opalescens, were found in 40% of all stomachs examined by Jones (1981). The tendency for harbor porpoises to affiliate with slicks and to mill while in slicks, suggests that porpoises may be feeding on higher localized prey densities as a result of concentrating properties of internal waves. It is likely that the porpoises were feeding on fish, squid, or other organisms attracted to zooplankton assemblages in the slicks. This paper was improved by comments from T. Jefferson, M. Newcomer, and B. Wursig. We thank the California Department of Parks and Recreation for the use of Sunset State Beach. This represents Contribution No. 10 of the Marine Mammal Research Program of Texas A & M University at Galveston. Literature Cited Bowman, M. J., and W. E. Esaias. 1978. Oceanic fronts in coastal processes. Springer-Verlag, Berlin, Heidelberg. Cairns, J. L. 1967. Asymmetry of internal tidal waves in shallow coastal waters. J. Geophys. Rev., 72:3563-3565. Chereskin, T. K. 1983. Generation of internal waves in Massachusetts Bay. J. Geophys. Rev., 88: 2649-2661. Ewing, G. 1950. Slicks, surface films and internal waves. J. Mar. Res., 9:161-187. Goetz, B. J. 1983. Harbor porpoise (Phocoena phocoena (L.)) movements in Humboldt Bay, Cali- fornia, and adjacent waters. M.S. Thesis, Humboldt State University, Arcata, Calif., 53 pp. Hendrickson, J.R. 1973. Study of the marine environment of the northern Gulf of California. NTIS Publication No. N74-16008, 95 pp. Jones, R. E. 1981. Food habits of smaller marine mammals from northern California. Proc. Cal. Acad. Sci., 42:409-433. Kingsford, M. J., and J. H. Choat. 1986. {nfluence of surface slicks on the distribution and onshore movements of small fish. Mar. Biol., 91:161-171. LaFond, E. C. 1959. Slicks and temperature structure in the sea. U.S. Naval Electronics Lab. Res. Rep., 937:1-27. Lepley, L. K., S. P. Vonder Haar, J. R. Hendrickson, and G. Calderon-Riveroll. 1975. Circulation in the northern Gulf of California from orbital photographs and ship investigation. Ciencias Marinas, 2:86-93. Norris, K. S. 1966. The functions of temperature in the ecology of the percoid fish Girella nigicans (Ayres). Ecol. Monogr., 33:23-62. Pryor, R. A., K. Pryor, and K. S. Norris. 1965. Observations on a pygmy killer whale (Feresa attenuata Gray) from Hawaii. J. Mamm., 46:450-461. Sekiguchi, K. 1987. Occurrence and behavior of the harbor porpoise (Phocoena phocoena) at Pajaro Dunes, Monterey Bay, California. M.S. Thesis, Moss Landing Marine Laboratories, Moss Landing, Calif., 49 pp. Shanks, A. L. 1983. Surface slicks associated with tidally forced internal waves may transport pelagic larvae of benthic invertebrates and fishes shoreward. Mar. Ecol. Prog. Ser., 13:311-315.' . 1988. Further support for the hypothesis that internal waves can cause shoreward transport of larval invertebrates and fish. U.S. Fish. Bull., 86:703-714. ,and W.G. Wright. 1987. Internal-wave-mediated shoreward transport of cyprids, megalopae, and gammarids and correlated longshore differences in settling rate of intertidal barnacles. J. Exp. Mar. Biol., 114:1-13. Shea, R. E., and W. W. Broenkow. 1982. The role of internal tides in the nutrient enrichment of Monterey Bay, California. Estuarine, Coastal and Shelf Sci., 15:57-66. Silber, G. K. 1990. Occurrence and distribution of the vaquita (Phocoena sinus) in the northern Gulf of California. U.S. Fish. Bull., 88:339-346. Smith, R. C., P. Dunstan, D. Au, K. S. Baker, and E. A. Dunlap. 1986. Distribution of cetaceans and sea-surface chlorophyll concentrations in the California current. Mar. Biol., 91:385—402. Watkins, W. A., and W. E. Schevill. 1976. Right whale feeding and baleen rattle. J. Mamm., 57: 58-66. 142 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 1979. Aerial observation of feeding behavior in four baleen whales: Eubalaena glacialis, Balaenoptera borealis, Megaptera novaeangliae, and Balaenoptera physalus. J. Mamm., 60: 155-163. 1982. Observations of right whales, Eubalaena glacialis, in Cape Cod waters. U.S. Fish. Bull., 80:875-880. Watson, A. P., and D. E. Gaskin. 1983. Observations on the ventilation cycle of the harbour porpoise Phocoena phocoena (L.) in coastal waters of the Bay of Fundy. Can. J. Zool., 61:126-132. Zelids, J. R., and J. B. Jillett. 1982. Aggregation of pelagic Munida gregaria (Fabricius) (Decapoda, Anomura) by coastal fronts and internal waves. J. Plankton Res., 4:839-857. Accepted for publication 7 March 1990. Gregory K. Silber, Institute of Marine Sciences, University of California, Santa Cruz, California 95064; present address: Friends of the Sea Otter, P.O. Box 221220, Carmel, California 93922, and Mari A. Smultea, Moss Landing Marine Labo- ratories, Moss Landing, California 95039; present address: Marine Mammal Re- search Program, Texas A&M University at Galveston, Galveston, Texas 77553- 1675. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 143-145 © Southern California Academy of Sciences, 1990 The Aquatic Dryopoid Beetles of Pinnacles National Monument: Optioservus canus Revisited (Coleoptera: Dryopoidea: Elmidae) Early in 1985 I was contacted by the Sacramento, California offices of the U.S. Fish and Wildlife Services. They were interested in what species of California riffle beetles might warrant listing on the national register. One potential candidate was Optioservus canus Chandler, 1954, the type locality of which is Pinnacles National Monument (PNM), in San Benito County, California. During the sum- mer of 1946 Harry Chandler collected four specimens of O. canus from Chalone Creek, in PNM. In his description of the species Harry also included two additional specimens collected by F. E. Winters in Santa Barbara and Riverside. Thus the type series included only six specimens. Despite careful collecting in California by such noted aquatic coleopterists as Hugh Leech and Harley Brown, no one recollected the species. Obviously this was a potentially endangered species. In August 1985, I traveled to PNM to obtain a collecting permit and to begin searching for this elusive species. The first two attempts to collect O. canus were fruitless despite searching the few permanent water sources and along the mostly dry bed of Chalone Creek. Several other species of dryopoids were collected however. On discussing things with the rangers, one recalled that Chalone Creek had been channelized to control erosion. It appeared that O. canus had perhaps gone extinct due to man’s lack of attention to “‘lesser’’ organisms. The one mitigating circumstance was that while traveling to and from the PNM area, I had collected a new and quite distinct species of Optioservus in Tres Pinos Creek, about 30 miles to the north. This new species was also found in the Arroyo Seco River some 20 miles to the south of PNM. Since it was unkeyable in the recent revision of Optioservus (White 1978), I set about describing this species. After examining many specimens among which the elytral coloration varied great- ly (from all dark to almost all light), I began to notice some were similar to White’s illustration of O. canus. This led to reexamination of the types of O. canus. Happily, I can report that O. canus still exists, although under a different ap- pearance than furnished by the type series and published illustrations. One last collecting trip to PNM, in December 1985, disclosed that O. canus still existed in Chalone Creek, the type locality. This population occurs in a very small area and is well hidden under an overstory of watercress. At this site surfacing bedrock forces water in Chalone Creek out of the bed sediments and over the bedrock for a few feet. In the original description and the first inclusion of O. canus in a key to Op- tioservus (Leech and Chandler 1956) the elytral maculae were noted but their shape was not even described. The most distinctive feature for the species was that the elytral pubescence gave a “‘grizzled look”’ to the specimens. Collier (1969) was the first to refer to the shape of the maculae. In his key he called them vittae, but in his description he called them spots. His illustration shows an elongate vitta in the lateral half of each elytron (Plate II, fig. 15). Collier examined three specimens from Pinnacles (all of which were in the California Academy of Sciences 143 144 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES collection). His couplet in which O. canus is separated does include the “grizzled appearance.” Brown (1972) repeated Collier’s couplets and illustrations in his manual on the United States’ species of dryopoids. In the latest revision of Op- tioservus (White 1978) O. canus is again illustrated with a vitta in the lateral half of each elytron. However, the couplet separating O. canus omits mention of the ‘grizzled appearance’”’ even though it is mentioned in the description given later. White notes having seen the holotype and his questioning of the paratypes col- lected by F. E. Winters (which by then were also in the California Academy of Science collection) implies that he examined them also. White concluded the Winters paratypes were teneral O. divergens (LeConte), 1874. After examining five of the six specimens in the type series and many other specimens, I am unable to explain the vittate pattern illustrated by Collier, Brown, and White. As mentioned earlier, the elytral coloration varies from entirely brassy- black to almost entirely yellow or dark orange. Humeral spots may be absent, very small, or fairly large. However, when vittate the maculae extend along the medial half of each elytron (not the lateral half as had been earlier illustrated). Sometimes the elytral suture is involved so the elytral disc is all light in color. However, the sides of the elytra are always dark. The type specimens examined appear to me to be of the bimaculate form and not at all vittate. The grizzled appearance, a product of the long pubescence, is characteristic of all color morphs and is the most distinctive character of this species. Chandler is vindicated! Specimens from PNM have a higher frequency of bimaculate forms than other populations. This may be an effect of a genetic bottleneck that perhaps occurred during a severe summer which killed all but a few individuals. Alternatively, it could be a founder effect. The aquatic dryopoid fauna of Pinnacles National Monument has thus been found by this author to include the following species: Psephenidae Eubrianax edwardsii (LeConte), 1874 Dryopidae Helichus productus LeConte, 1852 Helichus striatus LeConte, 1852 non H. columbianus Brown, 1931 Helichus suturalis LeConte, 1852 Elmidae Optioservus canus Chandler, 1954 Elsewhere I have found O. canus associated with Psephenus falli Casey, 1893, Zaitzevia parvula (Horn), 1870, Ordobrevia nubifera (Fall), 1901 and Microcyl- loepus similis (Horn), 1870. Streams in which I have found O. canus include: Monterey Co.—Piney Creek, Juan Higuera Creek, Grimes Canyon Creek, Par- tington Creek, and Arroyo Seco River; San Benito Co.— Tres Pinos Creek, Chalone Creek, and San Benito River. RESEARCH NOTES 145 Acknowledgments I thank the staff of Pinnacles National Monument for their help and comments regarding water sources within PNM. Dr. Harley P. Brown provided many helpful suggestions on this taxonomic snarl. Literature Cited Brown, H. P. 1972. Aquatic dryopoid beetles (Coleoptera) of the United States. Biota of Freshwater Ecosystems Identification Manual No. 6. Water Pollution Control Research Series, Environ- mental Protection Agency. Washington, D.C., 82 pp. Brown, W. J. 1931. New species of Coleoptera (II). Can. Entomol., 63(5):115-122. Casey, T. L. 1893. Coleopterological notices V. Ann. New York Acad. Sci., 7:281-606. Chandler, H. P. 1954. New genera and species of Elmidae from California. Pan-Pacific Entomol., 30(2):125-131. Collier, J. E. 1969. A taxonomic revision of the genus Optioservus (Coleoptera: Elmidae) in the nearctic region. Ph.D. Thesis, University of Minnesota. University Microfilms, Inc., Ann Arbor, Michigan, 59 pp. Fall, H.C. 1901. List of the Coleoptera of southern California, with notes on habits and distribution and descriptions of new species. Occ. Pap. Cal. Acad. Sci., 8:1-282. Horn, G. H. 1870. Synopsis of the Parnidae of the United States. Trans. Amer. Entomol. Soc., 3: 29-42. LeConte, J. L. 1852. Synopsis of the Parnidae of the United States. Proc. Acad. Nat. Sci. Phil., 6: 41-45. . 1874. Descriptions of new Coleoptera chiefly from the Pacific slope of North America. Trans. Amer. Entomol. Soc., 5:43-72. Leech, H. B., and H. P. Chandler. 1956. Aquatic Coleoptera, Chapter 13 in Aquatic insects of California. (R. L. Usinger, ed.), University of California Press, Berkeley, 508 pp. White, D.S. 1978. A revision of the Nearctic Optioservus (Coleoptera: Elmidae), with descriptions of new species. System. Entomol., 3:59-74. Accepted for publication 15 December 1989. William D. Shepard, Department of Biological Sciences, California State Univer- sity, Sacramento, 6000 J Street, Sacramento, California 95819. DESERT ECOLOGY 1986 A Research Symposium Twelve papers from the Desert Studies Consortium at the Academy 1986 An- nual Meeting comprise a new publication now available. Subjects include the Coachella Valley Preserve, Water Rights, Late Pleistocene Mammals, Chemical Defense Patterns of Certain Desert Plants, Off-Road Vehicle disturbances, Desert Pupfish, Plant Communities, Desert Bats, etc. Send name, address, and $29.00 per copy in check made out to The Southern California Academy of Sciences, 900 Exposition Blvd., Los Angeles, CA 90007. Bull. Southern California Acad. Sci. 89(3), 1990, pp. 147-148 © Southern California Academy of Sciences, 1990 INDEX TO VOLUME 89 Ampelisca brachyclaus n. sp., 124 Arneson, A. Charles, see Ronald J. Larson Austin, Jorja, see George W. Cox Blake, James A. and Brigitte Hilbig: Revision of Two Dorvilleid Species from the Pacific Coast of North America (Annelida: Polychaeta), 109 Blood, Brad R.: Taxonomy and Distribution of Sigmodon in California, 86 Breen, Robert T. and Mary K. Wicksten: Movement and Habitat Selection in Tagged Rock Crabs (Cancer antennarius) in Intertidal Channels at James V. Fitzgerald Marine Life Refuge, California, 10 Brasher, Anne M., see Daniel L. Castleberry Bright, Donald: Management of Hazardous Substances: An Overview, 49 Bursey, Charles R., see Stephen R. Goldberg Castleberry, Daniel L., Jack E. Williams, Georgina M. Sato, Todd E. Hopkins, Anne M. Brasher, and Michael S. Parker: Status and Management of Sho- shone Pupfish, Cyprinodon nevadensis shoshone (Cyprinodontidae), at Sho- shone Spring, Inyo County, California, 19 Cox, George W. and Jorja Austin: Impacts of a Prescribed Burn on Vernal Pool Vegetation at Miramar Naval Air Station, San Diego, California, 67 Davis, Stephen D., see Roy Stoddard Frost, Patrick, see Ingo H. Gaida Gaida, Ingo H. and Patrick Frost: A New Host in the Northern Hemisphere for the Parasitic Marine Isopod Ceratothoa gaudichaudii (Crustacea: Isopoda: Cymothoidae), 94 Goldberg, Stephen R. and Charles R. Bursey: Prevalence of Larval Cestodes (Mesocestoides sp.) in the Western Fence Lizard, Sceloporus occidentalis bi- seriatus (Iguianidae), from Southern California, 42 Hilbig, Brigitte and R. Eugene Ruff: Remarks on the genus Pettiboneria (Poly- chaeta: Dorvilleidae) with Description of Two New Species, 115 Hilbig, Brigitte, see James A. Blake Hopkins, Todd E., see Daniel L. Castleberry Larson, Ronald J. and A. Charles Arneson: Two Medusae New to the Coast of California: Carybdea marsupialis (Linnaeus, 1758) a Cubomedusa and Phyl- lorhiza punctata von Lendenfeld, 1884, a Rhizostome Scyphomedusa, 130 Mason, Waynelle, see Harrington Wells 148 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Parker, Michael S., see Daniel L. Castleberry Pettiboneia bathyalis n. sp., 115 Pettiboneia brevipalpa n. sp., 115 Reynolds, Robert E., see Barry Roth Roney, James D.: A New Species of Marine Amphipod (Gammaridea: Ampelis- cidae) from the Sublittoral of Southern California, 124 Roth, Barry and Robert E. Reynolds: Late Quaternary Nonmarine Mollusca from Kokoweef Cave, Ivanpah Mountains, California, 1 Ruff, R. Eugene, see Brigitte Hilbig Sato, Georgina M., see Daniel L. Castleberry Shepard, William D.: The Aquatic Dryopoid Beetles of Pinnacles National Mon- ument: Optioservus canus Revisited (Coleoptera: Dryopoidea: Elmidae), 143 Silber, Gregory K. and Mari A. Smultea: Harbor Porpoises Utilize Tidally-induced Internal Waves, 139 Smultea, Mari A., see Gregory K. Silber Stewart, James R., see Harrington Wells Stoddard, Roy and Stephen D. Davis: Comparative Photosynthesis, Water Re- lations, and Nutrient Status of Burned, Unburned, and Clipped Rhus laurina after Chaparral Wildfire, 26 Wells, Harrington, Waynelle Mason, and James R. Stewart: Prairie Dog Food Preference and the Photosynthetic Pathway-Selective Herbivory Hypothe- sis, 97 Wicksten, Mary K., see Robert T. Breen Williams, Jack E., see Daniel L. Castleberry Wilson, Edward C.: Mass Mortality of the Reef Coral Pocillopora on the South Coast of Baja California Sur, Mexico, 39 Wilson, Edward C.: The Tropical Colonial Stony Coral Tubastrea coccinea at Cabo San Lucas, Mexico, 137 hia The BULLETIN is published three times each year (April, August, and December) and includes articles in English in any field of science with an emphasis on the southern California area. Manuscripts submitted for publication should contain results of original research, embrace sound principles of scientific investigation, and present data 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. MANUSCRIPT PREPARATION The author should submit at least two additional copies with the original, on 8/2 x 11 opaque, nonerasable paper, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. 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All changes in galley proof attributable to the author (misspellings, inconsistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with the printer, not the Editor. CONTENTS Prairie Dog Food Preference and the Photosynthetic Pathway-—Selective Herbivory Hypothesis By Harrington Wells, Waynelle Mason, and SOMOS AR: SSCCWALE ge tanita aE Na een eee 97 Revision of Two Dorvilleid Species From the Pacific Coast of North Amer- ica (Annelida: Polychaeta) By James A. Blake and Brigitte Hilbig ...... 109 Remarks on the Genus Pettiboneia (Polychaeta: Dorvilleidae) with De- scriptions of Two New Species’ By Brigitte Hilbig and R. Eugene Ruff 115 A New Species of Marine Amphipod (Gammaridea: Ampeliscidae) from the Sublittoral of Southern California By James D. Roney |... 124 Two Medusae New to the Coast of California: Carybdea marsupialis (Lin- naeus, 1758), a Cubomedusa and Phyllorhiza punctata von Lendenfeld, 1884, a Rhizostome Scyphomedusa By Ronald J. Larson and A. Charles: AVne@SOn: js leo oe 130 Research Notes The Tropical Colonial Stony Coral Tubastrea coccinea at Cabo San Lucas, Mexico By Edward CAGES ORs Be BLO i Be seh RO BAL ace ET IO ye 137 Harbor Porpoises Utilize Tidally-induced Internal Waves By Gregory K. Silber and Mari A. SSUES Nae EA ys ENE ey ee a 139 The Aquatic Dryopoid Beetles of Pinnacles National Monument: Optioservus canus Revisited (Coleoptera: Dryopoidea: Elmidae). By William D. Shepard ............:.:.:cccceccsesec cece cece eee 143 | Kn (=), Semen eee eae NON RANE AI OREM Ts uM ey Ala chs ce ent RO 147 LIBRARY a¢R - 2 1996 MEW YORK SUIANICAL GARDEN COVER: Black-tail Prairie Dog (Cynomys ludovicianus). Page 97.