I( /'/W ISSN 0038-3872 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES ULLETIN Volume 115 Number 1 Ui iSl m iimu SI mmm fliw mmmi sn mmi NON 61001 OQ NQiONIHSVM NOliniliSNI PiNQSHiflilS m m NoiiniiiSNQOQNf is hioi ilHNIN iQNVHOIi/SNOlilSinOOlf UiiOO iMIiSNI mNOSHillNS m oQfosm.***.***.*** 115(1) 1^84 (2016) April 2016 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 © Southern California Academy of Sciences, 2016 2015-2016 OFFICERS Julianne Kalman Passarelli, President David Ginsburg, Vice-President Edith Read, Recording Secretary Ann Dalkey, Treasurer Daniel J. Pondella II and Larry G. Allen, Editors - Bulletin Brad R, Blood, Newsletter Shelly Moore, Webmaster ADVISORY COUNCIL Jonathan Baskin, Past President John Roberts, Past President John H. Dorsey, Past President Ralph Appy, Past President Brad R. Blood, Past President BOARD OF DIRECTORS 2013-2016 2014-2017 Ann Dalkey David Ginsburg Julianne K. 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All other communications should be addressed to the Southern California Academy of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 90007-4000. Date of this issue 26 April 2016 @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Bull. Southern California Acad. Sci. 115(1), 2016, pp. 1-14 © Southern California Academy of Sciences, 2016 The Return of the King of the Kelp Forest: Distribution, Abundance, and Biomass of Giant Sea Bass [Stereolepis gigas) off Santa Catalina Island, California, 2014-2015 Parker H, House*, Brian L.R Clark, and Larry G. Allen California State University, Northridge, Department of Biology, 18111 Nordhoff St., Northridge, CA, 91330 Abstract. — It is rare to find evidence of top predators recovering after being negatively affected by overfishing. However, recent findings suggest a nascent return of the critically endangered giant sea bass {Stereolepis gigas) to southern California. To provide the first population assessment of giant sea bass, surveys were conducted during the 2014/2015 summers off Santa Catalina Island, CA. Eight sites were surveyed on both the windward and leeward side of Santa Catalina Island every two weeks from June through August. Of the eight sites, three aggregations were identified at Goat Harbor, The V’s, and Little Harbor, CA. These three aggregation sites, the largest containing 24 individuals, contained a mean stock biomass of 19.6 kg/1000 vc? over both summers. Over the course of both summers the giant sea bass population was primarily made up of 1.2 - 1.3 m TL individuals with several small and newly mature fish observed in aggregations. Comparison to historical data for the island suggests giant sea bass are recovering, but have not reached pre-exploitation levels. The giant sea bass {Stereolepis gigas) is the largest teleost to inhabit nearshore rocky reefs and kelp forests in the northeastern Pacific (Hawk and Allen 2014). Though previously taxono- mically classified as a sea bass (Serranidae), the giant sea bass is actually a wreckfish, in the family Polyprionidae (Shane et al. 1996). Unlike most wreckfishes, they are a relatively shallow water species, inhabiting depths from 3 » 40 m. Their historical range is from Humboldt Bay, CA to Baja Mexico (Point Abreojos) and into the northern Gulf of California. However, they are primarily found south of Point Conception. Although, the giant sea bass is the largest mem- ber of the southern California rocky reef and kelp forest fish community, very little is known about its basic biology and life history (Allen and Andrews 2012). These fish have been docu- mented to grow over 250 kg (Domeier 2001) and live up to 76 years old (Hawk and Allen 2014). However, there are reports of giant sea bass living as old as 90 - 100 years and over 270 kg (Fitch and Lavenberg 1971), and even possibly reaching sizes of 360 kg as noted by author Charles F. Holder at the turn of the twentieth century (Holder 1910). These early reports of giant sea bass size and age remain unverified. Along with being a long-lived and slow growing species, with the exception of growing rapidly within the first year of life (Hawk and Allen 2014), they are also relatively late to mature. It is believed that giant sea bass mature between 11-13 years of age (Fitch and Laven- berg 1971). However, there have been no studies explicitly confirming age at sexual maturity. To maintain their large body mass, giant sea bass feed on a wide variety of demersal and con- spicuous rocky reef fishes as well as cephalopods and crustaceans. They have been documented to feed on rays, guitarfish, skates, flatfish, small sharks, barred sand bass, kelp bass, blacksmith, * Corresponding author: parker.h.house@gmail.com 1 2 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES ocean whitefish, sargo, sheephead, octopus, spiny lobster, cephalopods and squid (Domeier 2001, Love 2011). They are likely capable of feeding on nearly any species inhabiting nearshore rocky reefs and kelp forests off southern California, as they are the apex, tertiary megacamivore of this system (Cross and Allen 1993, Horn and Graham 2006). Like many slow growing, late maturing, large bodied marine predators worldwide (Pauly et al. 1998, Jackson et al. 2001, Dayton et al. 2002, Myers and Worm 2003), the giant sea bass popu- lation has historically been depleted due to overfishing and has been rare off southern California (Domeier 2001, Pondella and Allen 2008). During most of the twentieth century, they were highly sought after throughout the Southern California Bight and Mexico by both commercial and recreational fishermen. During the early twentieth century, the commercial fishery which began using hand lines had switched to gill nets providing peak landings during the early 1930’s at over 114mt before the crash of the commercial fishery off southern California in 1935 to under 10 mt (Crooke 1992). The commercial fishery of giant sea bass taken from Mex- ican waters had greater landings and durability than those off southern California. Peaking in the early 1930’s at over 362 mt with a steady decrease throughout the 1960’s (Crooke 1992). The recreational fishery for giant sea bass off southern California peaked in 1963, and in Mexico in 1973. That these peaks in recreational landings were after the crash of the commercial fishery is due to the later development of the recreational fishery, and not the population size itself (Domeier 2001). By the mid 1970’s, several boats would target presumed spawning aggrega- tions sites throughout the month of July off southern California and Mexico, consistently land- ing high numbers (Crooke 1992) and in one case up to 255 fish in three days (Domeier 2001). Likewise, during the 1960’s and 70’s the practice of spearfishing grew in popularity. The gregar- ious and bold disposition of giant sea bass did not help this apex predator against the increasing numbers of spearfishers, as they were easy targets and landed at high frequencies (Fitch and Lavenberg 1971, Crooke 1992). This combination of various fishing pressures led to their near disappearance during the 1970’s (Pondella and Allen 2008), and by 1981 both southern California and Mexico landings dropped below 5 mt (Crooke 1992, Domeier 2001). In 1981, a law was passed prohibiting the take of any giant sea bass off California, with the exception of two fish per vessel trip for com- mercial fishermen using gill or trammel nets, and the moratorium was put into effect in 1982. This law was later amended in 1988, allowing one incidental fish per commercial fishing trip off California waters. However, though this amendment limited the number able to be sold in California by commercial fishermen, it still allowed fishing via gill and trammel nets over near- shore rocky reefs and kelp forest habitat (Pondella and Allen 2008). These nearshore habitats that were targeted are those used by giant sea bass, especially during aggregation months from May - October, and the incidental bycatch of giant sea bass was discarded at sea (Crooke 1992, Domeier 2001), or rumored to be shared among commercial fisherman. Due to concerns over the viability of the giant sea bass population off southern California, this species was red listed in 1996 by the International Union for Conservation of Nature (lUCN) as a critically endangered species (Cornish 2004). It is rare to find evidence of a long-lived, slow growing, and late maturing species recovering after being strongly affected by overfishing (Hutchings 2000). However, after the gill net fishery was banned within three nautical miles of the mainland and one nautical mile of the islands with Proposition 132 in 1994, the population began to recover (Pondella and Allen 2008). After being seldom seen in southern California from the 1970s - 1990s (Domeier 2001), and not being observed in quarterly surveys by the Vantuna Research Group of Occidental College off the Palos Verdes coast between 1974 - 2001, giant sea bass began to be observed in 2002, and have been seen to the present day (Pondella and Allen 2008). Likewise, incidental commercial RETURN OF GIANT SEA BASS TO CATALINA 3 catch and CPUE from the Ocean Resource Enhancement Hatchery Program (OREHP) scientific gill net surveys showed a significant positive increase from 1995 - 2004, an increase that was not correlated to fluctuations in environmental factors (Pondella and Allen 2008) . These findings allude to a nascent return of giant sea bass within the Southern California Bight. Giant sea bass frequented yearly site-specific aggregations for presumed spawning purposes in the past, as fishermen targeted and depleted these areas during the 1970’s (Crooke 1992). Due to the elimination of previous spawning aggregations and the majority of the southern California giant sea bass population, modem day locations of aggregation sites are largely unknown. For conspicuous aggregation sites to reappear it is likely that population numbers would have to reach a certain abundance (Domeier 2001). However, anecdotal reports by the recreational dive community today suggests that historical spawning aggregations are returning primarily off La Jolla, Santa Catalina Island, and Anacapa Island, California. Surveying spawning aggre- gation sites allows for a unique opportunity to access a larger percentage of the reproductive population that would otherwise be spread over a greater geographic distribution (Johannes et al. 1999, Whaylen et al. 2004, Heppell et al. 2012). Furthermore, with information on a spawning aggregation biomass, through a length-weight relationship for the species and an estimate of total abundance, the spawning stock biomass of a species can be estimated (Jennings et al. 1996). Our study applies underwater visual censuses (UVC) using length calibrated lasers for more precise size estimation (Gingras et al. 1998, Colin et al. 2003, Heppell et al. 2012) to provide the first population assessment of the endangered tertiary carnivore, the giant sea bass, of the rocky reefs and kelp forests off southern California at Santa Catalina Island, CA. The objectives of this study were to 1) identify and document spawning aggregation sites and peak aggregation peri- ods throughout the summers of 2014 and 2015; and 2) establish baseline mean densities, stock biomass, and length/biomass distribution frequencies to compare with historical fish surveys. Materials and Methods Study Sites Eight sites were surveyed off Santa Catalina Island, CA during the summer of 2014 (6/9/14 to 8/13/14) and 2015 (6/11/15 to 8/11/15) (Fig. la). In an attempt to get both a windward and leeward representative sample for the island, the eight sites were located at Johnson’s Rock (33°28'37.08" N lat. 118°35'22.57" W. long.). Little Geiger (33°27'27.62" N lat. 118°30' 51.03" W. long.). Empire Landing (33°25'59.96" N lat. 118°26'52.44" W. long.), between Twin Rocks and Goat Harbor (33°25'04.49" N lat. 1 18°23'38.24" W. long.), Italian Gardens (33°2439.92" N lat. 118°22'32.50" W. long.). Casino Point (33°20'58.68" N lat. 118°19' 30.56" W. long.), The V’s (33°18'45.94" N lat. 118°22'11.38" W. long.), and Little Harbor (33°23'08.10" N lat 118°28'48.94" W. long.). Sites averaged a distance of 7km apart. Each of the eight sites contained habitat presumed suitable for giant sea bass aggregations based on characteristics of the Long Point State Marine Reserve (SMR) put into effect to protect the best known site for giant sea bass in southern California (CA MLPA South Coast Project 2009) . Each site consisting of deep (>18 m) rocky reefs, and reef edges, where Macrocystis kelp forests were present. Of the eight sites, four are thought to be possible giant sea bass aggre- gation sites based on historical records and reports from the recreational diving community (The V’s, Casino Point, Italian Gardens, and Goat Harbor). The Vs site was not surveyed in 2015 due to high surge and low visibility throughout the season. 4 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 1. Sampling sites and methods: A) Location of the eight sites surveyed off Santa Catalina Island, CA in the summers of 2014 and 2015; B) Image of a dive propulsion vehicle (DPV) with mounted length calibrated lasers and GoPro Hero 3 Black Edition video camera used for giant sea bass surveys; and C) Image of a giant sea bass showing broadside length calibrated laser markings at 10.2 cm. Survey Methods Surveys at each site were conducted every two weeks for two months in 2014 and 2015 for a total of 64 survey days. Each visual survey was conducted from 10:00 to 14:00 and consisted of five, three-minute, 100 m x 10 m SCUBA transects (1,000 m^) using Sea Doo Vs Supercharged Plus Sea Scooter diver propulsion vehicles (DP Vs). DP Vs were outfitted with two parallel waterproof length-calibrated lasers, set at 10.2 cm apart, and a mounted GoPro Hero3 Black Edition video camera (Fig. lb). Before surveys began, divers trained using the DPVs in combi- nation with timed fin kicks to cover 100 m in three minutes. The five timed transects per site were spaced at least 50 m apart with each transect randomly stratified in depth from 28 - 6 m to extensively survey the reef at each site. Along each transect, giant sea bass occurring in front of divers and within the transect area were counted, and their size estimated to the nearest 25 cm. Surveys for giant sea bass covered a large area during a short amount of fixed time to RETURN OF GIANT SEA BASS TO CATALINA 5 aid in reducing biases that may arise during non~instantaneous UVC of large mobile fishes (Ward-Paige et aL 2010, McCauley et al. 2012). In addition, transects were video recorded to help identify separate individuals between sampling periods by size, differences in morphology, physical markings, and spot patterns. Upon the conclusion of each transect, size-surveys were done by video recording individuals observed at a 90° angle to the video camera with the parallel lasers spaced 10.2 cm apart (Gingras et al 1998, Colin et al 2003, Heppel et al. 2012). To reduce possible size estimation error, giant sea bass recorded during size surveys were measured within 2.5 - 3 m of the indivi- dual fish. Images of fish from the size-survey videos that displayed broadside and perpendicular to the video camera with visible measurement laser markings (Fig. ic) were digitally captured and length (cm SL and TL) were estimated using the software program Image! (http://im.agej. nih.gov/ij/). The lengths obtained from, the size surveys were used to validate size estimations taken during transects. Lengths were converted to biomass (kg/ 1000 m^) using the length-weight relationship recently published for this species: kg = (0.0000001)*(SL (Williams et al. 2013). Age of sized individuals was back-calculated using the inverse of the published von Bertalan% growth curve (von Bertalanffy 1938) for giant sea bass: It = 2026.2(1 (Hawk and Allen 2014). The 2014 surveys were conducted during 6''9 6/24, 6/28 - 7/12, 7/15 ■■ 8/2, and 8/ 4 - 8/13/2014 while the seven sites in 2015 were conducted during 6/1 1 - 6/20, 6/22 - 7/10, 7/1 1 -7/31, and 8/4-8/11. In order to provide a historical perspective on the population off Santa Catalina Island, giant sea bass recorded on subtidal surveys conducted from 1965-2013 by the Channel Islands Research Program (CIRP) were generously provided by Dr. Jack Engel (UCSB). CIRP surveys consisted of divers visually surveying the reef between 4 - 2 1 m in depth for all algae, macro- invertebrates, and fishes within a timed period. Organisms were identified and abundances were estimated on a relative scale from 1 (rare) to 4 (abundant). On surveys where giant sea bass occurred the number of individuals was noted. Statistical Analyses The abundance (#/transect) and biomass (kg/ 1000 m^) estimates of giant sea bass included many zeros and did not fit the assumptions of nomiality required for parametric analyses. Numerical and biomass densities of giant sea bass for Site (fixed factor: 8 levels), Year (fixed factor: 2 levels), and Sampling Period (fixed factor: 4 levels) were compared using permuta- tioiial analysis of variance with PERMANOVA+ for PRIMER-E ver. 6 (Anderson 2001, Anderson and Millar 2004) with individual transects used as samples. The Dwass-Steel- Chiitchlow-Fligner Test for All Pairwise Comparisons was then used to test for differences between sites. For length frequency and biomass distribution analysis, lengths (mm TL) were grouped into 100 mm increments to investigate the length, biomass, and age class frequency distributions of the surveyed giant sea bass population off Santa Catalina Island for both the summer of 2014 and 2015. Length Frequencies of giant sea bass encountered in 2014 and 2015 were compared with a Noe-parametric Kolmogorov-Smimov Test using SYSTAT 13 (SYSTAT Software, Inc). Results Giant sea bass numerical densities (no. fish/1000 m^) were not statistically significant among the four sampling periods {Pseudo-F— 0.92, P(perwi)=0AK}. Despite this lack of significance, total number of individuals observed during surveys in 2014 peaked in late-Iuly and in early July in 2015 (Figure 2) Similar to numbers, biomass (kg/1000 m^) was not statistically different 6 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Number per Transect P(perm) -0.47 late-June early-July late-July early-Aug Biomass Density (kg/1,000 m^) 40 35 tn 30 fN 25 m E 20 g 15 ro ^ 10 ■ 2014 □ 2015 P(pennr0.45 \ 1^ I A late-June early-July late-July Sampling Period early-Aug Fig. 2. Mean number of giant sea bass per two-week sampling period (top), and (bottom) mean spawning stock biomass densities (kg/ 1000 m^) of giant sea bass per two-week sampling period. Error bars represent 2 standard errors. No significant differences {P(pemi) = 0-47; 0.45) were found in temporal distribution of giant sea bass numbers or biomass in either year. among the four sampling periods in either year {Pseudo-F = 0.92, P(perm) = 0.45). However, biomass density (Fig. 2) was consistent among the survey periods in both years with the excep- tion of early August when biomass density peaked in 2014 and decreased in 2015. Giant sea bass were observed at seven of the eight sites around the island (Little Geiger, Empire Landing, Goat Harbor, Italian Gardens, The V’s, and Little Harbor). No giant sea bass were observed at Johnson’s Rock. Numbers (Figure 3: Pseudo-F = 5.88; P(perm) < 0.001) and biomass (Fig. 3: Pseudo-F = 5.87; P(perm) < 0.001) differed significantly among sites over both summers of sampling. In the summer of 2014, aggregations were found at Goat Harbor, The V’s, and Little Harbor. The site containing the largest number of giant sea bass was The V’s, where 23 were seen on the second sampling and 24 on the third sampling in 2014. Little Harbor and Goat Harbor had the next highest numbers and spawning stock biomass. RETURN OF GIANT SEA BASS TO CATALINA 7 Fig. 3. Mean number of giant sea bass per transect at each sampling site (top) and (bottom) mean spawning stock biomass densities (kg/1000 m^) of giant sea bass per site during the summers of 2014 and 2015. Sites are arranged the NE end clockwise around the island. Letters (a-e) denote sites not statistically different from one another. (* - no transects conducted in 2015 at the Vs). An aggregation of ten fish was observed on transects at Little Harbor, while at Goat Harbor an aggregation of six was found. Mean biomass was higher at Goat Harbor (81.2 + 29.8 kg/ lOOOm^) than Little Harbor (34.0 + 18.67 kg/ lOOOm^) due to larger individuals aggregating at Goat Harbor. In 2015, Goat Harbor was the only site to contain an aggregation. The Goat Harbor aggregation averaged 8 giant sea bass per sampling period (3, 19, 7, and 4 individuals). Biomass density at Goat Harbor ranged from 11.03 to 66.67 and averaged 39.67 kg/1000 m^ in 2015. The remaining sites where giant sea bass were surveyed contained solitary individuals or a single pair. Size of surveyed giant sea bass ranged from 0.9 - 2.75 m TL. According to the established age-length curve for giant sea bass (Hawk and Allen 2014), the smallest individual (0.7 m TL) was estimated to be 8 years old. The length frequencies of separate individuals that were 8 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 4. Length frequencies (mm TL) of separate giant sea bass observed during survey transects in the summers of 2014 and 2015 off Santa Catalina Island. Arrows indicate presumed size at maturity based on estimate size/age estimates from Fitch and Lavenberg (1971). not significantly different in 2014 and 2015 (K-S test; p = 0.258) showed the typical giant sea bass at Santa Catalina Island in 2014 and 2015 to be 1.2 - 1.3 m TL (Fig. 4). Flowever, a large portion (25%) of the population’s biomass was found in individuals between 1.9 and 2.1 m TL (Fig. 5). The largest giant sea bass observed occurred in late June in both years. The 1994 to 2003 year-classes dominated the giant sea bass population observed in the summers of 2014 and 2015 (Fig. 6). Based on ages back-calculated from measurements of total length, these eight year-classes constituted 60% of all the giant sea bass observed. Another 16% of the individuals recruited between the years 1982 and 1993 with the remainder recruiting sporadically back to 1954. Overall mean biomass of giant sea bass off Santa Catalina Island during the summer was 25.14 + 6.57 kg/1000 m^ in 2014 and 11.96 + 6.28 kg/1000 m^ in 2015, with an overall mean biomass of 19.57 ± 4.64 kg/1000 for both summers. The historical CIRP survey RETURN OF GIANT SEA BASS TO CATALINA 9 Biomass by Siie 2014-15 Siie Class fmmTL| Fig. 5. Total biomass (kg) distribution per size class (mm TL) of giant sea bass observed during survey transects in the summers of 2014 and 2015. data from 1965”2013 (Figure 7) show one giant sea bass being observed during surveys in 1966 with a 29-year absence until 1996. After 1996, giant sea bass were observed in 1997, 2000, 2001, 2002, 2003, 2006, 2007, 2010, and 2011. The highest number of giant sea bass on CIRP surveys occuired in 2001 with 11 individuals observed. All Giants by Year Class, 1950-2015 30 25 ^ 20 « m 1 3 z 10 ■ 5 - Gill Net Ban Moratorium 1 I nri CT10101010101010101010101010101CTIOOOOOO Year Fig. 6. Year-class strength for giant sea bass for 1950 to 2015 back-calculated from in situ measurements of total length converted to age after (Hawk and Allen 2014). Arrows indicate the year that the fishing moratorium was declared (1982) and the year of the Proposition 132 Gill Net Ban (1994) from coastal waters in southern California. 10 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES momomomomo ^r^r^wooo1moor^ oiOTOioimoimooo Year Fig. 7. Giant sea bass observed on SCUBA fish surveys conducted by the Channel Islands Research Program (CIRP) at Catalina Island from 1965-2013. Data courtesy of Dr. John Engle (UC Santa Barbara) funded by the Tatmae Foundation. We observed 44 separate individuals on transects in 2014 and 32 in 2015 based on differences in size, morphology, physical markings, and spot patterns of giant sea bass. If we assume the estimated numerical densities per are accurate over the depth range of the eight established sites, then about 87 linear km (86,905 m) of coastline around Santa Catalina Island held about 49 giant sea bass in the summer of 2014 and about half that number, about 24, in summer 2015. Similarly, biomass densities estimated that 2.1 metric tons (mt) of giant sea bass occurred around Catalina in 2014 with about 1.0 mt occurring in 2015. The number of individual giant sea bass identified by divers and the number of giants estimated from transect densities were remarkably similar in both years of the study. Discussion Altogether, this study provides evidence of the return of giant sea bass to the rocky reefs and kelp forests off Santa Catalina Island, and possibly the Southern California Bight, by document- ing new spawning aggregation sites, considerable stock biomass, newly mature individuals recruiting to aggregations, and a large community presence at the island. Overall, abundance and biomass of giant sea bass did not differ greatly among the four sampling periods in 2014 and 2015. The large variation in numbers and biomass during the four sampling periods for both years can be largely attributed to the patchy distribution that resulted in the high number of transects where no giant sea bass were observed. Of the eight sites, at least three were identified as giant sea bass aggregations off Santa Cat- alina Island, CA. These sites were located on both the leeward (Goat Harbor) and windward (The V's and Little Harbor) side of the island. Goat Harbor is the only of these three sites where aggregations were encountered in both years of this study. Goat Harbor is also the only aggre- gation residing in a Marine Protected Area (MPA) as of 2012. The placement of the Long Point State Marine Reserve (SMR) was to protect the best-known aggregation area for giant sea bass off southern California from Long Point to Goat Harbor (CA MLPA South Coast Project 2009), and is a popular site for recreational divers. However, though this site had a consistent RETURN OF GIANT SEA BASS TO CATALINA 11 aggregation during each of the four sampling periods, it did not possess the largest giant sea bass aggregation. The largest suspected spawning aggregation was found at the V’s in 2014 with a total of 23 and 24 individuals occurring on transects during the second and third sampling period at 1 8 m depth. The individuals at the Vs were typically larger, 1 .2 - 2.3 m TL. Through- out the summer of 2014 human presence was minimal at this site, as the V’s is located in a more remote area of the windward side of the island. However, commercial squid fishing vessels were observed in close proximity to the reef where the giant sea bass aggregation was observed (P.H.H., personal obs.). Unfortunately, the V’s site was largely inaccessible to our divers in 2015. The third aggregation site was located on the reefs just outside and west of Little Harbor. The consistency of this aggregation varied. On the third and fourth sampling periods during 2014, 6 and 10 giant sea bass were observed respectively. The aggregation at Little Harbor in 2014 consisted primarily of smaller individuals (eight individuals under 1.2 m TL) compared to the other two aggregation sites at Goat Harbor and the V’s. No giant sea bass were seen dur- ing surveys at Johnson’s Rock in either summer. The Little Geiger and Empire Landing sites contained solitary individuals that were observed sporadically over the two years. Italian Gar- dens which is also inside the Long Point SMR had either solitary individuals or a single pair of giant sea bass that were likewise only observed sporadically. If the giant sea bass population off southern California is indeed recovering, then there is likely to be a larger proportion of smaller and younger fish within the population, which could manifest as a positive skew in length frequencies of the population (Heppell et al 2012). In the case of a spawning aggregation, smaller size classes represent newly mature fish entering the reproductive population. Our results do not show a strong positive skew as the majority of reproductive giant sea bass off Santa Catalina Island were -1.3 m TL and were estimated to be 18 ” 19 years-old. However, smaller individuals were observed during surveys in presumed spawning aggregations off the island. These individuals were estimated to be 10 - 11 years old. Age at sexual maturity has not been adequately explored for giant sea bass, however, Fitch and Lavenberg (1971) estimated sexual maturity to begin between 11 and 13 years of age. Our find- ings of young giant sea bass within presumed spawning aggregations support the Fitch and Lavenberg (1971) estimates. Based on year-class strength estimates (Fig. 6), these young fish are likely new recruits to the reprc*ductive population off Santa Catalina Island that were bom after the 1994 Proposition 132 gill net ban in coastal waters. Our results also suggest that these young individuals were able to find site-specific suspected spawning aggregations that were likely once decimated by overfishing. Although a large portion of the presumed reproductive population censused in the present study was made up of individuals 1.2 - 1.3 m in total length, this size class did not account for the largest portion of the stock biomass. The size class with the peak biomass was older (esti- mated to be 32 " 35 years old) and larger (1.9 - 2.0 m TL) individuals. This skew in total biomass distribution toward the larger size classes was also due to several behemoth individuals. In 2014, the largest individual on transect was measured at 2.3 m TL (1.9m SL) with a back- calculated age of 67 years old and 177.9 kg. However, this was not the largest giant sea bass measured in 2014. An individual that was measured during underv/ater observations, but did not occur within a survey transect was seen ar Goal Harbor and measured 2.70 m TL (380 kg). Similarly in 2015, at the same site (Goat Harbor) and sample period (late June) an indivi- dual was measured at 2.75 m (381 kg), and was obsewed on transect. It is possible that these two observations in 2014 and 2015 could eithei be ul ihe same individual or two separate indi- viduals. In either case, these would be the largest giant sea bass ever measured, and supports early, unverified accounts of much older and larger giant sea bass (Holder 1910). Giant sea bass in this size range are over the L» presented in Hawk and Allen (2014). Although their 12 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES age cannot be predicted accurately, it is not inconceivable that fish of this size are over 100 years of age. Similar to Pondella and Allen (2008), fish survey data collected by the Channel Islands Pvesearch Program (CIRP) beginning in 1964 suggests a similar trend to the Palos Verdes coast in number of giant sea bass sightings off Santa Catalina Island. From the CIRP surveys only one giant sea bass was obseiwed until the late 1990's and early 2000's to present day. However, although these data suggest a recent return of giant sea bass, historical accounts document fish- erman consistently taking 70 - 100 giant sea bass from summer aggregations (Domeier 2001), suggesting that present day aggregation densities are still well under historical levels. The two aggregation sites containing the highest abundance (the V’s) and younger individuals (Little Harbor) of the three spawning aggregation sites are currently in unprotected areas where fishing is allowed. Pre-exploitation biomass for the entire southern California population of the giant sea bass has been estimated to be 1,179 mt (Ragen, 1990). For comparison, our biomass estimates of 2.1 mt and 1.0 mt of standing stock biomass off Catalina Island in 2014 and 2015 were a Ml three-orders of magnitude lower. If our current estimates of biomass of Catalina Island are extra- polated to the entire southern California coastline, it appears that the current standing stock of the giant sea bass population off southern California, although returning, falls far short of what the natural stocks were prior to exploitation. As others have often cited (cf., Domeier, 2001), it may well be decades before the giant sea bass population recovers to levels appropriate for renewed commercial exploitation. Despite giant sea bass being a protected species they are often susceptible to barotrauma when caught incidentally. Schroeder and Love (2002) estimated how incidental catch and release mortality of giant sea bass could affect population sizes. Their estimates suggest that 1 00 giant sea bass, at a standard catch and release mortality rate of 20%, could be completely eradicated through incidental catch and release in just 16 years assuming no immigration. With the aggregation sizes found in our study, the largest being an aggregation of 24 fish, this incidental catch and release mortality rate could decimate the reproductive population off Santa Catalina Island during the summer spawning months. Seasonally established MPAs at identified giant sea bass spawning aggregation sites, similar to those set in place to protect Nas- sau grouper spawning aggregations in the Caribbean, could aid in reducing the incidental catch of giant sea bass near these areas. Furthermore, monitoring of aggregations after baseline esti- mates would allow temporal tracking of numerical densities, biomass, and population dynamics of giant sea bass off Santa Catalina Island and other sites within the Southern California Bight. Our study provides an effective way to survey these aggregations, and ftirther surveys of the kelp forest community are needed to document what potential influences a return of a long absent top predator may have to the dynamics of this ecosystem. Acknowledgments We want to express our gratitude to the following people for making this research possible. Thank you to Drs. Mark Steele, Mia Adreani, and Peter Edmunds for their invaluable input, insight, and review of this research. Our upmost gratitude goes to those who helped with the challenging field work schedule including Michael Abernathy, Matt Jelloian, Juan Aguilar and the staff at the USC Wrigley Institute for Environmental Studies. Special thanks to Kelcie Chiquillo for field assistance and support. We are also gratefiil to Drs. Steve Dudgeon (CSUN), Jack Engle (UCSB), Milton Love (UCSB), Douglas McCauley (UCSB), Ed Parnell (SIO), and three anonymous reviewers for their advice and input to various aspects of this study. RETURN OF GIANT SEA BASS TO CATALINA 13 This research was supported by funds from the CSUN Nearshore Marine Fish Research Pro- gram (NMFRP), CSUN Research and Graduate Studies, Sigma Xi GIAR, and USC Wrigley Summer Fellowships. Literature Cited Alien, L.G., and A.H. Andrews. 2012. Bomb radiocarbon dating and estimated longevity of giant sea bass (Stereoiepis gigas). Bull. So. Calif. Acad. Sci., 11 1(1): 1-14. Anderson, M.J. 2001. A new method for non"parametiri.c multivariate analysis of variance. Austral. Ecol, 26:32M6. Anderson, M.J. and R.B. Millar 2004. Spatial variation and effects of habitat on temperate reef fish assemblages in northeastern New Zealand. J. Exp. Mar. Biol. Ecol, 305:191-221. Colin, P.L., Y.J. Sadovy, M.L. Domeier. 2003. Manual for the study and conservation of reef fish spawning aggre- gations. Soc. Conserv. Reef Fish Aggregations, Special Publication, 1:9-70. Cornish, A. 2004. Stereoiepis gigas. The lUCN Red List of Threatened Species. Version 2014. . 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Appl, 22:385-392. doi: 10. 1890/1 1-1059.1 Myers, R.A., and B. Worm. 2003. Rapid worldv/ide depletion of predatory fish communities. Nature, 423 (6937):280-283. Pauly, D., V. Christensen, J. Dalsgaard, R. Froese, and F. Torres. 1998. Fishing down marine food webs. Science, 279(5352):860-863. 14 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Pondella, D.J. II and L.G. Allen. 2008. The decline and recoveiy of four predatory fishes from the Southern California Bight. Mar. Biol, 154:307-313. Ragee, T. J. 1990. Pre-exploitation abundances for the white seabass of (Airactoscion nobilis), yellowtail (Serioia lalandei), and giant sea bass {Stereolepis gigas) off southern California. In The estimation of theoretical population levels for natural populations. Doctoral dissertation, University of California, San Diego, 176 pp. Schroeder, D.M., and M.S. Love. 2002. Recreational fishing and marine fish populations in California. CalCOFI Rep., 43:182-190. Shane, M.A., W. Watson, and H.G. Moser. 1996. Polyprionidae: Giant sea basses and wreckfishes. Pp. 873-875 in The early stages of fishes in the California Current Region (H.G. Moser, ed.) Coop. Fish. Invest. Atlas No. 33. Allen Press Inc., Lawrence, Kansas. 1505 pp. von Bertalanffy, L. 1938. A quantitative theory of organic growth. Human Biol, 10:181-243. Ward-Paige, C., J.M, Flemming, and H.K. Lotze. 2010. Overestimating fish counts by non-instantaneous visual censuses: consequences for population and community descriptions. PLoS One, 5(7):ell722. Whaylen, L., C.V. Pattengill-Semmens, B.X. Semmens, P.G. Bush, M.R. Boardman. 2004. Observations of a Nas- sau grouper, Epinephelus striatus, spawning aggregation site in Little Cayman, Cayman Islands, including multi-species spawning information. Environ. Biol. Fishes, 70:305-313. Williams, C.M., J.P. Williams, J.T. Claisse, D.J. Pondella II, M.L. Domeier, and L.A. Zahn. 2013, Morphometric relationships of marine fishes common to central California and the southern California bight. Bull. So. Calif. Acad. Sci., 1 12(3):2 17-227. Bull. Southern California Acad. Sci. 115(i), 2016, pp. 15^0 © Southern California Academy of Sciences, 2016 Nudibranch Range Shifts Associated with the 2014 Warm Anomaly in the Northeast Pacific Jeffrey H. R. Goddard,^* Nancy Treneman,^ William E. Pence^^ Douglas E. Mason,^ Phillip M. Dobiy,^ Brenna Green, ^ and Craig Hoover^ ^Marine Science Institute, University of California, Santa Barbara, CJ 93106^6150 ^Oregon Institute of Marine Biology, Charleston, OR 97420 ^Alameda County Office of Education, Hayward, CA 94544 ^Science Department, California High School, San Ramon, CA 94583 ^California State University, East Bay, Hayward, CA 94542 ^Department of Invertebrate Zoology and Geology, California Academy of Sciences, San Francisco, CA 94118-4503 '^Biological Sciences Department, California State Polytechnic University, Pomona, CA 91768 Abstract-^Th.Q Northeast Pacific Ocean was anomalously warm in 2014, despite ENSO“ neutral conditions in the tropical Pacific. We document northern range shifts associated with this anomaly for 30 species of nudibranclis and other shallow-water, benthic lieterobranch gastropods from southern California to southern Oregon, Nine of these (Placida cremoniana, Trapania velox, Doriopsilla fulva, Janolus anulatus, J. barbarensis, Flabellina cooperi, Anteaeolidiella chromosoma, A. oliviae, and Noumeaella rubro- fasciata) were recorded from new northernmost localities, while the remainder were found at or near northern range limits which we show were established mainly during El Nino events. All 30 species have plankto trophic laival development, and six were observed spawning at northern localities, increasing the likelihood that their ranges will continue to shift poleward as the strong 2015-16 El Nino develops. Notable among these m^as Okenia rosacea, usually found south of San Francisco and last observed in Oregon as a single specimen found during the 1997-98 El Nino. In 2015 this bright pink nudibranch reached high densities and was observed spawning throughout northern California and into southern Oregon. Okenia rosacea is therefore poised to exploit abundant prey resources previously out of its reach in northern Oregon and Washington. Our results not only demonstrate a striking biological response to the 2014 warm anomaly in the North Pacific Ocean, but also support early physical indications of a larger regional climate shift, one reinfoiced by long-term global warming. Combined with historical data, these results highlight how shallow-water nudibranchs, with their planktotrophic larvae, short life cycles, conspicuous coloration, and accessibility are excellent biological indicators of ocean climate in the region. * Corresponding author; goddard@lifesci.ucsb.edu 15 16 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES NOAA/NWS/NGEP/EMC yarine Modeling and Analysis Branch Oper H.R. RTG_SST_HR Anomaly (0.083 deg X 0.083 deg) for 01 Feb 2015 160W HOW 120W 100W SUN FEB 1 2015 Fig. 1. SST anomalies in the NE Pacific Ocean, 1 Febraaiy 2015. Source; NOAA The winter of 2013-14 in the NE Pacific Ocean was unusually warm, with peak sea surface temperature (SST) anomalies greater than 2.5°C observed over a large part of the Gulf of Alaska in February (Bond et al. 2015). By May the warm SSTs had spread to the coastal zone and southward, where they merged with similarly warm anomalies resulting in part from reduced upwelling off southern California and northern Baja California (Leising et al 2014). Despite the lack of even a moderate El Nino, the remainder of 2014 and early 2015 was marked by warmer than normal SSTs along the entire west coast of North America (Fig. 1) (Leising et al 2014; CCIEA 2015). As indicated by the Pacific Decadal Oscillation (PDO) index, multivariate El Nino Southern Oscillation (ENSO) index (MEI), and the North Pacific Gyre Oscillation (NPGO), the entire region had undergone a major phase shift in ocean conditions (Leising et al 2014), potentially similar to the 1976-77 climate shift in the North Pacific Ocean (Miller et al 1994; Mantua and Hare 2002). Indeed, the value of the PDO index for December 2014 was the highest ever recorded for that month (Heberton 2015). The California Current Ecosystem encompasses the coastline from Oregon to Baja California Sur. In this system positive values of the MEI and PDO are associated with warmer SSTs and reduced upwelling, and negative values of the NPGO are associated with weakened southerly flow and reduced nutrients and primaiy production (reviewed by CCIEA). As demonstrated by Schultz et al. (2011), these conditions (which include moderate to strong El Nino events) are correlated with increased intertidal abundance of nudibranch gastropods in California, NUDIBRANCH RANGE SHIFTS m NORTHEAST PACIFIC 17 especially southern species, and can sometimes also force long-term shifts in their northern range limits (Goddard et al. 2011). Our observations in 2014 and early 2015 of nudibranchs and other sea slugs in long-term intertidal study sites in southern and central California, combined with dive reports from south- ern California, and posts on various photo and observation-sharing websites, indicated that simi- lar range shifts and increases in abundance of southern species of sea slugs were occurring again in California. Most conspicuous among these was the dramatic increase in intertidal density in northern California of the bright pink dorid nudibranch Okenia rosacea (Kraybill-Voth 2015; Stephens 2015). We therefore intensified sampling of sea slug populations in northern Califor- nia and southern Oregon, and alerted colleagues from northern California to Washington about changes they might expect to see in the composition of the nudibranch fauna. Here, we sum- marize our findings, not just to document changes in distribution and abundance associated with the unusual, non-El Nino related warming of 2014, but also because we predict that many of the species observed in this study will likely be carried to unprecedented northerly lati- tudes by the strong 2015-16 El Nino in the Pacific Ocean (Climate Prediction Center/ NCEP 2015). Materials and Methods To quantify the abundance of nudibranchs and other sea slugs we conducted timed counts in the low intertidal zone at 28 sites from Los Angeles County, California to Coos Couiify, Oregon (Fig. 2, Table 1). Additionally, CH used SCUBA to sample four subtidal sites in Santa Barbara, Ventura, and Los Angeles Counties, and BG used SCUBA to sample Point Cabrillo, Monterey and the Santa Cmz Wharf. We also qualitatively sampled the sides of floating docks in Charles- ton, Oregon and the Santa Cmz and Monterey Harbors in California. Inteitidally we focused on pools, the under-rock microhabitats supporting the sessile prey of nudibiandis, and green macro-algae known to support sacoglossan sea slugs. Intertidal surveys usually started approxi- mately 1 h before low tide and lasted 2-3 h depending on the size of the site and the numbers of observers, which varied from one to seven. Subtidal surveys lasted approximately 60 minutes each, with the number of trained observers vaiying between one and three. For analyses and pre- sentation, data for each species from the timed counts were converted to number of individuals h”^ observer”” ^ When possible we collected vouchers specimens and deposited them in the Invertebrate Zoology Collection at the California Academy of Science (CAS). These are refer- enced below by CASIZ followed by the collection number. For additional reports of unusual occurrence, we monitored posts on Flickr, iNaturalist, OCDiving, SoCal Uiideiwater Photogra- phers on Facebook, and Bodegaliead.blogspot.com. From these internet-based sources we used only records accompanied by an image and the date and locality of observation. With two exceptions, we chose a cut-off date of 31 August 2015 for inclusion of new^ obser- vations from all sources. The exceptions were (1) large Aplysia vaccaria observed subtidally on the Monterey Peninsula in mid-September 2015 (see Results), and (2) a large specimen of the nudibranch Janolus barbarensis found on 12 September 2015 (see Results). These individuals would have recraited months earlier to the benthos, prior to the Ml impact in the region of the developing 2015-16 El Nino. While examining our results it became apparent that many of the species we observed in this study have only been observed at some of our northerly study sites during previous warm water events, especially moderate to strong El Ninos. Where appropriate we describe these patterns of occurrence, utilizing historical records and following Null’s (2015) classification of El Nino events as weak, moderate or strong. Additionally, our study sites include those described by 18 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Charie^osi Squaw b. & Gregory P«nt Cape Arago ■ five mile Poirt Cape Blanco CapeSeba^an Whfakey Creek House Rock Oregon Schultz et al. (2011) and Goddard et al. (2011), and where appropriate we follow their geo- graphic classification of nudibranch species in the region as southern, northern, or widespread. We obtained historical records of occurrence from: (1) published literature, (2) the online database for the Invertebrate Zoology Collection at the California Academy of Science (CAS), (3) historical time series provided by Schultz et al. (2015) and Goddard et al. (2015), and (4) unpublished field accounts of California eudibranchs by James R. Lance, Richard A. Roller, and Gary R. McDonald. The field accounts of Lance and Roller are housed at CAS, NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 19 Table 1. Location of study sites shown in Figure 1. All sites intertidal unless indicated otherwise. Site Geographic coordinates Charleston Boat Basin (docks) 43.3453, -124.3220 Gregoiy Point 43.3400, -124.3749 Squaw Island 43.3375, -124.3774 Cape Arago North Cove 43.3094, -124.3986 Middle Cove 43.3026, -124.4007 South Cove 43.3026, -124.3988 Five-mile Point 43.2199, -124.4003 Cape Blanco, north side 42.8401, -124.5634 Hunters Cove, Cape Sebastian 42.3205, -124.4261 Whiskey Creek (= Boardman SP of Goddard, 1990) 42.2227, -124.3830 House Rock 42.1130, -124.3550 Lone Ranch 42.0997, -124.3493 Harris Beach 42.0641, -124.3087 Chetco Point 42.0436, -124.2899 Wilson Creek 41.5947, -124.1051 Luffenholtz Beach 41.0407, -124.1209 Punta Gorda 40.2747, -124.3640 Glass Beach, Fort Bragg 39.4513, -123.8139 Coleman Beach 38.3632, -123.0708 Pillar Point 37.4938, -122.4994 Pigeon Point, north side 37.1847, -122.3969 Scott Creek 37.0455, -122.2380 Santa Craz Wharf (subtidal) 36.9587, -122.0178 Santa Craz Harbor (docks) 36.9642, -122.0018 Point Cabrillo (subtidal) 36.6214, -121.9016 Monterey Harbor (docks) 36.6043, -121.8912 Asilomar 36.6282, -121.9421 Sand Dollar Beach 35.9216, -121.4716 Hazard Canyon, Montana de Oro SP 35.2899, -120.8845 Tarantula Reef, Jalama 34.4954, -120.4968 Naples 34.4320, -119.9493 Carpinteria Reef (subtidal) 34.3930, -119.5400 Tarpits Reef, Carpinteria SP 34.3869, -119.5166 County Line Reef (subtidal) 34.0472, -118.9710 Big Kelp Reef (subtidal) 34.0046, -118.7925 Point Dume, south side 34.0031, -118.8037 Big Fisherman’s Cove (subtidal) 33.4445, -118.4847 and the data from the Lance accounts for outer coast sites in San Diego County are publicly accessible online (California Academy of Sciences and Goddard 2013). The McDonald data, which cover the years 1967-2010, primarily in central California, are in an unpublished spread- sheet sent to the senior author (G. R. McDonald, personal communication to JG, March 9, 2010). We reference specimens from CAS using the collection number, prefaced by CASIZ, or entire groups of conspecific specimens simply as “CASIZ collection records”. As indicated above, those records are publicly available online. Results Significant locality records of nudibranchs and other sea slugs in California and Oregon in 2014-15 are listed systematically by species and documented below. Nine of these represent 20 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES new or previously unpublished northernmost locality records and are marked with an asterisk (*). The remaining 21 species included below have previously been recorded from at least as far north as we found them in 2014-15, usually during strong El Nino events, but typically occur significantly farther south. For each of these we note their usual and extreme northerly range limits and describe the anomalous patterns of distribution and abundance we observed from late 2013 through August 2015. Locality records obtained from non-peer reviewed sources or websites (excluding museum databases) are referenced in footnotes, as are images we posted online of specimens we found duri,ng this study. Sacoglossa Limapontiidae *Placida cremoniana (Trinchese, 1892) The single specimen of Placida cremoniana found on 10 Oct 2014 at 12 m at Stony Point, Santa Catalina Island^ is the first record of this widespread tropical sacoglossan in the northeast Pacific north of La Paz, Baja California Sur (Angulo-Campillo 2002). Another specimen was found on 23 May 2015 at about 27 m depth at Casino Point, Santa Catalina Island.^ Cephalaspidea Aglajidae Navanax inermis (Cooper, 1 863) Navanax inermis is a Californian species rare north of Point Conception. It has been recorded as far north as the Bolinas Lagoon (Behrens 1998; Behrens and Hermosillo 2005) and Bodega Bay (Gosliner and Williams 2007), where it was reported by J. Standing and colleagues as rare to common.^ The Bolinas record was linked by Behrens (1998) to the 1992-93 El Nino, and similarly the latter record from Bodega Bay can probably be linked to the strong El Nino of 1972-73, which coincided with sampling conducted by Standing et al. in the early 1970’s in Bodega Bay.^ On 4 June 2015 Grace Ha found one specimen of Navanax inermis in Bod- ega Bay."^ We found N. inermis crawling on the concrete wall below the Harbor Master's Office in the Monterey Harbor on 25 October 2014, 10 January 2015, and 21 May 2015, the first time we had seen this species in the Monterey Harbor since beginning observations there in 2008. On 14 May 2015 Robin Agarwal found a specimen in the Santa Cruz Harbor.^ Anaspidea Aplysiidae Aplysia californica (Cooper, 1863) ^ Peterson, B. 2014. Warmer California waters bring new opportunities for photographers. Retrieved 12 July 2015 from: califomiadiver.com/warmer-califomia-waters-briiig-new-opportunitiesl23/ ^Halstead, A. 2015. Photos from Aaron Halstead’s post in SoCal Underwater Photographers. Retrieved 11 September 2015 from: htlps://www.facebook.coiii/photo.php?fbid= 10153463203903054&set=gm.463220240513533 &type= 1 &theater ^ Standing, J., B. Browning, and J. W. Speth. 1975. The natural resources of Bodega Harbor, State of California, Department of Fish and Game. 224 pp. ^ Soees, J. 2015. Inhaling bubbles. The Natural History of Bodega Head, 4 June 2015. Retrieved 27 July 2015 from: http://bodegahead.blogspot.com/20 1 5/06/inhaling-bubbles.html ^ http://www.inaturalist.org/observatioEs/148890 1 NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 21 Aplysia californica rarely occurs on the outer coast north of Point Conception, but has been recorded in bays as far north as Yaquina Bay, Oregon, the latter during the strong 1982-83 El Nino (Pearcy and Schoener, 1987). On 1 December 2013, we found 20 individuals in the intertidal zone at Hazard Canyon Reef, the first we had seen in 14 years of approximately semi-annual sampling there. We did not find any at Hazard Canyon during two trips in May 2014, but observed two individuals on 18 May 2015. In temporally more limited sampling at Tarantula Reef beginning in 2009, we had not found A. californica until 2 February 2015, when the senior author counted 64 individuals. Similarly, two of us (WP and DM) have been sampling the Monterey Harbor since 2008 and found A. californica there for the first time on 23 September 2014, and through July 2015 had seen it there on five more visits. From August 2014 through April 2015 BG observed many A. californica, 8 to 15 m deep, off Monterey and Pacific Grove, and on 20 May 2015 found one specimen intertidally at San Dollar Beach, in southern Monterey County. By mid- 2015 high densities had also been reported from two sites on the outer coast of Sonoma County^’^, as well as from San Francisco and Tomales Bays (Anonymous 2015; Bay Nature Staff 20 15). On 14 August 2015, Dr. Troy Nash and his summer Invertebrate Zoology class from the Ore- gon Institute of Marine Biology (OIMB) found three specimens on a wave-protected rocky shore (43.3400° N, 124.3750° W) at Gregory Point, near Charleston, Oregon (T. Nash, personal communication to JG, 22 Oct 2015). Based on the image posted by OIMB^, one of the slugs was about 17 cm long and found on the red alga Neorhodomela larix (Turner, 1819). These specimens are to our knowledge the first ever found on the outer coast of Oregon. Examination of the dates of collection of the specimens at CAS of A. californica collected from Monterey Bay north since 1950 reveals that all but one were collected during El Nino events. CASIZ 57362, collected in San Francisco Bay in December 1984, may have been a 2”*^ generation holdover from the 1982-83 El Nino, one of the strongest on record. Aplysia vaccaria Winckler, 1955 We have been sampling Naples semi-annually to monthly since fall 2006 and on 20 Septem- ber 2013 io\xnd Aplysia vaccaria for the first time there. We recorded single specimens again in September, October, and November 2014. Aplysia vaccaria has been recorded as far north as Monterey Bay, California (Behrens 1991; Behrens and Hermosillo), and large specimens were observed subtidally by Cheryl Mitchell off the Monterey Breakwater on 18 September 2015'^. Nudibranchia Goniodorididae Okenia angelensis Lance, 1966 This species has been recorded once from as far north as San Francisco Bay, in September 1964 (Lance 1966), and from Monterey Bay in September 1963 (Lance) and again in October 1971 (CASIZ 9168). 1963-64 was a moderate El Nino, but 1971 was a moderate La Nina. DM ^ Sones, J. 2015. Munching at Miwok. The Natural History of Bodega Head, 23 May 2015. Retrieved 27 July 2015 from: http://bodegahead.blogspot.com/201 5/05/munchmg-at-miwok.html ^ http://www.inaturalist.0rg/0bservati0ns/l 1 83373 ® http ://oimb .uoregon. edu/ sea-hares/ ^ https://www.facebook.com/cheryl.mitchell.750/videos/888884974530765/; [Aplysia vaccaria in video at 0:36 and 5:05] 22 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 2. Comparison of abundance of Okenia rosacea at six sites in central California in early 2015 with his- torical highs recorded at same sites. 2015 Previous highs ^ Site Date No. inds.h ^ obs. ^ No. inds.h * obs. ^ ‘ Date Sampling period Duxbury Reef 01/01/15 Si 2“ 0.0 6/69-12/75 6/69-12/75 0.7 Nov 2010 12/07-12/10 Pillar Point 02/17/15 122.7 13.0 M 1993 9/88-2/95 0.2 Jan 2008 1/08-12/10 Scott Creek 01/20/15 65.0 11.4 Mar 1978 6/75-10/78 0.3 Mar 2009 12/07-10/10 Asilomar 01/22/15 13.5 5.8 Oct 1969 10/69-7/73 1.0 Nov 2007 11/07-10/11 Sand Dollar Beach 01/31/15 40.0 48.0 Dec 1997 Dec 1997 1.3 Mar 2009 3/08-3/14 Hazard Canyon 01/19/15 Sl50 ^4 Mar 1968 5/67-11/71 4.4 M 2012 11/99-5/14 “Data from: R. Agarwall (littp://www.maturalist.org/observations/l 156561) * Data from: Goddard et al. (2015) (Duxbury Reef); Schultz et al (2015) (Pillar Point, Scott Creek, Asilomar); R. Roller, unpublished California field accounts, CASE collection (Hazard Canyon 1967-71); Goddard, unpublished data (Sand Dollar Beach and Hazard Canyon 1997-2015). found one individual^^ of O. angelensis on the H dock in the Monterey Harbor on 30 September 2014 and more at the same locality in April and June, 2015. On 17 July 2015 Donna Pomeroy and Robin Agarwal found O. angelensis on floating docks in the Pillar Point Harbor in Half Moon Bay, San Mateo County* ^ Okenia rosacea (MacFarland, 1905) Okenia rosacea (Fig. 3A) was present in low abundance at two sites in central California in spring 2014 (Fig. 4). However, by fall, its abundance at both sites had increased an order of magnitude, and by winter 2014-15 had reached levels not seen before at our historical study sites in central California (Table 2). The density observed at Sand Dollar Beach, Monterey County during the strong El Nino of 1997-98 matched the densities we observed in 2015, and brief, qualitative observations by the senior author at Scott Creek on 28 December 1997 indicated similarly high abundance. On 3 January 2015 Jackie Sones of the Bodega Marine Laboratoiy (BML) reported finding 14 0. rosacea in a 2 m section of low intertidal shore at Bodega Head, the first specimens she had seen of this species in ten years of sampling at Bod- ega Head*^. On 21 January 2015 we counted 7.7 O. rosacea h”* observer”* at Coleman Beach, followed by 19.7 O. rosacea h“* observer”* at Glass Beach, Fort Bragg on 16 Feb 2015, the same day that Spencer Dybdahl Riffle (personal communication to JG, 17 February 2015) counted 46 indivi- duals at Patrick's Point SP near Trinidad in Humboldt County, and the day before David lAnderson (personal communication to JG 21 April 2015) found 12 individuals at False Klamath Cove in Del Norte County. In March BG observed 13 0. rosacea at Luffenholtz Beach, near Trinidad (CASIZ 204572) and another seven at Wilson Creek, False Klamath Cove ^ ® https ://www. flickr.com/photos/3 93 65 853 @N07/ 1 54062 1 4295 ^ ^ https ://www. flickr. com/photos/dpom 12/19 647657349 Sones, J. 2015. Finally! The Natural History of Bodega Head, 3 January 2015. Retrieved 28 August 2015 from: http.7/bodegahead.blogspot.com/2015/01/fmally.html NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 23 Fig. 3. Selected nudibranchs found in 2014-15 north of their usual geographic ranges. A Okenia rosacea and egg ribbons, Lone Ranch, Oregon, 18 July 2015. Image by NT. B Doriopsilla fulva. Whiskey Creek, Oregon, 16 June 2015. Image by NT. C Doriopsilla gemela, Tarpits Reef, Carpinteria, 19 May 2015. Image by CH. D Flabellina bertschi, Big Fishermans Cove, Santa Catalina Island, 2 January 2015. Image by CH. E Anteaeolidiella oliviae, Glass Beach, 16 February 2015. Image by DM. F Babakina festiva. Pigeon Point, 19 January 2015. Image by DM. G Noumeaelia rubrofasciata laying eggs, County Line Reef, Malibu, 21 February 2015. Image by CH. H Cuthona phoenix, Monterey Harbor, 24 July 2014. Image by DM. 24 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 4. Increase in abundance of Okenia rosacea at two sites in central California, spring 2014 to winter 2015. Sites were not sampled in summer. Image of O. rosacea by Gary McDonald. (CASIZ 2014576). On May 7, two weeks after we asked him to be on the lookout for O. rosacea at Cape Arago, Oregon, Richard Emlet of OIMB and his Invertebrate Zoology class found a specimen just south of Sunset Bay, Cape Arago (R. Emlet, personal commu- nication to JG, 8 May 2015). In May, June and July 2015 in Oregon we found O. rosacea at Harris Beach, Lone Ranch, House Rock, Whiskey Creek, and Middle Cove, Cape Arago, and Gregoiy Point. The specimen from Lone Ranch on 18 July was spawning (Fig. 3 A), and the highest number of O. rosacea we found at the Oregon sites was 21 (or 1.0 individuals h“^ observer”^) on 16 June 2015 at Whis- key Creek. We deposited specimens of O. rosacea from Whiskey Creek and Middle Cove, Cape Arago at CAS (CASIZ 204855 and 204854, respectively). On 2 July 2015 Spencer Dybdahl Riffle found abundant O. rosacea and egg masses intertidally at Patrick's Point SP, Humboldt County On 28 August 2015 NT found one O. rosacea at North Cove Cape Arago, the first specimen ever recorded from that site. The above specimens from Oregon are not only the most ever observed in the state, but the first recorded from Oregon since JG found one individual of O. rosacea at Middle Cove, Cape Arago on 24 July 1998, after 16 years of sampling for nudibranchs at Cape Arago, and during that year’s exceptionally strong El Nino (JG, unpublished data and photograph). The only other record of O. rosacea in Oregon is from Steinberg (1963) who, based on observations by L. Andrews, recorded it (as Hopkinsia rosacea) from Coos Bay. However, Integripelta Mia- Mata, the sole prey of O rosacea, does not occur inside Coos Bay, but rather on the open coast at nearby Cape Arago (JG, personal observations). Further, the specimen observed by L. Andrews was in an aquarium at OIMB and may actually have been collected in California (L. Andrews, personal communication to JG, 19 November 2009). Therefore, the northernmost verified locality for this species should be recorded as Gregoiy Point, Oregon. *Trapania veiox (Cockerell, 1901) https://www.flickr.eom/photos/riffle_Eatare__photos/l 95338972 1 5 NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 25 BG found one specimen at 5 m depth on sponges on a piling underneath the Santa Cruz Pier on 31 August 2015 (CASIZ 209038), extending the known range of this Californian species across Monterey Bay from Carmel, California (Behrens and Hermosillo). After one day in cap- tivity in Santa Craz, the specimen laid two egg masses. All records of Trapania velox from Car- mel (in 2004, 2006 and 2009)^^’^^ can be associated with weak to moderate El Ninos, or in the case of 2006, a short, weak La Nina following the 2003-04 and 2004-05 El Nino events. Onchidorididae Acanthodoris rhodoceras Cockerell in Cockerell and Eliot, 1905 This species has been found as far north as central Oregon (Goddard 1 997), during the mod- erate 1991-92 El Nino, and the only other published record of this species from Oregon (God- dard 1990) can be associated with the strong 1987-88 El Nino. In August 2015, NT found a total of eight specimens during five trips to Hunters Cove, on the south side of Cape Sebastian, Ore- gon. The specimens were the dull colored form as pictured in Figure 15 of McDonald and Nybakken (1980). Polyceridae Crimora coneja Marcus, 1961 Until recently Crimora coneja was known from only three mainland sites between Cape Axago, Oregon and San Diego County, California: Punta Gorda, Humboldt County (Goddard 1987), and Montana de Oro State Park and Moiro Bay in San Luis Obispo County^ On 18 June 2015 we found two specimens at Middle Cove, Cape Arago, and on 15 July two more at Whiskey Creek, and on 15 August one specimen at Lone Ranch. All of the specimens from Oregon were found on Hincksina min- uscuia, the only known prey of C. coneja. Additionally, three other new locality records were added in 2015: (1) Trinidad, Humboldt County by Cassidy Grattan^^, (2) Pillar Point by Matt Knoth^^, and (3) Palmer’s Point, Humboldt County by Cassidy Grattan, who on 2 August reported 10 specimens from Trinidad^^. Since these reports additional specimens have been found at Pillar Poinl^®, and NT found a single specimen at North Cove, Cape Arago on 28 August. During 14 years of observation at North and Middle Coves, Cape Arago from 1980 to 2008 Crimora coneja was found in seven years: three during the second year of strong El Nino events (1982-83, 1987-88, 1997-98), two during weak La Ninas (1984-85, 1985-86), and once during ENSO-neutral conditions (1981) (Goddard 1984, and unpublished data). Polycera atra MacFarland, 1905 This southern species has been found in Oregon, mainly in bays, only during strong El Nino events (Goddard 1984, unpublished data) and has been recorded once from Westport, just inside Grays Harbor, Washington, during the 1997-98 El Nino (Lamb and Hanby 2005; A. Lamb, per- sonal communication to K. Fletcher, forwarded to JG, 13 Sept 2012). On 17 June 2015 we found two specimens of P. atra on the sides of floating docks in the outer Charleston boat Basin. * ^ http ://www. seaslugforam.net/ shov/all/trapvelo http://www.ba11e.0rg/images/galleries/v/FieiciGuicie/Opisthobranchs/Trapa11ia_velox/; Goddard, J. and C. Hoover. 2011. Crimora coneja Marcus, 1961. Retrieved 11 September 2015 from: http://slugsite.us/bow2007/Eudwk758.htm https://www.flickr.eom/photos/l 28077533@N05/1 7 1 1 3424562 ^ * http://www.inaturalist.0rg/0bservati0ns/l 728 1 02 http://www.iiiaturalist.Org/observations/l 834 1 34 http://www.inaturalist.org/observations?taxoE_id= 50057 26 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 5. Number of years Triopha maculata present or absent on the southern Oregon coast by phase of ENSO, 1980-98 and 2006-2014. Data for Whiskey Creek, Cape Blanco, and all three coves of Cape Arago combined, from Goddard (1984) and JG and NT (unpublished data). Image of T. maculata by Gary McDonald. Triopha maculata MacFarland, 1905 Although recorded from Bamfield, British Columbia (Millen 1983) and Cape Arago, Oregon (Goddard 1984), Triopha maculata rarely occurs north of Cape Mendocino, California. From 18 May to 4 July 2015 in Oregon we found T maculata at Chetco Point, Harris Beach, Lone Ranch, House Rock, Whiskey Creek, Cape Blanco, all three coves at Cape Arago, and Squaw Island. One of us (NT) has been sampling four of these sites for nudibranchs since 2008 and never observed this species before, and records of T maculata from southern Oregon between 1980 and 2010 show that it occurred mainly during El Nino events (Fig. 5). Large specimens have also been observed in the Charleston boat basin during El Nino events (JG, personal observations), and on 28 August 2015 NT found one specimen there 6 cm long. Dorididae Thordisa bimaculata Lance, 1966 Thordisa bimaculata has been reported from Isla Natividad, Baja California to Carmel, California (Lance). Although it can be locally abundant on rocky shores in San Diego County (Sphon and Lance 1968; California Academy of Sciences and Goddard 2013), there are only a few records of this species from Santa Barbara County (Sphon and Lance 1968; CASIZ 98793), none from San Luis Obispo County, and besides the single specimen reported by Lance from off Carmel, there have been few sightings from Monterey County^ \ On 5 July 2015 Regina Roberts found a specimen laying an egg mass on a brown sponge at an unspecified depth off Point Joe, on the Monterey Peninsula^^. Bauder, C., 2001 Thordisa bimaculata from Carmel, California. Retrieved 7 September 2015 from: http://www.seaslugforam.net/fmci/388 1 . https://www.flickr.coiii/photos/reginadiver/l 9265753508 NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 27 We found a total of eight T. bimaculata on 6 November 2014, 30 January 2015, and 17 March 2015 at Tarpits Reef, Carpinteria. These were the first specimens we had found at that site since beginning semi-annual to quarterly sampling there in May 2008. At Naples we recorded our first specimens on 3 May and 5 November 2010, during that year’s moder- ate El Nino, and did not find any more there until finding one specimen on 19 February 2015 and another on 22 April 2015. On 3 July 2015 CH found one specimen at 7 m at Carpinteria Reef, and on 1 September 2015 CH found one specimen of T. bimaculata at 7 m depth at Punta Bocana, Bahia Magdalena, Baja California Sur, a new southernmost locality for this species. Chromodorididae Felimare californiensis (Bergh, 1879) CH found three Felimare californiensis at 8 m on Carpinteria Reef on 3 July 2015, and five more at this same site on 18 July 2015. This species appears to have gone regionally extinct in southern California in 1984 (Goddard et al. 2013), and these specimens are the first to be reported on the mainland of the Santa Barbara Channel since its recovery in the Southern Cali- fornia Bight began in 2003, near the end of the moderate 2002-03 El Nino (Hoover 2015). Felimida macfarlandi (Cockerell, 1901) This brilliantly colored chromodorid nudibranch has been recorded from Bahia Magdalena, Baja California Sur (Bertsch 1978) to Monterey, California (MacFarland 1966) but is usually found south of Point Conception. On 6 March 2015, Jon McNeill found one individual subtid- ally at Point Lobos (J. McNeil, personal communication and image to JG, 9 October 2015). This was followed by Dave Baessler’s subtidal observations at the Monterey Breakwater of one spec- imen^^ on 7 July 2015, and two more^"^ on 29 July 2015. These are the first specimens docu- mented from the Monterey Peninsula since the strong 1998 El Nino, when Gary McDonald found one subtidally at Del Monte Beach^^. Gary McDonald also found one individual intertid- ally at Carmel Point in February 1986, during ENSO neutral conditions following a weak La Nina. CAS has six lots of specimens of F. macfarlandi collected from the Monterey Peninsula in the following years: 1906, 1908, 1909, 1941, 1963 and 1978. Based on the values of the extended multivariate ENSO Index presented by Wolter and Timlin (2011), moderate El Nino events occurred in 1906, 1941, and 1963, and 1978 was a weak El Nino following the 1976- 77 decadal climate shift. 1908-09 experienced weak to moderate La Ninas, indicating that either F. macfarlandi is not always dependent on El Nino conditions to reach the Monterey Peninsula, or that once arrived (in this case, in 1 906) populations can persist locally for a few years through self-recruitment. Dendrodorididae *Doriopsilla fulva (MacFarland, 1905) Hoover et al. (2015) reinstated Z)ono/?.s'i7/a fulva as distinct from D. albopunctata. The lat- ter, which is more common intertidally in southern California, can be recognized externally https://www.flickr.eom/photos/73739720@N00/19380471779 https://www.flickr.eom/photos/73739720@N00/l 9702924544 http://www.inaturalist.org/observations/844 135 28 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES by its white spots around and between the dorsal tubercles, compared to the single, apically- located white spot per tubercle on D.fulva. Doriopsilla fulva is also usually bright yellow in color and is more common intertidally than D. albopunctata north of Point Conception (Hoover et al. 2015). We found one specimen of D.fulva, 8 mm long, at Whiskey Creek on 16 June 2015 (CASE 208751; Fig. 3B). This is the first record of this species in Oregon, and extends its known range from Abalone Beach in Humboldt County, California (Jaeckle 1984). NT found an additional specimen, 11 mm long, at Lone Ranch on 15 August 2015, and we found another specimen^^. 14 mm long, at Punta Gorda, California on 15 June 2015. On 22 Jan 2015 at Asilomar we recorded 28.2 D.fulva obs“’' h“^, the highest recorded since Nybakken and colleagues counted 24.7 obs“^ h~^ in April 1973, during that year’s strong El Nino (see Schultz et al. 2015). Addi- tionally, D.fulva was reported to be more abundant at Bodega Head in 2015 compared to pre- vious years^^. Doriopsilla gemeia Gosliner, Schaefer and Millen, 1 999 After seven years of approximately semi-annual sampling at Tarpits Reef, Carpinteria we found this species for the first time on 20 April 2015 and again on 19 May 2015. On both dates we found three large, mature adults (Fig. 3C), all with the combination of yellow gills and densely packed, opaque white spots concentrated in the middle of the dorsum characteristic of this species. At Naples we had observed single specimens on 5 February 2008, 29 November 2009, and 24 November 2012. However, on five of 11 sampling trips from April 2014 to May 2015 we found a total of nine specimens. BG found one specimen at 15 m off Point Cabrillo, Monterey on 13 October 2014. There is only one other verified record of this species in central California (also from 2014, off the Monterey Peninsula; see Hoover et al.), and the southern coast of Santa Barbara County, which includes Naples and Point Conception, appears to mark its usual northern range limit. Hancockidae Hancockia californica MacFarland, 1923 There are only two historical records of this species from north of Marin County: (1) Jaeckle’s (1984) record from Trinidad, Humboldt County, which contrary to Behrens and Hermosillo is the northernmost record of Hancockia californica, and (2) Behrens (2004) record for Fort Bragg, Mendocino County. However, the record from Fort Bragg was based on a misidentified specimen^^ of Dendronotus subramosus MacFarland, 1966 (confirro.ed by D. Behrens, personal communication to JG, 4 April 2008). On 23 May 2015 Jackie Sones found a specimen of Hancockia californica at Coleman Beach, Sonoma County^^, and on 2 August 2015 Spencer Dybdahl-Riffle found a specimen, 5 mm long, at Trinidad^®. https://wwm/.flickrxom/photos/34486353@N07/l 8524 191074 Sones, J. 2015. Sea salt The Natural History of Bodega Head, 10 February 2015. Retrieved 8 August 2015 from: http://bodegahead.blogspot.com/20 1 5/02/sea-salt, html ^^Behrens, D. 2003. Hancockia californica. The Slug Site. Retrieved 5 January 2016 from; http://slugsite.us/ bow/nudwk3 7 3 .htm ^^Sones, J. 2015. Be still my... The Natural History of Bodega Head, 25 May 2015. Retrieved 14 September 2015 from: http://bodegahead.blogspot.eom/2015/05/be-still-my.htmi ^®https://www.flickr.com/photos/riffle_nature_photos/20326335435 NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 29 Dotoidae Doto form A of Goddard (1996) Doto form A is the most common Doto in the southern California bight and is commonly observed subtidally on campanularid hydroids growing on kelp and other macroalgae. It has been found as far north as Drake’s Estero, Point Reyes National Seashore (Goddard 1996) but rarely occurs north of Point Conception. Goddard (1996) argued for the separation of this form from D. amyra Marcus, 1961 based on morphological and developmental evidence, and the genetic evidence presented by Shipman and Gosliner (2015) corroborate this. Compared to Doto amyra, Doto form A has cerata with brighter colored cores and longer, distinctly white papillae. It also has smaller eggs than D. amyra and planktotrophic development. Since mid- 2014 Robin Agarwal and Donna Pomeroy have made numerous sightings of this species (cited as Doto amyra), frequently with its egg masses, in Mono and Monterey Bays, and at Pillar Point^^ BG collected two specimens from the Santa Craz Harbor on 28 August 2015 (CASIZ 207370) Dironidae Dirona picta MacFarland in Cockerell and Eliot, 1905 Dirona picta has been reported from as far north as Cape Meares, Oregon (Goddard 1 997) but has rarely been observed on the outer coast of Oregon (Goddard 1984: 159, 1990), and the only specimen recorded from Cape Arago was during the strong 1997-98 El Nino (JG, per- sonal observations). We found single specimens of it on 17 June 2015 at Cape Blanco and in the Charleston outer boat basin, and another specimen at 5 -Mile Point on 1 August 2015. *Jano!us anulatus Camacho-Garcia and Gosliner, 2006 On 13 August 2015 one of us (CH) found one specimen of Janolus anulatus at Tarpits Reef, the first seen there since the first specimens were recorded from this site on 9 May, 20 June, and 4 July 2012 (CASIZ 189420)^^, following a transition from a weak La Nina to moderately posi- tive values of the MEL Previously, this species was known from La Jolla, California to Costa Rica (Behrens and Hermosillo; Camacho-Garcia et al 2005). Based on data collected by James Lance and colleagues from 1964 to 2002, J. anulatus (dis- tinguished from J. barbarensis and referred to by Lance first as Antiopelia sp. and later as Jano- lus sp.) peaked in abundance in La Jolla, California during strong El Nifio events (excluding the 1987-88 event), as well as following the 1976-77 climate shift (Fig. 6). * Janolus barbarensis (Cooper, 1 863) This Californian and Panamic species has been recorded from as far north as San Francisco Bay (Jaeckle 1983; Behrens and Hermosillo 2005) but is rare north of Morro Bay. On 28 July 2015 Benson Chow of the Tiburon Romberg Center collected one specimen in the Sausa- lito Marina, inside San Francisco Bay (CASIZ 207372). On 25 August 2015 Robin Agarwal found an additional specimen on the side of a floating dock in San Francisco Bay^^, a day after ^^https://www.flickr.com/photos/303 14434@N06/1 9 1 14409924; Mtps://www.flickrxom/photos/dpoml 2/ 18492397230 ^^Goddard, J. 2012. Janolus anulatus Camacho-Garcia and Gosliner, 2006. Retrieved 28 2015 from: http:// slugsite.us/bow2007/iiudwk7 85 .htm ^^http://www.inatoralist.org/obseivations/l 89205 1 30 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES finding one in the Santa Craz Harbor^"^, where she had first sighted one on 15 October 2014^^, On 1 1 September 2015 Shawn Brumbaugh and Chris Kwan found a J. barbarensis at the Spud Point Marina, Bodega Bay, California and informed Jackie Soees and Eric Sanford from BML, who the next day found one specimen 63 mm long on Bugula neritina on the side of a floating dock at the same locality^^. Flabellinidae Flabellina bertschi Gosliner and Kuzirian, 1990 This Panamic species ranges from the northern Gulf of California to Panama (Gosliner 1994), and has also been reported from the outer coast of Baja California (Goddard and Schickel 2000), as well as Catalina Island (Behrens, 2004; Behrens and Hermosillo 2005). Behrens (2004) represents the northernmost record of this species, but did not include an image or reference to voucher specimens. CH found one specimen (Fig. 3D) at 2 m depth at Big Fisherman’s Cove, Santa Catalina Island on 2 January 2015, feeding on a species of Eudendrium similar to that shown with F. bertschi in Mexico in Fig. 4B of Millen and Hermosillo (2007). * Flabellina cooperi (Cockerell, 1901) Known mainly from southern California, the northern range limit of Flabellina cooperi has been Elkhom Slough, Monterey Bay since 1970 (McDonald 1983). Green and Gosliner (2016) presented molecular genetic evidence that specimens of a previously unidentified Fiabeb Una with smooth to slightly wrinkled rhinophores and a notum irregularly covered with opaque white pigment collected by JG from Tarpits Reef, Santa Barbara County (CASIZ 195988) and Coleman Beach, Sonoma County (CASIZ 195990) are identical to F. cooperi from La Jolla, and (see below) Santa Cmz, California. Coleman Beach is therefore now the northernmost locality known for F. cooperi. The two specimens of F. cooperi from Coleman Beach (CASIZ 181322 and 195990) were collected on 29 April 2009 and 25 March 2010, respectively, at the beginning and end of the moderate 2009-10 El Nino. On 28 August 2015 BG found one specimen of F. cooperi (CASIZ 207369)^^ in the Santa Cmz Harbor, and based on the morphology of the above specimens included by Green and Gosliner, two speciro.ens of Flabellina found on 27 April 2015 by DM on the sides of floating docks in the Monterey Harbor^^ can now also be assigned to F. cooperi. Additionally, we have been sampling Naples regularly since 2006 and from 28 Febmaiy to 3 May 2010 found a total of 30 F. cooperi (e.g., CASIZ 182720, 195986, as Flabellina sp.) our first specimens of this species at that site. More recently, we found our first specimens of F. cooperi at Tarantula Reef on 3 January 2015, and additional specimens at Tarpits Reef on 30 January and 19 May 2015. The original sightings of Flabellina cooperi at Elkhom Slough consisted of a total of three specimens collected on 16 November and 7 December 1970, during a moderate La Nina (McDonald 1983). Two years later, during a strong El Nino, Gary McDonald (unpublished field data) found an estimated total of 30 individuals on 18 and 19 October 1972. ^\ttp://www.ioaturalist.org/observations/l 888084 ^^https://www.flickr.com/pliotos/30314434@N06/14938263624 ^^Sones, J. 2015. Straight outta Santa Barbara. The Natural History of Bodega Head, 12 September 2015. Retrieved 12 September 2015 from: http://bodegahead.blogspot.eom/2015/09/straight-outta-santa-barbara.html ^\ttp;//www.maturalist.org/observations/ 1 906734 ^®https://www.fiickr.com/photos/39365853@N07/16697361803 ^^https://www.flickr.com/photos/34486353@N07/5424958579 NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 31 Flabellina iodinea (Cooper, 1863) Although long known from as far north as the west coast of Vancouver Island (Bernard 1970) and Puget Sound, Washington (Bergh, 1879), this highly conspicuous species rarely occurs on the outer coast north of Monterey Bay (Gosliner and Williams 1970; Bertsch et al 1972; God- dard et at. 2015; Schultz et al. 2015). There are no published records from Oregon, and the only recent record from Washington is from subtidally near Cape Flattery during the strong 1997-98 El Nino (Lamb and Hanby 2005; A. Lamb persona! commmunicatioe to K Fletcher, forwarded to JG, 13 Sept 2012). On 15 May 2014 we found our first specimen at Hazard Canyon Reef since beginning our sampling there in 1999. We found another specimen at the same site on 18 May 2015. Further north, we found a specimen of F. iodinea on 20 January at Scott Creek, and by May 2015, specimens had been found in Bodega Bay"^'®, Coleman Beach (J. Sones per- sonal communication to JG, 23 May 2015), and Trinidad, Humboldt County'^^ Flabellina iodb nea was first reported from Trinidad by Jaeckle (1984). On 4 July 2015 Spencer Dybdahl Riffle counted 23 F. iodinea in the tidepools at Trinidad^^. Aeolidiidae *Anteaeolidiella chromosoma (Cockerell and Eliot, 1905) On 21 May and 7 June 2015 we found a total of four specimens on the H dock in the Mon- terey Harbor. TTiis species was previously known from as far north as Mono Bay (Behrens 1980) and ranges south to the Galapagos Islands (Camacho-Garcia et al. 2005). On 15 July 2015 Robin Agarwal found a specimen in the Santa Cmz Harbor, on the north side of Monterey Bay"^^, and on 25 August 2015 found two specimens and their egg masses in San Francisco At Naples, A. chromosoma was more abundant in the first half of 2015 than at any iiine 111 the past ten years, with a lesser peak in abundance during the 2009-10 El Nino (JG, unpublished data). *Anteaeolidiella oliviae (MacFarland, 1966) The northern range limit of this species has long been Duxbury Reef, Marin County, California (Gosliner and Williams 1970). On 21 January 2015 we found one specimen at Coleman Beach, Sonoma County, and on 16 February 2015 we found one specimen at Glass Beach in Fort Bragg, Mendocino County (Fig. 3E). 'The latter specimen had unusually pale cer- ata, probably reflecting an atypical complement of the anthozoans normally consumed by this species (Beeman and Williams 1980). Facelinidae Emarcusia morroensis Roller, 1972 This small (to 15 mm) cryptic species has been found only a few times since its original description and has been reported from Mission Bay, San Diego to San Francisco Bay (Roller 1972; Gosliner 1990), It is known mainly from bays (Roller 1972; McDonald 1983; CASIZ ^“Sones, J. 2015. Fiery and flamboyant. The Natural Histor/ of Bodega Head, 19 May 2015. Retrieved 28 August 2015 from: littp://bodegaliead.blogspot.com/2015/05/fiery"3nd"flamboyaiit.html '^'*https://www.flickr.com/pliotos/riffle__iiatare_photos/l 6S9S 1 66880; http://v/wvi/.inaturalistorg/observatioEs/ 1387158 J/www. flickr.com/photos/riffle_natore_photos/ 19287584140 “^^https ://wwv/. iEataralist.org/observatioEs/ 1767403 '^\ttps://www.flickr.com/photos/30314434@N06/20876669342 32 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 6. Number of Janolus anulatus observed per sampling trip, La Jolla, California, 1964-2001. Red arrows indicate strong El Ninos, and black arrow indicates 1976-77 climate shift. From data collected by James Lance and colleagues during 5 1 total sampling trips to intertidal sites at South Casa and Windansea Reefs (California Academy of Sciences and Goddard 2013). Image of J. anulatus by JG. collection records) and has been previously observed on the outer coast only twice: in September 1972 from a buoy anchor chain in Monterey Bay (CASIZ 69806), and in December 2007 from 15 m depth off Redondo Beach"^^. On. 20 May 20 1 5 the senior author found six specimens intertidally on the south side of Point Dume, Malibu^^. Robin Agarwal found a specimen in Morro Bay in September 2014, and another on 26 July 2015 in the Monterey Harbor^^. Two specimens were found in the Santa Cmz Harbor in August 2015, one by BG on the 28* (CASIZ 207371)^^, and another on the 3R‘ by Robin Agarwal^^. Babakina festiva (Roller, 1972) Babakina festiva is known from Nayarit, Mexico to Duxbury Reef (Behrens and Hermosillo 2005). However, it has been reported previously only twice from north of Point Conception (Gosliner 1990) and both sightings can be linked to the strong El Nino of 1987-88. In 2015 we found two specimens on the north side of Pigeon Point, one on 19 January 2015 (Fig. 3F), and the other on 3 July 2015. Three more specimens were observed between February and July 2015 in the Fitzgerald Marine Reserve in San Mateo County^®. "^^Kopp, K. 2007. Emarcusia morroensis Roller, 1972. Retrieved 26 August 2015 from: http://slugsite.us/ bow2007/nudwk58 1 .htm ^%ttps://www.flickr.com/photos/34486353@N07/l 77246 10780 ^’^http ://www. inaturalist . org/observations/ 1 8 06084 ‘^^https://www.flickr.coiii/photos/lemurdillo/2 1 0 1 1780632 '^\ttps://www.flickr.com/photos/303 14434@N06/204289883 14 ^®http ://www. inaturalist. org/observations?taxoii_id = 5 048 9 NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 33 *Noumeaella rubrofasciata Gosliner, 1991 On 21 February 2015 CH found a single specimen laying an egg string (Fig. 3G) at 18 m depth at County Line Reef, extending the known range of this species north from Santa Catalina Island (Gosliner 1991). CH observed two more at County Line Reef on 19 April 2015, and five specimens at Big Kelp Reef, Malibu on 7 March 2015. These spe- cimens are the first CH has observed at these sites in four and five years of observation, respectively. Tergipedidae Cuthona phoenix Gosliner, 1981 This slender, distinctive aeolid has reported once from Monterey Bay (Behrens 1991) and also been observed in Morro Bay and a few sites total in southern California, the Gulf of California, and Costa Rica (Behrens 1980 [as Tergipes sp.]; Gosliner 1981; Camacho-Garcia et al. 2005; CASIZ collection records). Cuthona phoenix has usually been found among hydroids growing on flotsam, floats, or giant kelp, Macrocystis. On 14 September 2014 Robin and Marisa Agarwal found two specimens on Macrocystis next to docks in Morro Bay^\ on 24 July 2014 one of us (DM) found four specimens on Macrocystis at the H dock in the Monterey Harbor (CASIZ 199397) (Fig. 3H), and on 24 August 2015 Donna Pomeroy found specimens on Macrocystis in the Santa Cruz Harbor^^. Discussion We documented northward range shifts related to the 2014 warm anomaly in the NE Pacific for 30 species of nudibranchs and other benthic sea slugs. Nine of these were recorded from new northernmost localities (Fig. 7), and the remainder, including Okenia rosacea, which reached unprecedented densities in northern California and Oregon, were found at or near their known northern range limits. Only the strong El Ninos of 1982-83 and 1997-98, as well as the 1976-77 climate shift are known to have forced similar shifts in the nudibranch fauna of the region (Pearcy and Schoener 1987; Engle and Richards 2001; Goddard et al. 2011; Schultz et al. 2011; see Goddard 1984, p. 157; and specific results above). During the 1976-77 climate shift, when the PDO shifted from cold to warm phase, the total abundance of southern species of nudibranchs in central California increased as northern species declined (Fig. 2 in Schultz et al. 2011). We observed a similar transition in 2014-15 at sites we have been monitoring regularly since at least 2008 (Fig. 8). Our results demonstrate a striking biological response to the 2014 warm anomaly in the North Pacific Ocean. They also (1) reinforce indications that the anomaly was part of a regional cli- mate shift, and (2) further demonstrate, as originally proposed by Schultz et al. (2011), that intertidal populations of nudibranchs - with their short life cycles and planktotrophic larvae - closely track nearshore ocean conditions. Range shifts of these brilliantly colored species there- fore constitute useful biological indicators of regional ocean climate. In fact, for much of the Oregonian Biogeographic Province, which stretches from Los Angeles to the northern end of Vancouver Island (Briggs and Bowen 2012), the population fluctuations of one species alone, Okenia rosacea, may serve as a valuable indicator. Integripelta bilabiata, its encrusting bryozoan prey, is locally abundant to British Columbia, and as we predicted for 2015 (see Kray- bill-Voth 2015; Stephens 2015; and Appendix, reference 21), when populations of this ^ ’ http ;//www. inaturalist. org/observations/8 78045 ^^http ;//www. inaturalist.org/observations/ 1889538 34 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Cape Arago Cape Blanco — Pacific Ocean Punta Gorda 100 km 1. Placida cremoniana \ cf C 2. Noumeaella rubrofasciata A j; ■ is 3. Janolus anulatus ^ 4. Anteaeolidiella chromosoma 5. Trapania velox 6. Flabellina cooper! 7. Janolus barbarensis ^ ^ Conception 3 8. Anteaeolidiella olivlae 9. Doriopsilla fulva Fig. 7. Range extensions in 2014-15 from previous northernmost known localities of the sacoglossan Placida cremoniana and eight species of nudibranchs in the NE Pacific Ocean. The previous northernmost locality at La Paz, Mexico for P. cremoniana is not shown, and the yellow part of the line for Flabellina cooperi indicates a range extension in 2009-10 (see Results). conspicuously pink dorid increase north of San Francisco, other southern species follow. Trio- pha maculata, which historically has occurred more frequently and at more sites than O. rosa- cea in Oregon, especially during El Nino events (Fig. 5), would also appear to be a good indicator of elevated SSTs and increased poleward transport of coastal waters. Similarly, in the Californian Province, the appearance of Janolus anulatus in the San Diego area has histori- cally been highly correlated with strong El Nino events (Fig. 6). The mechanisms by which southern species of nudibranchs expand their ranges northward include increased poleward and onshore transport of planktonic larvae from southern source NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 35 ■■■■■ ■■northern species ——southern species Pillar Point Hazard Canyon 2007 2008 2009 2010 2011 2012 2013 2014 2015 Naples Fig. 8. Total abundance of southern and northern species of nudibranchs at three intertidal sites in California, ¥/inters only, 2007-15. Data plotted as anomalies (deviation from mean over years shown, in units of standard deviation). Missing values (2009 at Naples, 2010 at Hazard Canyon, and 2014 at Pillar Point) were filled in by interpolation. populations, especially during periods of reduced upwellieg and with the late summer develop- ment in the Southern California Bight of the poleward inshore countercurrent, which north of Point Conception in the autumn becomes the slxong poleward Davidson Current (Stmb and James 2002; Schultz et a,L 201 1). Upweiling in 2014 off northern Baja California and southern California was anomalously lov/ (Leising et al. 2014), and episodes of strong poleward transport of surface waters were measured by High Frequency Radar (HFR) in Fall 2014 and Winter 2015 off California by the Central and Northern California Ocean Observing System (CeNCOOS). 36 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES For example, surface flows of at least 50 cm sec“^ were observed from Eureka to Cape Blanco from 4 to 9 Feb 2015 (Fig. 9), fast enough and over a long enough period to transport entrained larvae that entire distance. In fact, it was after observing Okenia rosacea spawning during early winter 2015 at Scott Creek, and then checking the surface flows recorded by CeNCOOS, that we predicted the occurrence of Okenia rosacea and other southern nudibranchs in southern Oregon by mid-20i5 and first contacted colleagues there to be on the lookout for them. With a strong El Nino developing on the heels of the 2014 warm anomaly (Climate Prediction Center/NCEP 2015), we expect that nudibranchs currently reproducing in their new northerly ranges (e.g., Okenia rosacea and Anteaeolidiella chromosoma) will be transported (as larvae) even farther north in the coming year. Some, including both specialists like O. rosacea and rela- tive generalists like Triopha maculata and Dirona picta, will find abundant encrusting prey, for- merly beyond their geographic reach. Local recruitment and reproduction of subsequent generations, combined with the accelerating rise in ocean temperatures owing to global warm- ing (Blunden and Arndt 2015; Gleckler et al. 2016), may then result in long-term persistence of some of these species in their new ranges, a contrast to the ephemeral appearances associated with previous warm-water events, particularly El Ninos (reviewed by Lluch-Belda et al. 2005). The effects of these newly arrived, specialized, fast-growing predators will then gradu- ally ripple through the benthic epifauna, altering species interactions and potentially changing community composition as they consume large patches of their prey, some of which, such as sponges, can be very long-lived. All of the species listed above have free-swimming, planktotrophic larvae [Schmekel and Portmann 1982 (for Placida cremoniana); Goddard 2004; Goddard & Hermosillo 2008; God- dard & Green 2013; JG and BG, unpublished data], and most, if not all, were carried northward and onshore into their new ranges as larvae. However, with no records of the widespread saco- glossan Placida cremoniana from between Santa Catalina Island and La Paz, Mexico, we can- not rule out a human-mediated introduction of this species to southern California, nor for that matter be certain of its region of origin. Therefore, excluding P. cremoniana, the average north- ward range shift for the nudibranchs found at new northernmost localities in 2015 was 151 km (SD =113 km, n = 8). This includes Flabellina cooperi moving north from Elkhom Slough to Santa Cmz, but not to Coleman Beach, where it was found in 2009-10. Notably absent from the 30 species listed above is Phidiana hiltoni. The northward spread, starting in 1977, of this large aeolid nudibranch from Monterey appears to have been stalled since 1 992 by a combination of oceanographic and geographic features in the vicinity of Dux- bury Pveef and Point Reyes and the short duration of its lecithotrophic larval stage (Goddard et al. 2011). Further spread north of this species should be closely monitored and may depend on a different mechanism, such as chance rafting of adults or egg masses on drift macro-algae or other floating substrata supporting growths of it hydroid prey. Goddard (1987) surmised that the rarely observed dorid nudibranch Crimora coneja “may... be primarily sublittoral, with rare intertidal outbreaks.” Its appearance intertidally in northern California and Oregon in 2015, especially at new sites, combined with the timing of much of its historical occurrence at Cape Arago (see Results), is consistent with increased onshore transport during warm-water events dri.ving larval recruitment from subtidal populations, part of the overall mechanism driving intertidal recruitment of the southern nudibranchs we observed in this study. The same may apply to Emarcusia morroensis, another rare species, which prior to this year, had to our knowledge never been observed on open coast rocky shores. NUDIBRANCH RANGE SHIFTS IN NORTHEAST PACIFIC 37 Coos Bay - * -■t" ;' ' ' ^ ‘ Deschuljes Naf/onaf Fores ( ^ •: .. .'. * ■ .r ‘ 'lo ‘.rJ'!- ^ -H ■ ' ■■■< Cap^Arago t ■/ ■ t t fl t ! f / ,■ ' ! t f t : ! ? m ♦ttttt + tttl'n. V/, ^ ^ tt t tf^ 'll >! 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O t ♦♦♦♦♦<♦ ^ ’, 4' ♦♦♦♦♦< H TlTi S 'll ? tt ♦ ♦ n ♦ n 7 Ytttm HI •!♦♦♦♦♦♦♦♦ .-T.';;ttttttttt1'l'i‘i'|7|'i yyfy 7 ?♦♦♦♦♦♦♦♦ 1 'll Tl 'll 7l ♦ ♦♦♦♦♦♦♦♦‘I'I'I'I'I'ITI Sacramento tl° Grove oVacaville Antioch Stanfs/^ ]$|^:,.^^ancisco' . Modesto o Fig. 9. Surface current flow, 6 February 2015, San Francisco, California to Cape Arago, Oregon. Source: Coastal Ocean Currents Monitoring Program (COCMP). Acknowledgements We thank Will and Ziggy Goddard for their expert assistance in the field, Jackie Sones and Eric Sanford of the Bodega Marine Laboratory for sharing their observations and knowledge of the intertidal fauna of Bodega Head and Sonoma County, and David Anderson of the National Park Service and Richard Emlet of the University of Oregon for sharing their respec- tive observations of Okenia rosacea in northern California and southern Oregon. We also thank 38 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Kate Wayne, James Treneman and Dana Garves for assisting us at Cape Arago, Emily Anthony for her assistance in southern Oregon, Craig Comu and Anne Donnelly for their hospitality in Coos Bay, and Karin Fletcher for her persistence in helping obtain the dates of observation of southern nudibranchs pictured in Lamb and Hanby (2005). BG thanks Allison Gong and Josh Hallas for support and assistance in the field. We also thank Liz Kools for her assistance at CAS, Gary McDonald for sharing his field data and for permission to use two of his images, and three anonymous reviewers for their helpful comments which improved the manuscript. Finally, we would like to acknowledge Robin and Marisa Agarwal, Spencer Dybdahl-Riffle, Cassidy Grattan, Matt Knoth, Donna Pomeroy, and Ken-ichi Ueda. 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Roses bloom in tide pools; warm water brings southern sea slug to central coast. UC Santa Cruz Newscenter: http://news.ucsc.edu/2015/01/sea-slugs.html. Accessed 12 July 2015. Strab, P.T. and C. James. 2002. Altimeter-derived surface circulation in the large-scale NE Pacific Gyres. Part 1. seasonal variability. Prog. Oceanogr., 53:163-183. Wolter, K. and M.S. Timlin. 20 1 ! . El Nieo/Southem Oscillation behaviour since 1 87 1 as diagnosed in an extended multivariate ENSO index (MEI.ext). Int. J. Climatology, 31; 1074-1087, doi: 10.1002/joc.2336. Bull. Southern California Acad. Sci. 115(1), 2016, pp. 41-71 © Southern California Academy of Sciences, 2016 Seed Collection and Germination Strategies for Common Wetland and Coastal Sage Scrub Species in Southern California Michelle L. Barton^*, Ivan D. Medel^, Karina K. Johnston^, and Christine R. Whitcraft^ ^California State University Long Beach ^Santa Monica Bay Restoration Commission Abstract. — There is a need for a consolidated source of information on native vegetation seed collection and germination strategies in southern California. Published literature on these methods is often experimental, species-specific, and widely scattered throughout online and print media. Planting and restoration strategies may need to be site-specific; however, similar methodological approaches are often utilized allowing for the development of general strategies for seed collection, storage, and germination methods. A better understanding of species-specific seed attributes and growth processes will help restoration ecologists collect high-quality, viable seed, thereby increasing the potential success of the restored vegetation community by reducing plant mortality, project costs, and effort. This paper synthesizes seed collection and germination strategies for native vegetation common to southern California estuarine wetland, coastal dune, and coastal sage scrub systems. Current estimates affirm that over 70% of coastal wetlands in the Southern California Bight have been lost since the 1800’s, with estimates increasing to over 95% for highly urbanized areas, such as Los Angeles County (Stein et al 2014). The magnitude of these losses and the continued degradation of coastal wetland systems, and adjacent upland and coastal sage scrub habitats, threatens the ecological integrity and sustainability of these habitat types and their watersheds. To address these issues, a number of restoration and mitigation projects aimed at restoring lost ecosystem services, increasing biodiversity, boosting resilience, and in the case of mitigation, creating new wetland habitat, are currently in the planning process in southern California (Noss 2000, Zedler 2000). The majority of wetland restoration or mitigation projects develop a site-specific framework of protocols and management strategies outlining a planting and re-vegetation strategy. Planting strategies designed to establish self-sustaining plant communities identify both the species to be included in the restoration and the source of plant material (i.e. nursery stock or local seeds) (Zedler 2001). Restoration plant palettes should be designed to mimic reference or historic site diversity and be composed of an appropriately broad range of species (Zedler 2001, Johnston et al. 2012). Because of their unique location in the landscape as the connecting habitat between marine, terrestrial, and freshwater ecosystems, coastal wetland complexes naturally support a variety of salt marsh, brackish, and freshwater plant species (Lichvar et al. 2014). Species from each of these habitat types should be incorporated into an appropriate plant palette. Evidence also suggests that the wetland-upland ecotone should be considered an extension of wetland habitat for conservation and restoration purposes (James and Zedler 2000, Wasson and Woolfolk 2011). Thus, coastal sage scrub, dune, and transitional species commonly found in the wetland- upland ecotone should be considered in wetland restoration re-vegetation strategies. * Corresponding author: mlbartonl4@gmail.com 41 42 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Once a restoration plant palette has been developed, species-specific plant material (e.g. seeds and seedlings) acquisition and propagation methods must be determined. While plant material can be obtained from local nurseries, collection and propagation of native seed from local sites is considered the most cost-effective and ecologically-sound method of sourcing germplasm for restoration and mitigation projects (Zedler 2001, Broadhurst et al. 2008). Site-specific or nearest neighbor collections are prefen'ed to distant collections and use of nursery stock, as locally- collected individuals are better adapted to community environmental conditions, maintain local genetic integrity, ensure persistence of local eco-types, prevent unintended gene flow, may improve the long-term sustainability of the site, and may enrich the diversity of the wetland plant community (Guerrant 1996, Montalvo et al 1997, Bowler 2000, Zedler 2001, Mitsch and Gosselink 2010, Vander Mijnsbmgge et al. 2010). Non-local genotypes may be maladapted to local site conditions, leading to improper establishment, or negative impacts to plant and animal communities through competition or species hybridization (Bischoff et al 2006, Vander Mijnsbmgge et al. 2010). The retention of local eco-types and genetic information is gaining importance in the field of restoration biology, reflected by the recent inclusion of required onsite and/or near neighbor collections by regulatory agencies overseeing restoration and mitigation work (Bowler 2000). It is also important to note that making collections in many nature preserves requires permits and/or express permission from the regulating agency. A number of techniques exist to propagate plant material for wetland, coastal sage scmb, and dune species. Seeds are often the primary means of reintroducing native plant species to restora- tion sites in a number of habitat types (Montalvo et al. 2002, Merritt and Dixon 2011). Restora- tion sites may be seeded using a variety of techniques (e.g. broadcast seeding, drilling, imprinting, or hydroseeding) or collected, cultivated in a greenhouse, and transplanted to the site (Bowler 2000, Montalvo et al. 2002, Merritt and Dixon 2011). Simple seeding experiments generally are performed with limited success, especially at lower elevations or within tidal wet- land habitats, as seeds often fail to germinate or float away with rising tides (Broome et al. 1988, Zedler 2001). Techniques like hydroseeding that involve mixing seed with water and either mulch, soil, or organic matter prior to application, tend to work well for many wetland and coastal sage scmb species [e.g. Salvia melUfera (black sage) and Eriogonum fasciculatum (California buckwheat)] and may enhance seedling establishment (Zedler 2001, Montalvo et al 2002, Montalvo and Beyers 2010). Transplanting greenhouse-grown seedlings is an effective re-vegetation strategy that may increase the potential establishment success when compared to direct seeding for some species. In one experiment, survivorship of 2-4 month old marsh seedling transplants was over 95% for all but one treatment, much higher than the success rate of direct seeding (Zedler 2001). Seedlings of a variety of halophytic marsh species including Suaeda esteroa, estuaiy seablite, and Salicornia bigeiovii, dwarf pickleweed, and a variety of coastal sage scmb species like A triplex canes cens, four- wing salt bush, have been successfully grown in greenhouses and transplanted for restoration purposes (Zedler 2001, Francis 2009). While use of seeds and seed- lings has been successful for many species [e.g. Achillea millefolium (common yarrow) and Astragalus tener var titi (coastal dunes milk vetch)], effective propagation techniques are spe- cies-specific and other species, like Batis maritime, saltwort, do not readily grow from seed and require use of alternate methods (Zedler 2001). Other common approaches to generate plant stock include use of cuttings, root division, and direct transplantation of seedlings or mature plants to the site of interest (Zedler 2001, Baskin and Baskin 2014). Direct transplantation of coastal sage scrab seedlings [e.g. Artemisia californica (California sagebmsh), Salvia melUfera, Encelia catifornica (California brittlebush), and Eriogonum fasciculatum] and mature plants salvaged from donor sites have been used with SEED COLLECTION AND GERMINATION IN S. CALIFORNIA V/ETLANDS 43 great success in mitigation efforts (Bowler et at. 1994, Bowler 2000). Similarly, use of transplants, sod, and small plugs of wetland soil, have been effective in introducing a number of wetland species, including Spartina foliosa, California cordgrass, to sites (Tmka 1998, Zedler 2001, Mitsch and Gosselink 2010). Use of cuttings is documented to work well for other spe- cies; cuttings of Salicornia pacifica, common pickleweed, for example, have been successMly propagated by Tree of Life Nursery in San Juan Capistrano, California. While each of these approaches has merit, the discussion in the remainder of this paper (and the accompanying appendices) focuses on the use of seeds and greenhouse-grown seedlings to target a data gap in peer-reviewed literature. Udiile general techniques for successfully establishing common wetland and coastal sage scrub species described in the preceding paragraphs are understood (Broome et al. 1988), the field of restoration biology is still developing and could benefit greatly from additional research. More specifically, the field could benefit from research regarding species-specific collection and propagation techniques because cultivation and planting strategies are often species-specific, highly variable, proprietary, or experimental. Information for many native species of interest does not exist, or is not publically available, forcing restoration managers and ecologists to rely on general information about the genus or costly and time-intensive exploratory studies (Dreesen and Harrington 1997). Publically available sources are scattered throughout a variety of peer-revicAved and non-peer-reviewed resources. With over a dozen wetlands in southern California considered candidates for large-scale wetland restoration projects, a compilation of literature summarizing re- vegetation strategies for the region is needed (SCC^^RP 2001). This paper synthesizes basic seed characteristics, as well as collection and germination strate- gies for vegetation species common to estuarine wetland and adjacent upland habitat types, spe- cifically coastal salt marsh and coastal sage scrub habitats in southern California. Materials and Methods Common seed collection, germination, and propagation techniques are described in the text of this paper. General species information (e.g. scientific name, common name, and habitat type) is included in Appendix I. Detailed species-specific data and recommendations are included in Appendix II, which summarizes available information for 66 native plant species commonly used in southern California coastal restoration projects. Species-specific details were compiled using available literature. While the majority is derived from peer-reviewed publications, some non-peer reviewed literature was included to fill data gaps in published infor- mation. As many data gaps exist, and gray literature was used throughout the article text and the accompanying appendices, the authors have chosen not to distinguish gray literature with footnotes and this was approved by the editors. Instead, these sources are listed, with all peer- reviewed sources, in the Literature Cited section. In instances where duplicate information was identified, the source with the most extensive experimental results was cited. Field observa- tions from the Ballona Wetland Ecological Reserve, Los Angeles, CA, were used to determine some seed collection windows. Appendix II is not intended to be comprehensive; instead, it focuses on common coastal wetland and upland species in southern California for which there was available literature. Priori^ was given to infonxiation specific to southern California coastal habitats, but species-specific information from other geographic areas was included as needed for completeness. Implementation of specific methods may vary slightly by site or project. A number of resources exist that provide general species profiles of the plants described in Appendices I and IL Three websites in particular, S&S Seeds (http://www.ssseeds.com), the Theodore Payne Foundation (http://theodorepayne.org), and Tree of Life Nursery 44 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1 . Suggested field, lab, and greenhouse equipment for seed collection, cleaning, and germination. Field equipment Lab/greenhouse equipment Collecting bins/paper bags Sieves of varying sizes (500 um-2 mm) Ziploc bags Paper envelopes Pens/pencils/markers Freezer Paper clips/binder clips Refrigerator Field data collection sheet Oven Clipboard Growing medium* Background documentation (recommended) Sterile petri dishes* Mesh screens/sieves"^ Hydrogen peroxide (H2O2)* Tarp(s)^ Nail clippers* Gloves'^ Mothballs* Gardening shears'^ Jepson manuaf Ethylene (ethephon or sliced apple)* * = species specific = optional (http://www.califomianativeplants.com), are recommended for supplemental information relating to life history and planting recommendations. Materials Equipment and supplies needed for seed collection, cleaning, and germination are highly variable based on the specific vegetation species. Recommended field, laboratory, and green- house equipment are listed in Table 1 . In addition to the field equipment listed, available back- ground information (e.g. reports, vegetation maps, taxonomic keys) should be brought into the field to aid correct taxonomic identification of species. Seed Collection Seeds should be collected within seed zones, geographic zones in which genetic exchange naturally occurs. Practitioners are advised to use life history traits, landscape context, and avail- able genetic studies to correctly determine seed zones (Krauss and He 2006). It is important to note that due to extensive urbanization and fragmentation in southern California, historic areas of seed exchange have been diminished. In addition to considering provenance of seeds, care should be taken to ensure that seed collections contain sufficient genetic diversity (Vander Mijnsbmgge et al. 2010) as diversity safeguards against disease, environmental fluctua- tions, and inbreeding depressions (Smith et al. 2007). To maximize the range of genetic diver- sity represented in the collection, seed should be collected fi-om 10-50 individuals per population (Lippitt et al. 1994, Vander Mijnsbmgge et al. 2010). Local adaptations and site- specific variability should also be taken into consideration, but site-specific recommendations are outside the scope of this product. When collecting seeds, less intense and more frequent seed harvests are preferable to infrequent and intense harvests (Wall 2009). Negative impacts on the seed source population must be considered (Krauss and He 2006). A general safe harvest- ing recommendation is to take no more than 5% of seed from a given species and geographic area (Zedler 2001). Once plant identity has been confirmed, carefully examine the seeds to assess maturity. Avoid collection of immature seed, as premature collection may result in low seed viability (Bonner and Karrfalt 2008, Baskin and Baskin 2014). In general, it is good practice to begin collecting seeds around the time that natural dispersal begins (Baskin and Baskin 2014). Seeds are consid- ered ripe if seed capsules are dry and tan or brown in color, rather than yellow or green (Lippitt SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 45 Table 2. General seed collection method based on plant anatomy (Wall 2009). Frait/seed type Collection techniques Moist fruitsdjerries Hand-pluck fruits. Dehiscent species Collect entire inflorescences prior to dispersal. Alternatively, secure cloth bags around ripening stalks to capture dispersed seed. Inflorescences Strip inflorescences. Seed heads Shake ripe seed directly onto a tarp or collection bag underneath the target plant. Seed clusters Remove entire seed cluster from plant. et al. 1994, Bonner and Karrfalt 2008, Baskin and Baskin 2014). Frequent visits to collection sites are suggested to repeatedly assess seed stage within the recommended collection time win- dow. For species with insufficient published seed collection data or infomiation, e.g. Artemisia dougiasiana, detailed fi,eld notes are essential to pinpoint the ideal collection window and suc- cessfully collect seeds. Once the seeds of target species are deemed ripe, the collection process can begin. Collection/ isolation of seed varies based on plant anatomy. Observe the plant and note if the species has berries or dry fruits, dehiscent or iedehiscent seeds, and note if seeds are in seed heads or seed clusters as collection methods vaiy for each categoiy (Table 2). Addirionally, if a species is known to be dioecious [e.g. Croton californicus (California croton), Baccharis spp., Saiix spp.], care should be taken to ensure that sufficient seed quantities are collected from both male and female plants (Clarke et al. 2007). Vouchering specimens from, collected seeds is a good practice and should be considered during the planning phase. Seed Cleaning Seed cleaning removes floral parts, seed coats, pods, fleshy fruit material, and other debris from seeds (Jorgensen and Stevens 2004). Machinery, including aspirators, hammermills, fan- ning mills, and blowers, exists to aid large-scale seed enterprises. Hammermills, fanning mills, and blowers help isolate seed and remove chaff and floral parts (Shaw 1975, Jorgensen and Stevens 2004). Although seed cleaning machinery is useful, cleaning for small-scale projects can be efficiently performed by hand (Bonner and Karrfalt 2008). To isolate seeds and remove excess chaff, remove seeds from branches and large floral parts. Then, rab remaining seeds and floral parts over a sieve. Once seeds are isolated from chaff, only retain seeds that look healthy and ripe (i.e. dark brown/tan in color, flilly-formed). For some species, chaff does not present a huge problem, and it may be more efficient to seed with some chaff Discard seeds that appear sickly or defnnned. If the seed is contained in a capsule, gently crush the capsule by hand or with a rolling pin. Removal of woody capsules, as seen in Abronia spp., may also be aided with the use of generic nail clippers (P.M. Drennan, personal communication). Seed Storage For the greatest germination yield, storage time should be minimized, and use of newer seeds should be prioritized. While native seed longevity varies by genus and species, a number of seeds are knov/n to be short-lived. For example, seeds of Lycium californicum, California box-thom, are viable for up to one year at most. While seeds of other species [e.g. Atripiex spp.. Astragalus spp., and Lupinus chamissonis (dune bush lupine)] will remain viable for much longer (i.e. 4-10 years), the germination rate of seeds in long-term storage will likely decline over time. In addition to reducing germination rate, long-term storage will often induce seed coat or embryo dorniaiicy, and stored seeds may need to be treated prior to planting. For example, the hard seed coat of Astragalus tener van titi seeds may require scarification, 46 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES or mechanical scraping with sandpaper, a file, or a knife, to initiate germination if stored for an extended period of time (Baskin and Baskin 2014, USFWS n.d.) The longevity of certain seeds can be increased if best management practices for storage are followed for the species and/or general seed storage procedures are applied. Most dry seeds should be stored at low temperatures, 10-15.6°C (50"60°F), and low humidity, less than 40% relative humidity (Jorgensen and Stevens 2004, Recon Native Plants Inc. 2015). Substandard storage in conditions with fluctuating temperatures or high humidity may result in significant seed loss (Merritt and Dixon 2011). Germination Considerations Successful propagation of southern California coastal plant species requires a thorough understanding of seed germination ecology. Seed germination is dependent upon a number of evolutionary and ecological factors which generally must be observed, and often replicated, in the laboratory or greenhouse to successfully grow propagules. These, often species-specific, factors include, but are not limited to: germination timing/seasonality, environmental condi- tions, such as temperature, soil texture, soil moisture, soil salinity, light availability, presence of smoke, and seed age, and dormancy state, both at the time of maturation and dispersal (Baskin and Baskin 2014). Germination Timing Seeds are adapted to germinate under favorable environmental conditions (Deberry and Perry 2000). An understanding of natural germination timing is helpful in determining the environ- mental conditions that best promote germination of a particular species in the greenhouse or laboratory. This is particularly true, as in both the greenhouse and laboratory, environmental conditions can be manipulated to mimic natural seasonal variation. Temperature, moisture, and light are generally controlled for this purpose (see Temperature’ and ‘Light’ sections below) (Noe and Zedler 2001). Temperature Understanding germination timing under natural conditions will often indicate what range of temperatures best promote germination. Temperature influences germination directly through regulation of enzymatic reactions, or indirectly by controlling the synthesis of hormones that alter seed dormancy. While temperature is an important determinant in the regulation of both germination and dormancy, response to temperature in freshwater wetland species seems to be dependent on habitat, not phylogenetic relatedness. Temperature interplays with other envir- onmental conditions to promote germination (Brandel 2006). Further, the germination rate of certain species is enhanced with simulated temperature fluctuations, rather than constant tem- peratures. WTiile response to fluctuating temperatures depends both on specific species and habitat, a few generalities exist. Both small-seeded species and forbs tend to respond well to fluctuating temperatures while larger-seeded and graminoid species do not show as marked a preference for temperature fluctuations (Liu et al. 2013). Soil Texture To grow seedlings, clean, viable seeds should be planted in mixtures of sand, top soil, and peat moss or vermiculite (Broome et al. 1988). To achieve the greatest germination rate, the exact composition of the mixture should be tailored to the individual plant species of interest. Life history and preferred habitat of the species should be considered when determining optimal SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 47 soil coEditions. For instance, Abronia maritima, which naturally occurs on sandy dunes, should be sown in soil consisting largely of sand, or other coarse grains. Soil Moisture Soil moisture must also be considered when sowing seeds (Noe and Zedler 2000, Noe and Zedler 2001). Most mature seeds must imbibe in the early stages of germination to activate enzymes (Deberry and Perry 2000). After seeds imbibe, sufficient, and relatively constant soil moisture is needed to ensure proper germination (Bonner and Karrfalt 2008). Most species in southern California salt marsh systems germinate well in moist soil at low salinity (Zedler 2001). Experiments suggest that Distichlis spicata grows best with a fluctuating inundation regime, where inundation was varied over time, but the soil surface was never completely dry (Elsey-Quirk et al. 2009). Germination of other high marsh plant species is highest with 41-51% soil moisture (Zedler 2001). It is important to note that while seeds of wetland species are adapted to wet conditions with limited oxygen, coastal sage scrub and upland transition species are more sensitive to inunda- tion. For these species, excessive exposure to water can be problematic, causing seeds to become waterlogged (Fenner 1992, Deberry and Perry 2000). Following germination, water regimes, that specify both the quantity and frequency of water application, both in the green- house and in natural environments, may influence growth rates and should be carefully considered. Soil Salinity Another major factor that influences germination is soil salinity (Noe and Zedler 2000, Noe and Zedler 2001). Certain halophytic species, like Salicomia bigeiovii germinate to higher per- centages under somewhat saline conditions (0.05-0.09 M). In general, although halophytes are salt-tolerant, high perceiiiages of halophyte seeds will germinate in distilled watei. Pxsults of salinity experiments suggest that seeds will often germinate to higher percentages in distilled water, as seeds tend to be sensitive to salt concentrations, and exposure to excessive salt can drastically decrease germination yields. Still, much variation exists in the germination of halo- phyte species in saline environments (Baskin and Baskin 2014). Light Light is another environmental factor that affects germination. Exposure to light is often required for germination to occur. Exposure to light has been documented to improve germina- tion rates for certain species [e.g. Eriogonum fasciculatum (California buckwheat), Baccharis salidfoUa (mule fat)] (Zedler 2001, Boiinei and Karrfalt 2008). Still, exposure is not always sufficient to ensure the successful occurrence of germination mechanisms. Duration of exposure to light (i.e. day length, or photoperiod) also plays an important role in seedling emergence and growth of southern California natives (Sprague 1944, Noe and Zedler 2000, Gieiner and Kohl 2014). For instance, long-day conditions (16 hours of light for eveiy 8 hours of darfjiess) are necessary to successfully culture Oenothera species (Greiner and Kohl 2014). Photoperiod may also influence other processes, such as flowering. Melica imperfecta and Stipa lepida have been shown to flowei 10-20 weeks faster with constant light (i.e. 24-hour photoperiod) when compared to an B-hour photoperiod (Ashby and Helliners 1959). Smoke Treatments Southern California, like most regions with Mediterranean climates, is subject to frequent and intense wildfires, and certain species have adapted to be fire-tolerant (Keeley and 48 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fotheringham 1998, Crosti et al. 2006). Germination of fire-tolerant species is generally enhanced by exposure to fire or smoke (Crosti et al. 2006, Baskin and Baskin 2014). Smoke- stimulated germination, via exposure to liquid or aerosol components of smoke, may be useful for many coastal sage scrub species. For instances, exposure of Salvia mellifera seeds to smoke or other components of fire, like charred wood or potassium nitrate (KNO3), may help stimulate germination (Montalvo and Beyers 2010). Other Considerations In some instances, information regarding the necessary conditions or procedures to promote germination is not readily available for a particular species. In such situations, it is advisable to consult local experts that may have species-specific knowledge. Alternatively, simple tests or experiments manipulating a variety of the environmental factors discussed above may be performed. Germination Testing If a seed lot requires germination studies, it is preferable that they are conducted shortly after seed collection, within 7-10 days, to ensure seeds are viable and have not entered seed dormancy. Germination trials can test outcomes of various pre-treatments and/or growing con- ditions. They are often also used to express the quality of a seed lot (Lippitt et al. 1994). The results of germination trials are typically reported as percentage germination or germination rates. Percentage germination is the percentage of seeds that germinate under the specified set of conditions. Comparing germination rates of a variety of treatments allows easy determination of the most effective combination of germination conditions. While germination rates are useful, the industry will often use other terms to describe the percentage of seed that will germinate under a given set of conditions. Pure Live Seed (PLS) is a common way to express viability. PLS is calculated by multiplying the percentage of pure seed by the percentage of total viable seed and dividing the product by one hundred (S&S n.d., Showers 2010). Other measures include specification by purity, bulk pounds, or PLS pounds (S&S). Dormancy Considerations Seeds for a number of wetland plants are known to be dormant. In these species, seed dor- mancy must be broken to promote growth and germination (Baskin and Baskin 2014). The process is generally moisture and temperature dependent, but varies both with species and type of dormancy. Three types of dormancy should be considered: physical (or seed coat) dormancy, internal dormancy, and morphophysiological dormancy. Seeds with physical dormancy have seed coats or other structures that are impermeable to water and/or oxygen (Lippitt et al. 1994, Baskin and Baskin 2014). This form of dormancy is generally broken by penetrating/ opening the seed coat or specialized structure that excludes water or oxygen. This can be achieved through scarification, cold and warm stratification, or exposure to dry heat, charate, fire, acid, and light. Internal dormancy, caused by a physiological mechanism that inhibits ger- mination, is generally broken through use of warm and/or cold stratification. Morphophysiolo- gical dormancy is similar to physiological, but seeds with this type of dormancy also have an underdeveloped embryo. A variety of methods can be used to break morphophysiological dormancy, including: scarification, submersion in hot water [82-93°C (180-200°F)], treatment with dry heat, exposure to fire, acid, mulch treatment, cold stratification, warm stratification, and exposure to light (Emery 1988, McClure 1997, Baskin and Baskin 2014). Common dormancy breaking methods are detailed in Table 3). SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 49 Table 3. Detailed methodology for techniques commonly employed to break seed dormancy. Method General description Scarification Hot water treatment Dry heat Charate Fire Water Cold stratification Warm stratification Mechanically scar seed coat with sandpaper, knives, files, or clippers. Alternatively soak seed in acid or hot water (Emery 1988, Lippitt 1994, Bonner and Karrfalt 2008). Place seeds into hot water (1 80--200°F) and leave them to soak as the water cools (Emery 1988, Bonner and Karrfalt 2008). Expose seeds to 180"212°F heat. Use of an incubator, rather than oven, preferred (Emery 1988). Expose seeds to ash from burned plants. This may neutralize germination inhibitors in species that naturally germinate when exposed to fire (Emery 1988, Baskin and Baskin 2014). Expose seeds to direct flame. This may be effective as a means to spur germination in species that naturally germinate when exposed to fire (Baskin and Baskin 2014). Soak seeds in water to leach out water-soluble inhibitors (Baskin and Baskin 2014). Store seeds in cold conditions (35-4 1°F) for 1-3 months to simulate winter conditions (Bonner and Karrfalt 2008, Elsey-Quirk et al. 2009, Baskin and Baskin 2014). Store seeds in warm conditions (65°F or higher) (Baskin and Baskin 2014). Unfortunately, as indicated by the variety of conditions listed above, there is not one prevail- ing standardized method to break seed dormancy. Again, methods vary based on the life history of the species. Species-specific life histories, available at the growers’ websites listed above, can be a good indicator of the required conditions for that species. For example, species that typically germinate in early spring after a cold and/or rainy winter, such as Platanus racemosa, western sycamore, often require cold, moist stratification mimicking natural wintering to break dormancy. Other species, such as Acmispon glaber, common deerweed, require heat treatment to break dormancy which also correspond with the life history of that species; A. glaher does particularly well after wildfire events. However, treating seeds to break dormancy is not enough to guarantee germination. Germination requirements must also be considered. Methods and information should be supplemented by experimentation when necessary. Mycorrhizae Establishing functional ecosystems also requires consideration of subsurface components of the system. Many plants have symbiotic relationships with soil-inhabiting microorganisms, yielding root systems that are more effective at extracting water and nutrients from the rhizo- sphere (i.e. soil profiles influenced by root secretions and soil fauna). The fiingus-root system is called mycorrhizae (Gerdemann 1968, Tree of Life Nursery n.d.). Research has shown that mycorrhizae can increase plant growth and are essential in successfully establishing vegetation during restoration and mitigation projects (Reeves et al. 1979, Allen and Allen 1980, Cooke and Lefor 1990). If planting areas are severely disturbed and lack a healthy rhizosphere, steps should be taken to ensure presence of mycorrhizae, or to increase the potential for natural development. As the presence of mycorrhizae is important in establishing many wetland and coastal sage scrub species, container plants are often inoculated prior to planting (Cooke and Lefor 1990, Bowler 2000). Seedlings can be inoculated with a spore suspension or via introduction of small amounts of collected soil from sites with a healthy rhizosphere to a sterile soil (van de Voorde et al. 2012). Starter-cultures are also available commercially. Discussion Southern California has lost a significant portion of its coastal ecosystems due to urban devel- opment, agriculture, invasive species, and in the case of coastal estuarine wetlands, severely 50 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES modified hydrology resulting from both channelization and deposition of fill sediments (Westman 1981). Loss of these ecosystems is concerning because they provide valuable ecosys- tem services including supporting important fisheries, filtering water, sequestering carbon, and providing habitat for a diversity of plant and animal life, including a number of threatened and endangered species. Wetlands are buffered by transition habitats, and many wetland-associated species also require adjacent upland habitat areas to breed, roost, or to have the highest likeli- hood of survival. Plant species in Southern California also display a high degree of endemism and the Southern California coast is considered a global biodiversity hotspot (SCCWRP 2001). Although wetlands in southern California have attained protected status and efforts are being made to restore degraded habitats throughout the Southern California Bight, the increasing human populations along the California coast will continue to impact these coastal ecosystems (Callaway and Zedler 2004). To preserve the spectrum of ecosystem services coastal wetlands and their adjacent upland habitats provide, managers throughout the southern California region need to work collectively to conserve remaining high quality coastal wetland habitat and to restore lower quality, degraded habitats. Clearly, there is a significant and ongoing regional need for restoration projects to recover lost habitats and preserve the unique communities. Increased reliance on ecological restoration of vegetation assemblages emphasizes the need for sound, scientifically-tested techniques to ensure the successful reestablishment of plant communities. While this document is not com- prehensive, and there is still a practical need for land managers to compile detailed site informa- tion and evaluate site-specific experiments prior to implementing a restoration scheme, this literature review compiles available seed collection and germination information for the southern California region and provides an initial assessment of published methods for common wetland, dune, and coastal scrub plants. Many unknowns remain in restoration ecology theory, and understanding of the most effective restoration practices remains incomplete. Knowledge gaps regarding the collection and germination requirements of integral species [e.g. Hazardia squarrosa (saw-toothed goldenbush)] and other species with limited research available [e.g. Elymus triticoides (creeping wild lye)] precluded their inclusion in this review. Planners are encouraged to conduct regular site monitoring and employ adaptive management strategies. In this way, progress can be evaluated and unexpected outcomes and shortcomings can be corrected. Still, there is a regional need for additional research regarding seed phenology and maturation of southern California species. Although a number of wetland, dune, and coastal sage scrub restorations are planned in southern California, information regarding seed collection and ger- mination for many naturally occurring species is not readily available. Therefore, the field of plant community restoration could benefit greatly from additional research regarding seed phe- nology and maturation, both in the form of species-specific experimentation and literature and broader-scale, regional or ecosystem-based reviews. Filling in existing knowledge gaps and developing a better understanding of seed processes will help restoration ecologists collect high quality, viable seed, thereby increasing the potential success of the restored vegetation community by reducing seed/seedling mortality, restoration cost and human effort. Perhaps more importantly, the region could benefit from the development of a coordinated network of restoration ecologists. Compilation of this literature review suggests that information regarding the restoration of wetland plant communities is abundant, but it is dispersed, produced by various sources, and often proprietary. Intentional withholding of information by nurseries or private environmental consulting firms inevitably leads to duplication of efforts by groups working in the southern California region and surely impacts both the overall quality of restored habitats and project efficiency. Engagement and cooperation of existing private industry groups SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 51 aEd public sector regulators with a vested interest in restoring coastal wetland plant commu- nities would be a major victory and a tangible step forward for the threatened coastal ecosystems in the region, \¥hile establishing vegetation in restored wetlands is a vital component to the overall restora- tion scheme, it is just a small part of the overall restoration process. Restoring wetland ecosys- tems is complex; plans must incoiporate vegetation, hydrology, substrate, and marine and terrestrial animals. To fulfill restoration aims, well-informed, inter-disciplinary approaches that incorporate ecologists, engineers, managers, lawyers, and practitioners from other technical fields are needed (Zedler 2000, Kiehl 2010). Inter-disciplinary approaches will best foster creativity and progress knowledge and understanding in the field of restoration. 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Wetlands of the southern California coast: Historical extent and change over time. i+50. Stettler, R. F., H. D. Bradshaw Jr., P. E. Heilman, and T. M. Hinckley. 1996. Biology of Populus and its implica- tions for management and conservation. NRC Research Press, Ottowa, Ontario, Canada. 539 pp. Stevens, M. e.d. Plant guide: Pacific siiverweed. Baton Rouge, Louisiana: USDA NRCS National Plant Data Center. . 1994. Plant guide: White sage. Baton Rouge, Louisiana: USDA NRCS National Plant Data Center. . 2003. Plant guide: California bulrush. USDA NRCS National Plant Data Center & Idaho Plant Materials Center. Tilley, D. J., D, Ogle, L. St. John, and B. Benson, n.d. Plant guide: Big sagebrash, USDA, Natural Resources Conservation Service, Idaho State Office. Boise, Idaho. Tree of Life Nursery, n.d. Mycorrhizae, Accessed February 24, 2015. http://v/ww^. califomianativeplaets.com/. Tmka, S. 1998. Environmental and parental height form effects on the growth of Spartina foliosa in southern Cali- fornia. Master’s Thesis. San Diego State University, San Diego, California, USA. U.S. Fish and Wildlife Service, n.d. Astragalus pycnostachyus var. lanosissimus (Ventura Marsh Milk- Vetch): 5- year review; Summary and evaluation. Ventura, California. van de Voorde, T. F. J., W. H. van der Putten, and T. M. Bezemer. 2012. Soil inoculation method determines the strength of plant-soil interactions. Soil Biol. Biochem., 55:1-6. doi:10.1016/j.soilbio.20T2.05.020. Vander Mijnsbragge, K., A. Bischoff, and B. Smith. 2010. A question of origin: Where and how to collect seed for ecological restoration. Basic Appl. Ecol, 11(4):300-311. doi:10.1016/j.baae.2009.09.002. Walker, J. 2005. Potentiiia anserina spp. pacifica. Univeristy of Washington, http://depts.washmgton.edu/ propplEt/Plants/Potentilia anserina.htm. Wall, M. 2009. General seed collection guidelines; For California native plant species. Rancho Santa Ana Botanic Garden. Accessed February 24, 2015. http://www.rsabg.org/. , and J. Macdonald. 2009. Processing seeds of California native plants for conservation, storage, and restoration. Claremont, California; Rancho Santa Ana Botanic Garden Seed Program. Wasson, K., and A. Woolfolk. 2011. Salt marsh-upland ecotones in central California: vulnerability to invasions and anthropogenic stressors. Wetlands, 31(2):389^02. doi: 10. 1007/sl3 157-01 1-0153-z. Westman, W. E. 1981. Diversity relations and succession in Californian coastal sage scrub. Ecology, 62(1 ): 170-84. Young, B. 2001a. Propagation protocol for production of container A Irfpfex californica moq. plants (treeband #5). San Francisco, California: Native Plant Network. . 2001b. Propagation protocol for production of container Croton californicus muell.-arg. plants (2 inch pot). San Francisco, California: Native Plant Network. . 2001c. Propagation protocol for production of container Diplacus aurantiacus w. curt, aurantiacus (w. curt.) Jepson plants (deepot 16). San Francisco, California. . 200 Id. Propagation protocol for production of contmnei Hordeum brachyantherum eevskii plants (leach tube). San Francisco, California: Native Plant Network. . 200 le. Propagation protocol for production of container Jaumea carnosa (less.) gray plants (treeband #5). San Francisco, California: Native Plant Network. . 200 If. Propagation protocol for production of container Limonium californicum (boiss.). San Francisco, California: Native Plant Network. 56 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES . 200 Ig. Propagation protocol for production of container chamissonis eschs. plants (deepot 40). San Francisco, California: Native Plant Network. . 200 Ih. Propagation protocol for production of container Melica imperfecta trin. plants (leach tube). San Francisco, California: Native Plant Network. . 200 li. Propagation protocol for production of container Rosa californica Cham. & Schlecht. Plants (dee- pot 40). Moscow (ID): University of Idaho, College of Natural Resources, Forest Research Nursery, Native Plant Network. . 200 Ij. Propagation protocol for production of container Salicomia virginica 1. plants (leach tube). San Francisco, California: Native Plant Network. . 2001k. Propagation protocol for production of Frankenia salina (molina). San Francisco, CA: Native Plant Network. . 2002. Propagation protocol for production of container Triglochin maritimum 1. plants (treeband #5). San Francisco, California: Native Plant Network. Young, J. A., and C.D. Clements. 2003. Seed germination of willow species from a desert riparian ecosystem. J. Range. Manage., 56:496-500. , B. L. Kay, H. George, and R. A. Evans. 1980. Germination of three species of Atriplex. Agron. J., 72:705-709. , and C.G. Young. 1986. Collecting, processing, and germinating seeds of wildland plants. Portland, Ore- gon: Timber Press. Young-Mathews, A. 2010. California sagebrush. USD A- Natural Resources Conservation Service, Plant Materials Center, Lockeford, CA. Zafar, S. L, and W. H. Shah. 1994. Studies on achene germination, transplantability, salinity tolerance, and cultivation of gumweed {Grindelia camporum) in hot and semi-arid conditions. Field Crop. Res., 37 ( 1 ):77-84. doi: 1 0. 1 0 1 6/0378-4290(94)90083-3 . Zedler, J. B. 2000. Progress in wetland restoration ecology. Trends Ecol. EvoL, 15:402-407. doi:I0.1016/S0169- 5347(00)01959-5. . 2001. Handbook for restoring tidal wetlands. Boca Raton, FL: CRC Press EEC. 1^64. Appendices Appendix 1. Species-specific habitat associations for wetland, coastal sage scrab, and upland transition species common in southern California. This table includes scientific and common names from Jepson eFlora (http://ucjeps.berkeley.edii/, accessed June 4, 2015). Habitat association infonnatioii is derived from Jepsoe and finther refined with information available from the Manual of California Vegetation (2“*^ edition), the S S Seeds Plant Database (www.ssseeds.com/plaEt-database), and the species-specific literature cited in Appen- dix n (Baldwin et al. 2012, Sawyer et al. 2009, S&S Seeds, n.d.). SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 57 ^ 2 M 2 2 I O S d I I ^ I M s ffi g o 5^ ^ B X X X K XX XXX X X X XXXX XXX X XX X X XXXX X X XXX XX X X XXX XX X X X XX X X X 1 1 •"S C -a I S S; S •S -2 S 5 g g I o Q 2 -S 2 5 8 2 i'il o ^ -c ^ g S “I o o 5 X "y Q Q Q ■a -S =3 P P P t V fr ^ ^ ^ 0 K »2 ii« a > y 1 1 1 S; I & ass 8, a.§ ■b 2 ’P ^ ^ ^ a - -8 k I 111^ g I II, 1 S J -s -g -g J J J J J a, .o, .Cl, .o, o o o ^ Qq ^ Distichlis littoralis Shore grass Distichlis spicata Salt grass Encelia califomica California brittlebush Eriogonum fasciculatum California buckwheat Frankenia satina Alkali heath Low Mid High Salt Low High Fresh Salt Scientific name Common name marsh marsh marsh pan transition transition Grass Scrub water tolerant 58 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES XX X XX X XXXX X XXX XX X XX XX XXX X XX X X XX xxxxx XXX XX XX X X X 5 o S 5 is P « 5 ^ 5 s ^ S s I ^ I s L P S’ J ^ •2 I 5 ^ I g g I "S ja 8 8 O 5 A § •S -S . ^ s ^ d a a f I 9^ 5 S ^ 'S I i g s ^ S Q »S “S O S o I S -s K a ^ 'C Q a, a. a § a K _.13 0. ^ « 5 K 9 “C ;i I ^ S sS « ^ '5 a ? ^ -2 i “i 2 I I g 8 ^ O O Q ^ a, ^ DO ^ -a X sc, J o •2 -2 5 ^ § ;2 ^ ;S 5 Do Co &0 I .c ^ I I .C3 .Q S' S- O Q K K o o Co Co SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 59 X ’’B g &, < 3 I o fa ^ ■V ^ -3 I 5 § -g 1 1 I e ^ fa W ffl ^ U M g a, ^ « ^ &0 Co « 5 2 I I sS a em California. leformation is sorted alphabetically by scientific name. 60 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 2 I I ^ 2 eo -a I s “S s -5 « &o TO ^ irt O ^ « I I « O K ed d S d S S ts! .s s 1 ^ 1 2 a ^ 5 n - 2 M Is s g to ’-g S p e S « ft 60 o ft ^ g p ^ -i ^ ft 60 s ^ ^ O 3 O 6fl !r o 1 S-.S X ^ o S O ^ feo d ft P to 03 o ^ rife flit iSB e 2 -S c 'd S « s « c c« ^ CD 2 '3} ^ d 'S &D « S 2 s ^ .s S d o I s g I w .g -a I 8 c*_ . 4, C 'S| I s I CO 4 b . 'd S a> 2 e- •c 2 QO ^ s 2 § I I I 03 S -o 2 ft •s § ■d « « d ^ 2 2 8 !! M _ « o > a o ed rt s I . O « -rt ^ d ^ ed *-> o v> d At ii I i ffl 1 § o C-1 6 d a 5 •S § 5 § •e 2 ® 2 .2 c« § ^ U ^ CN g 2 J « 0-1 ao p < S 2 .s - o 2 2 c o 2 « o o ^ o CNI §1 !■§> I ^ 5 2 S, ffl -5^ d « I s 2 S, ® ^ d d I ^ o ffl o w M 2 I, ‘I “F 2 I ® •2 of ^ a .3 ^ a « II "B o ^ _c ^ s 4 "d M to 2 p £ *- «fc .d o M c ™ to ft >4 o to ft ^ •*"* o to 2 to eS 03 d o e .2 I ^ 03 d ca 2, « ft o 4) d rd o ^ d d 4) « c M 6 ” tao d I o 2 O ft &a o S 2 E ^ “ •l^-2 ^ O O ^ ' e o CM •S s •a 3 .£ § I -a > o to LoEgevity ScicEtific name Start End Frait/seed characteristics Seed collection details Seed germination (yrs.) Artemisia douglasiama < ! mm, glabrous fruit. Small, Seed is ready to harvest when it Germinates naturally at relatively cool temps. 2-5 Eiidiom Slough National ellipsoid, hairless achenes can be easily removed from the Estuarine Research without ribs or angles. heads by shaking. Clip seed SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 61 -a d o 1 i 5 o O « i '-e 1 1 S G W © > m 2 iaO ^ o o c o -S 8 o © 60 tta u s > 2 m d o e S m © C c 1 t © -a § ,d c 2 U '3 C/3 g I M td I ^ B d ^ 2lt • S Rj rt RJ c -S ca « s ^3 J B I ^ s B J fl 2 .S « i_ g .3 mo o 03 r j m a 03 '3 OJ o •S ’3 ts,, © o > a > CM 2 © o ^ © o I I ^ « I g s 'S _ 0 fr O. 'O “ § 1 - d > S) ^ B S d a ^ a U -5 « o ^ 3 ^ c ■s g § s 3 - a > S -3 CM - g to p B a I B ^ 8 ^ o g *0 ^ 9 1 »n B ^ 3 rri “ "S B 43 'O 2 B © 3 d “ B tc S)^ ,5 o ^ d 2 Ig B ® o B i> S © Cd 03 m ^ & m o © CN s ^ I ^ ^ m *3 ^ --- S ry >- © o ^ 2 a 5 u 8 a e o .A i|®| S I "S s f g US 4; S 4 15 w 05 a © .S d I I 5 2 © ed S ffl “3 O ^ 2 fej 2 ^ m > 2 &. o ’-' © »3 JL, Cl "?« ^ ,d & M 05 ^ rs| i§ LoEgevity Scientific name Start End Frait/seed characteristics Seed collection details Seed germination. (yrs.) Atriplex caiifomica (Young Sep Oct Mature fruit is an utricle with 1 Gently seeds mb over #18 sieve. Pre-planting: soak in water for 24 hours, rinse. 10 2001a) seed. Seeds are black, shiny, Remove as much chaff as 86% germination rate after sowing in peat 62 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES &, .-s ca I ca O 1 2 S ^ 3 ^ ^ S o s ^ ‘3) « .S g ^ 3 -2 c U ^ S rM cfl “ § r< M I . O .s Sw o o O II 2 'g 3 9S ^ « W ^ > 5 -I g- E g 6 E .3 ■g 'd 3 2 S ^ M s B U w- 0) !- ..C « m « — ' o « -a S 6 .S o U < M ts m ■ „ d -d d 13 « « &o d •s g c ca E “ c o p. o ^ «E oa cb 'm -d B ^ ts d 0) &o .S sS o, m O « &, g ‘g ^ I g &0 “ I ^ s ° e 2 ^ • C M .£1 ed cfl o ^ g § 2 p ” ■fi 'O S s £ M ^ rS a - < ® d -e o «5 ’.P P O ^ « « 2 "O S •§ ® a -*’^11 ^’S ^ 6- a ° 1 g I g aj i .s o ^ a I < C3 2 .s I ^ ^ -c 'S I e ^ 6 2 2 DO a 2 M ^ -a m « p « o '« 5-< r- 0) ^ S o « -)- ^ -d 0 c « 2 1 I « « &i « ^ ^ « -o § I §1 Ic 00 hs ^ « & ^ P d "o p -o M -g ^ «5 - -o S « e « ao &o c c S S, g g g S *C *0 fd 2 . ^ ao £ J “ g ^ s 4) ^ feb E E a\ 5 I I .s' 3 ^ « a 2 ^ g ’B a, „ ail ^ a 2 g s 2 I i 3 c P O o S ^ “c p '2 g g, w 2 O i 00 &, < &, 4) C/3 « m « w < CM o' o o^ a *3 !P ^_> ^ d m -p c ^ •C -d « s I fs m m i •«^ K) c 1 11 ■w .S •s J S 2 S d .© m ■ft O fNl s ^ “ 3 fe, I I il s a a •& 5 « N o tP Cs! O CN i 5 5 ^ ^ ^ : ^ o g S,t2 Q < I ^ cN 5 ..3 .! ^ J 1 1^1 ! t N ^ I g S C ^ gq I S ffl S .g S S o I g ^ «S rs| ed .o a SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 63 s o & .S s 19 O I ■§ I I •3 ^ o| Q a ^ |i 1 1 « flj g « o ^ 2 g S *^3 « ® O VD ^ w a ^ a g. &a s g p ^ 3 S t- ^ 3 6 e y a o “ M 'O ’O « 2 ^ a g § 9 p o So g- o S 1 1 S ^ .9 'a ^ .2 S I El® •goo m o w CN s ^ 3 O « « ^ I s ^ I f^. ^ il.^ J 1^ fe o c 0^0 d S "S S « t| 8 ^ w O U » .S I -s a P S 8 * * a-s 2 E 8 I •§ s £ S I ls< « o I p* o o •S VO d H o c^ g ti o fNl ^ g S « I 8 ^ S- M g. « I ft ^ 3 f— I o S i d •’^ -2 a -g g fe» w f -o A « s « N a of 'q a 8 ss P c« .3 i'-s . &il 5 ^ T I 8 §§ i S' M 60 >* s I a r o ■g TO ^ I s ll§ as •s ■« e o « a a Li •s g ^ 09 'O *0 W) p i €U ^ ^ « o -r- 09 m a “ a ^ I e 09 t -a -a 5 ^ i I o S s ^ 09 _J 'O o fl &, w w _ > ^ g ^ 43 * - o e S c<3 ta II ’S O S p « O o a s ^ a 2^0 a « I o « S a ^ 2 ^ _c« eS 2 "S 03 S W 'O > « W ? « O M 09 E M d I I « C o fl m in cn a 0 09 l2 a ^ a c .g 1 6 M S ” o <» >. 'o I = S I i-^ a g 4' i .9 •g 03 o. o m ■a E « o L $3 ® S. a s a' S I ^ M ® o « :S S ^ • “ — a o lil •S CN S3 ^ ® ^ a « o senescence. 64 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES < W 2 ^ “ S I d s ^ ca ■S M d «D § § 23 ^ g ^ ? ^ « O -g ^ ! •S w « ca ? A s-i - ^ ^ IS I -2 I g ge^ ^ QQ .2 « . w ^ P M ^ S 2; 1 1 »o « _'§ o 3 S 5 g .1 ^ o ^ « ^ g s g ^ 3 M « ^ 3 S ^ 53 00 OJ 2 C ^ « s &0 -g M p 'd 4) 2 M §1 s ■S o s I 0) -a ^ ^ £S I 2 o c & S -S g S S d o, b s I a 5 S « 3 ^ ^ ^ e 5 n '^ ' ^ 2 i- s 3 ^ ^ o d Os p O 0\ •g 3 ’S is‘i fe 1 » s ^ ^ 2 o M ^ ^ in ^ II s ■s s s ^03 3 4) S^ ^ ^ .2 ‘S. ^ C/D s 2 S I 'd §" s ,Efl d 31 S CQ 3 l.l w ^ M 'd W S . ffl o^ o o o o CM rs| -d « C S 0) ^ S .S' l| •d o U M « s I 3 ^ • 2 d 1 I M 3' I fa « •c ^• a 3 S d « p &a '-^ I g 11 a «, d p o o O e - S g « +«. g ^ 3 d o « 3 a e f p a> S 4^ bb r ^ y d &0 S « « ^ M ffl w I “ a « 0 I 3 ^ 3 2 t a 1 I o U s « g 1^ a I g 1] &o es d g il “ c « .a d « § y ^ a a a ® s Cfl «C 'S ^ ^ (li d ^ g 2 a &, d II 5 O 3 o a I o ^ f g O vs a> «« .S > o r O Cfl b ^ S 2 'O d s' a 2-- 3 I ^ < 'a 3 2 a d Q _4) S Q »d [=■< S a a a &b « d ^2 w Q 05 3 O d ta O'! 3" d 4) ^ w 3, 3 A' ^ a-c I II s a E s ^ a ^ ^ I •303 p a S Is 5 M P < ?| S. 6 i ^ 00 o a ^ ^ a M S a 05 & a a a. g e o d g M a M C >T-! d « 05 d a s c (U -p a « 2 « a c d a ± fi I a ^ a a ^ ^ S d d d § « M I ^ g I 3 aa g ^ )-i .& Q m 8 3^ 04 « m o 0/ ^ ^ ^ 0 a 8 ^ ^ d I ^ ^ ^ d' S' ? a © o a s ^ a - ® d J ^ CN -a d 4) 1 a ^ a S e^ § I- g « © d .|>5 s5 ca 15 w d d N^ i g . Q d 5- © f 05 ^ s (N d ’— »« c3 O a d ^ ^ w d .2 •| § S SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 65 § & < ^ - o o U o o m 'C a, U 'S o >r» 1 fO .£ cd 00 ■S c M o £ -ta L., 00 S si c -a (D « o QO a cd CN ij (D "S •bee s •= o cr E "-S (U « e bO •§ S o c O 0) „ o w ■o fN - M w " .S 0} ^ r- g ^ Crt c« ^ O I 3 O (U S'? s I =!: a o ^ S 2 M ^ I .-S I * * ^ XI ■g ’m ^ O s «• ^ S ^ o o ? S S S cx « ’« M bO i5-g M flj 3 ^ Cl, O Cl, S'! = t 4> ,5 ^ a ^ s I e 3 o I « S X) Cl 0) 00 M :s oC « o o — ' Cn! C ? g -o ^ ^ o _s _ § ^ g > 2 ,3 -§ 3 S' S TS o a o o IM ^ ^ IBs a ^ S 1 si M ^ ^ > « flj lot. « c ^ .-G 05 g 2 e .2 .2 2 2 « 2^ ,2 2 S i o 5 § OT M O o ■^1 s ^ .S « « - & & o OQ o k3 E 'a « Pm 2 ^ ^ M “ d 'O Dm ^ « .S' >. S P Q 2 2 i I H § B ^ 1 1 0) if •s^ ■S c m g I S '+- o U « .2 S o O 4) ^ o § 2 •c I IE 2 S ^ I « « ^ d te s o d -o « ^ cd o d .S 5 2 ^ d o 2 § « I M > u "S s 2 g s I =tfc I s 1 I u a, •£ 2 M d o >% • S -a ^ ifi o ^ c II I <^b0 « « M — I- ^ E -S « o S ^ ^ o, 'O • M 2* 3 « d p « •p d C« 2' 1 0) « « OJ ca s & 2' I « ^1 o 2 o, c « E r W ^ & o S 2 ca p- •c O, M < (50 M) S .S .J-. {a S Q BO d 1 ^ &)l > o« 6 2 C/3 p . O II B o o e O 2 3 3 I 2 6 ^ I 0) « « p m C 'rt 1 ^ 1 2 ^ A rd o O 05 aj 05 •13 « o S B i « o s « 3 ^ &, -o ^ ’O I-. g 4J « .S “ 2 .S .s o ^ ^ d »- y « 3 ^ Oi O O •— .p I g o ,Q flj d > ' .2 'm S 3 o g 3 -d S' g“ O y « *- »- (50 3 2 « ro _ — -' « O -O o ■a p ..5 « M § ’S m '« 2 cd tS 3 ^ - w 4) W .P mC .&s it I s ^ o 'S f s s ^ « -§ 3 w « s I > 3 s § « » &b M -P w (U GO 03 « e ^ 2 o 9 3 ^ o 3 ^ 3 »_ - o 2 &o _ S c 3 c ■p o 4J ^ > M O O CN eo c S c 2 9 ffl o ^ ^ o ® CN 3 § N J o ^ 5 5 CN 3 « '« 3 a ^ I § B ed td d 2 g 5 I 6 2 &, 4) GO l.£ o 3 o S so S « J3 V5 "P en g CnJ cd -O M ^ 05 3 ■g g 8 t « S-2 I I 2 S g S '"' BO ^ ^ ^ g 2 9 •a 2 ,p m g) a < 6(5 /~s •S M i § I .S b Q o ^ CNJ ^ SO ^ d I I w « ’S % 1 = « .S ® 3 o 3 ^ « ^ y cd .1 S ^ « It o g « 3 I « S I S ^ o •§ ‘3) 3 w 'S •5^ « V S fc I M_. 4) «d a> s 2 t§ ^ &o e 2 3 o CnJ « . 3 & ^ ,Q "P D 4) 4> !« & II 4) VO M m N ^ ¥ 3 .S 3 I I "S ^ S m li SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 67 « a, a < W o S C g § S M CO II O g >- S oo a|g 1st 111 I >,w *** ^ ^ ^ w ® ^ ^ fN u w sr» O O « .< o^ o o « a « •=* 'B M M o d ^ _w 3 .1 ill ^ j I l-S-s ill II t s “ s II o B B d M ” Ifll O g M a g ^ a M ^ O O 0 « 11 \o ^ aJ w 3 1 s li u (O « ea ® (U ^ a « i M a o m p < •C a m a < -S ^ o ^ O CM fN| ^ o CM &0 § 3 b 3 ' W -fsa, ^ ® o ;Q ® ”11 !la - e3 ja g 3 d Q ^ 0\ O ^ o^ ^ J 3 3 I « « « 3 S S a-o P & b I fam a. ^ CM •i s ^ CM 1 2 g « 2 3 .2 M s ffl o ^ S I |g 12 |l S d i b m O CM 68 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES I > ^ W f I J N « M ^ ^ 0) c3 p > I g I P»> c « S 'S ** 'S w ■5 g .S w s M o oj S s ^ -a g ^ "C ° 9 'P •S ^ S g “ U O g P > o g (O p d OJ §■ s« 2 'S 8. M -O C e « ‘S S « D, S OT M tg « «5 o S 'S -g U o p o -a 'O tg « CO W « 0 4) « b B .i 1 I "s •S ^ -S “B ^ ^ I ^ I y ^ 8 o CO 4) >r1 -i-i 2 3 2 B g di M 'O P « i| a .2 •S £ P o. 8 rd 3 Q 0 “ ^ -o cfl 4> 'S 1 W S 0) P- P' 1 M ii 3 c s P O "5 3 ^ (S CO ^ 3 P 4) .2 i J « 3 -s ©23 5 4i W2 M 3 B B ^ it o g a g •g 4) Ii ^ I g o , USB g g ^ 3 g. e 3 « 2 O W td o •c a o u § CO ‘S "O o « «u 4) r— CO >3 r. © ? .s ^ E G o »3 el 2 I I* u I ® • • d -a “88 § ^ ^ "’g ^ -d ^ dj dJ I S S goo o >. 63 1 « M "m ^ O £ U-t Sfl p* P § ^ I 3 - S O 0 ffl ^ U S s S Q O 1 ^ ^1 I ^ I I a. . tY « o S 2 m 3 4) _3 a^ 3 P ^1 I G J I I s a 5 3 S 3 p o^ r<^ a\ M rsi d ^ o o > CNI ^ ^ S .a M e3 m 4i l| tl 'O 3 3 4) bh 3 o I tM 0 4-1 0 ii « 0 d 0 3 1 1 » CO 43 X 3. 3 S-I &, 43 3 8 u M P 9 M „ e 3^ B 3 43 p 1 ^ > 0 0 CO CO « &0 8 3 8 B 3 © g WQ 00 i 2 B 3 o O. ^ 3^ 3' 3 3 3 ^ CO Cq ^ s S s §■ |S^ 111 ^ ai) ei '3 CO o ^ rj CO ,0 B ^ ^ ■SOM S 6 ^ 3 © 3 S) m o 3 ^ ^ 3 E 3 U § o .& 0- 0 2 ca g C3 »rt W O cfl . « -O c« 1 SL III s Cl, O 00 & w w ^ p •a § S <§ ^ -a II t e g, E ■d 8 8 ^ z m U I S3 ^ n4 o 8 = § I S I sS.™ = "« c ® OPS 1^0 ^ w o o ^ fca ^ 3 o -© o N -3 & « S Longevity Scientific name Start End Fmit/seed characteristics Seed collection details Seed germination (yrs.) Saikomia pacifica Oct Nov Mature seeds pinkish white, Collect inflorescences when plant Variety S. utahensis grows best at 5% NaCl 2+ (Khan and Weber 1986, puberaleiit, and 0.5-1 mm tips are purple. Dry seeds on a treatment and under temperature regime of Young 200 !j) long. Sequentially screen for up to 3 months. 15-5°C. SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 69 is s On O ri c) s ° 2 VI 1 ^ ^ a ^ g| =• ® i §• 3 s a .2 ^ .S .2 ^ Q 0 1 I si W s ri -2 ^ -s o ^ M g I B ^ I s «D eS :g E « § o .2 e o b o S O & 'xi -I « s g o ..Q w ^5 'S s « 00 d ^ o o ^ . JL o d -.d d g (3 td ^ -P g ^ fa ^ S g ^ O S Q O ^ M cd ‘S § d 09 3 ® S ^ i s o ^ oo « N 'a cd . o d tS a § ° B M s xi «G ■■s « ! I 1 « ^ M ‘5b ^ « ^ « c/j ^ O' ^ M 4) ^ w •a s " f w © o „ u s p I & p C ed e 1 ^ P 0 3 'B f) fi ^ s & a. g & ^ a. .2 0 3 2 •S 1 > 1 1 e I ^ I i B 8 .s ^ ^ M -o o m I 'S o ..d W •d* ‘ri W fri ■I I ed o 'w ^ 8 fN •■fe 09 ^ ^ 2 a « a B S P ^ S B P' I S (3 S 'P “Is ^ M Q m p < &o P < a .fe« .2 i S o ^ g CM * « d I U I 1 -I fc - ^ o ^ CN ' S d aj Q S W W -ftf = . S p fn ^ d oo •a o p? .5 m o C CM O “I S &3 ^ O S W I 2 .s « Q_ d .« « .s § s ^ Cn! O CN 'S B 0) & 2 o d ^ 2 ^ -d ffl -- 'di -a 2 B ^ cd ed ® 2 > I ® to ^ eg TO TO U m z Q ’^koenoplectus acutm var, Aug Sep Wide, smooth fruits with 2 or 3 Because they are easily dispersed Physiological dormancy. Cold stratification occMemtalis (Lacroix and distinct sides. Fruit 2-3 mm by wind, it is important to breaks dormancy. Germination rates are low Mosher 1995, Johnson long and 1.2- 1.7 mm wide. collect seeds close to the time for the species due to the thick pericarp of 2004, Baldwin et al. 2012, of maturity. Seeds must be the achene. Germination rates increase with Baskin and Baskin 2014) overwintering in a pond or water source. Longevity Scientific name Start End Fmit/seed characteristics Seed collection details Seed germination (yrs-) separated from the panicle and Pre-treat seed with 0.05% solution of cleaned. sodium hypochlorite 5 days prior to 70 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES IS O 0} g § I 5 0) — < O O O 5 S 'a .2 « & o 2 'O & -s o, 4 o ^ & m •S d 1 VO o o 1 ‘B < CM d 2 CN 2 ffl 2 VI m a^ I § .SI §■ I § i I 8§ 5^ m O O s i 8 ^ « o ^ § rM « ^ > .S ed -S 4) ed ca ^ O « 2 s E "B cA i 5 S S' S o ® _ K e « s s 1, «& 5 of ^ m VI, p o g > « ^ -S .s-§ & m O ^ U U s d Q ffl c . m o, s 'B “g c« eS >« ^ O 'o I 'B I B o U I ^ s ^ 'O "S § s i-i o ^ __ O- II & m 'a I 8 05 s Longevity Scientific name Start End Frait/seed characteristics Seed collection details Seed germination (yrs.) dotted, warty, net-like, or Pass seed through a hammer prickly. mill or a sieve prior. Seeds should be spread out to dry SEED COLLECTION AND GERMINATION IN S. CALIFORNIA WETLANDS 71 cd I .9 . g § 'T3 s 1 1 02 8 4= S 8 a d M 8 S ■« g ^ o m g § s --g « 0) B s g S .S o B 2 S o ■S &o ;.s ^ d S § 3 o 1 1 -o 8 ^ C! W .S o ed -§ s ^ m . d -a la d ! o « to m •fe to i e ^ S § to ^ to U ^ ^ to 9 cri to to g I 1 1 1 1 S P S S to ’3 tri S S c« cu to 0*3 I ^ s .9 ^ I 'S o I « 8 I S S O CS| S ^ s ? i 7 « II “ ^ (E li » =■ o CNj *>© to oo ^ o\ .S m -a q ^ Sts M to S to ^ to 3 c « 2 to to 3 to ■2 I S B to S ^ 3 I 1 1 § S a ^ .2 § p .2 S d s .2 to •S o 3 S -a S d (S P to 'O to ^ o m d d d 'O ■ to d to 3 m Bull. Southern California Acad. Sci. 115(1), 2016, pp. 72-80 © Southern California Academy of Sciences, 2016 Redescription of Bathygyge grandis Hansen^ 1897 (Crustacea5 Isopoda, Bopyridae) from Southern California with Erection of a New Subfamilyj Bathygyginae John C. Markham Arch Cape Marine Laboratory, Arch Cape, Oregon 97102-0133, jmarkham@seasurf.net Hansen (1897), dealing with only fragmentary material, erected the genus Bathygyge with B. grandis as its type-species as one of the earliest bopyrid species known from the eastern Pacific Ocean. Bathygyge grandis was first recorded as a parasite of the deep-water crangonid shrimp Glyphocrangon spinulosa Faxon from off the coast of Acapulco, Mexico. It has since been reported from several different localities worldwide as a parasite of other species of Glyphocrangon, but it has never been properly described. Material that recently became avail- able from near the type-locality has made it possible to correct that situation. Order Isopoda Latreille, 1817 Suborder Cymothoida Wagele, 1989 Family Bopyridae Rafmesque-Schmaltz, 1815 Subfamily Bathygyginae, subf n. Genus Bathygyge Hansen, 1897 Type-species, by monotypy, Bathygyge grandis Hansen, 1897 Bathygyge grandis Hansen, 1 897 Figs. 1-2 Bopyrus - Faxon, 1895: 140 [Type-material later described]. Bathygyge grandis Hansen, 1897: 122-124; pi. V, figs. 2-2c [Pacific Ocean, off Acapulco, Mex- ico, 21°15'N, 106°23'W, 676 fm {= 1236m}; infesting Glyphocrangon spinulosa Faxon, 1893]. — Richardson, 1899a: 869. — Richardson, 1899b: 338.-Bonnier, 1900: 48, 221, 291- 292, 381; fig. 53. — Richard, 1900: 71. — Townsend, 1901: 527.-Richardson, 1905: 537-539; fig. 581.— Stebbing, 1908: 57-59; pi. XXXIII [Off Cape Point, South Africa, 800-900 fm {- 1463- 1646m}; infesting Glyphocrangon sculpta (S. I. Smith, 1882)].— Stebbing, 1910: 436. — Nierstrasz and Brender a Brandis, 1923: 86. — Barnard, 1940: 494, 721. — Dan- forth, 1963: 33, 37, 91, 92; pi. 5, figs. 1, 2.=§adoglu, 1969: 197.— Schultz, 1969: 312; fig. 496.— Danforth, 1970: 9, 43, 57-58, 149; fig. 5D, E.— Holthuis, 1971: 285.-Wenner, 1978: 1058-1061 [On continental slope of Middle Atlantic Bight; infesting G. sculpta and G. longirostris (S. I. Smith, 1882)]. — Bourdon, 1979: 510.-Markham, 1979: 771-772. — Markham, 1985: 19, 131 [Atlantic Ocean, off coast of Virginia, USA: infesting G. longiros- tris].— Markham, 1986: 155, 156; fig. 4B.— Kaufmann et al., 1989: 1882; tab. 4 [Magellan Rise, NE Pacific, 07°05'N, 176°55'W- 176°50'W, 3100m; infesting unspecified host, prob- ably G. vicaria Faxon] .-Salazar- Vallejo and Leija-Tristan, 1989: 429. — Leija-Tristan and Salazar- Vallejo, 1991: 1. — Markham, 1992: 3; tab. 1.^ — Espinosa-Perez and Hendrickx, 2001: 50. — Roman-Contreras and Soto, 2002: 279. — An, 2006: Abstract [on unnumbered p.], 73-74, 114, 117, 123, 131;fig.28 [East China Sea, 26°10'N, 126°00'E; miQsimg Glypho- crangon sp.]. — An et al., 2007: 1002, 1003; fig. 1 [Same material as An, 2006].— Yu and An, 2008: 691.-Stebbins, 2012a: 2.— Stebbins, 2012b: 2, 6, 16; 4 unnumbered figs. 72 REDESCRIPTION OF BATHYGYGE GRANDIS HANSEN (ISOPODA, BOPYRIDAE) 73 IGigantione bouvieri. — Bourdon, 1967: 857 [Canary Islands; infesting Glyphocrangon sp., hyperparasitized by Cabirops serratus Bourdon, 1967. Probably not Gigantione bouvieri Bonnier, 1900]. IBathygyge sp. — Bourdon, 1967: 857 [Same material tentatively called Gigantione bouvieri above].- — Bourdon, 1979: 510 [Azores, 1590-1665m; infesting Glyphocrangon longiros- tris]. — ^Lemos de Castro, 1970: 2.. — Holthuis, 1971: 339. — Restivo, 1971: 71; tab. 1. — Res- tivo, 1975: 153; tab. 3.— Bourdon et al., 1981: 498. — Rybakov, 1990: 415. — Roman- Contreras, 2008: 91. ?”bopyrid parasites.” — Holthuis, 1971: 339 [Off Atlantic coast of Nigeria, 04°15’N, 04°27'E -04°12'N, 04°28'E, 1280- 1320m; infesting Glyphocrangon longirostris']. Munidion sp. — Wicksten, 1979: 222 [San Clemente Basin, California, infesting Glyphocrangon vicaria Faxon, 1896: material examined herein, described below]. — Wicksten, 2009: 168. ?”branchial bopyrid.” — Chace, 1984: 11 [West of Halmahera, Indonesia, 00°16'30"N, 127°30'00"E, 497m; infesting Glyphocrangon faxoni de Man, 1918]. Bathygege [sic] grandis - Campos and Campos, 1989: 33; tab. 2. ?”Bopyrid isopod”-— -Moore et al, 2003: 368 [Bear Seamount, northwestern Atlantic, 39°55'N, 67°30'W, 1100 m; infesting ''Glyphocrangon'' {probably = G. sculpta}]. ?”bopyrid isopod.” — Ahyoeg, 2006: 68 [Tasman Sea, 32°04'S, 159°53'E, 1920- 1934m; infesting Glyphocrangon dimorpha Komai, 2004]. — Han and Li, 2007: 550 [East China Sea, 09°29'N, 123°41'E, 2000-2150m; infesting Glyphocrangon megalophthalma de Man, 1918] Material Examined Infesting Glyphocrangon vicaria Faxon, 1896. WV Agassiz Station M-7 Sta. 3, San Clemente Basin, eastern Pacific off California, USA, 32°28'N, 118°08'W, 1792m, 16 September 1971, 40-foot otter trawl. 29, 2(J, SIO (Scripps Institute of Oceanography) C3100. Redescription of Female Length 12.7 mm, maximal width 9.3 mm, head length L3mm, head width 2.8 mm, pleonal length 3.9 mm. Distortion 115° sinistrally. Body outline broadly ovate, widest across pereomere 5. All body regions distinct, pereomeres distinct but pleomeres medially fused; pleon strongly torsioned (Fig lA, B). Head deeply embedded in pereon, its anterior margin overreached by second oostegites. No eyes. Antennae (Fig. 1C) not extending beyond margins of head, first of 3 articles, second of 6 articles, setation obscure. Barbula (Fig. ID) with pair of unomamented slender falcate projec- tions on each end, slightly sinuous margin medially. Maxilliped (Fig. IE) of irregularly penta- gonal anterior article bearing subterminal articulating triangular palp (Fig. IF) densely setose along medial edge; and smaller subtriangular posterior article produced into long slender plec- tron (Fig. IG) directed anteromedially. Pereomeres separated dorsally by sinuate margins. Coxal plates well-developed on pereo- meres 1-4, those of first two pairs reflexed medially over dorsal surfaces of pereomeres (Fig. lA, J), other two pairs completely covering lateral margins of pereomeres. Pereomeres 5-7 lacking coxal plates but their lateral regions expanded and flat. Oostegite l(Fig. IH, I) slightly longer than broad, with nearly parallel sides; internal ridge lacking ornamentation; pos- terolateral projection about 1/3 width of posterior margin of oostegite, rather short and broadly rounded, turned slightly medially. Oostegites 2-5 of both pairs large and completely enclosing vaulted brood pouch. Pereopods (Fig. IK, M) all tiny, though slightly larger posteriorly, arrayed along lateral margins of pereon and extending little beyond those margins, pereopods 1 and 2 74 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. I. Bathygyge grandis Hansen, 1897, reference female. A. Dorsal view. B. Ventral view. C. Right antennae. D. Barbula, right side. E. Right maxilliped, external view. F. Palp of same. G. Plectron of same. H. Oostegite 1, external view. I. Same, internal view. J. Right pereopod 1 and attached coxal plate. K. Left pereopod 1. L. Dactylus and end of carpus of same. M. Left pereopod 7. N. Dactylus and end of carpus of same. Scale; 4.00 mm for A, B, D, E, H, I, J; 1.43 mm for K. M; 1.00 mm for C. F. G; 0.29 mm for L, N. clustered closely together, and pereopods 6 and 7 similarly clustered on both sides; all articles of all pereopods separate, dactyli reduced and blunt anteriorly (Fig. IL), bases (Fig. IM) longer and dactyli (Fig. IN) longer and sharper posteriorly. Pleon (Fig. 1 A) strongly torsioned and reflexed over pereon, so right (longer) side facing for- ward, of 6 pleomeres, final four pleomeres incompletely separated. Pleomeres 1-3 produced into broad blunt lateral plates on longer side. Pleomeres 4 and 5 produced into slender lateral plates on both sides. Pleomere 6 bearing widely separated uniramous uropods of structure and shape similar to that of lateral plates. No pleopods on any pleomeres. Other female quite similar. Distortion dextral, with pleon torsioned to left side. Length 10.5mm, maximal width 8.2mm. Reflexed coxal plates on left (shorter) side of pereomeres 1- 3, on opposite sides of pereomeres 1-2. Pereopods 1-4 with long dactyli, those of pereopods 5-7 smaller. Redescription of Male Length 6.8 mm, maximal width 1.6 mm, head length 0.6 mm, head width 1.1 mm, pleon length 1.8 mm, pleon width 1.2 mm. Head, pereomeres and pleon distinct. Sides of pereon par- allel from pereomere 2-6. No pigmentation (Fig. 2A). REDESCRIPTION OF BATHYGYGE GRANDIS HANSEN (ISOPOD A, BOPYRIDAE) 75 Head roundly quadrate, abruptly narrower than first pereomere and extending forward from it; anterior and posterior margins straight across, former somewhat shorter. No eyes. Antennae (Fig. 2B) of 3 and 7 articles respectively, minutely setose distally. Pereon narrowest across pereomeres 1 and 7, its sides nearly parallel between; all pereomeres separated by deep notches laterally, slightly ridged middorsally. No midventral tubercles. Pereo- pods (Fig. 2C, D) with all articles distinct, larger anteriorly; carpi sparsely setose on anterome- dial comers; all propodi enlarged, each produced into proximal lobe with socket receiving tip of reflexed dactylus; long sharply pointed dactyli on pereopods 1-5, shorter and blunter dactyli on pereopods 6-7. Pleon long and extended, markedly narrower than last pereomere, as tmncated oval, its sides nearly parallel, posterior margin broadly rounded. No trace of segmentation. No appendages. Other male very similar, its antenna 2 of 8 articles. Length 12.7mm, maximal width 3.8mm, head length 0.7mm, pleon length 1.5mm. Bathygyginae, new subfamily Diagnosis. Branchially-infesting bopyrid. Female: Body outline roughly circular, with no straight margins. Head oval, much broader than long, deeply embedded in first pereomere and overreached by second oostegites. Maxilliped with subterminal triangular palp and slender pointed plectron. Barbula bearing two long slender projections laterally, bare medially. All per- eomeres distinct dorsally, their margins irregularly curved. Pereopods all present, reduced. Coxal plates of pereomeres 1 , 2 and/or 3 extending medially over dorsal surface of body. Oos- tegites 2-5 on both sides well-developed and completely enclosing enlarged brood pouch. Pleon of 6 incompletely separated pleomeres, its central axis greatly rotated to one side, posterior-most point of body side of pleomeres 1 and 2, pleomeres bearing blunt lateral plates but lacking all pleopods and uropods. Male: Body about 4 times as long as broad, fusiform in outline. Head narrower than first pereomere and distinct from it. Pereomeres deeply divided. Pereopods all with enlarged propodi with sockets receiving tips of sharply pointed dactyli. Pleon separated from last pereomere, suboval, bulbous, completely lacking all traces of segmentation or appen- dages. Only one genus and species known, Bathygyge grandis Hansen, 1897. With a complete description of Bathygyge grandis, it now becomes possible to assess its sys- tematic placement. Characters unique to B. grandis, which indicate that it should be in a sub- family of its own are: Female: Second oostegites extending beyond anterior margin of head; coxal plates of first and second pereomeres extending medially over dorsal surface of pereon; pleon strongly torsioned and pointing forward over pereon. Male: All pereopodal propodi bear- ing sockets into which tips of sharply pointed dactyli retract. Unusual characters (though in rare instances known from members of other subfamilies of the Bopyridae) are: Female: Body broadly oval, almost circular; head lacking frontal lamina; pereopods proportionately tiny. Male: Body very long relative to width; pleon completely lacking all traces of segmentation and all appendages. Hosts: All known in genus Glyphocrangon A. Milne-Edwards, 1881 (Caridea, Crangonoidea, Glyphocrangonidae). Hansen (1897), whose description of Bathygyge grandis Richardson (1905) quoted verbatim and whose illustrations she reproduced, had “[ojnly a male, and the posterior part of a female.” He did not figure the female at all but remarked on it thus: “Abdomen: It is turned to the left in a startling degree...” That extreme rotation of the pleon, figured and described herein, is unique for bopyrid females. The present male closely matches the description and figures of the type-male by Hansen (1897) (reproduced by Richardson, 1905). The only illustration prepared of Bathygyge grandis since the original description is that of An (2006), of Chinese material in a dissertation, which has limited accessibility because it is entirely in Chinese and unpublished; 76 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 2. Bathygyge grandis Hansen, 1897, reference male. A. Dorsal view. B. Left antennae. C. Left pereopod 1. D. Right pereopod 7. Scale: 2.00 for A; 0.45 mm for B-D. REDESCRIPTION OF BATHYGYGE GRANDIS HANSEN (ISOPODA, BOPYRIDAE) 77 it only recently came to my attention. In her figure 28, she presents detailed drawings, the first known, of both sexes. Both the female and male illustrate the diagnostic characters of the spe- cies: the medially extending coxal plates of the first two pereomeres, the reduced pereopods and the strongly torsioned pleon of the female; and, in the male, the strongly separated head, pereo- meres and pleon; prominent proximal lobes on the propodi of the pereopods of the first pair; and complete lack of segmentation and appendages of its pleon. There are however, several differ- ences between the material herein described and that from China. In the Chinese female, the body is more nearly circular; the maxilliped's palp does not articulate, and its spur is much reduced; the first oostegite has a slightly more slender and longer posterolateral point, which is setose along its lateral margin; the posterior pereomeres bear tergal plates distinctly separated from the segments; and the lateral plates of the pleomeres are quite shaiply pointed. In the Chinese male, the body is slightly curved; tiny colorless eyes are present next to the posterior edge of the head; and the pleon is attached to the last pereomere by a nairow peduncle. It is uncertain whether the propodi of that male's pereopods bear sockets receiving the tips of the dactyli, as in the male herein described. An et al. (2007), who published mention of the same material as An (2006), included a photograph (their fig. 1) of the parasite in place on its host but did not present drawings of the specimens themselves. Few specimens assigned to Bathygyge grandis have been examined in detail (only the male in the type-collection, only two females and two males in the present collection and a single one of each sex in the report from China). Thus there remains some doubt whether the distinctions cited here are consistent among the populations from opposite sides of the Pacific Ocean. If they are, the individuals from China should quite probably be considered representatives of a separate undescribed species. The status of specimens reported from, elsewhere in the world is also uncertain. For now, however, I am retaining all of them in the synonymy of B. grandis presented above. Branchial bopyrid parasites of caridean shrimps are most commonly members of the subfam- ily Bopyrinae, wfrose species are not known to infest any hosts but carideaes. The small sub- family Argeiinae contains exclusively can dean-infesting parasites. In the large subfamily Pseudioninae, whose many species are typically parasites of anomurans, are a few species found as parasites of deep-water carideans. Bathygyge clearly does not belong in the Argeiinae, whose females, among other contrasting characters, have large rear-extending pleons of a very different shape. It has been dubiously assigned to the Bopyrinae (Shiino, 1965) or Pseudioninae (Markham, 1974) but does not fit well into either of those subfamilies for various reasons. Stebbiris (2012a, 2012b) expressed doubt about its proper assignment to subfamily. Its unique placement, emphasized by the present erection of a new subfamily, may be a reflection of its occurrence as the only known bopyrid species infesting any member of the family Glyphocran- goiiidae, of which Giyphocrangon is the sole recognized genus. Wicksten (1979) mentioned infestation of Giyphocrangon vicaria and called its parasite '"Munidion sp.” without any descriptive notes and later (Wicksten, 2009) repeated that record. I found her label in the container with the material herein redescribed, thereby confirming that I was dealing with the same specimens. Acknowledgments Joel W. Martin and Adam R. Wall of the Los Angeles County Museum of Natural History allowed me to borrow and study the material. Greg Rouse of the Scripps Institution of Oceano- graphy, University of California San Diego, provided information about its collection. Susan R. Gilmont of the Hatfield Marine Science Center, Oregon State University, obtained a needed 78 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES reference for me. Jianmei An of the School of Life Science, Shanxi Normal University, Linfen, China, provided a copy of her dissertation. Anonymous reviewers provided helpful suggestions. Literature Cited Ahyong, S.T. 2006. Crustacea: Galatheidae, Polychelidae and Glyphocrangonidae (squatlobsters, blind, deep-sea lobsters and deepwater shrimps). Results, Taxonomy. Explore lost worlds of the deep NORFANZ voyage. Final Report to the Department of Environments and Heritage (National Oceans Office). Chapter 3.1:51-83. An, J.M. 2006. Study on the taxonomy and zoogeography of the Family Bopyridae (Crustacea: Isopoda) in the China seas. Ph.D. Dissertation, Institute of Oceanology of Chinese Academy of Sciences, vii + 225 pp. [In Chinese] An, J.M., H.Y. Yu and X.Z. Li. 2007. One new record genus of Bopyridae (Crustacea, Isopoda, Epicaridea) from China. Acta Zootaxonomica Sinica, 32(4): 1002-1003. [In Chinese with English summary] Barnard, K.H. 1940. Contributions to the crustacean fauna of South Africa. 12. Further additions to the Tanaida- cea, Isopoda and Amphipoda, together with keys for the identification of hitherto recorded marine and fresh-water species. Ann. Sth Afr. Mus., 32:381-543. Bonnier, J. 1900. 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The caridean shrimps (Crustacea: Decapoda) of X\\q Albatross Philippine Expedition, 1907- 1910, part 2: Families Glyphocrangonidae and Crangonidae. Smithsonian Contr. Zool. 397: i-iv, 1-63. Danforth, C.G. 1963. Bopyridian (Crustacea, Isopoda) parasites found in the Eastern Pacific of the United States. Ph. D. Thesis, Department of Zoology, Oregon State University. 110 pp. . 1970. Epicaridea (Crustacea: Isopoda) of North America. Ann Arbor, Michigan, ii + 191 pp. Espinosa-Perez, M.D.C., and M.E. Hendrickx. 2001. Checklist of isopods (Crustacea: Peracarida: Isopoda) from the eastern tropical Pacific. Belgian J. Zool., 131(l):43-55. Faxon, W. 1895. Reports on an exploration off the west coasts of Mexico, Central and South America, and the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer Albatross dur- ing 1891, Lieut. Commander Z.L. Tanner, U.S.N., commanding. XV. The stalk-eyed Crustacea. Mem. Mus. Comp. Zool., Harvard College, 18:1-292. 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Pp. 81-114 in Crustaceos de Mexico: estado actual de su conocimiento (Alvarez Noguera, F., & G. A. Rodriguez Almaraz, eds.) Universidad Autonoma de Nuevo Leon. . and L.A. Soto. 2002. A new deep water genus and species of branchial bopyrid infesting the galatheid crab Munidopsis erinaceus from the southwestern Gulf of Mexico. J. Crustacean Biol., 22(2):279-286. Rybakov, A.V. 1990. Bourdonia tridentata gen. n., sp. n. (Isopoda, Cabiropsidae) a hyperparasite of Bopyroides hippolytes Kroyer from the shrimp Pandalus borealis. Parazitologiya 24 (5): 408-416. [PbiSaKOB, A.B., 1990. Bourdonia tridentata gen. n., sp. n. (Isopoda, Cabiropsidae) rnnepnapasHT Bopyroides hippolytes Kroyer h3 KpeTKH Pandalus borealis. HapasHxojiorHa 24(5):408-416.] §adoglu, P. 1969. Variations in eye degeneration and pigment in some parasitic Isopods during their life cycle. Pubb. Staz. Zool. Napoli, 37:173-209. Salazar- Vallejo, S.I., and A. Leija-Tristan. 1989. Progebiophilus bruscai n. sp., a new bopyrid isopod parasitic on the mud shrimp, Upogebia dawsoni Williams (Thalassinidea), from the Gulf of California. Cah. Biol. Mar., 30(4):423-432. Schultz, G.A. 1969. How to Know the Marine Isopod Crustaceans. Wm. C. Brown Company, vii + 359 pp. Shiino, S.M., 1965. Phylogeny of the genera within the family Bopyridae. Bulletin du Museum National d’Histoire Naturelle de Paris, (2)37(3):462-465. Stebbing, T.R.R. 1908. South African Crustacea (Part IV). Ann. So. Afr. Mus., 6:1-96. . 1910. General Catalogue of South African Crustacea, (part 5 of S. A. Crustacea, for the Marine Investi- gations in South Africa). Ann. So. Afr. Mus. 6(6):28 1-599. Stebbins, T.D. 2012a. California Bopyridae (Crustacea, Isopoda, Cymothoida, Bopyroidea). http://www.scamit. org/taxontools/toolbox/Phylum%20Arthropoda/Class%20Malacostraca/Order%20Isopoda/Suborder%20 Cymothoida/Family%20Bopyridae/Cal%20Bop5nids_Stebbins2012_Revl201 1 8.pdf pp. 1-6. . 2012b. California “Epicaridean” Isopods Superfamilies Bopyroidea and Cryptoniscoidea (Crustacea, Isopoda, Cymothoida). Presented to SCAMIT 13 February 2012. http://scamit.org/taxontools/toolbox/ Phylum%20Arthropoda/Order%20Isopoda/Suborder%20C5miothoida/Califomia%20Epicaiideans_Stebbms 2012_120118.pdf 35 pp. Townsend, C.H. 1901. Dredging and other records of the United States Fish Commission Steamer Albatross with bibliography relative to the work of the vessel. U. S. Fish Comm. Rept., 1900:387-562. 80 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Wenner, E.L. 1978. Comparative biology of four species of glyphocrangonid and crangonid shrimp from the con- tinental slope of the Middle Atlantic Bight. Canadian J. Zool, 56:1052-1065. Wicksten, M.K. 1979. New records of the species of Glyphocrangon in the northeastern Pacific Ocean (Caridea: Glyphocrangonidae). Proc. Biol. Soc. Washington, 92:217-224. . 2009. Decapod Crustacea of the Californian and Oregonian zoogeographic provinces. U. C. San Diego: Scripps Institution of Oceanography Library, http://escholarship.org/uc/item/7sk9t2dz, 418 pp. Yu, H., and J. An. 2008. Order Isopoda Latreille, 1817. Checklist of Marine Biota of China Seas. Pp. 690-699 in [Liu, R. (J. Y. Liu), ed.] Beijing: Institute of Oceanology, Chinese Academy of Sciences. Bull. Southern California Acad. Sci. 115(1), 2016, pp. 81-83 © Southern California Academy of Sciences, 2016 The Reef Gornetfishj Fistularia commersonii Riippell, ISSSj New to the California Marine Fish Fauna Milton S. Love Marine Science Institute, University of California, Santa Barbara, CA 93106, love@lifescL ucsb. edu I report here on several sightings in southern California of the reef cometfish, Fistularia commersonii Ruppell, 1838. These records mark the first time this species has been reported firom California marine waters. Mr. Bill Powers reported an unverified sighting on 7 November 2015 at Little Flower Reef, San Clemente Island (32°50.399*N, li8°22.136’W). Mr. Powers was diving over a sand-shell hash slope adjacent to a vertical rock wall in about 12 m of water. The approximately one- meter-long fish was more or less motionless and Mr. Powers was able to obseive it for several minutes, eventually approaching and touching it. Mr. Powers reports that the fish was green in color with blue spots and stripes and had a long filament extending from its caudal fm. This fish was not photographed. A second sighting was made by Ms. Sandy Dildine on 12 November 2015 within Crescent Bay, Laguna Beach (33°32.7'N, 117°48.3’W) in 7 m of water with surface water temperature at this site of about 20°C. This approximately one-meter-Iong fish was slowly swimming just above the bottom. Photographs and videos of this fish taken on 12 and 13 November 2015 show a green fish with blue spots and stripes. Ms. Dildine noted that when the fish was more or less motionless it had a series of dark bars along its body as well as bright, light blue spots and stripes (Fig. 1). When actively swimming, the bars of this individual quickly disappeared and the stripes and spots became darker (Fig. 2). Ms. Dildine also observed what was likely the same individual within the same circumscribed area (of about 10 m) on 19, 20, and 22 November 2015. In these instances, the fish was associated with an aggregation of blacksmith, Chromis punctipinnis (Cooper, 1 863). Three cometfish species live in the Pacific Ocean: Fistularia commersonii Riippell, 1838, reef cometfish; Fistularia corneta Gilbert & Starks, 1 904, deepwater cometfish; and Fistularia petimba Lacepede, 1803, red cometfish. All range widely in the IndO“Pacifi.c {E petimba is also found in the Atlantic Ocean). In the eastern Pacific, Fistularia corneta occurs as far north as Huntington Beach, southern California (Curtis and Herbinson 2001) and southwards to Callao, Peru (Chirichigno and Velez 1998), including the Gulf of California (Fischer et al. 1995) and Islas Galapagos (Grove and Lavenberg 1997). Fistularia petimba is absent from the eastern Pacific (Nakabo 2002). Of these taxa, F. commersonii is the only species that is green with blue spots and stripes (Tliomson et al. 2000, Robertson and Allen 2015) as F. corneta is orange or brown with pink dorsal, anal, and caudal fins (Robertson and Allen 2015) and F. petimba is red to orange-brown (Kells and Carpenter 2011). Thus, based on color and pattern, it is highly likely that the San Clemente Island and Laguna Beach individuals are F. commersonii. These California sightings extend the eastern Pacific range to southern California. The pre- vious northernmost range was Bahia Magdalena (Thomson et al 2000) to Iquique, northern Chile (Sielfeld et al. 2010), including the Gulf of California (Fischer et al. 1995) and Islas Gala™ pagos (Grove and Lavenberg 1997). They have been reported from surface waters to depths of 132 m (Mundy 2005). This species reaches a maximum length of 1.6 m (Fischer et al 1995). 81 82 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 1 . Dorsal view of a stationary reef cometfish, Fistularia commersonii, showing dark bars and lighter spots and stripes. Photographed on 13 November 2015 off Laguna Beach, southern California, by Ms. Sandy Dildine. These fish were observed during a very strong El Nino and were part of a wave of tropical reef fish species (including cardinalfishes and damselfishes) that were observed in southern California during 2015. It is interesting to note that the southern-most capture of F. commerso- nii, in northern Chile, also took place during an El Nino (Sielfeld et al. 2010) and that the Fig. 2. Dorsal view of a swimming reef cometfish lacking bars and having darker spots and stripes. Photographed on 13 November 2015 off Laguna Beach, southern California, by Ms. Sandy Dildine. REEF CORNETFISH, NEW TO CALIFORNIA 83 Huntington Beach records of E corneta, based on the capture of two small fishes, occurred dur- ing the waning months of the 1997-1998 El Nino (Curtis and Herbinson 2001). Notes Added in Proof. On 6 December 2015, Mr. Mike Couffer observed and photographed a small (4 cm TL) F. commersonii, swimming near the sea floor just south of the Newport Pier, southern California (33°36'N, 117°56'W). Similar to the San Clemente Island and the Laguna Beach individuals, this individual was identified based on its green color and blue spotting on the back. This specimen represents the northernmost record for this species. A paper by Jackson et al. (2015) postulates, based on F. commersoniVs rapid recent coloniza- tion of the entire Mediterranean Sea via the Suez Canal, that this species may be particularly well adapted to expand its geographic range when opportunities arise. Acknowledgments I thank Mr. Bill Powers and Ms. Sandy Dildine for bringing their observations to my attention and Mr. Spencer Salmon for providing the geographic coordinates of the Santa Clemente Island sighting. Literature Cited Chirichigno, F.N. and J. Velez D. 1998. Clave para identificaticar los peces marinos del Peru (segunda edicion, revisada y actualizada). Instituto de Mar de Peru. Publicacion Especial. Curtis, M.D. and K.T. Herbinson. 2001 . First record of the Pacific cometfish, Fistularia corneta Gilbert and Starks 1904, a new species to the Southern California fauna during the 1997-1998 El Nino. Bull. S. Calif Acad. Sci., 100:156-159. Fischer, W., F. Kmpp, W. Schneider, C. Sommer, K.E. Carpenter, and V.H. Niem. 1995. Guia FAO para la identificacion para los fines de la pesca. Pacifico centro-oriental. Vol. II, Vertebrados, Parte 1 . Vol. Ill, Vertebrados, Parte 2. FAO, Rome. Grove, J.S. and R.J. Lavenberg. 1997. The Fishes of the Galapagos Islands. Stanford University Press, Stanford, California. Jackson, A.M., K. Tenggardjaja, G. Perez, E. Azzurro, D. Golani, and G. Bemardi. 2015. Phylogeography of the bluespotted cometfish, Fistularia commersonii'. a predictor of bio invasion success? Mar. Ecol., 36:887-896. Kells, V. and K. Carpenter. A Field Guide to the Coastal Fishes from Maine to Texas. Johns Hopkins University Press, Baltimore, MD. Mundy, B.C. 2005. Checklist of the Fishes of the Hawaiian Archipelago. Bishop Museum Press, Honolulu, Hawaii. Nakabo, T. 2002. Fishes of Japan. Tokai University Press, Tokyo, Japan. Robertson, D.R. and G.R. Allen. 2015. Shorefishes of the Tropical Eastern Pacific: an Information System. Ver- sion 2.0. Smithsonian Tropical Research Institute, Balboa, Panama, http://biogeodb.stri.si.edu/sftep/en/ pages. Accessed 30 November 2015. Sielfeld, W., J. Laudien, M. Vargas, and M. Villegas. 2010. El Nino induced changes of the coastal fish fauna off northern Chile and implications for ichthyogeography. Rev. Biol. Mar. Oceanog., 45, S 1:705-722. Thomson, D.A., L.T. Findley, and A.N. Kerstitch. 2000. Reef Fishes of the Sea of Cortez. University of Texas Press, Austin. CONTENTS SMITHSONIAN LIBRARIES 3 9088 01866 6198 The Return of the King of the Kelp Forest: Distribution, Abundance, and Biomass of Giant Sea Bass (Stereolepis gigas) off Santa Catalina Island, California, 2014-2015. Parker H. House, Brian L.F. Clark, and Larry G. Allen.. 1 Nudibranch Range Shifts Associated with the 2014 Warm Anomaly in the Northeast Pacific. Jeffrey H. R. Goddard, Nancy Treneman, William E. Pence, Douglas E. Mason, Phillip M. Dobry, Brenna Green, and Craig Hoover 15 Seed Collection and Germination Strategies for Common Wetland and Coastal Sage Scrub Species in Southern California. Michelle L. Barton, Ivan D. Medel, Karina K. Johnston, and Christine R.Whitcraft 41 Redescription of Bathygyge grandis Hansen, 1 897 (Crustacea, Isopoda, Bopyridae) from Southern California with Erection of a New Subfamily, Bathygyginae. John C. Markham 72 The Reef Cornetfish, Fistularia commersonii Ruppell, 1838, New to the California Marine Fish Fauna. Milton S. Love 81 Cover: Giant Sea Bass, Stereolepis gigas, photo credit Parker House.