Biogeography of Hemisquilla ensigera californiensis (Crustacea: Stomatopoda) with Emphasis on Southern California Bight Populations

Lawrence V. Basch1 and John M. Engle2 1Biology Board of Study & Institute of Marine Sciences, University of California, Santa Cruz, CA 95064 2Department of Biological Sciences, University of California, Santa Barbara, CA 93106

distributed but containing few species, each Abstract. - The mantis shrimp Hemisquilla with relatively restricted ranges (Manning ensigera californiensis (Crustacea: Stomatopoda) 1980). Four species occur in the warm ranges from south of Point Conception in temperate waters off southern California. They southern California to the Golfo de Chiriqui, are Pseudosquillopsis marmorata (Lockington Panama. Hemisquilla populations occur off four 1877), Nannosquilla anomala Manning 1967, of the eight California Islands: Santa Cruz, Schmittius politus (Bigelow 1891) and Anacapa, Santa Catalina and San Clemente, as Hemisquilla ensigera californiensis Stephenson well as numerous islands off Baja California and 1967. This study focuses on H. e. californiensis, mainland Mexico. Insular and continental the largest and most common of the California populations occur in habitats usually mantis shrimps, which is distinctive among the characterized by: 1) generally stable, low Stomatopoda in its distribution, size, visual physical disturbance regimes; 2) silty sand characteristics, and aspects of ecology and sediments with a high organic content; 3) behavior (Basch & Engle 1989, in prep.). abundant food resources; 4) variable predation Hemisquilla ensigera californiensis is a large (to pressure and 5) fluctuating light regimes. 30 cm TL), compact, dorso-ventrally compressedmanti s shrimp with a heavily armored Mechanisms that may influence the observed telson, bright coloration and well developed patterns of distribution and abundance include: vision (pers. obs.; Schiff et al. 1986 and included 1) ocean currents favorable for dispersal of references). These stomatopods possess a very planktonic larval stages; 2) habitat suitable for powerful strike mechanism in the second successful recruitment; 3) sufficient food maxilliped or raptorial claw, the most rapid resources and 4) low levels of predation movement known (Burrows 1969; pressure and physical disturbance. Burrows & Hoyle 1972).

In this paper we discuss the relictual (sensu Introduction Ekman 1953) distribution of this species, focus particular attention on populations occurring in The order Stomatopoda representsthe Southern Californi a aprimitiv Bight,e angroud addresp (Holthuis s & Manning 1969; Schram 1977, 1979), whose center of aspects of population biology relevant to radiation occurs in tropical latitudes. The observed local distribution and utilization of phylogenetically separate position of stable sand habitats. stomatopods is reflected by Caiman's (1904) establishment of the superorder Hoplocarida, Methods which is postulated to have a distinct origin separate from the eumalacostracan Crustacea Biogeographic data on Hemisquilla ensigera (Schram 1969a,b, 1986). More than 350 species californiensis were compiled from direct in four super families are known, most dwelling observations and collections from populations in warm tropical seas throughout the world. off the California Islands and southern Many genera appear to be relicts, broadly California mainland, and from personal communications (including solicited surveys),

Third California Islands Symposium 211 museum collection records, and review of the morphological divergence. Details of global flats off the southeastern side of East Anacapa The typical bathymetric range extends from literature. Whenever possible, data were distribution and morphological variation in the Island in depths of 17-30 m, at two sites off 3 m in protected embayments (e.g. San Diego collected on: locality; latitude and longitude; genus Hemisquilla may be found in Manning Santa Cruz Island between 15-17 m and three Bay sand flats) to >30 m in many areas. depth; habitat; animal abundance and size; date; (1977) and Stephenson (1967), respectively. locations off San Clemente Island (Fig. 2) in Observed peak burrow densities at California time and method of sighting or collection and The geographic distribution of Hemisquilla depths of 13-17 m. Santa Catalina Island (Fig. Islands sites occur between 10-16 m depth. the source of information. ensigera californiensis (Figs. 1-2) ranges from 2) was surveyed most extensively. Populations Mainland sightings and collections are from Santa Barbara County, south of Point Population studies were conducted at several were observed in depths between 5-33 m at 19 10-70 m; appear to be locally common Conception (35° N latitude) in the Southern sites off Santa Catalina Island (33° 28' N, 118° stations (Fig. 2), where mantis shrimp were between 50 and 70 m, and peak abundances California Bight, south to the Golfo de 20' W; Fig. 2). The Willow Cove population common to moderately abundant. appear in the 15-25 m range. Extreme depth Chiriqui, Panama (5° N latitude) (present on the northern leeward shore of Santa Habitat Characteristics: Based on personal records range from an individual observed study; Hendrickx & Salgado-Barragan 1987; Catalina Island was studied most intensively. sightings and distributional records, it is foraging nocturnally in the rocky intertidal (<1 Manning 1977). The southern range limit is Population spatial and temporal distribution inferred that Hemisquilla populations may occur m) at high tide (M. Michael pers. comm.), to based on two specimens, a single juvenile from and abundance patterns were elucidated by whenever suitable oceanographic conditions 1,800 m for a trawl caught specimen in the Jicarita Island (Stephenson 1967) and an adult repeated diving surveys performed at various and habitats coincide. These environments Guaymas basin east of Isla Tortuga, in the Gulf male from the Golfo de Chiriqui (Manning times throughout several 24 hr periods. A 50 m include bays, lagoons and off protected low of California. x 60 m sampling grid (longest axis parallel to 1971). The next northern-most records are energy coastlines. Most of these areas have the The silty sand substrate most commonly shore) was established from 6-18 m depth. Five from Islas Clarion, Soccoro, and Tres Marias following habitat characteristics in common. inhabited by Hemisquilla is found along- transects were surveyed on numerous dates and off the Mexican mainland, south of Cabo Falso, The substrate consists of flat or gently sloping extended stretches of coast, throughout coves times in order to obtain data on all visible Baja California Sur (18° N latitude, Fig. 1). (0° to 30°) stable sand plains ranging from a and bays, or in patches (10's-100's m in burrows. All burrows sighted within and near Data obtained to date indicate that the minimum of 3 m depth inshore to ≥33 m depth diameter) within other, generally coarser the grid were marked and mapped. Data Californian Hemisquilla occur in mainland and seaward. A shallow zone with few or no resident sediment types. For example, Hemisquilla is collected included: time of observation; burrow insular populations in the Gulf of California at stomatopods usually is present from 5-8 m widely distributed in Santa Monica Bay from depth; diameter; whether the burrow was open least as far north as Isla San Nicolas, north of depth. The adjacent inshore area consists of north of Malibu to the Palos Verdes Peninsula or closed by a sand plug constructed by the Guaymas, Sonora, Mexico (28° N latitude, Fig. either physically disturbed sediments (Fig. 2). The local distribution pattern here resident; whether or not the animal was 1) (present study; Hendrickx & Salgado- (delineated by wave ripple marks), small appears to be primarily influenced by the observed; maturity and sex (juvenile, subadult, Barragan 1989). boulders, surf grass or rock reef. Sediments are presence of patches of suitable silty sands within female or male). Hemisquilla is known from numerous silty sands with a poor to moderately sorted areas of predominantly coarser sands. The peninsular and insular sites along the Pacific particle size distribution and a high organic overall distribution of the patches may change Results coast of Baja California (Fig.l). Populations content (Basch 1985). In some areas they are in some shallow areas as a result of seasonal appear to be common in or near embayments, stabilized by matrices of infaunal invertebrate Biogeography: Zoogeographic distribution onshore-offshore sand movements, sediment lagoons and off many south-facing points, tubes, and/or surface diatom or algal mats. in the Stomatopoda is just now becoming clear resuspension by storm waves and other coasts, and island shores as well as other areas These areas usually have a low to moderate as taxonomic revisions approach stability. The transport mechanisms (J. Cross pers. comm.). north to, and including, the Islas del physical disturbance regime, exhibiting vast majority of genera and species are shallow, Population Biology: Recruitment to the Coronados near the U.S.-Mexico border. protection from most heavy swells. tropical warm water forms (Schram 1986). Of stable sand community has been documented Observations from many points along the Predominant water movements result from the many extant genera of stomatopods only since 1983. Postlarvae are capable of rapid southern California coast range from San mixed-diurnal tides and fairly predictable tidal two, including Hemisquilla, may be considered burrowing and apparently build and occupy Diego Bay and Point Loma in the south to the and other long-shore currents. Exceptions to strictly warm-temperate in distribution burrows almost immediately upon settlement, area just south of Point Conception (Fig. 2). this pattern include strong surge induced by (Manning 1977). Hemisquilla (Hansen) includes since very small postlarvae have been collected California mainland populations occur in the powerful winter storm activity that causes two species (Manning 1969; Stephenson 1967), from burrows with entrance diameters of only same geographic and habitat types as indicated shifting sediments, scouring and/or destructive H, braziliensis (Moreira) from the western 3-4 mm (pers. obs.). At present we have by Mexican records (see above). Areas of high water movement. In more protected areas the Atlantic and H. ensigera (Owen), with limited data on Hemisquilla population abundance include San Diego and Mission bathymetric peak in abundance occurs in Californian, Chilean, and Australian subspecies parameters. Based on regression analysis of Bays, the Orange County coast and numerous relatively shallow water, 10-16 m depth. In (Stephenson 1967). The species H. ensigera burrow diameter vs. carapace width, harbor, bay and sandflat habitats from southern areas that regularly or intermittently appears to support the principle of taxonomic population size distribution ranges from about Los Angeles County (Seal Beach) north, at least experience high energy conditions, highest bipolarity (Ekman 1953) by formation of 3-65 mm (carapace width) (Basch & Engle in to Goleta, Santa Barbara County (Fig. 2). population densities may occur at greater relictual populations and subsequent prep.). As in other stomatopods (Reaka 1979) Hemisquilla recently was recorded from sand depths (approximately 16-30 m). the observed size distribution of Hemisquilla is

212 213 Figure 1. Distribution of Hemisquilla ensigera californiensis from Mexico to Panama. Figure 2. Distribution of mainland and island populations of Hemisquilla ensigera californiensis in the Southern California Bight. Blowup shows locations of survey sites on Santa Catalina Island. strongly skewed with a predominance of In addition to use of the burrow for mating extensive work on feeding ecology, widespread the vicinity of ocean sewage outfalls, below bait- adults over juveniles. The largest animals and reproductive purposes, the burrow also predation on stomatopods by fishes and other barges, off public fishing piers and other sampled are almost exclusively males. A undoubtedly functions as shelter from extremes potential predators has not been documented frequently fished areas). realistic estimate of population sex ratio is not of the external physical environment and in California (Hobson & Chess 1976, 1986; possible at present due to seasonal changes in predators. Protection from extreme physical pers. obs.). At present only one instance of Discussion observed abundance (Basch & Engle in prep.) disturbance provided by the burrow is predation on an adult Hemisquilla has been With a perspective on historical and present and the inability to determine the sex of some suggested by the absence of open burrows in noted in California (P. Haaker pers. comm.) individuals in the field. Of a total of 63 adults shallow (≤ 12 m) depths during storm-induced Hemisquilla forage from burrows to a variety distribution and aspects of the biology of collected at Willow Cove, Santa Catalina strong surge (pers. obs.). Several potential of habitats including subtidal sand flats and Hemisquilla and other stomatopods, we may Island in 1985 and 1986, the sex ratio predators inhabit stable sand communities. The sand channels, and infrequently onto rocky infer mechanisms contributing to biogeo- (female :male ) was: 12:15 (November 1985); predator guild changes throughout the substrata including kelp forests and the graphic distribution and evolutionary and 0:17 (May 1986); 15:4 (November 1986). The latitudinal range of Hemisquilla, with some intertidal zone. They may range a great ecological patterns relating to the dispersal, absence of females in May 1986 along with large piscine predators occurring more distance (to 60 m) from burrows and prey on a colonization and persistence of tropically relatively low apparent population density frequently in waters south of California. wide variety of invertebrates (Basch & Engle derived animals such as Hemisquilla in cold, strongly suggests that females remain in Several large black sea bass and grouper off 1989). Animals frequently have been observed high latitude environments. plugged burrows for extended periods in the Baja California are known to have ingested up and captured in soft bottom communities with Historical Events: The absence of spring, presumably while copulating or to 5-6 adult Hemisquilla (each ≥2 0 cm TL) per high infaunal biomass and in locations where Hemisquilla ensigera in equatorial latitudes in brooding eggs. predator (E. Sigler pers. comm.). Despite food abundance is artificially enhanced (e.g., in the eastern Pacific (Manning 1977) suggests a

214 215 case of taxonomic bipolarity (Ekman 1953) have several larval instars including few to many mass typically extends to San Clemente and transport of offshore water, increased where, in this instance, the same species may (e.g., 4-9) pelagic stages (Williams et al. 1985). Santa Catalina Islands and by September or downwelling and countercurrent intensification have initially been represented in high latitudes Hence, their dispersal is closely tied to currents. October it reaches north to Anacapa and Santa as the primary mechanisms which bring by two relict populations, and later by two Since Hemisquilla postlarvae and adults both fall Cruz Islands. This summary of water warming to southern California waters. subspecies, H. e. californiensis in the northern in the large size categories established by Reaka movement patterns illustrates the mechanism Atmospheric and oceanic forcing of warming hemisphere and H. e, ensigera in the southern, (1986), we infer that this species possesses for potential dispersal of larvae and the adult processes occurred together during the 1982- with a gap in distribution between the two. several pelagic life history stages which spend distributions of Hemisquilla as far north as the 1983 California El Nino (Norton et al. 1985). Present Biogeographic Distribution: As long periods (up to several months) in the Southern California Bight. It also may account Several lines of evidence indicate a recent indicated by Fig. 1-2, Hemisquilla ensigera plankton. It follows that Hemisquilla probably for the absence of Hemisquilla at San Nicolas large recruitment event for this warm water californiensis exhibits a broad distribution, has a high dispersal potential, and this may and the other northern Channel Islands, radiated species. Hemisquilla was never spanning 30° latitude. Possible limitations on explain its broad latitudinal range. A long despite the availability of suitable habitat in observed to be common or in high abundance knowledge of the detailed distribution patterns planktonic residence time likely is associated deeper water, because these islands are seaward off the California Islands from 1973-1983 south of the Southern California Bight may be with more larval mortality via predation and of the Santa Rosa-Cortez Ridge and, hence, are (pers. obs.; R. Given pers. comm.). Since the a reflection of a relatively coarse-grained starvation (Vance 1973) and less predictable not influenced by the north flowing shoreward early 1970's increased mainland trawl catches picture (i.e., comparatively few sightings in a recruitment to any given site (Thorson 1946). portion of the Southern California Eddy. This of adult Hemisquilla have been recorded large area made over many years). In contrast, High juvenile mortality suggests great longevity flow would be expected to cany larvae from approximately two years after warm water the image in southern California is fine-grained and iteroparity in adult stomatopods (Reaka mainland populations to those California events (R. Fay & S. Anderson pers. comms.). (i.e., frequent, accurate sightings with good 1979, 1986). Islands with known Hemisquilla populations, Observations of intense recruitment to several spatial coverage of the region over the past fifty Large Scale Mechanisms: The principal and few, if any, larvae would be expected to locations from 1983 to the present, as well as years). As a result, the biogeographical pattern large scale mechanisms underlying observed reach other islands. In addition, these flow numerous observations and records of large off southern California may be more indicative recruitment patterns and population patterns indicate that the Pacific coast of Baja increases in abundance of adult Hemisquilla of the actual distribution of Hemisquilla. distributions are currents. Seasonal and California may receive larvae both from throughout the region during the same period To understand the modern biogeographic aperiodic atmospheric and oceanographic southern California populations and from (unpublished data) indicate a strong influence distribution of Hemisquilla, recruitment and the conditions, including near shore eddies, permit populations south of 23° N. of warm water events, particularly the 1982- biological and physical factors influencing it northerly current flow from Panama to central The area just south of Point Conception is 1983 El Nino, and a continuing warm water must be considered on several scales. Large Baja California (Wyllie 1966; Norton et al. the northern limit of Hemisquilla and many influence on the distribution and abundance of scale factors (i.e., interacting biotic and abiotic 1985). Below 31° N nearshore current flow is other southern species of marine invertebrates. Hemisquilla. During this period the persistence mechanisms) affect the dispersal of larvae and predominantly southward throughout much of Except for the two small Santa Cruz Island of Hemisquilla populations has been stable in the overall pattern of larval availability over the the year. However, by late summer to early fall, populations, all known populations of place and time. Similar observations of distributional range. Small scale factors affect flow in this region is complicated by eddy Hemisquilla occur south of the 13° C surface anomalously high recruitment and numbers of larval settlement. Lastly, factors such as formation which may produce northward isotherm during most of the year. That adult and fishes also have been competition, predation and physical conditions nearshore currents (Wyllie 1966). In the populations occur near this environmental / noted by Cowen (1985). associated with early benthic life mediate post Southern California Bight, the southward flow latitudinal extreme may be explained by year to Small Scale Mechanisms: Acting in settlement survival of postlarvae and juveniles. of the California Current diminishes somewhat year variation in oceanographic conditions combination, near shore physical transport These processes may all interact on a variety of within 150 km of shore in August through (Jones 1971). In exceptional years warm mechanisms, larval behaviors and the local levels to produce a complex pattern of October. In late November through January it southern waters extend north of the latitude of availability of suitable habitat types are the recruitment and resultant adult distribution reverses to become a weak northerly flow Santa Cruz Island. These anomalous events, principal small scale factors influencing larval (Cowen 1985). (Hickey 1979). The Southern California Eddy, characterized by northward transport of warm settlement. We have seen that most stomato­ Hemisquilla Biology: At present there is similar to the California Current shows water throughout the Southern California pods, and probably Hemisquilla as well, have little information on life history in Hemisquilla, considerable, within-year variation (Reid 1965). Bight, are frequently associated with El Nino current dispersed larval phases. Nearshore however, based on the work of Reaka and Its axis corresponds roughly with the Santa events. These dramatic warm water incursions current patterns implicated in larval transport others (see Reaka 1986) on the relationships Rosa-Cortez ridge (Sverdrup & Fleming 1941). last occurred in 1972, 1977-1978, and 1982- can be influenced by within-year changes in between larval dispersal capability, postlarval Highest northward flow occurs shoreward of 1983 (Chelton 1981; Norton et al. 1985) and wind direction, bottom topography and and adult size, habitat and latitudinal this axis in the southern part of the eddy. dominated current patterns in the Southern anomalous oceanographic events (Jones 1971; distribution, we can infer several aspects of Warmer water is moved progressively California Bight. According to Norton & co- Norton et al. 1985). Shanks (1983) has Hemisquilla life history from patterns in other northward from July through October by the authors (1985) evidence points to weakening of demonstrated on a smaller scale, the stomatopod species. Stomatopods typically inshore flow of the eddy. By July this water the south flowing California Current, onshore association of surface slicks with shoreward

216 217 moving internal waves which are believed to Acknowledgments and R.L. Caldwell. 1972. Reproductive and Norton, J., D. McLain, R. Brainard and D. Husby, transport larval invertebrates and ichthyo- maternal behavior of the mantis shrimp We wish to thank the many biologists, divers, 1985. The 1982-83 El Nino event off Baja and plankton to nearshore environments. Larval Gonodactylus bredini Manning (Crustacea: museum curators, fishermen and journalists that Alta California and its ocean climate context. Pp. Stomatopoda). Biol. Bull. 142:417-426, 44-72. In: W.S. Wooster and D.L. Fluharty (eds.), invertebrates, particularly crustaceans, exhibit assisted in the compilation of detailed Ekman, S. 1953. Zoogeography of the Sea. Sidgwick El Nino North: effects in the eastern subarctic several behavioral mechanisms that would distributional records. The Tatman Foundation & Jackson: London, ENGLAND. 417 pp. Pacific Ocean. Washington Sea Grant. permit transport in these slicks (Ennis 1975; through the Channel Islands Research Program Ennis, G.P. 1975. Behavioral responses to changes Reaka, M.L. 1979. The evolutionary ecology of life Rice 1966; Thorson 1964). More specifically, provided support in all aspects of field work in hydrostatic pressure and light during larval history patterns in stomatopod Crustacea. Pp. Reaka (1986) and Dingle (1969) have shown including ship time aboard R/V Cormorant and development of the lobster Homarus americanus. J. 235-260. In: S. Stancyk (ed.), Reproductive Fish. Res. Bd. Can. 32:271-281. both diel and ontogenetic changes in the the able seamanship of Capt. Jerry Chomeau ecology of marine invertebrates. University of Hendrickx, M.E. and J. Salgado-Barragan. 1987. Los behavior of stomatopod larvae which may allow and crew. Illustrations by Kathy Langan, South Carolina Press: Columbia, SC. 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