ISSN 0038-3872 SOO LHERN CALIFORNIA ACADEMY OF SCIENCES LLETIN Volume 97 Number 1 BCAS-A97(1) 1-48 (1998) APRIL 1998 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 © Southern California Academy of Sciences, 1998 OFFICERS Robert S. Grove, President David Huckaby, Vice-President Jane A. Peterson, Secretary Margaret A. Neighbors, Treasurer Daniel A. Guthrie, Editor Hans Bozler, Past President David Soltz, Past President BOARD OF DIRECTORS 1996-1998 1997-1999 1997-2000 Kathryn A. Dickson Ralph G. Appy Robert S. Grove Margaret A. Neighbors Jonathan Baskin David Huckaby Jane R. Peterson Karen Martin Robert Lavenberg Robert E Phalen J.D. Stewart Kenneth E. Phillips Cheryl C. Swift Gloria Takahashi Susan E. Yoder Membership is open to scholars in the fields of natural and social sciences, and to any person interested in the advancement of science. 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Date of this issue 9 April 1998 © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Bull. Southern California Acad. Sci. 97(1), 1998, pp. 1-8 © Southern California Academy of Sciences, 1998 Temporal Changes in Diet and Foraging Habitat of California Killifish (Fundulus parvipinnis) in Marina del Rey, California Kristine Behrents Hartney and Lusine Tumyan Department of Biology, Occidental College, Los Angeles, California 90041 Abstract.—Feeding habits of the California killifish, Fundulus parvipinnis Girard, in Marina del Rey were examined relative to season and prey availability. Tem- poral changes in diet coincided with ontogenetic shifts in foraging habitats. Ju- venile fish fed selectively on relatively rare planktonic harpacticoid copepods and surface dwelling insects, whereas diets of adults were dominated by benthic prey taxa, principally tanaids. The seasonal availability of some prey taxa and the active selection of others associated with a particular foraging habitat affected the type and size of prey consumed. Our results indicate that patterns of prey use are not simple responses to seasonal changes in prey abundance. The California killifish, Fundulus parvipinnis Girard, is a common resident in shallow protected waters along the California coast from Morro Bay, California to Almejas Bay, Baja, Mexico (Miller and Lea 1976; Swift et al. 1993). While most abundant over muddy/sand bottoms of bays and estuaries, this small (less than 115 mm SL), short-lived (18 mo) fish may also persist in freshwater streams (Hubbs 1916; Miller 1939, 1943; Swift et al. 1993) and hypersaline ponds (Feld- meth and Waggoner 1972). Because of its ability to tolerate a broad range of environmental conditions, this species has been the focus of numerous physio- logical studies (Keys 1931; Wells 1935a, 1935b; Doudoroff 1945; Carpelan 1961; Hubbs 1965; Valentine and Miller 1969; Feldmeth and Waggoner 1972; Bagarino and Vetter 1992, 1993). However, far less consideration has been given to studies of its natural history, particularly feeding habits, despite its numerical dominance and important role in the structure and trophic dynamics of estuarine and bay communities (Fritz 1975; Allen 1980, 1982). Trophic analyses of killifish collected monthly from Anaheim Bay, California from November 1969 through January 1970 by Fritz (1975) and of fish sampled bimonthly from upper Newport Bay from January through November 1978 by Allen (1980) indicate that killifish are lower trophic level carnivores consuming primarily small crustaceans (e.g., ostracods, harpacticoid copepods, gammarid am- phipods), dipteran and hemipteran insects (Fritz 1975; Allen 1980), polychaetes, and gastropods (Allen 1980). However, changes in dietary patterns over time within a particular locale (Fritz 1975; Allen 1980) and differences in diets between these locations indicate that both temporal and spatial factors influence the type of prey consumed by these fish. While differences in diet within and between these sites may simply reflect a response by these fish to varying patterns of resource abundance (Moyle 1976), patterns of diet choice may also be driven by selective feeding behaviors that may change with age (ontogenetic Shifts) and prey characteristics (Kaiser and Hughes 1993). Therefore, the CALIFORNIA ACADEMY OF SCIENCES APR 2 8 1998 LIBRARY i) SOUTHERN CALIFORNIA ACADEMY OF SCIENCES study were to assess the feeding habits of Fundulus parvipinnis that reside in Marina del Rey relative to season and to examine the relationship of prey type, size, and availability to dietary choices. Methods All field work was conducted in water depths less than 3 m just offshore of the public swimming beach located at the extreme western end of Basin D within Marina del Rey. Killifish were collected following high tides on two occasions, March 30, 1995 (1300 to 1430 hrs) and October 13, 1995 (1130 to 1300 hrs) using a 30.5 m beach seine (4.8 mm bag mesh) that was initially deployed about 10 m parallel to shore and then slowly drawn up onto the beach. A total of 129 and 150 fish were captured and immediately fixed in 10% formalin during March and October respectively. On the October sampling date, prey resource samples were also collected from the water column and: substratum in the same general area as fish captures. To obtain an estimate of planktonic prey available to fish, the water column was sampled by SCUBA divers propelling a plankton net (0.33 mm mesh) with a 0.5 m diameter mouth opening for two minutes through midwater depths (0.5 to 2.5 m). Three replicate tows were made and all samples were immediately fixed in 4% formalin. Estimates of benthic prey available were obtained by driving a 12.5 cm long benthic core (10 cm internal diameter) completely into the substratum. Three replicate cores were taken and entire samples fixed in 10% formalin. Each benthic sample (982 cm?) was later washed through a 0.5 mm sieve and only the larger fraction was retained for further analysis. In the laboratory, the standard length of each killifish was measured to the nearest 0.1 mm using Vernier calipers and its age determined by reading scale annuli (Cailliet et al. 1986). The entire gut and gonads were then removed from a subsample of fish selected at random from March (N = 30) and October (N = 47) collections. The dietary analysis of 30 fish taken from the March sample was considered adequate for determining large scale differences in diet between sea- sons, however a greater number of fish (47) were analyzed from the October sample to ascertain fine scale patterns of prey selection within a single time period. All prey were removed from each gut and identified to the lowest taxon practi- cable. Whole animals or heads (when prey were not intact) were counted and the greatest length of up to 25 intact individuals per taxon (as randomly encountered) was measured to the nearest 0.01 mm using an ocular micrometer. Gender and reproductive status of killifish were determined by examining the gonads for pres- ence of eggs or sperm and scoring their stage of maturity according to the methods described by Cailliet et al. (1986). Reproductively immature fish less than one year old were considered juveniles. The contents of planktonic and benthic resource samples collected in October were identified, counted and measured using methods similar to those described for gut content analysis. However, each planktonic resource sample was split twice with a Folsom plankton splitter so that all organisms in only one quarter of the sample were completely identified, counted, and up to 25 individuals per taxon measured. Numbers and sizes of prey for the entire plankton sample were later extrapolated. Invertebrates (>0.5 mm) contained in each benthic resource sample were separated from sand by gently floating them away from inorganic materials FORAGING PATTERNS OF CALIFORNIA KILLIFISH 3 100 March 1995 October 1995 80 60 40 Number (%) 20 20 40 . harpacticoid copepods . tanaids gammarid amphipods . nematodes diptera . hymenoptera . hemiptera . diplura eggs 60 80 Frequency of Occurrence (%) A B Cc. D E. F G H I. 100 A B Cc D = F I A B Cc D E F G H Fig. 1. Percent composition of prey by taxon, number (%) and frequency of occurrence (%) in diets of 29 killifish collected in March 1995 (N = 2107 prey) and 40 killifish collected in October 1995 (N = 2097 prey). in distilled water. The remaining inorganic fraction was then examined for any misplaced organisms, which when found were combined with the organismal fraction that was analyzed in its entirety. To examine the relationship between prey consumed relative to prey available, electivity coefficients were determined for all available prey taxa collected in October resource samples using Ivlev’s Electivity Index (Ivlev 1961). While the sign and magnitude of resulting coefficients (from —1 to +1) were used as in- dicators of possible avoidance (negative values), random (zero values) or pref- erential (positive values) selection of specific prey taxa, the significance of these food preferences was statistically evaluated using a Chi Square analysis described by Pearre (1982). For these analyses, the total number of prey collected from the water column and substratum were combined and considered as a single resource base. Results Temporal Changes in Diet and Foraging Habitat Although the dietary spectrum of prey taxa consumed by fish collected in March and October was similar, there was a striking difference between them in the dominant types of prey eaten (Fig. 1). The diets of killifish in March were 4 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES dominated by benthic prey species, principally tanaids, and the nearly complete absence of planktonic or terrestrial prey (insects) in diets suggested that killifish foraged along the bottom at this time of year. In contrast, planktonic harpacticoid copepods were by far the most common and numerically abundant prey consumed by fish in October and insects were consumed with a relatively high level of frequency. The occurrence of these prey types relative to others suggest that in the fall of the year, fish fed in the water column or from the surface. The mean size of prey (+1 standard deviation) consumed by killifish in March (x = 1.85 + 0.96 mm) was significantly greater than the mean size of prey consumed in October (* = 0.55 + 0.52 mm; t = 54.7, df = 3244, P < 0.01). Overall, reproductively mature fish collected in March were significantly larger (x = 6.4 + 0.7 cm SL) and older (>1 yr) than the sexually immature juveniles (x = 3.9 + 0.9 cm SL) that were collected in October (t = 28.5, df = 277, P< 0.01). These size differences were maintained in subsamples of fish randomly selected for gut content analysis (March x = 6.3 + 0.8 cm SL; October * = 3.9 + 0.9 cm SL). Of the 30 fish collected in March that were selected for gut content analysis, 29 contained prey, whereas 40 of the 47 fish selected from October samples had prey in their guts. Availability and Selectivity of Prey In general, prey types were unique to a particular resource sampling area and taxa composing planktonic and benthic groups were easily separated from one another on the basis of relative differences in their numerical representation in samples taken from the water column and substratum (Table 1). Plankton samples were dominated by calanoid copepods, while organisms generally associated with the substratum such as tanaids, polychaetes, nematodes, and gammarid amphipods dominated benthic samples. Taxon specific differences in size and distribution strongly affected size frequency distributions of available prey (Fig. 2) such that the mean size of prey available from the water column (x = 0.82 + 0.18 mm) was significantly less than that of benthic (« = 3.75 + 2.35 mm) prey (t = 45.3, df = 1323, P < 0.1). Reflecting either sampling bias or low frequency of oc- currence, no insects were collected in resource samples. Killifish consumed few prey in direct proportion to their availability in the water column or substratum (Table 1). Significant electivity values indicate that in October, killifish preferentially selected relatively rare small harpacticoid co- pepods from the water column while they avoided relatively abundant calanoid copepods and numerically abundant benthic taxa (tanaids and polychaetes). Even though terrestrial insects were not represented in resource samples, the frequent appearance of insects in killifish guts indicate that they were probably selected whenever they were encountered. The choice of small planktonic harpacticoid copepods by fish in October strongly influenced the size range (0.2 to 5.9 mm) and mode (0.4 mm) of prey found in the guts relative to the range of sizes (0.1 to 3 mm) and mode (0.8 mm) of all planktonic prey available (Fig. 2). Although capable of consuming relatively large prey such as insects, they apparently avoid- ed benthic prey (e.g., tanaids) within a similar size range. Thus, differences in the size frequency distributions of prey consumed relative to those of available prey resulted from the active selection of some prey taxa over others rather than the selection of large or small prey regardless of taxon or habitat. FORAGING PATTERNS OF CALIFORNIA KILLIFISH 10';0> (Oma. 10:;0> (O'1)+ 10';0> (O'1)+ 10';0> (O'1)+ AVWANIITIS saouarajaid Adig 2K K sO;O< (F'0)0 S0'0> (F:0)= KK 10;0> (On) sO;O< (0 10 10°;0> (9°0)+ sO;0< (0°10 AVWATIIIIS saouasiajoid Adig s0';0< (O° L)0 10°;0> Onn KK 10';0> (O'1)+ 10°;0> (Or) — 10°0> OD — *>K d AWANDITA AWANIITAS sasouaiajoid Adig “S661 19q019Q UI pa}daT[O9 (67 = U) saydwies yn3 ysy pue “(¢ = U) SIYJUDQ ‘(¢ GOW c Seo (10) Z OOS 6S lt (r'0) 6 COWES SEES (O'l) IZ CG) 8'cr =Oreate] (CT) OF (WIUL) 9ZI§ Joquinyy C661 12q019Q—Ppoutnsuods Adig ok COM Ge TOE-91 (10) € (GOOG ae Cal (L'0) SI 7K * * (iOsEi7= Vers -Ci€ (SO) II * (WU) 9ZI§ JOquinNy C661 199019O—pe2ulnsuos Adld **K * 2K K (6°76) 0661 * QiO)SIOR SO T=€10 * KK (ac)* osuey (WIUL) 9ZI¢ C66I 19G019Q—PpeuInsuod Adig (%) oquiny ‘A1039}89 JdINOSaI Adid ayeulaiye ue UT poyIOddyY x.y. ‘soydures ut pajyuasoidal JON x. er us aS! kag (UMW) 9ZI§ JOquiInNy aovjins 1oyemM—oqeyiear Adig (00) 9° mes) (G Ci) sal (Ci ¢ Sa Dal! (3's) 30S (L'0) 9°7 €V-80 (S°S9) €L8 (€'0) 9°0 60-¢ 0 (GO). = (CC) 9 FS GiGi (8°07) LLZ (CHG (Gia | (CAO) CED De “BCI OC (C6) ECCT (0'0) 9°0 So O5oi0 (¢O) (UU) 9Z1§ Joquinyy wimensqns—osyqeyiear Adi GORE Ge Seie-O8l Go). ese (HOC ae eS GO (90) 76r CO) at A eo) Gio) T0Z1 (COED, 2 SI=Go (LO) 09S (GOGO! ~3'1-S'0 (0°86) P9l‘6L GONSO) He0-70 (yO) ZIE€ (Gi) 5 Sel che (10>) (dS) x osuvy (%) Joquinyy (WU) aZIg¢ uULUNTOS JayeEM— HI 8 Tr 9 DpIpuvI DANSSO) depLinsso‘y DIV] assa] DUILJaINUOp 8C I I 9 la I SI (a ZI ds auozojavy) pyouvaiqvdlds ‘f9 viusofisaly oep Tet Iv S ds snjspwojon sepypoude ds snyouvaqojsidy aeprpueasqoysidy a 8 —_—————— Ie} iv .0v Ge RE LE GSE CE TE IS CE GC 28Uale,9e CHC 26] Bib 2h OUI Vi ie ee eae ee uoneig/so1sedg 49) | ———————————————————— ‘keg unuingd ues ye saouepunge saiseds wyovysAjOg “¢€ IGRL 19 POLYCHAETES FROM SAN QUINTIN BAY E 3 x z LEO TE g Gee Vem AE OTR ag ae | Sew “Ve 9lS Ce Gl Eile eles "CS VeCenlS 29 Ol pan ae an “EcG Joquinu [eo], N N N I sepryPpueAqgoYpL |, avauogiyjad ods I S1]D1IOS DAOPA]Og I DAsafias1d (o1dsnuipy) oidsououg DIyIUDAGOAaJaYy O1dsOuoldg N — (oe) N ~ = pipiyouvaqyjnu o1dsouoiug aepruoids ds s1jjksouoig v SI1ID48 (S11]XG) S1]X5 I ADdSIp JUO80X7T QI I (© 194NO] FUOSOX] ds piupig aept]AS I ds auoy) I WNIDINIOIG DUNUO]TDS ap ds auoyony FePHIPdes SHUDIIXIOUL sojdojoosonaTy N \o piqnd sojdojoos 2EpIUIGI® pypydasosoiu siydnuosaaquiy I I - a I VS OVNOes 82 Wemce CEN ISAO CoC sca Le SC eCecGEol wil MeO SA cele Clea Sa eee Say mec uonrig/sotsadg -OL +S. 2 = SS Se eS ‘ponunuoy “¢ a1qRL 20 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Observations: In the studied organisms the transition from thorax to abdomen took place at setiger 25—30; different from the setigers reported by Hilbig (1996):24. Habitat: In shelf, on the continental slope and basin depths (Fauchald 1972). 11 to 2400 m; sand and sand mixed with silt-clay (Hilbig 1996). This species was collected in shallow waters, in both arms of the bay, in sandy-mud and muddy-sand, D = 1-4; Eh = —360 to —160; T = 20-22; O.M. = 0.05-3.00. Distribution: Southern California, Cedros Island, Acapulco, Zihuatanejo, in the Central American Trench (Fauchald 1972). México, Baja California; southern and central California (Hilbig 1996). Family Spionidae Grube, 1850 Prionospio heterobranchia Moore, 1907 Prionospio (heterobranchia) newportensis (Reish 1959) 13—1960:94. (Hartman 1969) 157, Fig. 1. Material examined: 22 Specimens: (12) 2: (13) 2: (16) 8; C26) 2: _@7) 2268 1 G2) 1G3) 33-67) 2. Characteristics: Incomplete organisms with 24 to 46 setigers. Rounded prosto- mium, slightly truncated. Two pairs of black eyes, anterior pair smaller, rounded and farther apart. The caruncule extends to setiger 2, without median antenna. Five pairs of branchiae from setiger 2; pairs 1, 4, 5 pinnate and 2, 3 cirriform. Capillary notosetae limbate, larger in the setigers with branchiae, becoming small- er posteriorly; hooded hooks from setiger 36. Habitat: Intertidal in sand and mud (Calder6n-Aguilera & Jorajuria-corbo 1986). Found in both arms of San Quintin Bay in sand, sandy-mud and muddy- sand, D = 2-5; Eh = —360 to —104; T = 20-—21.6; O.M. = 0.05-—3.00. Distribution: California (Macioleck 1985). From Newport, California to Pana- ma (Calder6n-Aguilera & Jorajuria-corbo 1986). In the Mexican Pacific: Bahia de Mazatlan, Sinaloa (Arias-Gonzalez 1984), Bahia Concepci6én, Baja California Sur (Salazar-Vallejo 1985), Isla Tiburén, Punta Arboleda, Sonora and Isla Maria Madre, Nayarit (Hernandez-Alcantara 1992). Prionospio (Minuspio) cirrifera (Wirén 1883; Macioleck 1985) 352-355, Fig. 10 a—-g Material examined: 1 specimen: 13 (1). Characteristics: Specimen with 23 setigers. Prostomium large and rounded, elongated posteriorly until setiger 2; two pairs of eyes in trapezoidal arrangement. Peristomium with lateral wings moderately developed, not overlaping the prosto- mium. Eight pairs of branchiae, all cirriform, anterior ones larger than posterior ones; not joined to the notopodial lobe. Multidentate hooded hooks present from neuropodia 13-18 and at the notopodia after setiger 26. Observations: In the studied organism the multidentate hooded hooks were present from neuropodia 19, however the reported rank by Light (1978) goes from 13-18. Habitat: Preference for silt and sandy silt, also found on sand and silty sand. Predominantly muddy bottoms, silty mud, off jeties, intertidal rock pools, sand. Eurybathyal, intertidal to 2500 m (Foster 1971; Light 1978). In marine and es- tuarine environments, from the intertidal to great depths (Calderén-Aguilera & Jorajuria-corbo 1986). From 11 to 2900 m (Maciolek 1985). Mud and sandy-mud, POLYCHAETES FROM SAN QUINTIN BAY 21 D = 30-150; T = 27-28; O.M. = 0.8—-1.65 (Rodriguez-Villanueva 1993; Mi- randa-Vazquez 1993). Collected only in the eastern arm, near Muelle Viejo, in sandy-mud, D = 5; Eh = —203; T = 20.5; O.M. = 0.80. Distribution: Arctic; Atlantic from Greenland to South America; North Sea and English Channel; Bering Sea to Gulf of California; Queensland, New South Wales and Victoria in Australia (Light 1978; Calder6én-Aguilera & Jorajuria-corbo 1986). Prionospio (Minuspio) multibranchiata (Berkeley 1927; Macioleck 1985) 365— 367, Figs. 15 a-e. Material examined: (7 specimens): (12) 2; (15) 5. Characteristics: Prostomium rounded, widest at level of eyes. Peristomium fused to setiger 1. Caruncule extending to 1-2 setiger. 4 eyes. Long cirriform branchiae from setiger 2, 7—9 pairs; first branchiae longer. Notopodial lamellae absent in first setiger, in the others well developed. All anterior setae capillary. Hooded hooks from setigers 12—18, in neuropodia from setigers 25—32. Sabre setae from neuropodia 12-16. Habitat: Intertidal (Maciolek 1985). In muddy sand, D = 104 m; T = 14.2; S = 35.26; O.M. = 7.2; DO = 2.40 ml/l (Hernandez-Alcantara 1992). This species was collected only in the middle region of the eastern arm, in silty-clay sediments; D = 4; Eh = —126 to —145; T = 19.1-19.9; O.M. = 3.33. Distribution: Vancouver Island, Canada; Washington, Florida, northern Gulf of Mexico (Macioleck 1985). Concepci6n River, Sonora (Hernandez-Alcantara 1992). Polydora socialis (Schmarda 1861) Polydora plena. Foster, 1971: 24—25, Figs. 22-29. Material examined: (7 specimens): (22) 1; (28) 1; (29) 5. Characteristics: Prostomium bilobed, with 4 eyes. Caruncule extending to se- tiger 4—8. Palps missing. Anterior setae all capillaries. Setiger 1 with notosetae; setiger 5 large, twice the size of preceding segments, with modified rounded hooks. Neuropodial bidentate hooks without manubrium from setiger 7, apical tooth diminishing in size in posterior segments. Branchiae from setiger 7—8 as small digitiform lobes, progressively increasing in size. Dorsal lamellae digiti- form. Pygidium with one large ventral lobe fused with two smaller dorsal ones. Observations: Hernandez-Alcantara (1992) found cirriform branchiae from se- tiger 8 and caruncule extending to setigers 4—9, we observed branchiae from Setigers 7-8 as Light (1978) and caruncule up to setiger 7. _Habitat: Intertidal to 70 m depth (Day 1973; Salazar-Vallejo 1981). In mud and silt, in lagoons (Hartman 1969). Forms silt tubes in a variety of substrates, pri- marily sandy silt (Reish 1968); also boring in living and dead shells (Johnson 1984). In sediments often forming large beds (Light 1978). This species was collected from middle area of eastern arm, as well as near the sea entrance, in sediments sand-silt; D = 2—3; Eh = —30 to —242; T = 18.9-21.6; O.M. = 0.3-— 1.68. Distribution: Cosmopolitan. California to Chile; North Carolina and Gulf of Mexico (Day 1973); San Francisco Bay south to Oceanside (Hartman 1969). East and west coasts of North America, Falkland Islands (Johnson 1984). New South pe SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Wales, Victoria, New Zealand, North and South America (Blake & Kudenov 1978). Spio pettiboneae (Foster 1971; Johnson 1984) 6-63, 6-65, Figs. 6-54 a-e. Material examined: 9 specimens: (4) 1; (13) 1; (30) 3; (32) 4. Characteristics: Incomplete organisms with 18 to 41 setigers. Head and anterior segments with brown pigment pattern. Prostomium anteriorly inflated, rounded, with caruncule. Two pairs of eyes, anterior pair larger and farther apart. Well developed peristomium, nucal organs extending on to setiger 3. Branchiae present, those on setiger | smaller than those on the following setigers. Anterior setae uni- or bilimbate, arranged in two rows. Ventral sabre setae. Tridentate hooded hooks first replacing posterior row of capillary neurosetae on setiger 11-15. Observations: Hooded hooks began at neuropodia 11—13; Johnson (1984) re- ported they begin at 11-15. Habitat: Intertidal to 120 m. Predominately in fine and medium sand; also in silty sand (Johnson 1984). In sand and sandy-mud, D = 28—46; T = 27-28; O.M. = 0.21-1.0 (Rodriguez-Villanueva 1993). This spionid species was collected in the head and the middle area of San Quintin arm and at the mouth of the bay, in sandy and sandy-mud sediments, D = 1—5; Eh = —203 to —94; T = 20.5—21.5; O.M. = 0.43-1.40. Distribution: North Carolina and Gulf of México (Foster 1971; Day 1973; John- son 1984), western Baja California, Mexico. Family Cirratulidae Carus, 1863 Monticellina tesselata (Hartman 1960; Blake 1996) 328-330, Figs. 8.27 a-f. Material examined: 9 specimens: (13) 4; (17) 2; (27) 1; (33) 2. Characteristics: Incomplete organisms with up to 42—63 setigers. Conical pro- stomium, rounded apically, without eyes. Long peristomium, no rings were ob- served. One pair of palps inserted dorsoventrally. Branchiae from the first setiger; inserted above the notopodial base. Notosetae long and slender, forming tufts in anterior and middle regions. Neurosetae shorter and fewer; those in posterior segments geniculate, with serrated edges. Observations: In the studied specimens the location of the tentacular palps was not clearly observed perhaps partly due to the bad state of the material; Blake (1996) said they insert at the posterior margin of the peristomium. The dorsal ridge along the thoracic region was not observed, but its presence is only men- tioned by Blake (1996). Hartman (1969) does not mention it. Habitat: On shelf and slope depths, from shallow water to great depths. In silty and muddy sediments (Hartman 1969; Blake 1996). In sand and muddy sand, D = 39-72; T = 21-30; O.M. = 11-0.94 (Gonzalez-Ortiz 1994). Eastern arm and near the mouth of San Quintin system, in sandy-mud and muddy-sand, D = 2-— 5; Eh = —285 to —200; T = 20.4—21.1; O.M. = 0.5-3.33. Distribution: Southern California (Hartman 1968) to western Mexico: Conti- nental shelf of the Gulf of California (Reish 1968; Van Der Heiden & Hendrickx 1982; Arias-Gonzdlez 1984; Lezcano-Bustamante 1989; Hernandez-Alcantara 1992) and Gulf of Tehuantepec (Gonzdalez-Ortiz 1994); central and southern Cal- ifornia (Blake 1996). POLYCHAETES FROM SAN QUINTIN BAY 23 Cirriformia spirabrancha (Moore 1904; Blake 1996) 361—363, Figs. 8.42 a-f. Material examined: 7 specimens: (8) 5; (14) 1; (35) 1. Characteristics: Incomplete organisms with 82—96 setigers, complete worms with 118—152 setigers. Rounded prostomium, without eyes. Long peristomium with 3 segments forming a protuberance which extends up to setiger 5, where both tuffs of tentacular filaments are already observed. Branchiae appear at setiger 5. Neuroacicular spines from around setigers 40—45, in the first 3—5 per fascicule, increasing to 5—6 from setigers 100 to 150; notoacicular spines beginning pos- terior to setiger 60—70. Observations: In the observed specimens neurospines begin at setigers 17—23 and notospines at setigers 26—40; in the first fascicles there are one or two spines, posteriorly we acknowledged 4—5 per fascicule, not in the number and order mentioned by Blake (1996). He also said that Cirriformia spirabrancha has been confused with Cirriformia moorei due to reports of the last species in muddy sediments, in estuaries and associated with Zostera marina as opposed to C. spir- abrancha, which has only been reported in hard substrates (rocks, gravel at ex- posed beaches). Habitat: Inhabits tide pools in the rocky intertidal on semi-exposed shores. In crevices and under rocks (Blake 1996). Our specimens were collected principally at sandy mud with some shell fragments, middle region of western and eastern arm, D = 2-5; Eh = —280 to —326; T = 20.4—21.1; O.M. = 0.58-2.21. Distribution: Northern, central and southern California (Blake 1996). Family Maldanidae Malmgren, 1867 Clymenura gracilis (Hartman 1969):439, Figs. 1—4. Material examined: 5 specimens: (13) 1, (6) 1, (38) 3. Characteristics: Organisms with 10—13 setigers. Cephalic plaque with smooth, wide flange almost all around. Long nuchal organs, extend through *% of cephalic plaque. Without eyes. Buccal segment as long as first and second setigers, fol- lowed by 5 shorter segments, setigers 9 to 13 longest. First setiger with few uncini increasing to 6 in a row in second setiger. Setigers 7 and 8 with glandular bands. Habitat: In shelf and canyon depths, in silty mud and green sand. Constructs a friable tube of silt and gravel (Hartman 1969). In San Quintin at 3—5 m depth, in the middle area of both lagoon arms, in sandy mud and muddy sand. D = 3-5 m; Eh = —203 to +103; T = 20-21; O.M. = 0.6—2.6. Distribution: Southern California (Hartman 1969). San Quintin lagoon, Baja California. Family Syllidae Grube, 1850 Exogone dispar (Webster 1879; Uebelacker & Johnson 1984) 30—42, 30—43, Fig. 30-36 a-e. Material examined: 15 specimens: (13) 1; (32) 1; (37) 9; (39) 4. Characteristics: Incomplete organisms with 44—48 setigers. Rectangular prosto- mium with two pairs of eyes in trapezoidal arrangement. Pharynx extends to setigers 3—5, with a subterminal tooth. Extended proventriculus up to setigers 3 to 6 or 4 to 7. Antenna rising from anterior part of prostomium, median antenna fusiform, short palps, lateral antenna small and digitiform. Palps completely fused 24 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES dorsally. Dorsal and ventral tentacular cirri digitiform; dorsal cirri present in all setigers. Compound setae with 1—2 superior spinigers per fascicle and one small bidentate seta. Compound falcigers, all distally bidentate with primary tooth ter- minal. Simple ventral setae bidentate. Habitat: Intertidal to 130m (Day 1973). From shallow waters to 5023 m; in sandy shell, sand and corals (Uebelacker & Johnson 1984). Middle area of eastern arm and western arm; in sand and sandy-mud, D = 1-5; Eh = —326 to —174; T = 19.8-20.6; O.M. = 0.53-—2.96. Distribution: Arctic, North Pacific, North Atlantic, Maine to Florida, Gulf of Mexico; Alaska to the Pacific coast of Mexico, Galapagos; South Japan and south- ern Africa (Day 1973; Uebelacker & Johnson 1984). Exogone lourei (Berkeley & Berkeley 1938; Kudenov & Harris 1995) 15-17, Fig. 1.3 a-f. Material examined: 2 specimens: (16) 1; (35) 1. Characteristics: One complete polychaete with 43 setigers, one incomplete with 25 setigers. Cylindrical body. Prostomium wider than long; two pairs of eyes, anterior pair largest, arranged trapezoidally. Antennae located in front of anterior pair of eyes. Medium antenna fusiform, twice as long as paired lateral antennae, with a distal papillae. Palps long and fused. Proventriculus cylindrical, shorter than pharynx. Parapodia with compound spinigers, falcigers and simple setae. Compound falcigers distally bidentate; secondary tooth larger than primary. Ven- tral setae curved and distally bidentate. Dorsal and ventral cirri digitiform; dorsal cirri on all setigers. Natatory setae present from setiger 17. Pygidium with a pair of cirriform anal cirri. Habitat: Lower intertidal to 40 m. Calcareous crusts, silty sand, coarse black sand, gravel with mud; mud. In the middle area of both arms of San Quintin system, in sandy-mud, D = 4-5; Eh = —360 to —320; T = 20-22; O.M. = 0.79-1.99. Distribution: Spain. Atlantic: Gulf of Mexico, Belize, Cuba, Canary Islands; Pacific: south of British Columbia to southern California (Nufiez et al. 1992); Maria Madre island, Nayarit (G6ngora-Garza 1984). Washington, Oregon, Cali- fornia, Gulf of Mexico: Texas, Louisiana, Mississippi, Alabama, Florida; Cuba; Spain (Kudenov & Harris 1995). Syllis (Syllis) gracilis (Grube 1850; Gardiner 1976) 139, Fig. 12 I-n. Material examined: 4 specimens: (6) 4. Characteristics: Complete specimens with up to 73 setigers, incomplete organ- isms with 37—63 setigers. Prostomium rounded with four small eyes; one median antenna with 7—22 articles and two lateral antenna with 7—12 articles. Proventricle extending two setigers. Triangular palps; dorsal and ventral tentaculat cirri artic- ulated. Ypsiloid setae between setigers 17—20. Pygidium with a pair of articulated anal cirri. Observations: Studied specimens presented ypsiloid setae between setigers 17 and 20, whereas Uebelacker (1984) mentioned they are present between setigers 14 and 21. Habitat: Shallow water to 235 m; among ascidians, algae, serpulid tubes, rocks, barnacles, oysters, hydroids, broken shells; on pillings. Coarse to fine-very fine POLYCHAETES FROM SAN QUINTIN BAY 25 sand, silty fine to very fine sand (Uebelacker & Johnson 1984). In the eastern arm, near Molino Viejo; silty clay, D = 3; Eh = +103; T = 20.5; O.M. = 2.80. Distribution: Cosmopolitan in temperate and tropical seas (Uebelacker 1984). Family Nereididae Johnston, 1845 Neanthes acuminata (Ehlers 1868; Taylor 1984) 31-15, Fig. 31-14 a-e. Material Examined: 26 specimens: (8) 1; (13) 4; (16) 7; (19) 4; (26) 2727) 7; Gr): I. Characteristics: Organisms 27 to 42 setigers. Prostomium short, wide posteri- orly. Tentacular cirri short. Oral ring of pharynx completely encircled by five or more rings of paragnaths. Group I = a line of four points or 8—12 in oval group, II = a wedge shaped group, III = an oval group of about 20, IV = a triangular group of about 20, V, VI, VII and VIII form a complete band of several irregular rows of roughly equal points. Parapodia similar in all body regions. Notopodia with well developed pre- and post-setal lobes. Neuropodia with longer presetal and shorter postsetal lobes. Neurosetae include homogomph spinigers and hetero- gomph falcigers. Habitat: Littoral zone to 100 m. Occurs in fine to coarse sediments, often associated with vegetation (Day 1973; Taylor 1984). In mud, sand and silty clay (Fauvel 1923). Species frequently found along the middle area of the eastern arm, in sand, sandy-mud and muddy sand, D = 1.5—5; Eh = —360 to —100; T = 20- 21.6; O.M. = 0.28—2.78. Distribution: Cosmopolitan in temperate and tropical seas. North Atlantic (En- glish Channel to Santander); Massachusetts to Florida; Mediterranean (France, Italy, Monaco); Southern California to Mexico; Tasmania and New Zealand (Day 1973; Taylor 1984). Family Goniadidae Kinberg, 1866 Goniada maculata (Orsted 1843; Hilbig 1994) 226—228, Figs. 7.5 a-l. Material examined: 2 specimens: (30) 1; (31) 1. Characteristics: Organisms with 116 and 128 setigers. Body slender, prosto- mium conical with up to 10 short rings, four biarticulate antennae. Eyes absent. Short proboscis; terminal jaws surrounded by 18 papillae; chevrons numbering 8 to 9 on each side, largest pieces in the middle of the group. Parapodia about twice as long as body width in posterior part of the body; uniramous through setiger 23 to 37. Notopodia with digitiform presetal lobe. Dorsal and ventral cirri digi- tiform and of subequal length. In anterior parapodia few falcigers; notosetae short, serrated capillaries; neurosetae compound spinigers and falcigers present in an- terior parapodia. Observations: The studied polychaetes presented 9 chevrons on each side, and were uniramous through setigers 35—38. Hilbig (1994) also mentioned uniramous through setigers 23 to 37. Habitat: Intertidal to 3000 m. Occurs in silt, sand and shelly sand (Gardiner 1976; Hilbig 1994). In sandy mud and muddy sand by D = 25—40; T = 28-31; O.M. = 0.40-0.60 (Gonzalez-Ortiz 1994). South region of San Quintin arm, in sand, D = 4; Eh = —94 a —104; T = 20-22; O.M. = 0.40-0.50. Distribution: Cosmopolitan (Gardiner 1976). Arctic Ocean: Greenland, Davis 26 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Strait; Atlantic Ocean: western Europe and Gulf of St. Lawrence, and U.S. coast to North Carolina; Pacific Ocean: Alaska to California and Japan, Gulf of Iran, Arabian Sea, Iran coast and South Africa (Hilbig 1994), and Gulf of Tehuantepec (Gonzalez-Ortiz 1994). Family Onuphidae Kinberg, 1865 Kinbergonuphis microcephala (Hartman 1944; Fauchald 1982) 24, Fig. 5a. Material examined: 1 specimen: (27) 1. Characteristics: Specimen with 36 setigers. Outer lateral antennae reaching se- tiger 2, inner lateral antennae reach about setiger 7 and median antenna reaches setiger 4. Ceratophores have four rings. No eyes were observed. Branchiae are first present from setiger 6; maximum number of branchial filaments is seven; branchiae absent on the second half of the body. Ventral cirriform cirri present in the first 2 setigers; postsetal lobes digitiform in first 10 setigers, tridentate compound hooks in the first 3 setigers, large hooks are present from setiger 5 to 22. Compound spinigers absent. Subacicular hooks present from setiger 23. Each pectinate setae posses about 10 teeth. The maxillary formula is 1 + 1,9 + 8,9 Tas Oro ange ltoele Observations: Our specimens differ from K. microcephala in that subacicular hooks begin at setiger 23 not at 26 and the large hooks appear from setiger 5 to 22. No eyes were observed; outer lateral antenna reach setiger 2, inner lateral antenna reach about setiger 6 and median antenna reaches setiger 4. Habitat: In intertidal sands (Fauchald 1968). Collected in the south half of San Quintin arm, in mud and sand, D = 4; Eh = —220; T = 21; O.M. = 2.23. Distribution: Gulf of California (Fauchald 1982) and San Quintin system, Baja California. Family Oenonidae Kinberg, 1865 Arabella iricolor (Montagu 1804; Uebelacker & Johnson 1984) 42-5, Figs. 42- 2 a-f. Arabella (Arabella) iricolor. Hilbig, 1995:320-—321, Figs. 12.1 a—g. Material examined: 3 specimens: (8) 1, (13) 1, (38) 1. Characteristics: Incomplete organisms with 86 to 146 setigers, complete ani- mals with 198 to 213 setigers. Elongated, cylindrical body; iridescent, brown or light yellow. Conical prostomium, longer than wider, without antenna or palpes. No eyes. Peristomium with two rings, the same size as other setigers. Parapodia with short dorsal cirri and without acicular spines. Aciculae distally spotted. Man- dibles dark brown or black with light points. MI = 1 + 1, short, falcated with 6—7 basal teeth; MII = 10-11 + 10—12 asymmetrical, right largest; MII] to MV symmetrical, MIII = 6 + 6 rounded, bearing 5 small teeth and 1 bigger; MIV = 4+ 4; MV = 1 + 1 single, long pointed tooth arising from small base. Maxillary support long and large in the superior part, branches slightly separated. Observations: Our specimens had no eyes, organisms described by Hilbig (1995) had four subequal eyes in a strait line along the posterior margin of the prostomium. Habitat: Intertidal to 85 m; among shells, oysters, among Zostera holdfasts, bryozoans and algae. In mud, sand, muddy sand and sandy gravel (Uebelacker POLYCHAETES FROM SAN QUINTIN BAY 4 | eevones: 1984)..Infine sand; D= 22.2=-1012T = 13:52=16:,6:=934.8235:5330M. = 2.4—5.7 (Hernandez-Alcantara 1992). In muddy sand, D = 22.2—101, T = 17.3— 27.2; O.M. = 0.12—5.7 (Gonzalez-Ortiz 1994). Intertidal to 90 m. Burrows deeply into mud, sand and gravel; it is also found under rocks; in oyster and mussel beds, among bryozoans and other colonial animals (Hilbig 1995). Middle region of both arms of San Quintin system, in muddy sand, D= 1-5; Eh = —280 to —110; T= 20.4-—21; O.M. = 0.79-2.78. Distribution: France, England, Mediterranean Sea, Massachusetts to Florida, western Mexico (Fauchald 1970). Cosmopolitan in temperate and subtropical wa- ters (Uebelacker & Johnson 1984). Baja California (Reish 1963; Fauchald 1970; Rioja 1947, 1962; Salazar-Vallejo 1985; Herndndez-Alcantara 1992); Sinaloa (Rioja 1962; Fauchald 1970; Hernandez-Alcantara 1992). Gulf of Mexico, Co- lombia and Venezuela; Vancouver Island to California; Mexico, Argentina; Japan, China, Persian Gulf, Red Sea, Indian Ocean; Strait of Magellan; West and South Africa (Hilbig 1995). Family Flabelligeridae Saint-Joseph, 1894 Brada villosa (Rathke 1843; Milligan 1984) 47-13, 47-15; Figs. 47-10 a—d. Material examined: 26 specimens: (8) 3; (11) 4; (17) 15; G3) 2; (41) 2. Characteristics: Incomplete animals with up to 26—93 setigers. Body cylindrical, slightly flattened ventrally. Dorsal surface encrusted with sand grains. Papillae cirriform densely distributed. Ventral papillae similar in shape but shorter and less dense. Filiform branchiae arranged in two lateral groups. Eyes absent. All setae simple capillaries, crossbared; those from setiger 1 longer, projecting forward forming the cephalic cage which is weakly developed. Neurosetae slightly shorter and stouter than notosetae. Conical nephridial papillae present ventrally. Habitat: From shallow waters to 2000 m. Occurs in mud, gravel, sand, silty sand, clayey silt and rocks (Milligan 1984). In silty sand, D + 37.2; T = 15.1; S = 35.5; O.M. = 7.2 (Hernandez-Alcantara 1992). Occurred in both arms of the bay, in sandy mud and muddy sand, D = 1-5; Eh = —285 to +110; T = 20.4- 21.6; O.M. = 0.05-3.33. Distribution: Cosmopolitan (Milligan 1984). Northern Gulf of California (Her- nandez-Alcantara & Solis-Weiss 1993). Family Sabellidae Malmgren, 1867 Megalomma bioculatum (Ehlers 1887; Uebelacker & Johnson 1984) 54-27, 54- 30, Figs. 54-22 a-g. Material examined: 5 specimens: (13) 1; (39) 3; (41) 1. Characteristics: Incomplete organisms 25—27 setigers. Collar bilobated, dorsal edges rounded and well separated; ventral edges prolonged as two triangular lobes. Radioles numbering 6—15 pairs, with 1—2 transverse brown bands. Dorsal- most pair of radioles bearing two subterminal, large rounded eyes. Palps long, triangular, brown. Thorax with eight setigers. Thoracic notopodia with numerous slender limbate setae; thoracic neuropodia with an anterior row of companion setae and a posterior row of avicular uncini with long handles and crest of small teeth. Abdominal notopodia with avicular uncini with short handles and neuro- podia with limbate setae. 28 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Habitat: In depth of 10—200 m. In sand, silt and clay (Uebelacker 1984); 90— 200 m (Perkins 1984); sandy mud, muddy sand, gravel and sandy gravel (Ueb- elacker & Johnson 1984). Found south to Muelle viejo in San Quintin arm and near the head of False Bay, in sandy mud and muddy sand, D = 1—6; Eh = —200 to +110; T = 19.8-21.8; O.M. = 0.53—2.57. Distribution: Veracruz (Rioja 1946). Gulf of Mexico, Florida, North Carolina. Tropical western Africa (Uebelacker & Johnson 1984). Discussion Of the 677 annelids collected, 16 new records of polychaete species and 4 records of new genera for the San Quintin lagoon system were found: Leitoscol- oplos mexicanus, Scoloplos rubra, Scoletoma tetraura, Monticellina tesselata, Ex- ogone dispar, Exogone lourei, Goniada maculata, Clymenura gracilis, Brada vil- losa, Prionospio multibranchiata, Spio pettiboneae, Polydora socialis, Cirrifor- mia cf spirabrancha, Syllis (Syllis) gracilis, Lysidice ninetta, Marphysa sanguinea, Praxillela sp., Apistobranchus sp., Pionosyllis sp., Euchone sp., and one uniden- tified species from the family Trichobranchidae. This last family together with Apistobranchidae are reported for the first time from San Quintin; unfortunately the organisms were damaged and could not be identified to species level. Of the 28 polychaete families previously reported for San Quintin, 17 were found in the present study and two were added. Of the 32 identified taxa, only twelve had been previously reported in the lagoon system (Reish 1963; Calder6én- Aguilera & Jorajuria-Corbo 1986). Until now 30 families and 81 species of poly- chaetes have been recorded in the area (Table 1). The differences among the three studies at San Quintin may indicate anthro- pogenic modifications and/or environmental changes which may have affected the lagoon system over the last 30 years. Nevertheless, they may also be due to insufficient sampling effort. The first reported sampling took place in the eastern arm (Reish 1963) and only one other sampling is reported in the literature, with a reduced number of stations (Calder6én-Aguilera & Jorajuria-Corbo 1986). Con- sidering the previous study of the area it may be that the new records reported here are due to a more balanced sampling effort in both arms of the lagoon system. However, it is possible that the previous authors found more species because they used a 0.5 mm mesh which allowed them to retain young worms and smaller species. Most species presented densities of 1-10 individuals/0.02 m7’, nevertheless five species corresponded to the class of 71—100 individuals/sample (3500—5000 ind./ m7’): Scoletoma tetraura, Brada villosa, Cossura candida, Neanthes acuminata, and Pionosyllis sp., these are consistent with an extremely rich fauna. When species richness per station was plotted we found that two stations (12 and 13), located in the middle region of the eastern arm presented the highest species richness with 12 and 14 species respectively. Nine stations showed intermediate values (6 to 8 species/station), whereas the 19 remaining stations presented low species richness values going from 1 to 5 species per station. Conditions in the western arm might favor the development of the more sen- sitive species, those which cannot tolerate low oxygen concentrations, because here water is exchanged with the sea at a higher rate. In this study most of the species were found in fine-grain sediments, predominantly muddy sand and sandy POLYCHAETES FROM SAN QUINTIN BAY 29 mud, with moderately negative Eh values. Only three stations presented positive Eh values (station 6, 103 mV, station 10, 168 mV and station 23, 152 mV). Only four stations (8, 12, 37, 38) were located in the edges of Zostera beds so we can not say much about polychaete composition in this habitat. In conclusion there is still a lot of work to be done in San Quintin Bay. Not only listing the species inhabiting the area but also analyzing their structure and organization. The fact that we found 20 taxa and two families not previously reported in this area is probably due to anthropogenic modifications in the last years. Nevertheless, after examining these results it is evident that there is a need for extensive research in the coastal lagoons of Baja California in order to ac- knowledge their biodiversity and also for the local authorities to adequately man- age their resources. Acknowledgements We would like to thank Ignacio Romero, Julio Villaescusa, Victor Martinez Magana and Dr. Efrain Gutiérrez for their help during field work and Rosa Ji- ménez for sorting the macrofauna. We are grateful to M. S. Pablo Hernandez- Alcantara, M. S. 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A quantitative study of the benthic polychaetous annelids of Bahia de San Quintin, Baja California. Pac. Nat., 3(14):399—436. . 1968. A biological survey of Bahia de Los Angeles, Gulf of California, México. II. Benthic polychaetous annelids. Trans. San Diego Soc. Nat. Hist., 15:67—106. Rioja, E. 1946. Estudios Anelidol6gicos. XIV. Observaciones sobre algunos poliquetos de las costas del Golfo de México. An. Inst. Biol. UNAM., 17:193—203. 32 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES . 1947. Estudios Anelidol6gicos XVII. Contribucion al conocimiento de los anélidos polique- tos de Baja California y Mar de Cortés. An. Inst. Biol. UNAM. 15:139-145. . 1962. Estudios Anelidol6gicos XX VI. Algunos anélidos poliquetos de las costas del Pacifico de México. An. Inst. Biol. UNAM. 33:131—229. Rodriguez- Villanueva, L. V. 1993. Los poliquetos (Annelida: Polychaeta) de la plataforma continental interna del sur del Golfo de México: abundancia, distribuci6n y diversidad. Tesis Prof. Fac. Ciencias. U.N.A.M., 128 p. Rosales-Casian, J. 1996. Ichtyofauna of Bahia San Quintin, Baja California, Mexico, and its adjacent coast. Ciencias Marinas 22:443—458. Salazar-Vallejo, S. 1981. Coleccién de Poliquetos (Annelida: Polychaeta, de la Universidad de Nuevo Leén, México. Tesis Profesional. Fae. Ciencias Biologicas, UANL. 156 p. Salazar-Vallejo, S. I., 1985. Contribuci6n al conocimiento de los poliquetos (Annelida: Polychaeta) de Bahia Concepcion, Baja California Sur, México. Tesis Maestria, CICESE, Ensenada. México, 311 p. Silva-Cota, S., and S. Alvarez-Borrego. 1988. Rhe “El Nino” effect on the phytoplancton of a north— western Baja California Coastal Lagoon. Est. Coastal and Shelf Sci., 27:109-115. Stoner, A. W., and R. J. Livingston. 1980. Distributional ecology and food habits of the banded blenny Paraclinus fasciatus (Clinidae): a resident in a mobile habitat. Mar. Biol., 56:239-246. Taylor, J. L., 1984. Chapter 1. Orbiniidae., Chapter 31. Nereidae., Chapter 35. Nephtyidae. In: Ueb- elacker, J. M. and P. G. Johnson (eds.). Taxonomic Guide to the Polychaetes of the Northern. Gulf of Mexico. Final Report to the Minerals Management Service, contract 14-12-001-29091. Barry A. Vittor and Associates. Inc., Mobile, Alabama. 7 vols. Uebelacker, J. M. 1984. Chapter 10. Heterospionidae; Chapter 17. Opheliidae; Chapter 24. Eulepeth- idae; Chapter 28. Hesionidae; Chapter 30 Syllidae; Chapter 41. Lumbrineridae; Chapter 42. Arabellidae; Chapter 51. Ampharetidae; Chapter 54. Sabellidae. In: Uebelacker, J. M. and P. G. Johnson (Eds.). Taxonomic Guide to the polychaetes of the Northern Gulf of México. Final Report to the Minerals Management Service, contract 14-12-001-29091. Barry A. Vittor and Associates. Inc., Mobile, Alabama. Uebelacker, M., and P. G. Johnson. 1984. Taxonomic Guide to the Polychaetes of the Northern Gulf of México. Vol. I to VII. Final Report to the Minerals Management Service, contract 14-12- 001-29091. Barry A. Vittor and Associates. Inc., Mobile, Alabama. 7 vols. Van Der Heiden, A. M., and M. E. Hendrickx. 1982. Inventario de la fauna marina y costera del sur de Sinaloa, México. Segundo Informe. Inst. Cienc. Mar. Limnol., UNAM. 135 p. Westheide, W. 1974. Interstitielle Fauna von Galapagos, XI. Pisionidae, Hesionidae, Pilargidae, Syl- lidae (Polychaeta). Mikrofauna Meeresb., 44:1—146. Accepted for publication 19 March 1997. Bull. Southern California Acad. Sci. 97(1), 1998, pp. 33-38 © Southern California Academy of Sciences, 1998 Cetaceans of Isla De Guadalupe, Baja California, Mexico Juan-Pablo Gallo-Reynoso' and Ana-Luisa Figueroa-Carranza? 'Centro de Investigacion en Alimentacion y Desarrollo, Unidad Guaymas. Apdo. Postal 284, Guaymas, Sonora 85480, México *Reserva Islas del Golfo de California, Instituto Nacional de Ecologia, SEMARNAP. Miramar 210, Col. Miramar, Guaymas, Sonora 85450, México Isla de Guadalupe (29°00'N, 118°15’30"W) is located 240 km west of Baja California, México. The island lies within the California Current, with an average sea surface temperature of 18°C (range: 16°C spring, 20°C summer; Lynn and Simpson 1987). Northwesterly winds predominate (Berdegué 1957). The orien- tation of the island and its elongated shape (35 km long and 6.5—9.5 km wide) acts as a barrier against the flow of the current which produces a series of swirls at different depths. The island is surrounded by depths of 3600 m or more. The island does not have a shelf around it with exception of the southern tip were a 4 km wide and 200 m deep shelf is found between Isla Guadalupe, Isla Zapato and Isla Toro. The coastline and nearshore physiography is composed of loose basaltic rocks and boulders bounded by towering cliffs (Pierson 1987). The only report of cetaceans near Isla de Guadalupe is by Fleischer (1978), who reported two minke whales (Balaenoptera acutorostrata) 500 m offshore and bottlenose dolphins (Tursiops sp.) on several occasions. Other reports on marine mammals in the area exist (i.e. Mangels and Gerrodette 1994), but they are far offshore from the island. Our study was carried out while studying Guadalupe fur seals, Arctocephalus townsendi, during 1991-1993. We made observations during winter (February 1991-1992), spring (June 1983 and 1991), summer (July—August 1991, 1992 and 1993) and fall (November—December 1991 and 1992), during a total of 189 days. Nearly all of our survey time (179 days) was on the east side of the island. Observations of cetaceans were conducted: a) from the camp, located at ‘‘Cor- ralitos’’, on the southeastern coast of the island (28°53’30"N), 30 m above sea level. Daily observations started at 0600 hr and ended at 2000 h; b) from the catwalk of Mexican Navy Coastguard boats (approx. 10 m above sea level) during the approach to or while leaving the island, and c) while conducting censuses of Guadalupe fur seals in the eastern side of the island (except in summer censuses, when the entire island was covered). These censuses were conducted in a 5—7 m fiberglass skiff with an observation height of =2 m, by two to four observers, along the shoreline at idle speed (2—3 knots). Observer bias was relatively consistent because we collected the data on all census days. We estimated group size, and recorded ventilation times and general behavior of the cetaceans sighted. Results are given in averages + standard de- viation (S.D.). Eleven species of odontocetes and two species of mysticetes were observed. 34 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1. Relative abundance of cetaceans of Isla de Guadalupe in 1991-1993, expressed as number sighted per hour of effort (n = 101 observations). Total number Spring Summer Fall Winter (% of sightings) Bottlenose dolphin 7 62 2, 2 T2286 Tursiops truncatus Baird’s beaked whale l 4 2 l SZ. Berardius bairdii Unidentified Bottlenose whale _ 5 — — 4.95 Cuvier’s beaked whale 1 1 —_ a 1.98 Ziphius cavirostris Unidentified beaked whale —- ] = — 0.99 Sperm whale — 2 — — L.9sS Physeter catodon Dwarf sperm whale —- ] — — 0.99 Kogia spp. ; Common dolphin ] -- — l 1.98 Delphinus delphis Risso’s dolphin = = — ] 0.99 Grampus griseus Short-finned Pilot whale -= — 1 | 1.98 Globicephala macrorhynchus Killer whales 1 | — —- 1.98 Orcinus orca Blue whale = = a l 0.99 Balaenoptera musculus Fin whale oe a a 1 0.99 Balaenoptera physalus Total number of cetaceans 364 1241 84 76 1765 Hours of effort 180 840 240 160 1420 Number cetaceans/hour 2.02 |e) 0.4 0.5 1.24 Bottlenose dolphin, Tursiops truncatus This was the most frequently observed cetacean year-round (Table 1). Group sizes were recorded almost daily, averaging 19 + 12 individuals (range: 1—50 individuals, n = 73 schools). Bottlenose dolphins moved daily early in the morn- ing (0600—0900 h), at a distance between 500 m and 3 km, parallel to the east coast heading north and returned moving south inshore in the evening (from 1600—2000 h). It was common to observe a very spread formation of dolphins moving slowly, apparently searching for prey. Upon finding a school of fish, aerial displays started, which congregated the dispersed dolphins and started the herding and encircling of fish schools. Immediately after this the dolphins chased the fish into the shallows and started feeding on them. The same behavior has been re- ported for T. truncatus from the Gulf of California (Gallo-Reynoso 1989). This behavior was observed when they were feeding on skipjack tuna (Katsuwonus pelamis) and yellowtail (Seriola lalandei). On other occasions, with the same behavior, they fed on chub mackerel (Scomber japonicus), Pacific golden-eyed tilefish (Caulolatilus affinis), red snapper (Lutjanus peru), and flying fish (Cyp- selurus sp.). On 18 February 1991, at 11:30, while traveling along the east coast of Isla de Guadalupe, we observed a group of.10 bottlenose dolphins that turned toward us, crossed our bow, moving rapidly between the four foot waves. The CETACEANS OF ISLA DE GUADALUPE 35 dolphins appeared to be fleeing from a 4.0—5.0 m great white shark (Carcharodon carcharias) that was 30 m away and moving toward them. Interactions between bottlenose dolphins and great whites have been reported by Connor and Heithaus (1996). The dolphins escaped to the south moving fast and porposing out of the water. Baird’s beaked whale, Berardius bairdii This was the second most observed odontocete in the study area. They were observed year-round in groups averaging 4 + 0.9, individuals (range: 2-5, n = 8). These whales were identified by their size (9-13 m), their elongated, cylin- drical beak and prominent melon. Calves (~5 m in length) were observed on three occasions, two in June and july 1991 and one in November 1992. A juvenile (~7 m in length) was observed in July 1991. These whales were on average sighted 3.2 + 1.5 km offshore (range: 1—5.5 km). Dive times for these whales averaged 26.5 + 8.5 min (range: 18-35, n = 8 dives). Twice on 8 July 1991, when the whales sounded, yellowfin tuna (Thunnus albacares) started to jump out of the water moving away from the whales. Unidentified bottlenose whale There were five observations of a large unidentified ziphid, similar to a bottle- nose whale. They were observed only in the summers of 1992 and 1993. Gallo- Reynoso and Figueroa-Carranza (1995) described them as Hyperoodon ampul- latus, but there is no authenticated record of this species in the eastern Pacific. In three occasions individuals of this species were breaching or partially breaching which facilitated their identification, showing a whitish head, buff colored body, squarish melon with short but well-defined beak, and falcate dorsal fin located two-thirds of body length all characteristics of H. ampullatus (Leatherwood and Reeves 1983). Size of breaching individuals was estimated at 7 m. Dive times averaged 17.6 + 6.1 min (range: 10—25, n = 3 dives). These whales were diving over waters 810 + 175 m (range: 600—1000, n = 5). During one observation, a partially breaching whale was accompanied by a school of 25 bottlenose dolphins, four of which were breaching with the whale for 10 min. Cuvier’s beaked whale, Ziphius cavirostris They were observed on two occasions. These whales were identified by their brown coloration, white head with distinctive shape without a “‘beak’’, with many white scar lines on the head and dorsum, and a triangular dorsal fin, situated far in the back. A group of 7 individuals were observed in June 1991 at 300 m off the coast. The largest individual was compared to the skiff (7 m) and estimated to be 7.5 m. These whales were diving repeatedly in the same spot during four hours of observation. Diving times averaged 21 + 10.2 min (range: 15-27, n = 16 divings). The group was accompanied by a school of 12 bottlenose dolphins which were also diving. The second observation (July of 1991), consisted of 5 individuals, 2 km offshore, slowly moving to the north while diving. The areas where these whales were diving averaged 565 + 49.5 m in depth (range: 535-— 600, n = 2). 36 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Unidentified beaked whale Was observed in one occasion from a distance of 100 m. It had a non bulbous, whitish head, with a distinctive beak (shorter than in young B. bairdii). The dorsum was dark grayish-brown, with a few long white scars, and a broad falcate dorsal fin situated far in the back (Leatherwood et al. 1988). The size of the animal was about 5 m. Sperm whale, Physeter catodon Two observations on 28 July, 1993, 6.5 h apart (1233 and 1915 h) were possibly the same individual. The sperm whale was at an average distance of 4.4 + 0.4 km (range: 4—4.8, n = 2), over waters more than 1000 m in depth. In the two observations the whale was moving to the north. Dwarf or Pygmy sperm whale, Kogia spp A group of 12 individuals were observed from the camp in August 8, 1993 at 1816 h, at about 3 km offshore. They were recognized by its small size, the lack of a conspicuous beak and their small but falcated dorsal fin found in the mid- dorsum. Dive times averaged 40 + 15.7 min (range: 45—28, n = 4 divings). The group was slowly moving north against the current. Common dolphin, Delphinus delphis This species was recognized by its small size, the presence of a short beak and their characteristic clear coloration, compared to D. capensis. A large school of about 250 individuals was observed in June 1983, moving fast to the south. A second group, in February 1991, consisted of 15 individuals. Risso’s dolphin, Grampus griseus A group of 12 individuals (including two calves) were observed in 6 February, 1992 at 12 km northeast of the island. The group surfaced from the depth and the younger individuals (judging by the size and darker coloration) rode the bow wave of the ship for two minutes, then returned to the group. Water depth was 2980 m. Short-finned pilot whale, Globicephala macrorhynchus This species was observed on two occasions, a group of six whales in February 1991, and a group of eight in November 1992, heading south. Killer whale, Orcinus orca Fishermen reported 10 whales (including a calf) hunting elephant seals at Playa Elefante (elephant seal beach) in June 1983. In August 1993, at 21:30 we regis- tered the possible presence of this species as we heard from the camp at night that the whales were slapping their flukes vigorously very close to shore, their spouting was very strong. The fur seals were unusually silent, and a young ele- phant seal, shaking violently was hauling out in rocks where they don’t normally climb. The slapping continued for several minutes, coming from at least four different locations. The slapping stopped when we searched with a light to identify the animals. Apparently the whales left after this and we did not hear them again. Although we never saw killer whales, the sound of their spouts and flukes slapping CETACEANS OF ISLA DE GUADALUPE 37 in the surface and the behavior of the other marine mammals suggest their pres- ence. Blue whale, Balaenoptera musculus We observed three individuals feeding close to Punta Sur (3 km), by doing sideways movements with their mouths open. They were slowly moving to the southwest at 1614 h on February 17, 1992. Fin whale, Balaenoptera physalus A large individual was observed 500 m offshore in February 1992. The whale was lunge-feeding and moving to the south. The cetacean fauna at Isla de Guadalupe is dominated by squid eaters (61.5%, eight species): four species of beaked whales (Ziphiidae), two larger dolphins (Delphinidae: Globicephalinae), the sperm whale (Physeteridae) and a species of pygmy sperm whale (Kogiidae). There were also two fish and squid eating dol- phins (Delphinidae) with 15.4%, and two zooplankton filtering whales (Balaen- opteridae) with the 15.4%. One observed species has omnivorous habits (Del- phinidae: Globicephalinae), with the 7.7%. The composition of this cetacean fauna was somewhat different during the El Nino event of 1992, with a higher sea temperature and sea level (Fahrbach et al. 1991). Sea surface temperatures we measured during summer was 5°C higher in 1992, and 3°C higher in 1993 than the mean of 20°C during 1991 (Gallo-Reynoso 1994). Probably the absence of Z. cavirostris and the presence of an unknown bottlenosed beaked whale species, that was observed only in the summers of 1992 El Nino, and 1993 a post-E] Nino year were related to the shift of squid species found during El Nino of 1992, in the diet of the Guadalupe fur seals. A greater proportion of southern squid species was found in the scats of these fur seals than in the previous year (Gallo-Reynoso 1994). é The few observations of blue and fin whales and the absence of other species of mysticetes may be related to the fact that the waters surrounding Isla de Gua- dalupe have a moderate primary production of ~36—84 gC/m’yr according to Koblents-Mishke and coworkers (in Berger 1989). Acknowledgments Direcci6n General de Intercambio Académico, Universidad Nacional Autonoma de México (UNAM), The University of California Education Abroad Program, and The National Geographic Society funded this research during 1991. The In- stitute of Marine Sciences, University of California, Santa Cruz, funded during part of 1992. UC-Mexus funded during fall 1992 and summer of 1993. The Di- reccién General de Asuntos del Personal Académico, UNAM, funded in part from 1991-1993. We thank A. Delgado-Estrella, A. Sanchez and M. Peralta for their help in the field. We thank Secretaria de Marina, and fishermen of ‘“Cooperativa de langosteros y abuloneros de Ensenada” who provided transportation and lo- gistic support. These observations were conducted under permits No. 0561 of Secretaria de Pesca, and Nos. 2538, 4933 and 2025 of Secretaria de Desarrollo Urbano y Ecologia (SEMARNAP), México. Finally we thank an anonymous re- viewer that improved our work. 38 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Literature Cited Berdegué, J. 1957. La Isla de Guadalupe, México. Contribucién al conocimiento de sus recursos naturales renovables. Secretaria de Marina, Direcci6n General de Pesca e Industrias Conexas, México. 67 pp. Berger, W. H. 1989. Global maps of ocean productivity. Pages 429-455 in Productivity of the Ocean: present and past. (W. H. Berger, V. S. Smetacek and G. Wefer, eds.). John Wiley and Sons Limited. Connor, R. C., and M. R. Heithaus. 1996. Approach by great white shark elicits flight response in bottlenose dolphins. Marine Mammal Science., 12(4):602—60S. Fahrbach, E., F Trillmich, and W. Arntz. 1991. The time sequence and magnitude of physical effects of El Nifio in the Eastern Pacific. in Pinnipeds and El Nifio, responses to environmental stress (F Trillmich and K. Ono, eds.). Springer-Verlag. pp. 8—21. Fleischer, L. 1978. The distribution, abundance and population characteristics of the Guadalupe fur seal, Arctocephalus townsendi (Merriam, 1897). Master in Science Thesis. University of Wash- ington, Seattle. 93 pp. Gallo-Reynoso, J. P. 1989. El bonito (Euthynnus lineatus) (Scombridae) como alimento para toninas (Tursiops truncatus gillii) (Delphinidae) y por lobo marino (Zalophus californianus) (Otariidae). Anales Instituto de Biologia. UNAM. Serie Zoologia., 60(1):125—127. . 1994. Factors affecting the population status of Guadalupe fur seal, Arctocephalus townsendi (Merriam, 1897), at Isla de Guadalupe, Baja California, México. Ph.D. dissert., University of California, Santa Cruz. 199 p. and A. L. Figueroa-Carranza. 1995. Occurrence of bottlenose whales in the waters of Isla Guadalupe, Mexico. Marine Mammal Science., | 1(4):573—575. Leatherwood, S., and R. R. Reeves. 1983. The Sierra Club handbook of whales and dolphins. Sierra Club Books, San Francisco, California. , R. R. Reeves, W. E Perrin, and W. E. Evans. 1988. Ballenas, delfines y marsopas del Pacifico Nororiental y de las aguas articas adyacentes. Comision Interamericana del Atun Tropical. La Jolla, California. Lynn, R. J., and J. J. Simpson. 1987. The California Current system: the seasonal variability of its physical characteristics. J. Geophys. Res., 92(c12):12,947—12,966. Mangels, K. F, and T. Gerrodette. 1994. Report of cetacean sightings during a marine mammal survey in the Eastern Pacific Ocean and the Gulf of California aboard the NOAA ships McArthur and David Starr Jordan July 28—November 6, 1993. U.S. Dept. of Commerce. NOAA-TM-NMEFS- SWFSC-211. 86 p. Pierson, M. D. 1987. Breeding behavior of Guadalupe Fur Seal, Arctocephalus townsendi. in Status, Biology, and Ecology of Fur Seals (J. P. Croxall and R. L. Gentry, eds.) NOAA TECN. Rep. NMES. S1:83—94. Accepted for publication 27 September 1997. Bull. Southern California Acad. Sci. 97(1), 1998, pp. 39-48 © Southern California Academy of Sciences, 1998 A General Allometric Model for Blade Production in Aostera marina L. Elena Solana-Arellano, Dora J. Borbon-Gonzalez, and Hector Echavarria-Heras Centro de Investigacion Cientifica Y Educaci6n Superior de Ensenada, Baja California, México Mail Address: P.O. Box 434844, San Diego, California 92143-4844 U.S.A email: hechavar@ CICESE.mx Abstract.—We introduce a generalized allometric model to express leaf dry weight in terms of leaf width and size in Zostera marina L. A formal justification of the derived model is presented. For the statistical validation we used data collected on two well defined strata over a period of one year. A comparison of the results using an independent data set was also performed. Applications of the model to estimate average leaf production are illustrated as well. The seagrass ecosystem such as Thalassia testudinum Baks ex KOnig and Zos- tera marina L. have an important role in shallow tropical and temperate waters. These kinds of ecosystems are among the most productive marine systems. Con- sequently many attempts to predict their response to changes in the environment have been made. Most of the methods used to study marine phanerogams are expensive, time consuming, and require destructive techniques such as leaf mark- ing (Sand-Jensen 1975; Jacobs 1979). Moreover it has been shown that excessive manipulation of raw material increases the error introduced in the data (Mandel 1964). Other techniques to measure seagrass production involve leaf cropping (McRoy and McMillan 1977) but such methods influence leaf growth and phys- iology (Hamburg and Homann 1986). In this paper we introduce a general allo- metric model which simplifies the estimation of blade production for Zostera marina. Our results could be applied to other marine phanerogams in a straight- forward way. The necessity to predict biomass and production for Zostera marina has stim- ulated the development of allometric models in productivity estimations. Patriquin (1973) used an allometric equation to predict leaf weight from leaf width for Thalassia testudinum. Jacobs (1979) gives a relationship between shoot length and density; McRoy (1970) relates leaf length and dry weight. Duarte (1991) presents an allometric study for marine phanerogams which considers the rela- tionship between the sizes of different parts of a plant. Nevertheless, he does not include the variation of leaf dry weight in terms of length and width. Our allo- metric model permits the reduction of the problem of blade production estimation to the measuring of linear variates, particularly leaf length and width. For the calibration of the model presented here we used data collected in two well-defined strata over a period of one year. We obtained a remarkable consistency of the model with the measurements. An alternate validation used independent data sets. 59 40 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES SAMPLE SITE 196m x 30m. —-— TRANSVERSAL TRANSECT —— COAST LINE APROXIMATION [_] STRATUM I 2 02 STRATUM II f R, “zy a ‘y Z) G UZ %Y Ryu % Fig. 1. Location of the study site. Two strata were sampled, one at —0.8 to —0.2 below (MLLW) (stratum I) and the other at —0.2 to O m below (MLLW) (stratum II). (Solana et al. 1991). We also show the usefulness of the general allometric model to estimate the time variation of average leaf dry weight. This is obtained in terms of the allometric parameters and the leaf length and width averages. An example of the referred estimation is illustrated. It uses the allometric parameters obtained from our data and measurements of length and width obtained independently by Ibarra-Obando (1992). ALLOMETRIC MODEL FOR ZOSTERA BLADE PRODUCTION 4] Field, Laboratory, and Statistical Methods The study was carried out in an area located in San Quintin Bay, Mexico, a coastal lagoon located in the Pacific Ocean waters of Baja California between 30°24’ and 30°30'N latitude and between 115°56’ and 116°01'W longitude. Ibarra- Obando and Huerta-Tamayo (1987) give a complete description of San Quintin Bay. The sampled site is a rectangular area 200 m by 30 m located in the arm of the bay (see Fig. 1). Ibarra-Obando and Huerta-Tamayo (1987) showed that there exist significant differences in the dynamics of Zostera marina L. between the intertidal zone and the transition intertidal zone. Hence we used a topographical study to determine two well-defined strata. Stratum I was from —0.2 m to —0.8 m below Mean Lower Low Water (MLLW) and stratum II from 0 m to —0.2 m below MLLW. From November 1992 through November 1993 we sampled monthly eight quadrants of 20 cm by 20 cm in each stratum following a stratified random sampling (Wonnacott & Wonnacott 1984). We collected a total of 10,000 complete leaves from stratum I and 9000 from stratum II. Each sample was placed in individually labeled plastic bags and refrigerated until processed in the laboratory. To produce our data each shoot was cleaned with distilled water. Measurements of length and width were taken for each leaf, and then its dry weight was deter- mined. All the basic statistics and the fit of the model by means of non linear regression was obtained using the STATISTICA package (STATISTICA 1993). Finally we applied a goodness of fit test to corroborate that for both strata the regression equation used was consistent with the data with a probability of 0.95. Theoretical Methods The model presented here generalizes the equation introduced by Hamburg and Homann (1986). This relates leaf dry weight simultaneously with leaf size and width. In their formulation these authors considered that leaf dry weight depended linearly on leaf length and non linearly on width. Besides this restrictive assump- tion they did not provide a formal justification to their model. Our theoretical exploration shows that when length and width determine simultaneously the vari- ation of leaf dry weight we must necessarily consider non linear dependencies in both variables. At time tf let w(x,t) (measured in mg) be the weight and /(x,f), h(x,t) (both measured in mm) respectively the length and the width of a typical Zostera marina L. leaf collected at a position x of the two dimensional sampling space. Consider also that the measurement of the width is taken at the middle of the longitudinal size of each leaf. According with Batschelet (1975) we have the allometric rela- tionships: L souer (ek el 5 9 SS SS gp = CD) w or [ot 1 oh ko al a @) hor bot Where k, and k, are positive constants. The allometric laws (1) and (2) permit us to obtain the direct dependency of 42 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES dry weight with respect to leaf length or width. The simultaneous dependency on both variables can be formally justified. To achieve this goal, we notice that according to the chain rule (Fulks 1978) we have; dw OW ol ow oh amma! ay an are @) Combining equations (1), (2) and (3) we obtain, k\w ol _ ow ohh Sly tel al ky “tat al ot oh ar Simplifying we can establish the partial differential equation; > te lah = kw. (4) In the appendix we have shown that a solution to equation (4) which satisfies the allometric relations (1) and (2) becomes, w(x,t) = Kh(x,t)*1(x,t)® (5) where k, a and B are constants that can be obtained from the data. The allometric model given by equation (5) can be used to estimate the dy- namics for average leaf production using measurements of leaf size and width. Let us consider a set of leave samples. Suppose that the mth sample contains N,,, complete leaves collected at a time ¢. Then for the ith leaf in that sample equation (5) gives W seas t) a KIM OG) Up Xyisk)” (6) where W,,(Xnist)s Am(Xmist), and L,,(x,,;¢) are respectively the dry weight, the width and length of the considered leaf. In our notation, the sub-index mi identifies a leaf collected at a position x,,,; and belonging to the mth sample for 1 = 7 S N,,,. Taking logarithms in (6) and averaging we have mi? 1 Nin OL Nin B Nin N. > (in Wmi(Xmis t)) = In K aE Na > In Has Mie t) air N,. > In L ni(Xmis t) (7) m t=) m t= The left hand side of equation (7) gives the natural logarithm of the geometric mean of the set of N,, dry weight values in the sample. This equation also permits the exploration of the validity of equation (5) for different data sets. It is worth to point out that the variability observed in the measurements makes the geometric mean of dry weight a better estimator of central tendency for leaf dry weight. On the other hand the use of logarithms in equation (7) will avoid the vanishing effect associated to the large product of small numbers required to obtain the referred mean. Results Using the collected data, we tested the model given by equation (5). We ob- tained very similar results for both strata (see Table 1). No spatial dependencies for the allometric parameters x, a and B along the strata where detected. For stratum I and stratum II the coefficients of determination were 0.91 and 0.87 ALLOMETRIC MODEL FOR ZOSTERA BLADE PRODUCTION 43 Table 1. Estimations for both strata of the fitted parameters and their respective standard errors. Stratum I Stratum II Std. error Stratum I Std. error Stratum II K 0.000005 0.000015 0.04(p < 0) 0.03 (p < 0.0) a 1.09 3 0.01 (p < 0.05) 0.01 (p < 0.0) B 0.469 0.45 0.04 (p < 0.05) 0.03 (p < 0.0) respectively. The standard error for the estimate was 0.43 for stratum I and 0.55 for stratum II. For both cases the parameters found in the regression have very small standard errors and very significant p-values (see Table 1). The residual analysis for both strata gave no indication of lack of fit of the model. Plots of the predicted and observed values of leaf dry weight are presented for each stratum to provide a visual interpretation of the adequacy of the fit (see Fig. 2). Finally a lack of fit F-test was applied to data in both strata. We obtained significant evidence that the expected value of the dependent variable w is indeed represented by means of the allometric equation (5) with a probability of 0.95. Using data collected on a preliminary sampling, Solana et al. (1991) found similar values for the parameters xk, a and B. This was corroborated using a Student-t test at a 95% confidence level. As a conclusion equation (5) gives the correct allometric relationship for the involved variables in our study site. An application of the present model was performed. Using the values of the parameters x, a and 6 corresponding to our data, we simulated the behavior of the average mean dry weight as given by equation (7). For that purpose we used an independent data set for leaf length and width collected by Ibarra-Obando (1992) in the same MLLW level of stratum I. Fig. (3) shows the temporal variation of the average dry weight obtained directly from our data and the predicted by quation (7) for the Ibarra-Obando (op. cit.) data set. Using analysis of variance and a LSD (Least significant difference) test, we found no significant differences between both means. This shows that the sample paths shown in Fig. (3) are both generated by the same stochastic process. This also implies the consistency of the considered allometric relationship. Discussion The characterization of productivity for the eelgrass Zostera marina both under laboratory and in situ conditions has been relevant in the study of the ecology of the seagrass ecosystem. The advantages of allometric relationships in these studies are pointed out by several authors e.g., (Jacobs 1979, McRoy 1970, Duarte 1991, Patriquin 1973, Hamburg and Homann 1986). An allometric description of dry weight in terms of direct in situ measurements could avoid tedious laboratory processing and destructive sampling. The model we tested here permits the ex- pression of leaf dry weight in terms of leaf length and width through a simple equation whose parameters are easily obtained using available nonlinear regres- sion methods. Although a form close to the proposed model was introduced by Hamburg and Homann (1986) no formal justification was provided. Furthermore the model proposed by these authors considers linear dependence of dry weight with respect to leaf length. That model fails to provide the generality of the allometric equations (1) and (2). On the other hand, as it is shown in the appendix, 44 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES stratum | 0.16 0.14 0.12 o = Ad = BS 2 0.08 >sG o> Hn & 8&5 0.06 0.04 0.02 0 ~s. Regression 0 0.02 0.04 0.06 0.08 0.1 0.12 0.14 95% confid. Predicted Values (Leaf dry weight) stratum II Observed Values (Leaf dry weight) “ea. Regression 0.14 95% confid. 0 0.02 0.04 0.06 0.08 0.1 OraZ Predicted Values (Leaf dry weight) Fig. 2. Predicted vs. observed values of leaf dry weight predicted by equation (5) in terms of leaf length and width. a) Stratum I. b) Stratum II. To produce these plots we selected a random sample of 1000 leaves. the model given by equation (5) can be justified using the results in the theory of first order partial differential equations. We conclude that an allometric relation in the form proposed here must necessarily be considered in order to express leaf dry weight in terms of length and width measurements. The model that we present here provides a simple procedure to identify the ALLOMETRIC MODEL FOR ZOSTERA BLADE PRODUCTION 45 0.014 0.012 0.01 0.008 0.006 0.004 natural logarithm of the geometric mean of leaf dry weight 0.002 TIME (months) Fig. 3. Variation over time natural logarithm of the geometric mean of leaf dry weights. The continuous line corresponds to our data. The dashed line represents the simulation of the mean leaf dry weight predicted by equation (7) using data on leaf length and width obtained by Ibarra-Obando (1992) and the allometric scaling parameters k, a and B found in the present study. allometric scaling factors for the considered variables. The high determination coefficient obtained for the data sets analyzed in this study corroborate the claim that our model is consistent with these observations. Our model was also tested against independent data sets. In a preliminary application of our model using only 21 shoots in a single sample, Solana et al. (1991) found similar values for the allometric parameters. A Student-t test gave no indication of statistical dif- ferences. Hence we can expect that the values obtained for the allometric param- eters k, a and B will not depend on the particular data set considered. As a conclusion the model given by equation (5) identifies unambiguously the allo- metric linkage which relates the variation of the leaf dry weight as a function of leaf length and width for Zostera marina L. in our study site. When we used the values of parameters xk, a and £6 found with our data to obtain by means of equation (7) the dry weight corresponding to data on length and width (Ibarra- Obando 1992) we observed some deviations with respect to the corresponding values measured in our study. As we have pointed out, these characterizations of the temporal variation for the leaf mean dry weight portrayed in Fig. (3) corre- spond to sample paths of the underlying stochastic process. Nevertheless these deviations could be explained by the occurrence of a strong “El Nino Southern Oscillation’? (ENSO) event which took place from June 1986 to January 1988; that is just before and during the year where the data were taken (Climate Di- agnostic Bulletin 1996). Solana et al. (1996) found that temperature is the most significant variable in the determination of leaf size dynamics in our study site. In the same study, dissolved nutrients were also found to be a determinant. The ENSO event was reported to induce an increase of water temperature and a re- 46 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES duction of dissolved nutrients in San Quintin Bay (Silva-Cota and Alvarez-Bor- rego 1988). By virtue of the tolerance law, the rise in water temperature could have produced sub-optimal leaf growth rates. High temperature stress has been claimed to induce a deleterious effect for Zostera marina (Rasmusen 1977). It has also been reported that a sizeable increment in temperature reduced shoot abundance and a change in the seasonal pattern of abundance of Z. marina in Chesapeake Bay (Penhale 1977; Wetzel and Penhale 1983; Evans et al. 1986). In our study site the reduction of nutrients could have also contributed to the pre- dicted reduction of average leaf dry weight. The conjunction of high temperature- stress and a reduction in dissolved nutrient availability could have created a de- layed response characterized by a smaller leaf production which was observable in summer of 1987. Given the values of the allometric parameters equation (7) becomes an adequate tool to estimate leaf production in Zostera marina L. (see Fig. 3). We claim that our model is a general tool which could be straightforwardly applied in other studies. This will simplify biomass estimations, eliminating tedious laboratory processing while avoiding destructive sampling. Acknowledgments We thank Fernando Escoto and Hector Atilano-Silva for their fine laboratory work. This paper is part of the research on eelgrass production partially funded by CONACYYT, by grant 0263-N9107. Roberto Iglesias-Prieto provided valuable suggestions and references. His orientation is greatly appreciated. Literature Cited Batschelet, E. 1975. Pp. 362—364. in Introduction to mathematics for life scientist. Springer Verlag. Duarte, C. M. 1991. Allometric scaling of seagrass form and productivity. Mar. Ecol. Prog. Ser., 77: 289-300. Evans, A. S., K. L. Webb, and P. A. Penhale. 1886. Photosynthetic temperature acclimation in two coexisting seagrasses, Zostera marina L. and Ruppia maritima. L. Aquat. Bot., 24:185—197. Fulks, W. 1978. Advanced Calculus. John Wiley and Sons Inc. New York. 500 pp. Hamburg, S. P., and P. S. Homann. 1986. Utilization of growth parameters of eelgrass, Zostera marina, for productivity estimation under laboratory and in situ conditions. Mar. Biol., 93:299-—303. Ibarra-Obando, S. E., and R. Huerta-Tamayo. 1987. Blade production of Zostera marina L. During the Summer-Autumn on the Pacific coast of Mexico. Aquat. Bot., 28:301-—315. 1992. Contribution a la connaissance de lherbier a Zostera marina en Bajja California (Mexique): Biologie et Production Primaire. Thése de Doctorat, Université d’ Axi-Marseille I, France, 286 pp. Jacobs, R. P. W. M. 1979. Distribution and aspects of the production and biomass of eelgrass, Zostera marina L., at Roscoff, France. Aquat. Bot., 7:151—172. Mandel J. 1964. The Statistical Analysis of Experimental Data. Dover Publications, Inc. New York. 410 pp. McRoy, C. P. 1970. Standing stock and other features of eelgrass (Zostera marina) populations on the coast of Alaska. J. Fish., 27:1811—1812. , and C. McMillan. 1977. Production, ecology and physiology of seagrasses. pp. 53-87 in Seagrass ecosystems as scientific perspective. (C. P. McRoy and C. Helfferich, eds.), Marcel Dekker New York. Patriquin, D. G. 1973. Estimation of growth rate, production and age of the marine angiosperm, Thalassia testudinum. Koenig Carib. J. Sci., 13:111—123. Penhale, P. A. 1977. Macrophyte-epiphyte biomass and productivity in an eelgrass (Zostera marina L.) community. J. Exp. Mar. Biol. Ecol., Vol. 26, pp. 211-224. Rasmussen, E. 1977. The wasting disease of eelgrass (Zostera marina) and its effects on environ- ALLOMETRIC MODEL FOR ZOSTERA BLADE PRODUCTION 47 mental factors and fauna. pp. 1-51 in Seagrass ecosystems, a scientific perspective, (C. P. McRoy & C. Helffrich, eds.), Marcel Dekker, New York. Sand-Jensen, K. 1975. Biomass net production and growth dynamics in an eelgrass (Zostera marina) population in Vellerupo Vig. Denmark. Ophelia, 14:185—201. Silva-Cota S., and S. Alvarez-Borrego. 1988. The ‘‘El nino” effect on the phytoplankton of a North- western Baja California coastal lagoon. Estuarine, Coastal and Shelf Science, 27:109—115. Solana-Arellano, M. E., S. E. Ibarra-Obando, and H. Echavarria-Heras. 1991. Calibraci6n de un modelo alométrico para evaluar produccion foliar de Zostera marina L. Hidrobioldégica, 1(2): 41-11. , H. Echavarria-Heras H., and S. E. Ibarra-Obando. 1996. Leaf size dynamics for Zostera marina L. In San Quintin Bay, Mexico: A theoretical study. Estuarine, Coastal and Shelf Sci- ence. (in press) STATISTICA. General Conventions and Statistics I. Copyright © 1984-1994 StatSoft Inc. Wetzel, R. L., and P. A. Penhale. 1983. Production ecology of an eelgrass community in the lower Chesapeake Bay. Mar. Tech. Soc., 17:22-31. Zachmanoglou, E., and D. W. Thoe. 1976. Introduction to Partial Differential Equations with Appli- cations. Williams and Wilkins Company, pp. 60-62. Wonnacott T., and R. Wonnacott. 1984. Introductory Statistics for Business and Economics. John Willey and Sons Inc. New York, pp 614-627. Appendix According to theorem 2.1 in Zachmanoglou and Thoe (1976) (pages 60-61), the solution w = f(h,/) of equation (5) is given implicitly by the relation: F(u,(h,Lw),u(h,lw) ) = 0 (Al) Where F is a function of class C! and u,, u, are two functionally independent solutions of the auxiliary system: dl_— dh dw —= = — (A2) l k,h kw From the system (A2) we can obtain by direct integration l ly a (A3) he he Ue ie ope (A4) W Wo Where wo, /) and hy are respectively the dry weight, the length and width of a leaf at the beginning of the growing process and a and 6 are constants defined in terms of the allometric scaling factors k, and k, (see equation (1) and (2)) trough the relationships = (— AS " a o=k, (A6) From equations (A3) and (A4) we see that the functions u, and uw, defined according to Ww u,(h, L w) = a (A7) he u,(h, l, w) = a (A8) satisfy grad(u,) X grad(u,) ~ O in the region / > O, h > O and w > O. Then in general u, (h, 1, w) = c, and wu, (h, 1, w) = c, where c, and c, are constants, define two functionally independent solutions of the auxiliary 48 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES system (A2). Hence for u, and u, as given by equations (A7) and (A8) the implicit relation (Al) gives the general integral to p.d.e. (4). From equations (A7) and (A8) it is easily seen that io = Ls voles (A9) he wiih, hence for u, and u, as defined above we have u, — cu, = O (A10) Where c is a constant that can be identified using equation (A9). Consequently equations (A1), (A10), the choosing E(u, Us) = Uy — Cl and the implicit relation (Al) permit to conclude that the function w = Khel® Where B = 6 — | is a solution to equation (4) which also satisfies the allometric equations (1) and (2). 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Photo by Juan-Pablo Gallo-Reynoso. 33 7. te Pee neERN. -CALIFORNIA ACADEMY OF: SCIENCES BULLETIN Volume 97 Number 2 BCAS-A97(2) 49-88 (1998) AUGUST 1998 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 © Southern California Academy of Sciences, 1998 OFFICERS Robert S. Grove, President David Huckaby, Vice-President Susan E. Yoder, Secretary Robert Cavenberg, Treasurer Daniel A. Guthrie, Editor Hans Bozler, Past President David Soltz, Past President BOARD OF DIRECTORS 1997-1999 1997-2000 1998-2001 Ralph G. Appy Robert S. Grove Kathryn A. Dickson Jonathan Baskin David Huckaby Donn Gorsline J.D. Stewart Robert Lavenberg Larry G. Oglesby Gloria Takahashi Kenneth E. Phillips Robert E Phalen Susan E. Yoder Cheryl C. Swift Membership is open to scholars in the fields of natural and social sciences, and to any person interested in the advancement of science. 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Date of this issue 3 August:.1998 © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). CALIFORNIA ACADEMY OF SCIENCES AUG 1 9 4998 LIBRARY Bull. Southern California Acad. Sct. 97(2), 1998, pp. 49-75 © Southern California Academy of Sciences, 1998 Range Extensions of Ten Species of Bats in California Denny G. Constantine 1899 Olmo Way, Walnut Creek, California 94598 Abstract.—Ten rare or uncommon species were among some 22,000 bats identi- fied after being tested for rabies in California: Macrotus californicus, Choeron- ycteris mexicana, Leptonycteris curasoae, Myotis velifer, Lasionycteris noctiva- gans, Lasiurus blossevillii, Lasiurus xanthinus, Nyctinomops femorosaccus, Nyc- tinomops macrotis, and Eumops perotis. Numerous geographic range extensions were among 194 new localities reported. Perceived range extensions of southerly species may be due to either increased sampling, global warming or other factors. Identification by professional taxonomists of host animals tested for pathogens is encouraged to save valuable data and as prerequisite to disease problem compre- hension and resolution. New geographic range records and associated data have resulted from the iden- tification of more than 22,000 bats tested for rabies in California. Distributional highlights of 425 bats of ten generally rare or uncommon species, nearly all supported by preserved specimens, are presented here, whereas additional distri- butional, population, ecological, and rabies epidemiological studies of these and common species are in progress. Methods The author periodically identified bats tested for rabies in California subsequent to his first bat rabies survey in the state in 1954 (Enright et al. 1955), but it was not until 1977 that he did so regularly, if on a voluntary basis. Counties that could cooperate shipped the carcasses of bats they had tested to the author, who was employed at the California Department of Health Services in Berkeley. Other workers participated in identifications in early years. Keith EK Murray made several determinations cited in Constantine et al. (1979). Beginning in 1973 and ending in 1981, former Los Angeles Department of Health Services biologist Loran M. Whitelock, who earlier had worked on bats with the author, performed bat identifications for that county. Personnel took relevant health precautions (Johnson 1979: Constantine 1988), including preexposure rabies immunizations. Nearly all carcasses were damaged and decomposed when received, and the cranium had been opened to remove the brain, destroying key characters needed to differentiate some species. Most rare or uncommon bats were salvaged as skeletons or in fluid preservatives, and a few were prepared by the author as study skins, despite some hair slippage. Measurements are presented to support certain critical identifications. Different measurement methods were sometimes used for different species in order to match techniques used by authors of relevant published studies and thus facilitate com- parisons by the reader. Most external measurements were made with a rule. Fore- arm, wing digit, and cranial measurements were made with dial calipers. 49 50 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Most new locality records are merely listed, in accordance with the limited purpose of this report, relevant details being reserved for other reports, but the more unexpected records are given in detail. Plant communities, cited in reference to especially unusual locality records to reflect ecological parameters, are from Kuchler (1977). Maps bear a single symbol for each new (closed circle) or previously published (open circle) locality. Symbols do not reflect the numbers of reports or specimens. Museum catalog numbers and other details are generally given only for specimens that represent peripheral range extensions, although a few of these bats have not yet been cataloged. Recipient museums are the Museum of Vertebrate Zoology, University of California, Berkeley (MVZ) and the Natural History Museum of Los Angeles County (LACM). Results California leaf-nosed bat Macrotus californicus Baird 1858 This insectivorous species has been reported from northern Sinaloa and south- western Chihuahua, Mexico northward through Sonora and Baja California to southern California, extreme southern Nevada, and southern Arizona; an appar- ently disjunct population has been reported in Tamaulipas, Mexico (Koopman, 1993). The species is included herein primarily to report recent observations at reported and unreported westernmost stations of known occurrence. Howell (1920b), referring to this species in a cave (see photograph in Howell’s paper), said to be 2 mi N of Owensmouth, Los Angeles Co., reported six of the bats present on 6 April, none on 4 June, and about thirty on 14 December (years not given). Owensmouth has been renamed Canoga Park. The cave, about 24 m deep and 9 m high in calcareous conglomerate, was called Vanowen Bat Cave by Halliday (1962) after the author took him there. It was found to be in the first canyon south of Bell Canyon, about 6.6 km W of Canoga Park and just inside Ventura Co. some 0.97 km W of the present end of Vanowen Street at Sunset Ridge Court. On 30 January 1949, I observed three Macrotus on the cave ceiling. Identified by their large ears, nose leaves, and “‘twirling’’ when hanging by one foot to watch the observer, they were not disturbed further. No bats were seen in the cave on subsequent visits made February 1953, 4 July 1989, and 27 April 1990: A second cave, this one in Los Angeles Co. about 10.31 km NNE of the first cave and about 3.32 km NNW of Chatsworth, was found by Thomas Cade and Gary Casey to contain a small group of these bats on 8 May 1947, when six were collected, prepared as skins, and given to the LACM. I joined them on a visit to the cave later that month, when several Macrotus skeletons were found on the cave floor. Mark Ryan collected a bat at this cave 15 June 1950 (MVZ 113637). The cave was a dugout under several large adjacent boulders on an extensive boulder-strewn property known as the Iverson Ranch. The area has since been bisected by a freeway and subdivided for housing, with much ground surface distortion. A search for the cave by the author and Kenneth E. Stager in April 1990 was unsuccessful. Only four Macrotus are known to have been submitted for rabies testing in RANGE EXTENSIONS OF BATS IN CALIFORNIA 51 Choeronycteris mexicana Oo published @® new 100 mi. Ee a | 100 km. Fig. 1. Known distribution of the Mexican long-tongued bat, Choeronycteris mexicana, in Cali- fornia. California, as expected for a species that prefers living far from human habitat. They were from within the known range in Imperial, San Bernardino, and San Diego counties. Mexican long-tongued bat Choeronycteris mexicana Tschudi 1844 This nectar, pollen, and fruit-eating bat has been reported from El Salvador and Honduras (Arroyo-Cabrales et al. 1987) northward through Mexico, including Baja California, to San Diego, California (Olson 1947), southern Nevada (Con- stantine 1987), southern Arizona (Hoffmeister 1986), southwestern New Mexico (Findley et al. 1975), and extreme southern Texas (LaVal and Shifflett 1971). The first observations of this species in California were made September to December 1946, during an “‘invasion’”’ by the species, in San Diego, San Diego Co. Fifty of the bats were collected, usually as individuals found hanging in dimly lit sites about buildings, but as many as 40 to 50 were seen at one place and time, with probably 75 total bats being detected (Olson 1947; Huey 1954b). Bond (1977) mentioned that subsequent San Diego specimens were obtained in October of 1947 and 1963 and in December 1947. Banks and Parrish (1965) reported another specimen taken 15 October 1963 from Lemon Grove, at the eastern edge of San Diego. I have received 15 additional bats of this species: 10 from five localities in San Diego Co., three from at least two localities in Orange Co., one from Los Angeles Co., and one from Ventura Co. Unfortunately, labels came off individual con- tainers within two frozen shipments, so data on two of these bats (one each from Los Angeles and Orange counties) were lost (Fig. 1, Appendix). The San Diego Co. bats do not represent remarkable range extensions. The first Orange Co. bat, a female (MVZ 181846), was taken at an unknown place and date between 1976 and 1978. The second, a male (MVZ 186389), was taken at San Clemente 28 September 1993. The third, a male (LACM 94031), was found at Tustin on 24 November 1995. The Los Angeles Co. bat, a female (MVZ 32 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 181845), was taken at an unknown locality and date between July and December 1988. The Ventura Co. bat, a male (LACM 94032), was found 15 November 1994 at Ventura, 260 km NW of San Diego. This species, one of three leaf-nosed bats now known in California, is readily differentiated from the other two. Macrotus has huge ears compared to the rela- tively short ears of Choeronycteris and Leptonycteris. The latter lacks an evident tail, whereas the short and exsert tail of Choeronycteris arises from the dorsal surface of the interfemoral membrane near its base. Southern long-nosed bat Leptonycteris curasoae yerbabuenae Martinez and Villa-R. 1940 These nectar and pollen eating bats are reported to appear in southern Arizona in spring, produce offspring in colonies within caves and mine tunnels, and dis- appear in fall, evidently returning south into Mexico (Hayward and Cockrum 1971; Hoffmeister 1986). Arita and Humphrey (1988) considered L. curasoae to be divisable into two subspecies: (1) L. c. verbabuenae, distributed from southern Arizona, New Mexico, and Tamaulipas, Mexico southward to Guatemala and El Salvador, and (2) L. c. curasoae, thus far known as a disjunct population in northern Colombia and Venezuela and nearby Caribbean islands. The species is reported herein for the first time from California, where two bats were found, each in a different county in the southern part of the state. The first bat was found in suburban Yucaipa, 799 m elev., San Bernardino Co., formerly characterized by coastal sagebrush, adjacent to chapparal-covered hills. At about 1100 h PDT, 3 October 1993, a male bat (MVZ 186390) was inadver- tently flushed from a shaded site behind a meter-high bush growing next to the outer side of a residence chimney, which was flush against the south side of a one-story house. The bat flew about 4.6 m through the back door and into the shade of a screened carport-patio, where it was captured. After brain removal, the carcass was frozen and sent to the author four months later, when it was made into a study skin with skeleton, and other tissues were saved. Dehydration pre- cluded getting reliable ear and weight measurements. The second bat, a male (LACM 94033), was found in urban Oceanside, 14 m elev., San Diego Co., an area of former coastal sagebrush adjacent to chapparal and southern oak forest. The bat was observed at 900 h PDT 18 October 1996 hanging outside under a canopy over the front door of a business concern. The frozen carcass was received by the author four months later, when it was prepared as a study skin and skeleton. The bat had considerable deposits of subcutaneous fat throughout but especially in the neck, interscapular, and leg base areas. The bats were identified as L. curasoae and not Leptonycteris nivalis by com- paring them with skins and literature. Measurements were made using methods described by Arita and Humphrey (1988), except bilateral measurements were taken, where applicable, and averaged. The hair coat was typically short and dense, and the hair fringe on the border of the uropatagium was inconspicuous. The presphenoid ridge was prominent and rounded. Recoverable measurements (Table 1) were consistent with those published for samples of bats designated L. c. verbabuanae from Morelos and Colima, Mexico by Arita and Humphrey (1988), from Baja California Sur, Mexico (as L. sanborni) by Woloszyn and Wo- Nn OW RANGE EXTENSIONS OF BATS IN CALIFORNIA Table 1. Leptonycteris Curasoae: measurements (mm) and weight (g).* Yucaipa, San Oceanside Bernardino Co San Diego Co Measurement 3-X-1993 18-X-1996 Total length 70 81 Length of foot 14.4 14.0 Length of ear from notch 7/0) Length of folded forearm 53.70 52295 Length of 3rd digit 99.45 oe )al 3) Length of 3rd metacarpal 49.05 49.65 Length of Ist phalanx of 3rd digit 14.58 13.75 Length of 2nd phalanx of 3rd digit 23.47 2375 Length of 3rd phalanx and cartilage of 3rd digit 35 12.00 Condylobasal length Pasytes0) 26s lg) Greatest cranial length excluding incisors 2ORS5 Zygomatic width 10.30 10.78 Least interorbitai width 4.35 4.40 Greatest width of braincase 10.09 9.98 Greatest width at mastoids 10.80 10.64 Width of rostrum at last maxillary molars 6.60 6.40 Length of palate excluding spine 14.14 14.18 Length of maxillary tooth row excluding incisors 9.00 8:95 Length of mandible 17.98 18.10 Length of mandibular tooth row excluding incisors 9.48 AE | Weight (less brain) DOD * Methods are from Arita and Humphrey, 1988. loszyn (1982), and from Arizona (as L. nivalis sanborni and later as L. sanborni) by Hoffmeister (1957, 1986). The present known distribution of L. c. verbabuenae in the United States and northwestern Mexico is indicated in Fig. 2. Yucaipa, the new northernmost locality where the species is known, is 463 km west and slightly north of the previous northernmost locality at Glendale, Mari- copa Co., Arizona (Constantine 1966), 420 km NNW of the previous westernmost locality in the United States at Agua Dulce Mountains, Pima Co., Arizona (Cock- rum and Petryszyn 1991), and 823 km NNW of the northernmost Baja California Sur, Mexico, locality at Santa Rosalia (Arita and Humphrey 1988). Oceanside is 98 km SSW of Yucaipa. Cave myotis Myotis velifer velifer (J. A. Allen 1890) This insectivorous species is known to occur from the southeastern California and Nevada borders at the Colorado River to Kansas and southward through mainland Mexico and Guatemala to El Salvador and Honduras (Hayward 1970; Helebuyck et al. 1985). The subspecies M. v. velifer reportedly occupies the west- ern and southern parts of the range. Hayward (1970) concluded that the smaller bat of the southern end of Baja California Sur, Mexico, classified as M. v. pen- insularis by Miller and Allen (1928), is a separate sibling species, M. peninsularis, as originally determined by Miller (1898). Found during warm months in California only near the Colorado River, aggre- 54 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Leptonycteris curasoae © published @new . Fig. 2. Known distribution of the southern long-nosed bat, Leptonycteris curasoae yerbabuenae, in the southwestern United States and northwestern Mexico. From Arita and Humphrey (1988), Cock- rum and Petryszyn (1991), Hoffmeister (1986), and the present study. gations of this presumably migratory subspecies previously were common in abandoned mine tunnels, where its numbers have been drastically reduced. Three bats of this species are reported herein from Los Angeles Co., a surprising distance westward from the former westernmost known boundary of the species. The first bat, a female (MVZ 186397), was captured after it flew through a door into a house during daytime 28 September 1992 at Florence, 49 m elev., an urban area originally of coastal sagebrush. The scalped carcass was received by the author the next month when it was salvaged as a skin and skeleton, although it was decomposed with extensive hair slippage. The second bat, a male (MVZ 186398), was found alive during daylight the morning of 28 November 1994 clinging to an overhang above a door at Valencia, 303 m elev., in a parklike suburban setting with adjacent chapparal and riparian vegetation. The scalped, decomposed and somewhat dessicated carcass was sent to the author some six months later when it was salvaged as a skin, legs and tail vertebrae contorted within the shriveled uropatagium, and skeleton with skull fragments. The third bat, a male (LACM 94035), whose canines were shortened by wear, was captured about 1900 h PST 27 March 1997 while hanging in a covered entryway outside the front door of a residence at Lancaster, 718 m elev. Areas representing three plant communities meet in the vicinity: Joshua tree scrub, Mo- jave creosote bush, and desert saltbush. I received the partially decomposed car- cass the following month and salvaged it as a skin and skeleton. Some external and recoverable cranial measurements of the three bats are pre- RANGE EXTENSIONS OF BATS IN CALIFORNIA =)5) Table 2. Myotis velifer from Los Angeles Co: measurements (mm) and weight (g).* Florence Valencia Lancaster Measurement 23-IX- 1992 28-XI-1994 3-XII- 1996 Total length 95 98 o5 Length of tail 4] 43 42 Length of foot a0 7 10.0 Length of ear from notch 1325 14.5 Length of forearm 42.75 44.05 42.40 Occipitonasal length 15:70 15.86 Condyle-premaxillae length (C) 15.05 14.94 Cranium breadth TD 8.10 7.78 Mastoid breadth 7.85 8.08 Rostrum breadth (H) 4.53 4.21 4.60 Least interorbital breadth 3.D> 4.05 3165 Braincase depth (H) 5502 (SHA Maxillary tooth row length 6:33 6.18 632 Length of mandible (C) PEAT 11.99 12.29 Mandibular tooth row length (C) 6.71 6.66 6.67 Weight (less brain) 6.9 * Methods are from Hayward, 1970, who cited Cockrum, 1955 (C), from Hoffmeister, 1986 (H), or from both where identical methods were used. sented in Table 2, and localities of collection are indicated in Fig. 3 and the appendix. The measurements of the three bats are consistent with those given by Grinnell (1918), Miller and Allen (1928), Stager (1939), Vaughan (1954), Hayward (1970), and Hoffmeister (1986), although there are some differences between these au- thors possibly referable to methods or specimen sexes and ages. The long forearms of these three bats separate them from all other members ORS SSS SSS CS SRS SSCS SSO e ay Myotis velifer ate ee rE A Soo tte eT a PCat a © published @ new 100 mi. 200 km. Fig. 3. Known distribution of the cave myotis, Myotis velifer velifer, in California and adjacent border areas. 56 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES of the genus known from the United States except M. volans, M. thysanodes, and the southeastern M. grisescens. However, like velifer they lack the rounded ears, furred basal wing membranes, and keeled calcar of volans, the fringed uropata- gium and larger ears of thysanodes, and the unicolored dorsal hairs and tarsal wing attachment of grisescens. Valencia, Los Angeles Co., is now the westernmost known locality for this species and is 377 to 482 km W of the previously reported westernmost localities at or near the Colorado River as follows: Jackass Flats, Clark Co., Nevada (Cockrum and Musgrove 1964) Needles, San Bernardino Co., California (Grinnell 1918) Parker, La Paz Co., Arizona (Hoffmeister 1986) Riverside Mountains, Riverside Co., California (Stager 1939) Blythe, Riverside Co., California (LACM 13626) Picacho, Yuma Co., Arizona (Hoffmeister 1986) Valencia is distant from the other two Los Angeles Co. localities as follows: 52 km WSW of Lancaster, and 62 km NNW of Florence. The three cave bats from Los Angeles Co. are the only bats of that species known to have been tested for rabies in California. Silver-haired bat Lasionycteris noctivagans (Le Conte 1831) This insectivorous and presumed migrant has been reported from forested areas throughout North America from southeastern Alaska across the southern half of Canada and southward to or near the southern border of the United States ex- cluding Florida, southwestern Arizona, and southern California (Hall 1981; Kunz 1982). Thus far, only one bat has been reported from Mexico, in Tamaulipas, south of Texas (Yates et al. 1976). The species is also known to occur during spring and fall in Bermuda, evidently as wind-blown migrants from the east coast of North America (Van Gelder and Wingate 1961). The distribution of this species in California is generally depicted on maps to include and extend from the northern third of the state southward along coastal mountains to Pacific Grove, Monterey Co. (Grinnell 1918) and, leaving the San Joaquin Valley blank, southward through the Sierra Nevada Mountains and, in the eastern part of the state, to Death Valley National Park (Grinnell 1937; Bradley and Deacon 1971). However, Bond (1977) reported a bat much farther south in eastern San Diego County. I have identified numerous bats of this species from nearly all counties in the northern two-thirds of the state, including the counties of Monterey, Kings, Tulare, and Inyo northward. Unfortunately, relatively few bats of any species were re- ceived during much of the period under consideration from the counties of San Luis Obispo, Kern, Santa Barbara, and Imperial, so the absence of silver-haired bats from those counties is of little significance. However, 23 of the bats were received from 15 new localities in the southern Californian counties of Los An- geles, San Bernardino, and San Diego during 1973-1997 (Fig. 4, Appendix). Of the 23 bats from southern California, seven (identified by L. M. Whitelock) were discarded before I was able to review and salvage specimens. Thus, voucher specimens were lost for the following localities ‘in Los Angeles Co.: Bellflower, Lakewood, and Torrance. RANGE EXTENSIONS OF BATS IN CALIFORNIA a7 Lasionycteris noctivagans © published @ new | 100 mi. 100 km. Fig. 4. Known distribution of the silver-haired bat, Lasionycteris noctivagans, in southern Cali- fornia. The southernmost California locality for this species on the Pacific coast is San Diego, represented by a male (MVZ 181863) taken 15 February 1978. San Diego is 615 km SE of the previously published southernmost Pacific coastal locality at Pacific Grove (Grinnell 1918) and 84 km WSW of the inland collection site at Agua Caliente Springs in eastern San Diego Co. (Bond 1977). Western red bat Lasiurus blossevillii frantzii (Peters 1871) This insectivorous, foliage-dwelling, migratory bat, recently regarded as taxo- nomically separate from the eastern red bat, L. borealis (Baker et al. 1988; Mo- rales and Bickham 1995), has been reported from southern British Columbia, Canada, Utah, and western Texas southward through the western United States, Baja California, mainland Mexico, Nicaragua, and south into South America (Hall 1981; Shump and Shump 1982; Eisenberg 1989; Redford and Eisenberg 1992). This species generally occurs in California’s central valley, foothills, and in similar areas of tree growth in southern California, preferring the dense foliage of trees for shelter, presumably avoiding tree-deficient deserts. Range maps often depict continuity of distribution through areas of unknown occurrence, although supportive data are unavailable. Such is true of this species in reference to Cali- fornian deserts. That limitation may be less misleading in reference to migrants like the red bat that may seasonally or periodically migrate or stray into or through these deserts. The following report may represent such an occurrence. On 21 October 1991, a female western red bat was found crawling on the ground at the Park Village government housing area near Furnace Creek Ranch, Death Valley National Park, Inyo Co. External measurements (mm) were: total length, 116; length of tail, 50; length of ear from notch, 12; length of foot, 7.5; length of tibia, 21.0; length of forearms, 41.0, 41.5. This record is within a broad void in the reported distribution of the species. It is from 183 to 432 km distant from the nearest knowns stations of occurrence: 5 mi SW Fallon, Churchill Co. Nevada (Hall 1946) 58 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Overton, Clark Co., Nevada (Hall 1946) Mouth of Bright Angel Creek, Grand Canyon National Park, Coconino Co., Arizona (Hoffmeister 1986) Big Sandy Creek, Mohave Co., Arizona (Hoffmeister 1986) Three Rivers, Tulare Co. (Grinnell 1918) Lancaster, Los Angeles Co., (based on a discarded female bat identified by L. M. Whitelock) Warren’s Ranch, San Bernardino Co., (Grinnell 1918) Santa Ysabel, San Diego Co. (Grinnell 1918) Grinnell (1937) suggested that the appearance of hoary bats, Lasiurus cinereus, and silver-haired bats at Furnace Creek Ranch, Death Valley National Monument, in spring and fall coincided with their anticipated migratory movements, a con- clusion that may be applicable as well to the present red bat record. The oasislike site, surrounded by desert, would be especially attractive to these tree-dwelling species. Western yellow bat Lasiurus xanthinus (Thomas 1897) Originally described as a subspecies of L. ega, this insectivorous bat recently was regarded as a separate species (Baker et al. 1988; Morales and Bickham 1995) whose range reportedly extends southward from southwestern New Mexico, southern Arizona, and southern California through Baja California, western Mex- ico and the Mexican plateau to Nuevo Leon and Morelos. Early locality records of this species in California include: Palm Springs, Riverside Co. (Constantine 1946) Cottonwood Spring, Joshua Tree National Monument, Riverside Co. (Loomis and Stephens 1964) Pomona, Los Angeles Co. (Stewart 1969) Borrego Springs, San Diego Co. (Bond 1970) Fourteen additional localities in the counties of Riverside, San Bernardino, San Diego, and Imperial were reported later (Constantine et al. 1979). Including the last-mentioned report, 214 bats of this species were received for identification after rabies testing during the period 1969-1997. They represented 59 localities within the aforementioned four counties and from Los Angeles and Orange coun- ties (Fig. 5, Appendix). Published northernmost and westernmost Arizona records for this species are Phoenix, Maricopa Co. (Constantine 1966) and Yuma, Yuma Co. (Constantine 1966), as mapped by Hoffmeister (1986). The present report extends the known northern and western limits of the range from Phoenix to Los Angeles County as follows: Blythe, Riverside Co., 7-XI-1980 (MVZ 181879) Twentynine Palms, San Bernardino Co., 13-VIII-1980 (MVZ 181943) Yucca Valley, San Bernardino Co., 25-IX-1985 (MVZ 181946) Muscoy, San Bernardino Co., 14-I[X-1992 (MVZ 186347) Azusa, Los Angeles Co., 12-XI-1987 (MVZ 181876) Glendale, Los Angeles Co., 27-IX-1984 (MVZ 181877). RANGE EXTENSIONS OF BATS IN CALIFORNIA 59 Lasiurus xanthinus © published @ new 100 mi. 200 km. Fig. 5. Known distribution of the western yellow bat, Lasiurus xanthinus, in California and ad- jacent Arizona. Pocketed free-tailed bat Nyctinomops femorosaccus (Merriam 1889) Long classified in the genus Tadarida (Shamel 1931), this insectivorous species recently was placed in the genus Nyctinomops by Freeman (1981). The species is known from southwestern Texas (Easterla 1973; Schmidley 1991) and southern New Mexico (Constantine 1958; Findley et al. 1975) through southern Arizona (Hoffmeister 1986) to southern California and southward in Mexico to Baja Cal- ifornia, Jalisco, and Nuevo Leon, as mapped by Hall (1981) and by Kumirai and Jones (1990). In addition to the type locality at Agua Caliente [Palm Springs], Riverside Co. (Merriam 1889), the species has been reported from only two ad- ditional California localities, both in San Diego Co.: Palm Canyon, near Borrego Springs (Neil 1940) and 3 miles southeast of Suncrest, now known as Crest (Krutzsch 1944, 1945). I have identified 47 additional bats of this species from 30 localities, 29 of which are new, from the Californian counties of Imperial, Los Angeles, Orange, Riverside, San Bernardino, and San Diego (Fig. 6, Appendix). Stations constitut- ing the northernmost tier of these localities, proceeding from east to west, are: Calexico, Imperial Co., 3-X-1995 (MVZ 186401) La Quinta, Riverside Co., 13-IV-1994 (MVZ 186386) San Bernardino, San Bernardino Co., 15-XI-1985 (MVZ 181965) Covina, Los Angeles Co., 30-IV-1985 (MVZ 181959) Inglewood, Los Angeles Co., 18-X-1994 (LACM 94036) Calexico is located at the Mexico border, 190 km SSW of the nearest Arizona record at Alamo Crossing, Mohave Co. (Cockrum and Musgrove 1965). Ingle- wood is 182 km NW of the previously reported westernmost locality at Crest, San Diego Co. Big free-tailed bat Nyctinomops macrotis (Gray 1839) This insectivorous species is known from British Columbia, Canada, in the Northwest (Cowan 1945), North Carolina on the Atlantic coast (Di Salvo et al. 60 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Nyctinomops femorosaccus oO published @ new 100 mi. 200 km. Fig. 6. Known distribution of the pocketed free-tailed bat, Nyctinomops femorosaccus, in Califor- nia and adjacent Arizona. - 1992), and the central United States southward through most of Mexico, the Caribbean islands of Cuba, Jamaica, and Hispaniola, and South America to Bo- livia, northern Argentina, and Uruguay (Hall 1981; Milner et al. 1990). Reports indicate that its greatest density in the United States is in the Southwest, reports dwindling northward and eastward. For example, there are no published reports between San Diego and British Columbia, there are only two reports each from Oklahoma, Kansas, and Iowa, and no reports between Texas and the single spec- imen from North Carolina, whereas 36 collection localities for the species were reported from Arizona, New Mexico, and Texas (Constantine 1958, 1961a, 1961b; Hoffmeister 1986; Findley et al. 1975; Schmidly 1991). Four of these bats were reported previously from California. No locality was given for the first bat (Shamel 1931). The other three were from San Diego, San Diego Co. (Huey 1932, 1954a; August and Dingman 1973). The present report concerns 26 additional bats from 25 new localities in the Californian counties of Contra Costa, Imperial, Los Angeles, Orange, Riverside, San Bernardino, San Diego, San Luis Obispo, San Mateo, and Santa Barbara (Fig. 7, Appendix). Only the northermost localities are listed below, proceeding from east to west: El Centro, Imperial Co., 31-II]-1982 (MVZ 181981) Palm Springs, Riverside Co., 4-IV-1994 (MVZ 181986) Pomona, Los Angeles Co., 23-XI-1987 (MVZ 181985) Azusa, Los Angeles Co., 2-X-1997 (LACM 94037) Burbank, Los Angeles Co., 19-XI-1987 (MVZ 181982) Santa Barbara, Santa Barbara Co., 27-XI-1996 (MVZ 186402) Morro Bay, San Luis Obispo Co., 18-XII-1981 (MVZ 181992) Pacifica, San Mateo Co., 3-I-1984 (MVZ 181993) Martinez, Contra Costa Co., 13-XI-1979 (MVZ 181980) El Centro is 200-302 km distant from the nearest published reports to the east at Eagle Tank, Yuma Co., Arizona (Simmons 1966); 1.5 mi SE Kingman, Mohave Co., Arizona (Cockrum et al. 1996); Henderson, Clark Co., Nevada (Bradley et al. 1965). Martinez is 1279 km S of Essondale, British Columbia, the northern- most published locality. Relatively few bats of any species routinely are tested RANGE EXTENSIONS OF BATS IN CALIFORNIA 61 Nyctinomops macrotis Oo published @ new Fig. 7. Known distribution of the big free-tailed bat, Nyctinomops macrotis, in California and adjacent Nevada. for rabies in Californian coastal counties north of the vicinity of San Francisco, rendering nearly meaningless the absence of specimens of this rare bat from those counties. Western mastiff bat Eumops perotis californicus (Merriam 1890) This insectivorous subspecies has been reported from California to Texas south- ward to at least central Mexico, whereas other subspecies are recorded from north- ern South America southward into Argentina, (Koopman 1978). The species is also known from Cuba (Koopman 1993). Eighty-six mastiff bats were received from laboratories between 1973 and late 1997. Forty-nine new California locality records from 13 counties resulted, which with 38 published records, are presented in Fig. 8 and the appendix. The new records generally fill voids in the known range but include the northermost lo- cality. This species has been reported earlier at or near the Colorado River from Yuma, Yuma Co., Arizona (Cockrum 1960); 38.6 km S of Palo Verde, Imperial Co. (Eger 1977); Parker, La Paz Co., Arizona (Sanborn 1932); and Las Vegas, Clark Co., Nevada (Bradley and O’Farrell 1967). However, it has not been re- 62 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Eumops perotis © published @ new 100 mi. a a | 200 km. Fig. 8. Known distribution of the western mastiff bat, Eumops perotis californicus, in California and adjacent border areas. ported heretofore from the Imperial Valley, Imperial Co., the Coachella Valley north of Mecca, Riverside Co. (Grinnell 1918), from elsewhere in Riverside Co. except 4 mi SW of Lakeview (Vaughan 1959), from San Bernardino Co. except Colton (Grinnell 1918; Howell 1920a) and Keys Ranch (Campbell 1931), which apparently was a sight record. Counties represented herein from which no published records exist are Ventura, Santa Barbara, Merced, Stanislaus, and San Joaquin. The Calexico locality (MVZ 186385) is about 33 km WNW the nearest Mex- ican locality record at Cerro Prieto, Baja California (MVZ 110877, a skull col- lected by S. B. Benson 19 April 1948), the only record of this species in Baja California. The El Mirage, San Bernardino Co. bat (MVZ 186400) is the first specimen from the western Mojave Desert north of the transverse San Gabriel and San Bernardino Mountain Ranges, although Vaughan (1959) reported hearing the char- acteristic high-pitched, piercing calls of this species in the same general area. The Durham, Butte Co. bat, a female (MVZ 186399) taken 27 February 1997, represents the northernmost specimen-substantiated locality for this species. Only 25.6 km NW of Oroville, the previous northernmost locality (Eger 1977), Durham RANGE EXTENSIONS OF BATS IN CALIFORNIA 63 is 692 km WNW of Las Vegas, the next northernmost locality, and 952 km NW of Calexico. Discussion The samples of bats from which these locality records were derived were char- acterized by numerous biases that varied in space and time. In general, the bats were dead, disabled through injury or disease, or very young when discovered, and the majority had been brought home by household cats. Categorized as rabies- suspect, they were tested for that infection, especially if people or pets had been bitten or otherwise exposed. Numbers of bats tested were proportional to the human population that discov- ered them, so many were from cities and towns, whereas few were from unpop- ulated areas. Most counties sent for taxonomic identification all of the bats they tested, but others sent bats intermittently if at all, a few counties sending only some of the bats that had proved to be infected. Relevant news stories temporarily increased public awareness and submissions for testing, and molestation of col- onies or illegal poisoning of bats had the same effect due to consequent increased human and pet contacts. The surprisingly great range extensions indicated for M. velifer and L. curasoae at first elicited thoughts that the bats may have been accidently transported by truck or other vehicle in which they had temporarily taken shelter and been closed inside. However, such possibilities seemed unlikely after the second and third velifer were found and after consideration that other southerly species (e.g., C. mexicana, M. californicus, and L. xanthinus) are now known or were recently known in the same general southern California areas. Whether all of the reported range extensions from more southerly population centers reflect true changes or our emergence from a less-informed state is unclear. The increase in sampling of bats subsequent to the discovery of bat rabies in the United States in 1953 and the ever-increasing human and cat populations to collect them are undeniable. However, as evident are factors that support a concept of invasion. Lasiurus xanthinus is now relatively common in much of southern Cal- ifornia, where it was undetected until 1945. That species typically lives in fan palms, with which human habitat in California has become increasingly forested for ornamental purposes, evidently encouraging this bat’s range expansion from the occasional desert oasis where fan palms naturally grow in California. The majority of the migratory nectar-drinking bats (Choeronycteris and Leptonycteris) reported herein were males found during fall, corresponding to the comparable range-expansion scenario typically observed in pioneering birds, especially mi- gratory species (Johnson 1994). Myotis v. velifer appears to be migratory as well, because it has been unknown during cold months along the Colorado River and in most of Arizona. Perhaps overriding influences effecting these perceived range expansions of bats from the south and southeast are climatic warming and in- creased summer moisture, as hypothesized by Johnson (1994) in reference to perceived avian range expansions. Six of the 10 species of bat dealt with herein are on the State of California Department of Fish and Game’s List of Mammal Species of Special Concern: M. californicus, C. mexicana, M. velifer, N. femorosaccus, N. macrotis, and E. per- otis. A seventh, L. blossevillii, is expected to be added, and an eighth, L. curasoae, 64 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES will probably be included after its presence in California is anounced herein. Of these eight species, M. californicus, C. mexicana, L. curasoae, and M. velifer are generally found in caves or mines. Caves are relatively few in California, and far fewer are suitably warm for these bats. Most are visited by recreational hikers, and human population encroachment on them is increasing. The widespread clos- ing of unused mine tunnels and shafts during hazard abatement programs, the destruction of entire mine-riddled mountains by heap-leach mining practices, and the establishment of toxic ponds and runoff by the latter must contribute to the loss and future extinction of these and other associated species of wildlife in California. Nyctinomops femorosaccus, N. macrotis, and E. perotis naturally live in crev- ices in cliffs or in spacious caves, but they sometimes live in suitably high human structures, such as older wooden buildings, most of which have been replaced by unsuitable substitutes. Like most other bats, these molossids have suffered as targets of exterminators, especially Eumops, the largest and more obvious of the three. Increasing destruction of old-growth forest is removing the attendant hollows and loose bark crevices used as shelter by L. noctivagans, a species also adversely affected by the increasing elimination of native forested riparian habitat, the tree foliage of which provides shelter for L. blossevillii. The distributional data reported herein represent only one of several advantages consequent to identifying to species the bats tested for rabies in California or elsewhere. Taxonomic criteria, delineated through the past and continuing work of scientists in field, museum, university, or laboratory, find increasing application in disentangling the epidemiologic web of each disease that escapes from natural ecosystems into invading human populations, and rabies is no exception. Despite early opinions of leading health authorities that determining the species of each rabid bat was superfluous and that there existed only a single natural strain of rabies virus, it is now acknowledged that each of various known natural host species of bat or carnivore may be afflicted by one or sometimes more strains of rabies virus specific to that species. Host species-specific rabies virus strains were first recognized by differences in either susceptibility or in pathological reactions to the strains following inoculations of virus into panels of other mammal species (Constantine 1967a), later by monoclonal antibody techniques (Smith et al. 1986; Rupprecht et al. 1987), and recently by nucleotide sequence analysis of the viral nucleoprotein genome (Smith 1996). It is evident that a factual, comprehensive awareness of this and other zoonosis problems, now better known through the contributions of vertebrate, invertebrate, and virus taxonomists, will precede man- kind’s ability to develop appropriate responses. Thus, other states are encouraged to utilize the services of professional taxonomists to salvage valuable data and voucher specimens of bats currently being discarded after rabies tests. Acknowledgments I am indebted to many local, county, and state personnel who have and continue to support the effort to identify the bats that have been tested for rabies in Cali- fornia. Although participants include animal and vector control personnel, sani- tarians, public health investigators and others, my contacts have been with health department directors, physicians, public health veterinarians, and especially lab- RANGE EXTENSIONS OF BATS IN CALIFORNIA 65 oratory directors, microbiologists, and laboratory assistants who have shared the unpleasant work of organizing, labeling, and packaging for shipment the usually decayed and often mangled carcasses. Thus, my records lack the names of many participants but represent primarily laboratory personnel, as follows: E. Aaron, A. Adams, R. Alexander, O. Armstrong, M. Ascher, B. Austin, R. Avedian, A. Back, L. Barkley, R. Barnes, L. Barrett, I. Bihl, Jr., M. Brashear, J. Bunter, A. Chandler, J. Cleaves, S. Coffey, D. Cottam, G. Cookson, M. Croghan, R. Crosby, T. E Cuculich, H. DeBoer, D. Dondero, L. Doughty, J. Earnst, C. Egan, H. B. Ehrhard, R. W. Emmons, J. Falcon, D. Ferrero, P Flanders, B. Fleischer, K. Flink, C. Flinn, W. Fontes, B. Freeman, B. Fujikawa, EK Gettman, M. Giles, D. Gillies, R. Greenwood, W. Gregory, G. Guibert, M. Hansen, H. Hanson, K. V. Hardy, M. Hartley, S. Harvey, T. P. Herbenick, D. Hird, N. Hirada, M. T. Holland, J. Horn- stein, E. Howard, S. Hull, G. L. Humphrey, R. R. Jackson, H. Johnstone, H. N. Johnson, S. Kaddas, G. H. Kellogg, N. S. Kelly, R. Koelewyn, S. Kwan, R. Lasater, W. Lawrence, E. H. Lennette, S. Liska, D. Lockford, K. S. Mahoney, J. Marron, M. J. Martins, S. Matson, W. H. McCarley, D. McFarlane, T. Meier, D. V. Miller, L. N. Miller, M. Miller, D. Moore, B. Mognus, K. EK Murray, D. Murrill, M. Nachtigal, L. Oliver, W. Ota, D. Perez, C. A. Peter, J. Peterson, E. D. Pierson, E. Portoli, A. Pruitt, R. Purves, W. E. Rainey, K. Riley, W. L. Rottman, R. Ruiz, J. Ruprecht, C. 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Southern California Acad. Sci. 97(2), 1998, pp. 76-81 © Southern California Academy of Sciences, 1998 Strategies of Predator Attacks on the Schooling Fish, Selar crumenophthalmus, in Academy Bay, Socorro Island, Islas Revillagigedo, Mexico Bayard H. Brattstrom Department of Biology, California State University, Fullerton, CA 92634-9480 Abstract.—One of the main reasons that fish form schools is that it serves to reduce the risk of being eaten. Single predators are most successful at capturing individuals not in schools. For every successful anti-predator strategy by a prey species there is usually a concomitant more successful strategy by the predators. I report here on the behavior of three species of predatory fish and two species of predatory birds toward a school of jacks, Selar crumenophthalmus which dem- onstrate that these predators use a variety of methods to obtain fish from a school. Fish school for a variety of reasons. The behavior, dynamics, and advantages of fish schools and other similar aggregations in birds, tadpoles, and mammals have been fully described by many authors (Brattstrom 1962, 1989; Breder 1959, 1967; Cushing and Harden Jones 1968; Elliott, et al. 1977; Hamilton 1971; Major 1978, 1979; Partridge 1982; Partridge and Pitcher 1980; Seghers 1974; Shaw 1970, 1978; Webb 1980). One of the main reasons that fish school is that it serves to reduce the risk of being eaten (Breder 1967; Partridge 1982). Single predators are most successful at capturing isolated, individual prey and less successful at capturing individuals in schools (Brattstrom 1989; Major 1978, 1979). Thanks to natural selection, there usually will be a concomitant more successful strategy by the predators for every successful anti-predator strategy by a prey species! Studies on the behavior of the predator in response to schooling fish are diverse (Katzir and Chamhi 1993; Major 1978, 1979; Parish, Strand, and Lott 1989; Schmitt and Strand 1982). The outcome of any interaction between the predator and the prey fish usually depends on three factors: relative performance, maneuvering, and timing (Webb 1980). In addition, Major (1978) showed that while single predators are most successful at capturing isolated prey and less successful at capturing individuals in schools, grouped predators were more successful at capturing schooled prey. Larger predators were also able to break up schools of prey, re- sulting in increased numbers of prey becoming isolated. These predators then attacked these isolated individuals (Major 1978). I report here on the behavior of three species of predatory fish and two species of predatory birds preying on a school of jacks, Selar crumenophthalmus. This behavior shows that these preda- tors have developed diverse strategies to prey on schooling fish. I observed the predation of three species of fish yellowtail, Seriola lalanderi, California needlefish, Strongylura exilis, and black-tipped shark, Carcharhinus limbatus) and two species of birds (masked booby, Sula dactylatra, and great frigatebird, Fregata minor) on schooling jacks, Selar crumenophthalmus, locally called cabalito. 76 PREDATORS ON FISH SCHOOLS We. LAVA FLOW ms MIDDLE BEACH Fig. 1. Diagram of Academy Bay, Socorro Island, Mexico of cliff from which observations were made. Observations were made from a cliff above Academy Bay, at the north end of Socorro Island, Islas Revillagigedo, Mexico. Socorro Island is 390 km SW of the southern tip of Baja California (see descriptions in Brattstrom 1955, 1963, 1982, 1990; Brattstrom and Howell 1956; Richards and Brattstrom 1959). Observations were made throughout the day of 14 April, 1978 by me and other members of the Carnegie Museum/Sea World, 1978 Expedition to the Islas Re- villagigedo. Notes and photographs (35 mm slides, 16 mm color film) were made of the schooling jacks (Fig. 1, 3). Predation by the fish was observed in the clear shallow water from the cliff (Fig. 1), predation by the birds was observed from the cliff, beach, and aboard the expedition ship anchored in Academy Bay. The temperature of the water in the bay as taken from under the ship was 25°C. Following Breder (1959), I define a school as a behavior in which the fish are oriented in the same direction and are more or less one fish-length apart, a Pod as a group of fish that are in contact, a Pod I (or “‘ball’’) when the fish show no orientation and a Pod II when the fish show orientation. The jacks were in two, large schools (Fig. 1), each estimated to contain tens of thousands of fish. These schools would often spontaneously change into a Pod I or Pod II. Even when the fish were oriented in one direction, the entire school or pod did not really move very far. Instead, the entire school or pod seemed to slowly ‘‘float’’ about the bay. School B (see Fig. 1) once divided in two and then recombined. The two main schools never joined, even though at one time they were within 3 m of each other. 78 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES The behavior of the fish in the school changed as the school approached the shore (Fig. 2; beach effect). Here the waves and/or the shallow water caused the fish nearest the shore to form a Pod II (dots in Fig. 2). This pod formation spread until the whole group was in a pod. This pod slowly moved away from the shore and switched back to a school. Three species of predatory fish were also observed in the bay and each at- tempted to feed on the jacks in a different way. Groups of from two to 12 yel- lowtail swam around the bay always some distance from the jacks. They would circle both schools or swim back and forth in front of the beach or below the cliff. Now and then, and very suddenly, the group would turn toward the school and swim side by side straight into the school of jacks (Fig. 2, top). The fish in the school would immediately form a Pod I or Pod II at the edge of the school where the yellowtail were attacking. With the continued forward movement of the yellowtail, a ripple-like effect of school changing into pod occurred until finally all the fish were in a ball or Pod I. As the predators left, the pod would change back into a school formation as diagrammed in Fig. 2. I could not deter- mine if the yellowtail actually caught any of the jacks, though this strategy is usually successful (Major 1978). Cooperative foraging in yellowtail has also been documented by Schmitt and Strand (1982). Two black-tip sharks were also swimming in the bay. Now and then a shark would circle around a school of jacks, moving closer and closer to the school as it circled (Fig. 2). When the shark was about 3 m from the school, the jacks on the outer edge of the school would begin to contact each other. As the shark got closer the pod formation continued from outside toward the center until all fish were in a Pod I or ball. At that time the shark turned directly into the ball, mouth opening and closing (Fig. 2). A single needlefish also occurred in the bay. It usually swam just below the tall cliff on which I was standing. It appeared not to notice the schooling fish, yet (and I saw this happen many times) the needlefish would turn, suddenly, directly toward the school. It swam fast, and just before reaching the school, would jump out of the water and almost sail, like a flying fish. It would land in the water in the middle of the schooling jacks with its mouth opening and closing (Fig. 2). Again, I could not determine if the needlefish was successful in obtaining a prey fish, but the density of jacks would suggest that there was a high degree of success. The schooling jacks were also preyed upon by two sea birds. For most of the day a single frigatebird and a single masked booby flew over the bay. Both fed on the jacks by plunging down on the fish from above (Fig. 2). Both birds were successful in catching fish. The booby, for example, caught three fish in three dives over a six minute period from 1432-1458 hrs. The school/pod concentration of the prey made escape responses of the fish less effective (Katzir and Camhi 1993) and presumably therefore increased the success of the birds. While schooling behavior in fishes may have many causes and advantages, the most documented advantage is, of course, the group effect against predators (Breder 1967; Partridge 1982). It is usually the individual, isolated or separated from the herd, flock, or school, that is taken by predators (Brattstrom 1989). Yet the mere presence of this school in this bay allowed the feeding on the school by these predators. In addition, two of the predators manipulated the school into a 79 PREDATORS ON FISH SCHOOLS ‘UONVULIOJ pod & OJUI SUISURYS SI [OOYSS dy} DIDYM PUL UDYM dJBdIPUT SJOP YORI ‘;OOYds dy) JO 9zIs pue adeys ay) smoys SUIPINO Ie[NSILy “yoeRoq ay} 0} syoef oy} JO asuodsoa1 puv ‘sJOJeposd asoy) 0} syoef oy) JO asuOdsa ‘s1OJepoid [VIDADS JO JOIARYOQ SUIMOYS SUILISVIGQ “7 ‘SI HOV3d SNIAVST q— Od Guid 31Vv9Id4a GNV ASOOd Ov; HSI4d 31034N NOILVWYO4 aod es 1034343 HOVW3a WIVLILV AWYVHS NOILVWHO4 GOd 100HOS HLIM GOd ~ L * / ~ oN "4 &, NS HICE 1OOHIS ——— GOd SHOLVG3Hd ON NOILYVWYO4 GOd 1OOHIS CO @ cG2@: eC STIVIMOTI3A 80 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 3. Photograph of a school of jacks, Selar crumenophthalmus, in Academy Bay, Socorro Island, Mexico. Photograph by Robert Pitman. pod formation where it would be expected that nearly any bite by a predator would be assured of striking a fish. Parrish, Strand, and Lott (1989) showed that while predation was highest on isolated stragglers, predation on a school of flat- iron herring, Harengula thrissina, due to predator strategies, was about equal on peripheral and central fish in the school. Thus, while there is an advantage to schooling by fish, there are also predators that take advantage of that schooling behavior. In addition, cooperative hunting can circumvent the schooling advantage to prey species (Schmitt and Strand 1982). Acknowledgments The 1978 trip to the Islas Revillagigedo was aboard the R/V Sea World and was sponsored by the Carnegie Museum of Philadelphia and Sea World of San Diego under the leadership of Drs. Kenneth Parkes and Joe Jehl and was specif- ically designed to study aspects of the biota of Socorro and allow me to continue my studies of the effect of Barcena Volcano, 1952 on the biota of San Benedicto Island (Brattstrom 1963). I am very thankful to those gentlemen and their insti- tutions for support. Identifications of fish were made by Dr. Richard Rosenblatt, Scripps Institution of Oceanography and an early draft of the manuscript was read by Dr. Michael Horn. Figure 1 was photographed by Robert Pitman and the copy made by Alan Fugleberg. The drawings were made by Mark Zolle with funds from the Department of Biology, California State University, Fullerton. PREDATORS ON FISH SCHOOLS 81 Literature Cited Brattstrom, B. H. 1955. Notes on the herpetology of the Revillagigedo Islands, Mexico. Am. Mid. Nat. 54:219-—229. . 1962. Thermal control of aggregation behavior in tadpoles. Herpetologica. 18:38—46. 1963. Barcena Volcano, 1952; its effect on the fauna and flora of San Benedicto Island, Mexico. Pp. 499-524. In Pacific Basin Biogeography. (J. L. Grissett, ed.), Bishop Museum Press: . 1982. The comparison of the social behavior of Chelonia mydas on the Islas Revillagigedo, Mexico. Herp: Rev: 13:71: . 1989. Predation of Bald Eagles, Habaeetus leucocephalus, on American Coots, Fulca Amer- icana. J. Raptor Research, 23:16—17. . 1990. Biogeography of the Islas Revillagigedo, Mexico, J. Biogeogr. 17:177-183. , Brattstrom, B. H., and T. R. Howell. 1956. The birds of the Revillagigedo Islands, Mexico. Condor 59:107—120. Breder, C. M. Jr. 1959. Studies on social groupings in fishes. Bull. Amer. Mus. Nat. Hist. 117:393- 482. . 1967. On the survival value of fish schools. Zoologica 52:25—40. Cushing, D. H., and EF R. Harden Jones. 1968. Why do fish school? Nature 218:918—920. Elliott, J. P, I. McTaggart-Cowan, and C. S. Holling. 1977. Prey capture by the African Lion. Ca- nadian J. Zool. 55:1811—1828. Hamilton, W. D. 1971. Geometry for the selfish herd. J. Theor. Biol. 31:142—-159. Katzir, G., and J. M. Camhi. 1993. Escape response of Black Mollies (Poecilia sphenop) to predatory dives of a Pied Kingfisher (Ceryle rudis). Copeia 1993:549-553. Major, P. EF 1978. Predator-prey interactions in two schooling fishes, Caranx ignobilis and Stolephorus purpureus. Anim. Beh. 26:760—777. . 1979. Piscivorous predators and disabled prey. Copeia 1979:158. Parrish, J. K., S. W. Strand, and J. L. Lott. 1989. Predation on a school of Flat-iron Herring, Harengula thrissima Copeia 1989:1089. Partridge, B. L. 1982. The structure and function of fish schools. Sci. Amer. 1982:114—123. Partridge, B. L., and T. J. Pitcher. 1980. The sensory basis of fish schools. J. Comp. Physiol. 135: 315-325. Richards, A. F, and B. H. Brattstrom. 1959. Bibliography, cartography, discovery, and exploration of the Islas Revillagigedo. Proc. Calif. Acad. Sci. Ser 4. 20:315-—360. Schmitt, R. J., and S. W. Strand. 1982. Cooperative foraging by yellowtail, Seriola lalandei (Caran- gidae) on two species of fish prey. Copeia 1982:714—717. Seghers, B. H. 1974. Schooling behavior in the guppy (Poecilia reticulata): an evolutionary response to predation. Evolution 28:486—488. Shaw, E. 1962. The Schooling of fishes: Sci Amer. June 1—10. . 1970. Schooling in fishes: Critique and review. in Aronson, L. R., E. Tobach, D. S. Lehrman, and J. Rosenblatt (eds.), Development and Evolution of behavior: essays in memory of T. G. Schneirla. W. H. Freeman Co. San Francisco. . 1978. Schooling of fishes. Amer. Sci. 66:166—175. Webb, P. W. 1980. Does schooling reduce fast-start response latencies in teleosts. Comp. Biochem. Physiol. 652A:231—234. Accepted for publication, 26 June, 1997. Bull. Southern California Acad. Sci. 97(2), 1998, pp. 82-85 © Southern California Academy of Sciences, 1998 Composition of the Helminth Community of a Montane Population of the Coastal Whiptail, Cnemidophorus tigris multiscutatus (Sauria: Teiidae) from Los Angeles County, California Stephen R. Goldberg,' Charles R. Bursey,” and Mei Q. Wu! 'Department of Biology, Whittier College, Whittier, California 90608 *Department of Biology, Pennsylvania State University, Shenango Campus, 147 Shenango Avenue, Sharon, Pennsylvania 16146 Abstract.—Two-hundred sixty two Cnemidophorus tigris multiscutatus from the San Gabriel Mountains of Southern California were examined for helminths. The helminth community consisted of two species of cestodes (Oochoristica scelopori and Mesocestoides sp.), two species of nematodes (Pharyngodon cnemidophori and Physaloptera sp.) and one species of acanthocephalan (Moniliformis monili- formis). The helminth with highest prevalence (15%) and greatest mean intensity (6.35) was Pharyngodon cnemidophori. Cnemidophorus tigris represents a new host record for Oochoristca scelopori and Moniliformis moniliformis. The western whiptail, Cnemidophorus tigris Baird and Girard, 1852 ranges from north central Oregon and southern Idaho, south to Baja California and south- ern Coahuila, México and east to western Colorado, New Mexico and west Texas (Stebbins 1985). There are reports of helminths from C. tigris from Arizona (Ba- bero and Matthias 1967; Benes 1985; Goldberg et al. 1997), California (Read and Amrein 1953; Telford 1970; Mankau and Widmer 1977), Idaho (Lyon 1986), Nevada (Babero and Matthias 1967), Texas (Specian and Ubelaker 1974a, b) and Utah (Grundmann 1959). The purpose of this paper is to report on the composition of the helminth community from a montane population of a subspecies of C. tigris, namely, the coastal whiptail, C. tigris multiscutatus Cope, 1892, from the San Gabriel Mountains of Los Angeles County, California. There are no previous reports of helminths from this subspecies which occurs in coastal California and Baja California (Stebbins 1985). Additionally, comparisons are made with hel- minth communities in other populations of C. figris. Materials and Methods Two-hundred sixty two Cnemidophorus tigris multiscutatus from the San Ga- briel Mountains, Los Angeles Co., California were examined for helminths. Two sites were sampled: 198 (112 female, 86 male) lizards were collected along Cal- ifornia Highway 39 at 1580 m elevation and 64 (31 female, 33 male) from along California Highway 2 at 1830 m elevation. These specimens were collected in 1971 and 1974 by shooting with 22-caliber dust shot, fixed in 10% formalin and stored in ethanol. They were deposited in the Natural History Museum of Los Angeles County (LACM 111193-110931). The body cavity was opened by a longitudinal incision from vent to throat, and the digestive tract was removed by cutting across the anterior esophagus and rectum. The lumen of the esophagus, stomach, small and large intestines and the 82 HELMINTH COMMUNITY OF THE COASTAL WHIPTAIL 83 Table 1. Helminths from 262 Cnemidophorus tigris multiscutatus from the San Gabriel Mountains, Los Angeles County, California (collected 1971 and 1974). Preva- Mean Number Infected lence! abun- Helminth species helminths — lizards (%) dance? Site Cestoda Mesocestoides sp. (Tetrathyridia) 388 3 l 1.48 Body cavity Oochoristica scelopori 6 5 2 0.02 Small intestine Nematoda Pharyngodon cnemidophori 1663 38 15 6.35 Large intestine Physaloptera sp. (larvae) 6 3 1 0.02 Stomach Acanthocephala Moniliformis moniliformis 9 3 l 0.03 Small intestine ' Number of hosts infected with one or more individuals of a parasite species divided by the number of hosts examined. ? Total number of individuals of a parasite species divided by the total number of hosts examined. surfaces of the liver and body cavity were examined for helminths. Each helminth was initially placed in a drop of glycerol on a glass slide. Nematodes were iden- tified from these temporary mounts. Cestodes and acanthocephalans were stained with hematoxylin and identified. Terminology usage is in accordance with Bush et al. (1997). Selected specimens were deposited in the U.S. National Parasite Collection, USNPC, Beltsville, Maryland: Mesocestoides sp. (87530); Oochoris- tica scelopori (87529); Pharyngodon cnemidophori (87531); Physaloptera sp. lar- vae (87532); Moniliformis moniliformis (87533). Results The helminth community of the San Gabriel Mountain population of C. t. multiscutatus was found to consist of two species of cestodes, Oochoristica sce- lopori Voge and Fox 1950 and Mesocestoides sp. (tetrathyridea only), two species of nematodes, Pharyngodon cnemidophori Read and Amrein 1953 and Physalop- tera sp. (3rd stage larvae), and one species of acanthocephalan, Moniliformis moniliformis (Bremser 1811). The number of helminths, number of infected liz- ards, prevalence, mean abundance and site of infection are given in Table 1. At 1530 m elevation, (18%) 35 of 198 lizards harbored helminths, at 1580 m, (14%) 9 of 64; there was no significant difference for prevalence of helminths between elevations (chi-square = 0.33, | df, P > 0.05). Likewise, no significant difference was found between prevalence of helminths between female, (18%) 26 of 143 infected, and male, (15%) 18 of 119, lizards (chi-square = 0.31, 1 df, P > 0.05). It should be noted, however, that the nine Moniliformis moniliformis were found only at 1530 m elevation; one female and two male lizards were infected. The occurrence of Oochoristica scelopori and Moniliformis moniliformis represent new parasite records for Cnemidophorus tigris. Discussion In the present study, C. t. multiscutatus served as a paratenic host for three of the five species of helminths found. These species were represented by juvenile forms only; tetrathyridia of Mesocestoides sp., 3rd stage Physaloptera sp., and juvenile Moniliformis moniliformis. Tetrathyridia of Mesocestoides sp. are known 84 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 2. Helminth communities (species reaching maturity) of Cnemidophorus tigris. Locality Arizona, Maricopa County Arizona, Mohave County Arizona, Pima County Reference Benes 1985 Babero and Matthias 1967 Goldberg et al. 1997 Helminth community Oochoristica sp. Alaeuris sp. Pharyngodon warneri Oochoristica bivitellobata Abbreviata terrapenis Pharyngodon warneri California, Los Angeles County this paper Oochoristica scelopori Pharyngodon cnemidophori Oochoristica bivitellobata Pharyngodon cnemidophori Skrjabinoptera phrynosoma Thubunaea iguanae Pharyngodon cnemidophori Oochoristica bivitellobata Oochoristica bivitellobata Thubunaea cnemidophorus Parathelandors texanus Pharyngodon cnemidophori Oochoristica bivitellobata Pharyngodon warneri California, Riverside County Telford 1970 California, San Bernardino County Read and Amrein 1953 Idaho : Lyon 1986 Nevada, Clark County Babero and Matthias 1967 Specian and Ubelaker 1974a Specian and Ubelaker 1974b Utah Grundmann 1959 Texas, Brewster County from a large number of lizard species (see McAllister 1988) and have previously been reported from C. tigris from California (Mankau and Widmer 1977), Arizona (Benes 1985) and Nevada (Babero and Matthias 1967). Mesocestoides sp. is thought to require an arthropod intermediate host (Webster 1949). Third stage larvae of Physaloptera sp. are commonly found in species of Cnemidophorus (see Goldberg et al. 1993). In North America, no species of Cnemidophorus is known to harbor adult Physaloptera. Helminths (USNPC # 80202) identified as Physa- loptera retusa in the report by Goldberg and Bursey (1989) were found to be Abbreviata terrapenis. Arthropods serve as intermediate hosts (Lincoln and An- derson 1975). Mammals, especially rodents, serve as definitive hosts for M. mo- niliformis; insects are intermediate hosts (Van Cleave 1953). Unidentified acan- thocephalans were reported in C. tigris from central Arizona by Benes (1985) and Centrorhynchus sp. was found in C. tigris from southern Arizona by Goldberg et al. (1997). These helminth species might be expected in any insectivore. Cne- midophorus tigris also serves as a paratenic host for another nematode, Angus- ticaecum sp. from Utah (Grundmann 1959). Cnemidophorus t. multiscutatus served as definitive hosts for two of the five species of helminths found: Oochoristica scelopori and Pharyngodon cnemido- phori. Oochoristica scelopori is known from a variety of North American lizards (see Goldberg et al. 1996). Pharyngodon cnemidophori has been reported only from teiid lizards: C. tigris in Texas (Specian and Ubelaker 1974b); C. tigris in California (as Cnemidophorus tesselatus in Read and Amrein 1953; Telford 1970). Other helminths for which C. tigris serves as a definitive host are Oochoristica bivitellobata, Abbreviata terrapenis, Pharyngodon warneri, Parathelandros tex- anus, Skrjabinoptera phrynosoma, Thubunaea cnemidophorus and T. iguanae (Ta- ble 2). Cnemidophorus tigris from central Arizona was reported to harbor Alaeuris HELMINTH COMMUNITY OF THE COASTAL WHIPTAIL 85 sp. and Oochoristica sp. by Benes (1985). We believe the identification of Alaeu- ris sp. to be incorrect and consider this instance to represent an oxyurid species, probably Pharyngodon sp. It is of interest to note that the helminth community for which C. tigris is definitive host is different in each population so far studied (Table 2). This suggests that distribution patterns of helminth species are often different from distribution patterns of hosts or potential hosts. Acknowledgments We thank Robert L. Bezy (Natural History Museum of Los Angeles County) for permission to examine Cnemidophorus tigris multiscutatus for helminths. Literature Cited Babero, B. B., and D. Matthias. 1967. Thubunaea cnemidophorus n. sp., and other helminths from lizards, Cnemidophorus tigris, in Nevada and Arizona. Trans. Amer. Microsc. Soc., 86:173— AT Benes, E. S. 1985. Helminth parasitism in some central Arizona lizards. Southwest. Nat., 30:467— 473. Bush, A. O., K. D. Lafferty. J. M. Lotz and A. W. Shostak. 1997. Parasitology meets ecology on its own terms: Margolis et al. revisited. J. Parasitol., 83:575-583. Goldberg, S. R., and C. R. Bursey. 1989. Helminths of the giant spotted whiptail, Cnemidophorus burti stictogrammus (Sauria: Teiidae). Proc. Helminthol. Soc. Wash., 56:86—87. , and H. Cheam. 1997. Helminths from the Sonoran spotted whiptail, Cnemidophorus sonorae, and the western whiptail, Cnemidophorus tigris (Sauria: Teiidae), from southern Ar- izona with comments on Abbreviata terrapenis (Nematoda: Physalopteridae). Great Basin Nat., SOS OAM ; , C. T. McAllister, H. M. Smith and Q. A. Truong. 1996. Oochoristica scelopori (Cestoda: Linstowiidae) in a grassland population of the bunch grass lizard, Sceloporus scalaris (Phrynosomatidae), from Arizona. Great Basin Nat., 56:180-—182. , and R. Tawil. 1993. Gastrointestinal helminths of the western brush lizard, Urosau- rus graciosus graciosus (Phrynosomatidae). Bull. So. Cal. Acad. Sci., 92:43-51. Grundmann, A. W. 1959. Parasites recovered from six species of Utah lizards. J. Parasitol., 45:394. Lincoln, R. C., and R. C. Anderson. 1975. Development of Physaloptera maxillaris (Nematoda) in the common field cricket (Gryllus pennsylvanicus). Can. J. Zool., 53:385—390. Lyon, R. E. 1986. Helminth parasites of six lizard species from southern Idaho. Proc. Helm. Soc. Wash., 53:291-293. Mankau, S. K., and E. A. Widmer. 1977. Prevalence of Mesocestoides (Eucestoda: Mesocestoididea) tetrathyridea in southern California reptiles with notes on the pathology in the Crotalidae. Jap. J. Parasitol., 26:256—259. McAllister, C. T. 1988. Mesocestoides sp. tetrathyridea (Cestoidea: Cyclophyllidea) in the iguanid lizards, Cophosaurus texanus texanus and Sceloporus olivaceous, from Texas. J. Wild. Dis., 24: 160-163. Read, C. P., and Y. U. Amrein. 1953. North American nematodes of the genus Pharyngodon Diesing (Oxyuridae). J. Parasitol., 39:365—370. Specian, R. D., and J. E. Ubelaker. 1974a. Parathelandros texanus n. sp. (Nematoda: Oxyuridae) from lizards in West Texas. Trans. Amer. Microsc. Soc., 93:413-—415. , and . 1974b. Two new species of Phryngodon Diesing, 1861 (Nematoda: Oxyuridae) from lizards in West Texas. Proc. Helm. Soc. Wash., 41:46—51. Stebbins, R. C. 1985. A field guide to western reptiles and amphibians. Houghton Mifflin Company, Boston, xiv + 336 pp. Telford, S. R., Jr. 1970. A comparative study of endoparasitism among some Southern California lizard populations. Amer. Midl. Nat., 83:516—554. Van Cleave, H. J. 1953. Acanthocephala of North American mammals. Illinois Biol. Monogr., 23:1— 179. Webster, J. P- 1949. Fragmentary studies on the life history of the cestode Mesocestoides latus. J. Parasitol., 35:83—90. Accepted for publication 22 December 1997. Bull. Southern California Acad. Sci. 97(2), 1998, pp. 86-88 © Southern California Academy of Sciences, 1998 Notes on the Late Prehistoric Extension of the Range for the Muskrat (Ondatra zibethicus) along the Ancient Shoreline of Lake Cahuilla, Coachella Valley, Riverside County, California Robert M. Yohe II Archaeological Survey of Idaho, Idaho State Historical Society Boise, Idaho 83702 In 1990, a series of archaeological excavations was conducted by the Archae- ological Research Unit (ARU), University of California, as part of environmental assessments prepared in anticipation of several proposed development projects in La Quinta, California. La Quinta, located in the northwestern Coachella Valley, is 24 km (15 miles) southwest of Palm Springs in central Riverside County (Fig. 1). Three archaeological sites in La Quinta, CA-RIV-3682, CA-RIV-3144 and CA- RIV-1182, yielded collections of subfossil vertebrate remains (interpreted as food refuse) in addition to an array of cultural materials. These three archaeological sites, as well as many others in the region, are believed to represent small fishing/ lacustrine resource gathering encampments located along the shore of prehistoric Lake Cahuilla (also known as Lake LaConte; see Wilke 1978). This lake filled the Salton Basin up to the Coachella Valley at various times until the latter part of the sixteenth century (Weide 1976; Wilke 1978; Figure 1). At maximum fill, Lake Cahuilla was a veritable inland sea with an estimated surface area of 1,256,550 acres (Weide 1976). The three prehistoric sites and associated subfossil assemblages are thought to date to A.D. 1300 to 1500 based on several radiometric analyses of fire-hearth samples (Arkush 1990; Yohe 1990), a period that represents the last stand of Lake Cahuilla. The analysis of the vertebrate faunal assemblages from the three archaeological localities (conducted by the author) revealed a wide range of aquatic and terrestrial taxa, the former confined to CA-RIV-3682. A summary of taxa identified at the sites include several species of freshwater fishes (Xyrauchen texanus, Gila spp., Mugil cephalus, Ptychocheilus lucius, Elops affinis), reptiles (Gopherus agassizi, Dipsosaurus dorsalis, Sceloperus sp., Crotalus sp.) some birds (Fulica americana, cf. Anas acuta, Passeriformes), and numerous mammals, with the Audubon cot- tontail (Sylvilagus audubonii) and woodrat (Neotoma sp.) dominating the mam- malian assemblage (Yohe 1990). Of particular interest among the mammals at these three sites are the remains of muskrat (Ondatra zibethicus). This is signif- icant since the present range for this species is 130 km southeast of La Quinta (Cockrum 1960; Grinnell et al. 1937; Ingles 1965; Willner et al. 1980). At all three sites the Ondatra skeletal elements are rare, consisting of isolated mandib- ular and maxillary molars (n = 8; CA-RIV-3682), assorted postcrania (1 ulna, 1 radius, 1 metapodial, 2 phalanges [CA-RIV-3144]); and a partial rostrum and palate with complete molar series identified from CA-RIV-1182 (Arkush 1990). In historic times, the range of the closest extant subspecies of Ondatra, the Colorado muskrat (Ondatra zibethicus bernardi); has been both sides of the Col- orado River and the New River to the south (Cockrum 1960; Grinnell et al. 1937; 86 RANGE OF MUSKRAT IN RIVERSIDE COUNTY 87 " MUTT SHR ULB CA-RIV-3682, -3144, and per WYTEAAON GS AC ; Ss, -7182 Moya ve Desert RU ALLA] eT or, : % ca Pricey) my « Ig Dees ce) Biggs Ying 4 ” = “4, Qs ae pees 5 je Mins. % sy Fy AE oes! : Eag el OS ; <7 nea au) %, a4 * ww aT ny RN Aa 4 a \ Z8 ZNSE watt Mins” MH ay, F % O. zibethcus bernardi a” IW NG ig, ier My, : BW [) = mn %, eas t Ine = Www“, WN ee 3 ~N*, Orocopia Mins. 2 %, ZY S - Mle A114 wer ~ "Maem oF 4, W®Y 22 £ " Ny ny why, Syith COAT] nie WN HR Ff iaviag nen aR aa nehlly wet a \ ° Wor: My, Zoe sane N aT | Pa (2 ca "eae OT " Vy “4 2, eZ S Zt SS _ me Fy i es, \\ RIZONA ‘ Fe gs. dg M1 GA 6 ‘ Ap SA AE % SN LAKE CAHUILLA SHORELINE Opi 5 “x fi ays = ty, z Ml, afl N (A.D. 900-1500) iF" y= B 7 7 Miyw z CALIF noni mms, ‘os ig ‘ LS Wo AD Bas Sus S= ae, oes 2 en {1° % -N- % My 0 30 ; ay (eS RNI. Kilometers CALIFO ie a —_—_™ Fig. 1. Regional map of the Salton Basin and Coachella Valley, California with general locations of archaeological sites discussed in text. Dark outline represents the approximate shoreline of prehis- toric Lake Cahuilla, stippling represents the current range for Ondatra zibethicus. Ingles 1965; Willner et al. 1980). It is also found along irrigation canals and sloughs in the south end of the Imperial Valley. Prized for their pelts, the Colorado muskrat was hunted and trapped extensively in the southern Imperial Valley in the earlier part of the 20th century where it occurred in large numbers (Grinnell et al. 1937). According to Grinnell et al. (1937), approximately 25,000 muskrats were trapped in the Imperial Valley alone between 1919 and 1920. The presence of late prehistoric muskrat remains in the northwestern Coachella Valley places this genus more than 100 km north of its present accepted range. Late Pleistocene/Early Holocene Ondatra fossils have been discovered in the east- ern Salton Basin near East Mesa (Reynolds 1989), but no other paleontological occurrences in southeastern California have been reported. Ondatra fossils are rare in California. The only other known occurrence is a single femur from Cos- teau Pit (Rancholabrean-age) in the Los Angeles Basin (Miller 1971). Archae- ological occurrences of the muskrat from southern California also are rare; a few Ondatra specimens have been recovered from San Joaquin Marsh in Newport Bay at CA-Ora-119 and -193, where they were found in both cultural and natural deposits ranging in age from ca. 6000 to 750 B.P. (Langenwalter 1986). The course of the Colorado River, which has been naturally diverted on nu- merous occasions Over the past several thousands of years to produce an inland sea in the Salton Basin (Wilke 1978), has provided an environmental setting 88 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES attractive to various lacustrine vertebrates in the past. The archaeological sites containing the Ondatra remains all appear to correspond temporally with the last stand of Lake Cahuilla (A.D. 900—1500; Wilke 1978). Muskrat remains occurring in the faunal assemblages of archaeological sites associated with ancient Lake Cahuilla suggest that these animals once frequented the cattail marshes (which were Clearly present based on paleobotanical studies of human coprolite contents from numerous archaeological sites along the shoreline [see Wilke 1978; Farrell 1988]) on the northwest end of the lake, and were a source of food and possibly pelts for the prehistoric occupants of the region more than 500 years ago. Acknowledgments I thank the following people for their comments on this note: Robert Reynolds, Department of Earth Sciences, San Bernardino County Museum; Brooke S. Ar- kush, Weber State University; Greg McDonald, Hagerman Fossil Beds National Monument, Idaho; Bill Akersten, Department of Vertebrate Paleontology, Idaho Natural History Museum, Idaho State University; Susanne J. Miller, Idaho Falls, Idaho. I also thank Mark Q. Sutton for his assistance with the graphics. Literature Cited Arkush, B. S. 1990. Archaeological investigations at CA-RIV-1182, CA-RIV-3143, CA-RIV-3144, CA-RIV-3868, and CA-RIV-3882, Tentative Tract 25429, La Quinta, Central Riverside County, California. Report on file, Archaeological Research Unit, Department of Anthropology, Uni- versity of California, Riverside. Cockrum, E. L. 1960. The recent mammals of Arizona: their taxonomy and distribution. University of Arizona Press, Tucson. Farrell, N. 1988. Analysis of human coprolites from CA-RIV-1179 and CA-RIV-2827. Pp. 129-142 in Archaeological investigations at CA-RIV-1179, CA-RIV-2823, and CA-RIV-2827, La Quinta, Riverside County, California (M. Q. Sutton and P. J. Wilke, eds). Coyote Press, Salinas, Cali- fornia. Grinnell, J., J. S. Dixon, and J. M. Linsdale. 1937. Fur-bearing mammals of California: their natural history, systematic status, and relations to man. Contr. from the Mus. of Vert. Zool., University of California Press, Berkeley. Ingles, L. G. 1965. Mammals of the Pacific States: California, Oregon, and Washington. Stanford University Press, Stanford. Langenwalter, P. E. 1986. Indigenous muskrats, Ondatra zibethicus, in coastal Southern California. Calif. Fish and Game 72(2): 121-122. Miller, W. E. 1971. Pleistocene vertebrates of the Los Angeles Basin and vicinity (exclusive of Rancho La Brea). Bull. of the Los Angeles Co. Mus. of Nat. Hist., Science, No. 10. Reynolds, R. E. 1989. Paleontologic Monitoring and Salvage, Imperial Irrigation District Transmission Line, Riverside and Imperial Counties, California: Final Report. Manuscript on file, Los Angeles Co. Mus. of Nat. Hist. Wilner, G. R., G. A. Feldhamer, E. E. Zucker, and J. A. Chapman. 1980. Ondatra zibethicus. Mam- malian Species, No. 141, American Society of Mammalogists. Weide, D. 1976 Regional environmental history of the Yuha Desert. Pp. 9-20 in Background to the prehistory of the Yuha Desert Region (P. J. Wilke, ed.). Ballena Press Anthropological Papers No. 5., Ramona, California. Wilke, P. J. 1978. Late prehistoric human ecology at Lake Cahuilla, Coachella Valley, California. Contributions of the University of California Archaeological Research Facility 38. Yohe, Robert M. II, 1990. Archaeological investigations at five sites located at One Eleven La Quinta Center in the city of La Quinta, Central Riverside County, California. 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Yohe II - COVER: Photograph of a school of jacks, Selar crumenophthalmus, in Academy Bay, Soccoro island, Mexico. Photograph by Robert Pitman. 76 82 86 ISSN 0038-3872 PerdeERN CALIFORNIA ACADEMY OF . SCIENCES BULLETIN Volume 97 Number 3 BCAS-A97(3) 89-126 (1998) DECEMBER 1998 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 © Southern California Academy of Sciences, 1998 OFFICERS Robert S. Grove, President David Huckaby, Vice-President Susan E. Yoder, Secretary Robert Lavenberg, Treasurer Daniel A. Guthrie, Editor Hans Bozler, Past President David Soltz, Past President BOARD OF DIRECTORS 1997-1999 1997—2000 1998-2001 Ralph G. Appy Robert S. Grove Kathryn A. Dickson Jonathan Baskin David Huckaby Donn Gorsline J.D. Stewart Robert Lavenberg Robert E Phalen Gloria Takahashi Kenneth E. Phillips Cheryl C. Swift Susan E. 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A paper by any combination of student and professional co-authors will be considered eligible provided that it represents work done principally by student(s). In the case of an award to a co-authored paper, the monetary award and a one year student membership to the Academy will be made to the first author only. ite ee — oe era | 1998 SCAS STUDENT AWARD WINNERS (and honorable mentions*) Janelle Johnson, Pacific Estuarine Research Laboratory, San Diego State University: Best Poster Award for her poster entitled ‘“‘Fish Use of Southern California Salt Marshes” with Joy B. Zedler. Rebecca Herrick, Department of Biology, California State University, Fullerton: Jules Crane Award for the Best Paper for her paper entitled ‘“‘Do Biochemical Indices of Aerobic Capacity Correlate with Swimming Speeds in Scombrid Fishes?”’ with K. A. Dickson. Mason Posner, University of Southern California, Los Angeles County Museum of Natural History, Fishes Section: Durham Memorial Award for Vertebrate Zoology for his paper entitled ‘‘Light Atten- uating Molecules in the Lens of the Fish Eye.” Ramon M. Valencia, Biology Department, California State University, Long Beach: ARCO Award for Environmental Sciences for his paper entitled “An Assessment of the Toxicological Effects of Ingested Copper and Tungsten-Tin-Bismuth (TTB) Bullets on the California Condor (Gymnogyps californi- anus)’ with A. Silverman, A. Z. Mason, D. Clendenen, and R. Risebrough. Steve Lonhart, Department of Biology, University of California Santa Cruz: Margaret Barber Award for the Best Paper for his paper entitled “‘Comparison of Consumption Rates Among Benthic Inver- tebrate Predators.” Mauricio S. Ramos, Biological Sciences Department, California State Polytechnic University: Southern California Academy of Sciences Award for the Best Paper for his paper entitled “‘Synthesis and Cloning of Hammerhead Ribozyme Construct Targeted to Non-Variable Sequence of Mouse Leukemia Virus Reverse Transciptase Gene”’ with B. K. Pal. Maress Lacuesta, Biological Sciences Department, California State Polytechnic University: Honorable Mention Award from the Southern California Academy of Sciences for the poster entitled ““Oxidative and Glycolytic Metabolic Activities in Neurons and Microglial Cells are Altered in Rats Exposed to High Doses of Ketamine”’ with W. J. Tulpinski, and G. H. Kageyama. Junior Academy: Participants in organizations belonging to the Southern California Junior Academy of Sciences will present their papers at sessions scheduled in a way to avoid conflict with Academy meeting symposia and contributed paper sessions, thus permitting professional Academy members to attend. High school students contributing papers through the Junior Academy should contact J. D. Stewart (213) 763-3318. Registration: All individuals attending the meeting including presenters at contributed paper, poster, and symposia sessions are required to register for the meeting. However, all presenters will be offered the early registration rate for Academy members or students, whichever applies. INSTRUCTIONS FOR THE ABSTRACT/INFORMATION Abstract Guidelines: Your abstract must not exceed 250 words of text, exclusive of title, authors, and author affiliations. You must submit abstract in Microsoft Word format (any version: DOS, Mac, or Windows) as an e-mail attachment to scas@dhvx20.csudh.edu by March 1, 1999. Do not send as text in the body of an e-mail message. If you do not receive an acknowledgment by e-mail within 5 days, please resubmit your abstract. If you still do not receive an acknowledgment within 5 days, phone (310) 243-3381. No revisions will be possible after submission and acceptance. All abstracts received by March 1, 1999 will be available for review on the web site http://www.csudh.edu/scas by April 1. If you cannot possibly send an e-mail attachment, contact: John Roberts, Local Committee Chair, at (310) 243-3381 or by e-mail, jroberts@dhvx20.csudh.edu. NOTE: use 12 point Ariel font if possible. 1. Using a continuous format, type the - ¢ TITLE in all capital letters. Italicize only species names. ¢ Author(s) in upper and lower case. List all author’s names by their initials followed by surname. Place the presenting author first. Underline the names of all authors and use superscripts if the affiliation of authors varies. ¢ Affiliation of each author including institution, department, or company together with the city, state, and zipcode. Superscripts should correspond to those assigned to each author. 2. SKIP ONE LINE leaving one line of blank space between the title information and text of the abstract. 3. INDENT 5 spaces and begin the text of the abstract. AFTER ABSTRACT, SKIP 5 LINES, AND, type the following information. 1. Full name of the presenter UNDERLINED, mailing address, and phone number with area code. Include a fax number and e-mail address if available. Indicate if presenter is an Academy member or nonmember. 2. Full name(s) and mailing address(es) of all co-authors as appropriate. Indicate for each co-author if he/she is an Academy member or nonmember. 3. Indicate the status of the presenter (first author), 1.e., whether student or professional. If a student presenter, indicate eligibility and desire to compete for a student award (e.g., ““Yes, I am eligible and wish to compete or no, I do not wish to compete’’). 4. Indicate the type of presentation you are submitting, i.e., contributed paper, contributed poster, or invited symposium paper. 5. Indicate the general subject area of your contributed paper or, if you have been invited to present a symposium paper, please indicate the title of the symposium session. 6. List any audiovisual equipment required. Submission of Abstract/Information: Abstracts that fail to conform to the guidelines or that are mailed after the deadline will not appear in the program. Facsimiles are unacceptable. ABSTRACT/INFORMATION FORMS MUST BE E-MAILED NO LATER THAN MARCH 1, 1999. SAMPLE ABSTRACT & INFORMATION MICROBIAL ACTIVITY IN THE DIGESTIVE TRACT OF THE HALFMOON, Medialuna californiensis. J. S. Kandel', J. R. Paterek* and M. H. Horn’, 'California State Univ. Fullerton, CA 92634 and ?Agouron Institute, La Jolla, CA 92037. We report the presence of a diverse microbial flora and of microbial fermentation products in the hindgut region of the halfmoon, Medialuna californiensis, a seaweed- eating fish from southern California coastal waters. Viable aerobic and anaerobic bacteria were found in all sections of the gut, but were of highest concentration (10°— 10*/ml) in the hindgut. Microscopy revealed vibrios, spirilla, rod-shaped bacteria and flagellated protozoa in the midgut and hindgut, but primarily vibrios and rods in the stomach and foregut. Acetic, isobutyric and butyric acids, the volatile products of microbial breakdown of carbohydrates, were found only in the hindgut, as was eth- anol, a nonvolatile product. These results provide the first evidence for microbial fermentation and its possible contribution to the energy supply in a north-temperate herbivorous fish. . Judy S. Kandel, Department of Biology, California State University, Fullerton, Fullerton, CA 92634, 714-773-2546. FAX 714-773-3426, jkandel @fullerton.edu. Nonmember . J.R. Paterek, Agouron Institute, La Jolla, CA 92037. Nonmember Michael H. Horn, Department of Biology, California State University, Fullerton, Fullerton, CA 92634, 714-773-3707. Member . Professional . Contributed paper . Marine Biology, Microbiology, or Ichthyology . Kodak 35mm slide carousel projector Bull. Southern California Acad. Sci. 97(3), 1998, pp. 89-95 © Southern California Academy of Sciences, 1998 Two New Stenosini Species in the Genus Araeoschizus LeConte from Baja California, Mexico (Coleoptera: Tenebrionidae) Charles S. Papp 7451 Albezzia Lane, Sacramento, California 95828 The first Araeoschizus species from the northern part of the Baja California Peninsula was described by Blaisdell (1943) as antennatus collected at Punta Prieta, by E. A. Michelbacher and E. S. Ross from the California Academy of Sciences, in 1938. The most expansive collecting was done in the 1970s by E G. Andrews, A. R. Hardy, T. D. Eichlin and M. Wasbauer from the California De- partment of Food and Agriculture; their material supplied most of the specimens for my revision of the genus (1981). Also, W. H. Clark, P. H. Blom, and others from the Orma J. Smith Museum of Natural History, Albertson College, Caldwell, Idaho and the University of Idaho, Moscow contributed generously to the further study of this genus. It was a puzzle for me to classify the material William H. Clark initially col- lected in the broader San Agustin area. Subsequent collecting supplied more ma- terial (over 700 specimens) from this area, where, according to I. L. Wiggins (1980), four distinct plant communities meet: (1) the Californian Region, (2) the Baja California Coniferous Forests, bordered to the west by (3) a Microphyllous Desert habitat, and to the south by (4) the Sarcophyllous Desert Region (Fig. 1). There are two recognized subspecies from this general area: Araeoschizus antennatus clarki Papp (1989:335—337) is characterized as the more slender form. Head narrower posteriorly (more so in many specimens). Ocular lobe posterior to eye flat, not well outlined; ocular ridge shallow, with row of dense, erect to semierect squamules. Prothorax similar to A. a. antennatus, except the squamules on the longitudinal median ridge (creating the groove) and those along the margin of the prothorax are goldish yellow, erect and long, longer than those squamules of the ocular ridge. Elytral costae with dense row of some- what shorter and erect squamules; rows of squamules in the elytral interspaces are much smaller, sparsely spaced and posteriorly decumbent, like those parallel with the tightly fused sutural line. Overall dark brown; prothorax slightly darker, appendages slightly lighter in color; surface shiny. Known to occur in the Rancho Santa Inez area (550 m elev.), found by W. H. Clark in foraging columns of the ant Neivamyrmex nigrescens Cresson. Araeoschizus antennatus blaisdelli Papp (1989:338) with much paler, less dense and generally narrower squamules than A. a. clarki. The squamules at anterior half of elytra slightly thinner and somewhat roundedly pointed; in posterior half narrow and club shaped, resembling that of A. a. antennatus. Squamules on the longitudinal ridges of the elytra are shorter, sparser; those in the interspaces shorter and more sparsely spaced. Uniformly lighter brown; surface shiny. From the Ran- cho Santa Inez area (550 m elev.); also found in Valle Montevideo La Laguna Wash, 18 km W of Bahia de Los Angeles. W. H. Clark and P. E. Blom collectors. On several collecting trips of William H. Clark and his collecting companions, 89 90 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Californian Region B.C. Coniferous Forest Microphyllous Desert Sarcophyllous Desert Sarcocaulescent Desert Magdalena Region Arid Tropical Region Sierra de la Giganta Region B.C. = Baja California B.C.S.= Baja California Sur Major Plant Communities in Baja California, Mexico After |. L. Wiggins: Flora of Baja California Stanford Univ. Press, 1980 (Redrawn by C.S. Papp, 1998) Fig. 1. The eight plant communities of Baja California. The area encircled is where four com- munities meet. (After I. L. Wiggins, 1980). they were able to collect more specimens in a wider area of, as we now call it, the Four Corners (Figs. | and 2). Araeoschizus agustinus Papp, n. sp. (Fig. 3) In some respects the species resembles squamulissimus Papp (1981) from Dia- blo Dry Lake east of the Sierra de Juarez, some 50 miles W of the Colorado River delta, but the latter species is far more squamulose, head longer than broad with deeper and longer ocular groove; prothorax longer than broad and edges heavily squamulose. Dark brown, shiny throughout. Head.—Slightly (one-tenth) longer than broad, about evenly rounded. Ocular groove shallow more so posterior to eye; ridge slightly elevated, with erect and short squamules; ocular lobe similarly squamulose. Occipital triangular impression shallow, occipital region roundly elevated. Surface minutely punctured and with forwardly decumbent short pale squamules. Frontal edge fairly straight, slightly serrated, with few longer, hair-like squamules. Eyes large, almost covering the width of the ocular groove, with 20 facets dorsally and 5 ventrally. Antennae NEW SPECIES OF ARAEOSCHIZUS FROM BAJA CALIFORNIA 9] BAJA CALIFORNIA, MEXICO From El Rosario to Punta Prieta souyEL ROSARIO Thm ae 30 40 MILES 50 60 KILOMETERS iE of /” PUERTECITOS mar Wess Guayaquil E/N amen { Pico Son San Agustin xo = bs Pongo N@Rancho Sonora | eee) Rogar > \ 4 : A Pico Son Miguel aoe — tifa” are) yy 2 Ss i, / Santa Catarina £=¢, | ae “@ Santa Inez x , is erro Ugorle @ nme (asa / ec 164 .,, Lodos Santos 3 aay @) = The Catavina-Santa Inez area the major collecting site of the Clark’s Fig. 2. The major localities mentioned in the text. robust; segments with row of forwardly decumbent narrow squamules on anterior margins of all segments with thinner squamules on sides. Prothorax.—About as long as broad (occasionally very slightly longer), ante- rior margin broader with well defined anterior pronotal angle; moderately con- stricted posteriorly with short pronotal angle. Longitudinal groove shallow and relatively broad; ridges with semierect narrow squamules with posterior end of ridges longer, rosette-like. Edge densely squamulose, about the size of squamules on longitudinal ridge. Surface granulose, with sporadically spaced forwardly de- cumbent squamules shorter than those on margin. Elytra.—About one-third longer than head and prothorax combined. Sides in middle two-thirds parallel; shoulders broadly, posterior end more narrowly round- ed. Longitudinal ridges prominent, sharply elevated, on ridge with posteriorly decumbent, curved, narrow squamules. Puncture lines very prominent; there are no secondary rows of squamules. Sutural line shallow, with a row of somewhat shorter and more sparsely spaced squamules than on longitudinal ridges. ® 92 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES hes Fig. 3. Araeoschizus agustinus Papp, n. sp. Underside Head: Basal groove of sublabial plate deep; proboscis very shallow, frontal margin straight. Gular impression very shallow (almost non-existent). Surface coarsely punctate and very sparsely covered with forwardly decumbent short squamules. Prothorax: Surface with larger punctures than head; prosternal ridge evenly rounded; prosternal process narrow and more squamulose than rest of prothorax. Hind body: Surface with large, deep punctures, each puncture with a posteriorly decumbent short and thin squamula; these somewhat longer toward posterior end of body. Legs: Medium pair smallest, posterior pair largest; sparsely squamulose throughout; tarsi with somewhat longer, hair-like squamules. Length: 4.1—4.5 mm. Distribution Holotype: San Agustin, elevation 580 m, in Ethylene Glycol Pitfall Trap (EGPT), VI. 16. 1991 to V. 27. 1992, William H. Clark, Paul E. Blom and Ellen M. Clark collectors. In the Orma J. Smith Museum. Paratypes: 12 specimens from the same location (in EGPT). Additional specimens (all in EGPT): 3 from the same location, VI. 20. 1990— NEW SPECIES OF ARAEOSCHIZUS FROM BAJA CALIFORNIA oS III. 10. 1991 by W. H. and Ellen M. Clark collectors.—14 from 1 mi N of Santa Catarina (Ranch), XII. 9. 1991—VIII. 3. 1992 by W. H. Clark and P. E. Blom collectors.—8 from 1.5 km SW from Guayaquil, elev. 600 m, VI. 16. 1991—V. 27. 1992 by W. H. & E. Clark, P E. Blom and David M. Ward collectors.—10 from 10 km SE Rancho Sonora, elev. 600 m, III. 12. 1991—VII. 16. 1991 by W. H., M. H., C. J. & K. D. Clark and Jane C. Luther collectors.—2 from 9 km NW Santa Inez, VII. 17. 1991—V. 26. 1992 by W. H. and E. M. Clark collectors.—1 from 2 km E Mission San Fernando, elev. 480 m III. 12. 1991—VII. 3. 1991.—7 from 11 km ENE El Rosario, elev. 140 m VI. 22. 1991 -III. 9. 1992.—1 from Valle Montevideo Wash, 18 km W Bahia de Los Angeles, elev. 380 m, III. 19. 1991-VIII. 19. 1991.—1 from Rancho La Ramona, elev. 500 m, III. 21. 1991— VII. 3. 1991.—4 from 2 km E Mission San Fernando, elev. 480 m, VII. 3. 1991— V. 20. 1992 by W. H. Clark collector. Araeoschizus blomi Papp, n. sp. (Fig. 4) Resembles antennatus Blaisdell (1943), however blomi can easily be differ- entiated by the robust antennae, the narrow posterior portion of head, the more prominent longitudinal groove of prothorax and the unique arrangement of squa- mules. Secondary row of squamules hardly detectable. Brown to blackish brown, shiny; also smaller. Head.—A\lmost twice as long as prothorax; occipital portion narrowly rounded with prominent, yet small, occipital impression. Ocular lobes only slightly ele- vated, rounded, inner ocular ridge angularly placed (parallel to margin of head in antennatus), short, slightly elevated with prominent row of decumbent squamules. Ocular groove short, abruptly flattened posteriorly. Vertex round, evenly elevated, a slight horizontal impression between ocular lobes separates it from the nearly flat frons. Surface finely punctured, with forwardly decumbent squamules. Sides with erect longer squamules, more sparsely spaced on anterior margin. Frontal margin almost straight, with several semierect spine-like squamules. Eyes with 14-16 facets dorsally, with 5—6 facets ventrally. Antennae more robust; joints are Squamulose, more densely so on anterior margin of each segment. Prothorax.—Anterior margin slightly curved inwardly, angles more narrowly rounded than that of antennatus, posterior third constricted. Longitudinal groove evenly deep, one third as wide as length of posterior margin of prothorax; finely punctured. Ridges with long, erect squamules (see top insert of Fig. 4), anterior and posterior end slightly decumbent, as long as squamules on margin of protho- rax, which are on sides horizontally, on anterior and posterior margin vertically erect, on the latter somewhat shorter, dense, more numerous. Surface finely punc- tured, with very few forwardly decumbent short squamules. Elytra.—Slightly longer than head and prothorax combined. Shoulders narrow- ly rounded, sides in mid-third almost parallel. Primary cordae prominent, on fron- tal fourth longer, erect, other places with shorter and posteriorly decumbent, slight- ly club-shaped squamules. Puncture lines are prominent, punctures deep, closely Spaced; secondary row of squamules between them hardly detectable, consists of Short, thin, sporadically spaced, posteriorly decumbent squamules. Sutural line Slightly elevated and with very short, thin, posteriorly decumbent squamules. 94 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Sint ad -F a 98 7) as * NogS a YY ' ax S yy v3. EAD S iS 7a WAP Pe § N BY "4 q ( 4 § yy; ie C73 yt SS Am) ye aays IA Abe ot RAPES alae AIK AY A\ te pie) d AS Fig. 4. Araeoschizus blomi Papp, n. sp. Underside Head: Densely punctured; with few forwardly decumbent squamules. Basal groove of sublabial plate deep, posteriorly extended into a rounded-triangularly shaped impression about half way to the very deep gular impression. Margin of sublabial plate straight; wide, proboscis long, sharply pointed, at base deeply carinate. Prothorax: Prosternal ridge highly elevated, with few, very short, erect squamules on ridge. Coarsely punctured. Prosternal process broad and with round- ed posterior margin; squamulose. Hind body: With large, closely fit punctures, each puncture with narrow, posteriorly decumbent thin squamules longer than diameter of punctures. Legs: Middle pair smallest, hind pair largest; covered with relatively long narrow squamules, these thinner and longer toward tip of tibia; longer and thinner on tarsi. Length: 3.8—4.1 mm. Distribution Holotype: 9 mi N of Rancho Santa Inez, elevation 550 m, in EGPT, III. 18. 1991—VII. 17. 1991, by W. H., Mary H., Cynthia J. & Caren D. Clark and James C. Luther collectors. In the Orma J. Smith Museum of Natural History. NEW SPECIES OF ARAEOSCHIZUS FROM BAJA CALIFORNIA 95 Paratypes: 12 specimens, same location, same collectors; in the same Museum. Additional specimens (all in EGPT): 45 from San Agustin, elevation 580 m, III. 10. 1991—VII. 16. 1991, same collectors; additional 21 specimens from the same location XII. 21. 1988—VIII. 29. 1989 by W. H. Clark collector.—2 from 2 km SE Rancho Sonora, VII. 16. 1991—V. 27. 1992, W. H. & E. M. Clark and P. E. Blom collectors.—3 from 5 km SW Guayaquil, 600 m elevation, III. 23. 1991— VIII. 3. 1991 and VII. 3. 1991—V. 27. 1992 by the same collectors.—3 from Santa Inez, III. 13. 1991—VII. 17. 1991 by W. H. and Mary H. Clark collectors; 11 specimens VII. 4. 1991-I. 4-5, 1992 by W. H. Clark and P. E. Blom collectors. It will be interesting to see what specimens the Clarks come up with south from Santa Inez. No doubt, antennatus was reported in the above mentioned general area (Papp, 1981:324) 13 mi E from El Rosario, collected by G. E. & E. S. Ross and V. L. Vesterby in 1938 and later by W. H. Clark in 1978 to recently. It would be interesting to find out the distribution of this species further to the south and behind the type locality, Punta Prieta. I believe antennatus is the dom- inant species in the center two-thirds of Baja California. In the collection of the California Academy of Sciences there is a specimen from Baja California Sur with spines on all femora and with secondary rows of squamules on elytra. The specimen was collected by S. C. Williams in an isolated area at San Migual de Comundu at 1500 ft. elevation on April 21, 1969. This area should be intensively collected. This unique specimen belongs to Group I in the key (Papp, 1981:295), the first ever collected in the southern portion of the Baja California Peninsula. Literature Cited Blaisdell, EF E. 1943. Contribution Toward the Knowledge of the Insect Fauna of Lower California. No. 7: Coleoptera, Tenebrionidae. Proc. Calif. Acad. Sci. 24(7):171—188, pls. 10 and 11. Papp, C. S. 1981. Revision of the Genus Araeoschizus LeConte (Coleoptera: Tenebrionidae). Ent. Arb. Mus. Frey, 29:273—420, 68 figs. . 1989. Notes on the Stenosini genus Araeoschizus LeConte from Baja California, Mexico (Coleoptera: Tenebrionidae). Entomography, 6:335—340, 3 figs. Wiggins, I. L. 1980. Flora of Baja California. Stanford University Press, 1025 pp., 970 figs. (see pp. 21-26). Accepted for publication 7 May 1998. Bull. Southern California Acad. Sci. 97(3), 1998, pp. 96-103 © Southern California Academy of Sciences, 1998 Distribution and ‘Taxonomic Remarks for Five Crab Species of the Family Grapsidae (Crustacea: Sesarminae and Varuninae) of the Mexican Pacific Ernesto Campos and Alma Rosa de Campos Facultad de Ciencias, Universidad Aut6noma de Baja California, Apartado Postal 2300, Ensenada, B.C., 22800 México Abstract.—The present report updates the distribution of Armases magdalenense (Rathbun, 1918), Hemigrapsus oregonensis (Dana 1851) and Goetice americanus Rathbun 1923 along the Baja California coast. Previous records of Tetragrapsus jJouyi (Rathbun 1893) to the rocky intertidal of Punta Pelicano, near Puerto Pef- asco, Sonora, are rejected. These were based on misidentifications of specimens of G. americanus. Tetragrapsus jouyi is known from salt marsh areas of Guaymas, Sonora, and Bahia de Los Angeles, Baja California (new locality). The presence of Hemigrapsus nudus in the Gulf of California is not confirmed. It undoubtedly occurs on the west coast of the Baja California Peninsula northward to Alaska, U.S.A. An identification key to the Varuninae of the East Pacific is provided. Resuimen.—E\ presente trabajo actualiza la distribuci6n de Armases magdalenense (Rathbun, 1918), Hemigrapsus oregonensis (Dana 1851) and Goetice americanus Rathbun 1923. Se rechazan los registros previos de Tetragrapsus jouyi (Rathbun 1893) para el intermareal rocoso de Punta Pelicano, cerca de Puerto Penasco, Sonora. Estos fueron identificaciones incorrectas de especimenes pertenecientes a G. americanus. Tetragrapsus jouyi se conoce para saladares de Guaymas, Sonora y Bahia de Los Angeles, Baja California (nueva localidad). No se confirma la presencia de Hemigrapsus nudus en el Golfo de California. Esta especie ocurre, con certeza, en la costa occidental de la peninsula de Baja California y hacia norte hasta Alaska, E.U.A. Se provee una clave para identificar las especies de la sub- familia Varuninae del Pacifico Oriental. The study of specimens collected in the Gulf of California and of others bor- rowed from several institutions allows us to correct and update the distribution of 5 species of grapsid crabs (1 Sesarminae and 4 Varuninae) of the East Pacific. Information presented herein update those reported by Hendrickx (1995). For each species listed, some taxonomic and ecological remarks based on the new material are presented. In addition, a comparative morphological analysis allows us to provide a key for the members of the subfamily Varuninae of the East Pacific. Material and Methods This study is largely based on material collected by the authors along the Baja California Peninsula and Sonora coast, north to parallel 31°N. Additional material came from the Invertebrate Collection (Crustacea) of the Scripps Institution of Oceanography, University of California, La Jolla, California (SIO) and the In- vertebrate Collection of the Peabody Museum of Natural History, Yale University, 96 GRAPSIDAE OF THE MEXICAN PACIFIC o7 ah en ), wreagheat y “et Fig. 1. Armases magdalenense (Rathbun 1918); A, male holotype dorsal view; B—C dactyli of the fourth walking leg (A—C from Abele 1992). New Haven, Connecticut (YPM). The Baja California records of Hemigrapsus oregonensis provided by the late John S. Garth, are based on material of the Allan Hancock Foundation, from collections made primarily by the “‘Velero IV”’, “Searcher”? and *““The Kenyon-Williams”’ expeditions. This material is in the Nat- ural History Museum of Los Angeles County (LACM). Voucher specimens are deposited in the Laboratorio de Invertebrados, Facultad de Ciencias, Universidad Aut6noma de Baja California (UABC). Other abbreviations used are: Gulf of California (GC); Baja California (BC); Baja California Sur (BCS); Sonora (SON); Departamento de Investigaci6n Cientifica y Tecnologica, Universidad de Sonora (DICTUS). Taxonomic Account Family Grapsidae MacLeay 1838 Subfamily Sesarminae Dana 1851 Armases magdalenense (Rathbun 1918) Fig. 1A—C Previously known distribution.—Bahia Magdalena, west coast of BCS and from Bahia Altata, Altata, Sinaloa (GC), to Agua Brava, Nayarit (Abele 1992; Hen- drickx 1993, 1995; Villalobos-Hiriart et al. 1989). Material examined.—20+ males and ovigerous females, Estuary of Mulegé River, Mulegé, BCS (GC), 30 Jul 1996. Remarks.—Four species of the genus Armases Abele 1992 have been recorded along the East Pacific: A. angustum (Smith 1870) (Mexico to Ecuador); A. occi- 98 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES dentale (Smith 1870) (El Salvador to Ecuador); A. gorei (Abele 1981) (Peru) and A. magdalenense (Rathbun 1918) (Mexico), (see Abele 1992). Armases magda- lenense can be separated from other species for its carapace (Fig. 1A), distinctly wider than long (cl/cb = 0.83), the extensor margin of the dactyl of the last walking leg is also armed with black spines (Fig. 1B—C) and the palm of the chela is smooth (Abele 1992). The present record extends the northern distribution limit of A. magdalenense in the GC approximately 300 km. It is a common but inconspicuous crab along banks of estuary of the Mulegé River and its habitat agrees with that reported by Hendrickx and Salgado-Barragan (1992): under dead plants on sandy substrate, in the shade of mangrove trees above the water line. Two species of fiddler crabs, Uca latimanus (Rathbun 1893) and U. crenulata crenulata (Lockington 1877), and the grapsoid crab Geograpsus lividus (H. Milne-Edwards 1837) were col- lected in the same habitat. Subfamily Varuninae H. Milne-Edwards, 1853 Goetice americanus Rathbun 1923 Fig. 2A—B Previously known distribution.—GC, Bahia San Luis Gonzaga, BC, and Guay- mas, SON; west coast of the BC Peninsula at Bahia Tortugas (=San Bartolomé), BCS (Rathbun 1923). Material examined.—100+ males and females, San Felipe and vicinity, BC, Puerto Pefiasco, SON, Bahia de Los Angeles, BC, and Bahia Concepcion, BCS, 1985-1995. Remarks.—Goetice americanus is the most abundant brachyuran crab of the highest rocky intertidal of the GC. It is a common species from Bahia de Los Angeles, BC north to San Felipe, BC and Puerto Pefiasco, SON, but is rare at Bahia Concepcion and southward along the BC peninsula coast. Hendrickx (1994) reported G. americanus to Guaymas. However, his collecting efforts along the tropical Pacific (see Hendrickx 1995) and that of ours on the west coast of the BC peninsula (1985-1997) had failed to produce specimens of this species. Our findings suggest that Rathbun’s (1923) record of G. americanus to Bahia San Bartolomé (=Bahia Tortugas) on the west coast of the BC Peninsula is extra- limital. Goetice americanus can be easily recognized by its coloration. The carapace has a marble color with a great deal of variation of white, red and gray. Ovigerous females have been collected in January, April and November. Additional remarks on this species are under Hemigrapsus nudus (Dana 1851) and Tetragrapsus jouyi (Rathbun 1893). Hemigrapsus oregonensis (Dana 1851) Fig. 3A Known distribution.—From Resurrection Bay, Alaska to Bahia San Juanico, BCS (Campos and Campos 1989); San Felipe and Bahia de Los Angeles, BC (Luke 1977). Material examined.—14 juveniles, Estero Uno, north of Campo Don Abel, San Felipe, BC, 17 Mar 1995 (UABC); 20 males, 20 females, Laguna Percebt, San Felipe, BC, several dates (UABC); one ovigerous female, Guerrero Negro, BCS, GRAPSIDAE OF THE MEXICAN PACIFIC 99 C Fig. 2. A-—B, Goetice americanus Rathbun 1923; C, Hemigrapsus nudus (Dana 1951). A, C, frontal view; B, dorsal face of the fourth walking leg. 21 Mar 1956 (LACM); 12 males, 12 females, Bahia Todos Santos, BC, 16 Apr 1980; 22 Nov 1996 (UABC); one male, two females, Bahia Tortugas, BCS, Jan- Apr 1987 (UABC); number and sex not available, Bahia San Juanico, BCS, 8 February 1955 (LACM). Remarks.—See remarks under H. nudus. Hemigrapsus nudus (Dana 1851) Fig: 2€ Known distribution.—From Yakobi Island, Alaska to Bahia Tortugas, BCS. Mexico (Garth and Abbott 1980); presumably Bahia de Los Angeles (Luke 1977). Material examined.—1 male, 1 female, Bahia Todos Santos, Ensenada, BC, no date available. 100 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 3. A, Hemigrapsus oregonensis (Dana 1851); B—C Tetragrapsus jouyi (Rathbun 1893). A, C, frontal view; B, dorsal face of the fourth walking leg. Remarks.—Rathbun (1923), Brusca (1980) and Garth and Abbott (1980) were hesitant about records of H. oregonensis and H. nudus in the Gulf of California. Hendrickx (1995), based on Luke’s (1977) account, recorded these species to San Felipe, BC and Bahia de Los Angeles, BC respectively. The study of the material on which Luke (1977) based the records of H. oregonensis (SIO: C373, C375, C379, C1334) permits us to confirm the presence of this species in the upper Gulf of California. However, part of this material belongs in Goetice americanus. The presence of H. nudus is not confirmed by the present study. The only lot of specimens (SIO—C366) that supports Luke’s report was not found in the SIO Crustacean Collection (C366). On the northeast Pacific coast Hemigrapsus nudus lives under rocks in the middle and higher intertidal. Hemigrapsus oregonensis, a burrower species, inhabits open mud flats, under rocks in muddy habitats, or in mats of Enteromorpha and beds of Zostera, high to low intertidal (see Garth and Abbott 1980; Bonfil et al. 1992). GRAPSIDAE OF THE MEXICAN PACIFIC 101 Tetragrapsus jouyi (Rathbun, 1893) Fig. 3B—C Known distribution.—GC, Guaymas, SON (Rathbun, 1918); ‘‘quiet waters from Puerto Pefiasco to Mazatlan, Espititu Santo Island and San Francisco and La Paz,”’ Mexico (Vogel 1966; Brusca 1980). Material examined.—40+ specimens, Bahia de Los Angeles, BC, Mar 1987 and Jul 1996. Remarks.—Tetragrapsus jouyi was originally recorded in Guaymas, SON (Rathbun 1918). Later, Vogel (1966) reported this species in Punta Pelicano, a locality close to Puerto Penasco, SON. Brusca (1980) pointed out that it is a common and abundant species throughout the Gulf of California, living under rocks. We have examined the 25 males and 13 females (YPM 5698) on which Vogel (1966) based her report and they belong in Goetice americanus. The spec- imens reported by Brusca (1980) were not found in the Puerto Pefiasco Laboratory of the DICTUS or elsewhere. Their identity remains uncertain. Brusca (in litt.) informed us that many of his records, including that of T. jowyi, were based solely on field identifications, which further complicates this inquiry. We believe that the grapsoid crab Brusca (1980) recorded as common and abundant under rocks along the Gulf of California is either G. americanus, H. oregonensis or both. Our conclusion is supported by the fact that 7. jouyi occurs intertidally in salt marsh habitat. In Bahia de Los Angeles, it burrows among the pickle weeds (Salicornia pacifica) and grasses (Distichiis spicata). Tetragrapsus jouyi never occurs under rocks as do G. americanus and H. oregonensis. Varuninae of the Mexican Pacific.—Six species of Varuninae occur along the East Pacific coast. Except for Glyptograpsus impressus Smith 1870 (Acapulco, Mexico to Panama) and Euchirograpsus pacificus Tiirkay 1975 (Galapagos) (see Tiirkay 1975; Hendrickx 1995), the remaining species G. americanus, H. nudus, H. orego- nensis and T. jouyi occur intertidally in temperate and sub-tropical waters of the Mexican Pacific. They inhabit the SON and BC coast, in the upper GC and the west coast of the BC Peninsula along the Californian Province. The Varuninae of the Mexican Pacific are morphologically similar, particularly in the general shape of the carapace, third maxilliped, and chelipeds. This similarity among these species has resulted in misidentifications. A detailed morphological comparison of these species allowed us to recognize several features of the carapace, abdomen and walking legs that permit easy recognition of each species. These features have been summarized in the key below. Regarding habitat, H. oregonensis and T. jouyi prefer salt marsh areas. However, the former may also live under rocks in muddy habitats. Hemigrap- sus nudus and G. americanus inhabit rocky intertidal areas. Key to the Grapsidae-Varuninae of the East Pacific 1. First segment of male abdomen covering entire sternum between legs of TAS DAMME re PA a RE NER ae os oo ss ent one Pian em Sale oe Biel Baw eee 3) 1’. First segment of male abdomen not entirely covering the sternum heres Fe SemO lel Aste caliypes yao aN. cu acer ao Biel ae eae Sub RS ae wee ee 2 2. Walking legs 1—4 stout, bare or with scatter, short tufts of hair setae (Fig. OED ee verre Pete REIN ooo cae ds Ae Oe Bole eh wie MlAew team tied 3 Zee Walking legs 1-4 slender and hairy (Fig. 3B). .........0.0.20205..7.5-. 4 102 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 4. A, Euchirograpsus pacificus Tiirkay 1975, male holotype; B, Glyptograpsus impressus Smith 1870, male dorsal view (A from Tiirkay 1975; B, Rathbun 1918 respectively). 3. Antero-lateral margins of carapace straight and parallel, front deeply emarcinate, (Fil. DA) Wi Ale he ee, Goetice americanus Rathbun 1923 3’. Antero-lateral margins arcuate, front gently emarginate (Fig. 2C) ..... MA EIR La Pet See a: a Hemigrapsus nudus (Dana 1851) 4. Front deeply emarginate, with two prominent dorsal lobes (Fig. 3A) .. S Loaweeaiah, See saeets Wire SEES SE AEG. RH Hemigrapsus oregonensis (Dana 1851) 4’. Front gently emarginate, without prominent dorsal lobes (Fig. 3C) .... ste Gees AP AEA ES Tetragrapsus jouyi (Rathbun 1893) 5. Carapace squarish, lateral margins straight (Fig. 4A) ................ iteyen etek: th Mie ch aes, eee ee Euchirograpsus pacificus Tiirkay 1975 5’. Carapace subrotund, lateral margins arcuate (Fig. 4B) ............... ih deat cox Dede oe ene Glyptograpsus impressus Smith, 1870 GRAPSIDAE OF THE MEXICAN PACIFIC 103 Acknowledgments Our great appreciation is due to Eric Lazo-Wasem, Manager Collection, Pea- body Museum of Natural History, Yale University, for making available part of the reported material; to Richard C. Brusca, Michel Hendrickx, Raymond B. Man- ning and Michael Tiirkay for supplying pertinent literature and sharing important information; to José Delgadillo for the identification of the salt marsh plants; to Eduardo, Carmina, Lalito and Paulina Aguirre y Gonzalez, and Tony Resendis for their invaluable support during an academic sojourn at Puerto Pefiasco Lab- oratory, DICTUS and The Archleon campus, Bahia de Los Angeles respectively. This work was supported by the agreement CONACyT-UABC 3587-N9311. Literature Cited Abele, L. G. 1992. A review of the grapsid crab genus Sesarma (Crustacea: Decapoda: Grapsidae) in America, with the description of a new genus. Smith. Contr. Zool. 527:1—60. Bonfil, R., A. Carvacho, and E. Campos. 1992. Los cangrejos de la Bahia de Todos Santos, Baja California. Parte II. Grapsidae, Pinnotheridae y Ocypodidae (Crustacea: Decapoda: Brachyura). Ciencias Marinas 18(3):37—56. Brusca, R. C. 1980. Common intertidal invertebrates of the Gulf of California. University of Arizona Press, Tucson, 513 pp. Campos, E. and A. R. de Campos. 1989. Range extension of Decapod Crustaceans from Bahia Tor- tugas and vicinity, Baja California Sur, Mexico. Calif. Fish and Game 75:174-177. Garth, J. S., and D. T. Abbott. 1980. Brachyura. The true crabs, chapter 25. Pp. 594-630 in Intertidal Invertebrates of California. (R. H. Morris, D. P. Abbott, and E. C. Haderlie, eds.), Stanford University Press, Stanford, California. Hendrickx, M. E. 1993. Crustaceos decapodos del Pacifico mexicano. Pp. 271-318 in Biodiversidad Marina y costera de México. S. I. Salazar-Vallejo and N. E. Gonzalez (eds.). CONABIO- CIQRO, México. . 1994. Catalogo de crustaceos estomat6podos y decapodos. Coleccién de referencia, Estacion Mazatlan. Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Aut6noma de México-Comisi6n Nacional para el Conocimiento y uso de la Biodiversidad, 134 pp. . 1995. Checklist of brachyuran crabs (Crustacea: Decapoda) from the eastern tropical Pacific. Bull. Inst. Roy. Sci. Nat. Belgique (Biologie) 65:125—150. , and J. Salgado-Barragan. 1992. New records of two species of brachyuran crabs (Crustacea: Decapoda: Brachyura) associated with tropical coastal lagoons on the Pacific coast of Mexico. Rev. Biol. Trop. 40(1):149—150. Luke, S. R. 1977. Catalog of the benthic invertebrate collection, I. Decapod Crustacea and Stomato- poda. University of California, Scripps Institution of Oceanography. Reference Series 77—9: 72 pp. Rathbun, M. J. 1918. The grapsoid crabs of America. Bull. U.S. Nat. Mus. 97:1—461. . 1923. Scientific results of the expedition to the Gulf of California by the U.S. Fisheries Steamer ‘“‘Albatross”’ in 1911. XIII. The brachyuran crabs collected by the U.S. Fisheries Steam- er “Albatross” in 1911, chiefly on the west coast of Mexico. Bull. Amer. Mus. Nat. Hist. 48: 619-637. Tiirkay, M. 1975. Zur Kenntnis der Gattung Euchirograpsus mit Bemerkungen zu Brachygrapsus und Litocheria (Crustacea: Decapoda). Senckenbergiana Biol. 56(1/3):103—132. Villalobos-Hiriart J. L., J. C. Nates-Rodriguez, A. Cantu-Diaz-Barriga, M. D. Valle-Martinez, P. Flores- Hernandez, E. Lira-Fernandez, and P. Schmidtsdorf-Valencia. 1989. Listados Faunjisticos de México. I. Crustaéceos estomatépodos y decapodos intermareales de las Islas del Golfo de California, México. Instituto de Biologia, Universidad Nacional Aut6noma de México, 114 pp. Vogel, B. R. 1966. A report on a collection of crabs from the Gulf of California. Southwest. Nat. 11(1):139-140. Accepted for publication 29 December 1997. Bull. Southern California Acad. Sci. 97(3), 1998, pp. 104-109 © Southern California Academy of Sciences, 1998 A Dietary Analysis of Hippoglossina stomata Eigenmann and Eigenmann, 1980 (Pisces: Bothidae) along the Western Coast of Baja California, Mexico Ricardo R. Murillo, A. A. Ortega-Salas, and Marco A. Martinez-Mufioz Instituto de Ciencias del Mar y Limnologia, UNAM, México 04510, D.F. Abstract.—A benthic trawl survey was conducted at depths of 38 to 218 m in September of 1990 along the western coast of Baja California on board the R/V E] Puma. A dietary analysis of 67 Hippoglossina stomata stomachs was made in order to contribute to the knowledge of the diet of this species. Crustaceans, principally Pleuroncodes planipes (44.8%), and stomatopods, Hemisquilla ensi- gera californiensis (41.3%), were the most important prey items. Small crusta- ceans such as Malacostracea, Penaeidae, Decapoda not identified represent 5%. Fish and others were also consumed (8.4%). The geographical and bathymetric range of the bigmouth sole, Hippoglossina stomata spans from Monterey Bay, California, USA to Cabo San Lucas and into the Gulf of California, México (Roedel 1953; Berdegué 1956; Eschmeyer et al. 1983) at depths from 30 to 240 m (Martinez and Ramirez 1992). The bigmouth sole is a relatively abundant flatfish inhabiting soft sediments on the continental shelf of southern Baja California, Mexico. It could be of com- mercial importance (300 mm) due to the excellent quality of its meat (Berdegué 1956). Ecologically it is important since it preys on mysids, gammarideans and amphipods (Allen 1982), as well as the red crab, Pleuroncodes planipes (Ramirez- Murillo 1995). In turn, this species serves as food for the California sea lion, Zalophus californianus (Aurioles et al. 1984). The taxonomy of the genus Hippoglossina was studied by De Buen (1961). Leonard (1971) studied the larvae of Hippoglossina oblonga. Yany et al. (1977) surveyed the food intake of Hippoglossina macrops in Valparaiso. Goldberg (1982) studied the seasonal spawning cycles of Hippoglossina stomata in Mag- dalena Bay, Mexico and Ramirez-Murillo (1995) examined the age and growth of H. stomata in Baja California. The purpose of the present study was to provide a preliminary knowledge of the food intake of H. stomata, off the western coast of Baja California, Mexico. Materials and Methods During September of 1990, a demersal trawl survey was conducted along the western continental shelf off Baja California from the northern portion of Bahia Vizcaino to the southern part of Magdalena Bay, between 24° and 28°30’N latitude and 111°30’ and 114°30’W longitude (Fig. 1). The samples were collected by the R/V El Puma, at depths between 38 and 218 m using a shrimp otter trawl] net, with a mouth opening of 21 m and 3 cm 104 DIETARY ANAYLSIS OF HIPPOGLOSSINA OFF BAJA 105 Hippoglossina stomata 114° Fig. 1. Sampling stations from the EP9009 cruise, September, 1990. mesh size. The catch was discharged on the deck and its contents counted and identified. The H. stomata specimens were separated into plastic bags and fixed in 10% formalin for later transport to the laboratory. The dietary analysis involved the following: 1. Stomachs were extracted, fixed and maintained in 10% formalin. 2. Food items were identified and counted. Those too difficult to identify were considered unknown remains and were not considered in the analysis. 3. State of digestion of prey was noted according to the method of Banner (1948a, b) and Brusca (1980). 4. Numeric, volumetric, frequency of occurrence and relative importance (IRI) of each taxon to the diet of these fish were determined using the methods of Pinkas et al. (1971): IRI = (7 N + % V)(FO) 106 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES where: N: numeric percentage; V: volumetric percentage; FO: frequency of occur- rence. Results Of the 30 trawling operations carried out off the west coast of Baja California, H. stomata was present in 25 hauls (83% FO), rendering a total of 450 specimens with the greatest abundance in the area between the 24th and 27th N parallel. The minimum standard length of the fish sampled was 70 mm and the maximum 258 mm with an average of 171 mm. The main species associated with H. stomata were Prionotus stephanophrys, Citharichthys xanthostigma, C. fragilis, Merluc- clus augustimanus, and Synodus lucioceps. Among the total of 67 H. stomata stomachs sampled, 11 were empty (16.4%). The remaining 56 stomachs belonged to individuals of standard length between 87 and 125 mm, with an average of 106.8 mm. Analysis of the diet indicates that H. stomata feeds on a benthopelagic fauna as well as an epibenthic one. The stomachs analyzed are from relatively small and immature individuals. Crustaceans were the most important prey groups over- all in the diet of this species, followed by osteichthyes and others (Table 1). Among the 105 prey items observed, 7 families, 6 genera and 2 species could be identified by means of the analysis. The most abundant food item was the red crab, Pleuroncodes planipes with 45.4% V, 29.8% FO and 44.8% IRI, while in some stations the stomatopod, Hemisquilla ensigera californiensis was more abundant, with 29.8% V, 30.3% FO and 41.34% IRI. It does suggest some trends, when the distribution of Pleuroncodes is juxtaposed with the shift in dominance of the diet from Pleuroncodes to stomatopods. Preference for eating the red crab, Pleuroncodes planipes (44.8% IRI) occurs when this species is most abundant. When P. planipes is absent, it is replaced in the diet by other species, like Hemisquilla ensigera californiensis (41.34% IRI), which live in shallower waters, and by other species, including fish, which total 13% IRI. Thus, diet diversifies based on food availability. As indicated by its large mouth and eyes and by its tooth type, Hippoglossina stomata feeds by settling to the bottom and waylaying its food. Discussion Studies of ecological communities are based on the organisms and their envi- ronmental relationship, which could be observed by analysing the feeding habits, selection of prey, transportation of energy, and nutrients. Methods and habits of food intake are highly related to internal and external morphology of the organism (Cailliet et al. 1986). Frey (1971) states that the young flatfish settle on the bottom, eat small crus- taceans, polychaetes, molluscs and fish, but, as they grow, they eat larger food items of the same groups. In this paper, more than 91.5% of the crustacean, Pleuroncodes planipes (44.8%) are the food intake of H. stomata on the Pacific coast of Baja California. Allen (1982) found that bigmouth sole, H. stomata and California halibut, Par- alichthys californicus eat mysids, gammarideans and amphipods. He did not men- 107 DIETARY ANAYLSIS OF HIPPOGLOSSINA OFF BAJA 001 6 VEEP O01 60'OF OO COI K It's CLT8E v'6l OE! v9 86°C ao vl 13y10 Z1'0 EGS el OT CG 01 Ol I ‘dds sdysyoroyny AVGIAHLHOIIVuvVd SHAHLHOISLSO OL'0 91 O£ 97 O; Or 091 OL 8 ‘dds uowavjvd AV CINOWAY 1Vd OMT 883° LP SE OS 8°8 ES BE v ‘dds piuo4Kr1g AVGIAVNdd O£ 0 brit 97 07 Cl 090 67 € ‘dds saouvy AVCIMONVO VdOdv0Odd ETE 00°LrI Cyl xs 90 S70 06 07 VACGIONLITVL VdOdIHdNV 08 tr € SENT 8°67 0°07 vSP 07’ 8I 6°77 v7 sadiunjd sapoouosnald AVdIAHLV IVD VUNWONV per 0'9S8I €O€ OW? 8°67 C61 C67 I€ SISUaIUOfI]DI viasisua vyjinbsiuay AVGITIINOSINGAH VdOdOLVWOLS VAOVLSNYO VdGOdOUHLYV Tel % Tal OA % Od A % A ‘ON % ‘ON ‘(Qouvyodull sATe[I1 JO XOpul TY] pue sous1mn9900 jo Aouonbory :Q4 ‘[WE UT oUINTOA :A ‘suIsTURZIO JO JaquINU :ON) INS eIUIOFTeD eleg ul VIDWOIS DUISsOo]soddiy Kq peuinsuods AdIq “[ IGeL 108 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES tion, however, that they eat red crab and stomatopods as well, possibly because his samples were taken in shallower waters. Also Allen (op. cit) established 3 groups according to length: A, within the range of 6.3 to 10.0 cm; B, 11.4 to 18.9 cm and C, 19.2 to 30.8 cm. The first and second group consisted of immature fish and the main food intake were mysids and gammarideans. He also mentioned that most of the food was taken from the bottom. In the present paper the size of the fish could be compared to groups A and B, but here they prefer to eat anomurs and stomatopods. Haig in 1955 (in Yany 1977) mentioned that Pleuroncodes monodon is found on the coast of Chile which spans to Ancud, and Pleuroncodes planipes appears on the Mexican coast off Baja California. In the study area P. planipes was found between 24° and 27°N latitude, although it was more abundant at 27°N (Aurioles- Gamboa 1995). According to Yany et al. (1977), Hippoglossina macrops has as preferred food the crustaceans, “‘langostino amarillo,’? Cervimunida johni in San Antonio, Punta Gallo, Laguna Verde, Concon, Quintero and Papudo; and Pleu- roncodes monodon in Mejillones, Chile (Tomicic 1973 in Yany et al. 1977). This could indicate a close trophic relationship with similar taxa between the feeding habits of these flatfish representative of the Galatheidae family, which could be due to the distribution and abundance of these species. In either case, when the red crab is abundant in the coastal areas of Mexico and Chile, flatfish feed pref- erentially on this resource and on other additional groups when red crabs are scarce. The main food source for H. stomata in Mexico is the red crab, Pleuroncodes planipes, and for H. macrops in Chile, the closely related P. monodon, due to the great abundance of these crustaceans. However, to the south of the 27th North parallel, Hemisquilla ensigera californiensis, which is less abundant than the red crab, is always found in the stomach content of H. stomata together with other species. Acknowledgments This study was partially supported by the ‘‘Consejo Nacional de Ciencia y Tecnologia de México,”’ grant: P22OCCOR880518, by the “‘Universidad Nacional Auténoma de México” for providing the R/V “‘E] Puma”’ from 1988 to 1991, by the “Instituto de Ciencias del Mar y Limnologia’’ for the analysis of the infor- mation, and by the ‘‘Centro de Investigaciones Biolégicas del Noroeste,’’ which processed the raw material. Literature Cited Allen, M. J. 1982. Functional structure of soft bottom fish communities of the southern California shelf. PhD dissertation, Univ. Calif. San Diego, La Jolla, Ca. 577 pp. (Univ. Microfilm Inst. Ann Arbor, MI. ref. 830091). Aurioles-Gamboa, D. 1995. Distribucién y abundancia de la langostilla bent6nica (Pleuroncodes pla- nipes) en la plataforma continental de la costa oeste de Baja California. In Aurioles-Gamboa, D. y E. E Balart (eds). La Langostilla: Biologia, Ecologia y Aprovechamiento. Centro de Investigaciones Bioldgicas del Noroeste, S.C. 59-78. Aurioles, D., C. Fox, E Sinsel and G. Tanos. 1984. Prey of the California sea lion (Zalophus califor- nianus) in the Bay of La Paz, Baja California Sur, México. J. Mammal. 65:519-521. Banner, A. H. 1948a. A taxonomic study of the Mysidacea y Euphausiacea (Crustacea) of the North- eastern Pacific. Part I. Trans. Royal Canada Inst. (26): 347 pp. DIETARY ANAYLSIS OF HIPPOGLOSSINA OFF BAJA 109 . 1948b. A taxonomic study of the Mysidacea y Euphausiacea (Crustacea) of the Northeastern Pacific. Part II. Trans. Royal Canada Inst. (27):65—1235. Berdegué, A. J. 1956. Peces de importancia comercial en la costa noroccidental de México. Dir. Gral. Pesca e Industrias conexas. Cop. Fon. Pisc. Rural. Secretarfa de Marina. 345 pp. Brusca, R. C. 1980. Common intertidal invertebrates of the Gulf of California. 2nd Edition. The University of Arizona Press. Tucson, Arizona. 513 pp. De Buen, FE 1961. Peces Chilenos. Estaci6n de Biologia Marina de Montemar, Chile. Rev. Bidl. Marina, 1:1—52. Cailliet, M. G., M. S. Love and A. W. Ebeling. 1986. Fishes: a field and laboratory manual on their identification and natural history. Wadsworth Publishing Co. USA. 194 pp. Eschmeyer, N. M., E. S. Herald and H. Hammann. 1983. A field guide to Pacific Coast Fishes of North America. Houghton Mifflin Co., Boston, MA. 336 pp. Frey, H. W. 1971. California’s living marine resources and their utilization. State Calif. Res. Ag. Dep. Fish and Game. 61-69. Goldberg, S. R. 1982. Seasonal spawning cycles of two California flatfishes, Pleuronichthys verticalis (Pleuronectidae) and Hippoglossina stomata (Bothidae). Bull. Mar. Sci. 32(1):347—350. Haig, J. 1955. The Crustacean Anomura of Chile, Reports of Lunds Univ. ARSSKR, 51(12):1—60. Leonard, S. B. 1971. Larvae of the Fourspot Flounder, Hippoglossina oblonga (Pisces: Bothidae), from the Chesapeake Bight, Western North Atlantic. Copeia, 4:677—681. Martinez-Munoz, M. A. y Ramirez-Cruz, J. C. 1992. Distribuci6n y abundancia de pleuronectiformes (TELEOSTEID), en la costa occidental de Baja California Sur, México. Tesis de Licenciatura. Univ. Nal. Aut6n. México, Fac. de Ciencias. 133 pp. Pinkas, L., M. S. Oliphant y I. L. K. Iverson. 1971. Food habits of albacore bluefin tuna and bonito in California waters. Calif. Fish and Game, Fish. Bull. 152, 105 pp. Ramirez-Murillo, R. 1995. Edad y crecimiento de Hippoglossina stomata Eigenmann & Eigenmann, 1890 (Pisces:Bothidae) en la costa occidental de Baja California Sur, México. Tesis Profesional. U.M.S.N.H. 48 pp. Roedel, P- M. 1953. Common ocean fishes of California Coast. Calif. Dept. Fish and Game. Fish Bull., (91):1-184. Tomicic, K. J. 1973. Alimentacién de Hippoglossina macrops Steindachner en Mejillones (Pisces, Bothidae). Not. Mens. Mus. Nac. Hist. Nat., Santiago 205:3-7. Yany, G. G., C. Moreno y P. Ramirez. 1977. Alimentacion de Hippoglossina macrops Steindachner 1876 (Pisces: Bothidae), en la zona de Valparaiso. Cienc. y Tec. del Mar, Cona 3:23—36. Accepted for publication 29 December 1997. Bull. Southern California Acad. Sci. 97(3), 1998, pp. 110-114 © Southern California Academy of Sciences, 1998 SEM and Histological Evidence of Enlarged Nephridial Papillae in Loandala Monro (Polychaeta: Pilargidae) Sergio I. Salazar- Vallejo Depto. Ecologia Acudtica, ECOSUR, Apdo. Postal 424 Chetumal QR 77000 MEXICO Loandalia Monro and Parandalia Emerson & Fauchald are two closely allied pilargid genera. Both have a rather cylindrical body with reduced parapodia an- teriorly and enlarged parapodia posteriorly. The prostomial appendages are also similar since they both lack antennae and have bifid, often biarticulated palps. Setae are also similar; notosetae are simple spines sometimes with one or two smaller companion setae dorsally and neurosetae are spinulous capillaries. When establishing Parandalia, Emerson and Fauchald (1971) set the differ- ences between both genera. Loandalia was restricted to the type-species (L. aber- rans Monro), described from one specimen collected off Angola which lacks notopodial spines and has unusually well-developed branchiae in posterior setig- ers. Parandalia was separated from Loandalia by possessing notospines and lack- ing branchiae in posterior setigers. These authors noted, however, that Monro (1936) had described notospines though they did not find any when the type specimen was examined. The branchiae on Monro’s specimen are unusual since they are directed ventrolaterally and free from neuropodial lobes. The original designation of these structures as branchiae has been retained by other authors. The second species of Loandalia (L. maculata) was described by Intes and le Loeuff (1975) from several specimens collected off the Ivory Coast. They noticed the species was intermediate between both genera since it had emergent noto- spines and branchiae in posterior setigers. It also had an emergent ventral spine in setiger 1. These authors noted that the branchiae of L. maculata were smaller and started more posteriorly (setiger 50; body length 50 mm) than in L. aberrans (setiger 33; body length 35 mm). Salazar-Vallejo (1987) described the third spe- cies, L. riojai, from several specimens collected off Western Mexico. It has no- tospines from setiger 7 and branchiae from about setiger 22 (body length 59 mm), but it lacks emergent ventral spines in setiger 1. The fourth species, L. salazar- vallejoi, was described by de Leén-Gonzalez (1991), from three specimens col- lected off Western Baja California. It has notospines from setigers 10—13 and branchiae from setigers 31—40 (body length 67 mm); this species also lacks any emergent ventral spines in setiger 1. Thus, in the above species the presence of emergent spines in anterior neuro- podia cannot be used to set apart Loandalia Monro from Parandalia Emerson & Fauchald. Therefore, the only distinguishing feature is the presence of ventrolat- eral branchiae in Loandalia (Salazar-Vallejo 1990). Branchiae have been employed in polychaete taxonomy to set apart closely allied genera. Although there are some arguments against branchiae as a generic character (Orensanz 1990; Fauchald 1992), its utility still has some support. How- ever, if these enlarged ventrolateral structures were nephridial papillae instead of 110 NEPHRIDIAL PAPILLAE IN LOANDALIA 111 ; os ay ) a a ees Fig. 1. SEM of a posterior fragment of Loandalia riojai. A. Panoramic view of the ventral portion; B. Close-up of neurosetae (the oblique thread is cotton); C. Close-up of closed nephridial papillae; D. Close-up of open nephridial papillae (scale in D is the same as in C). branchiae, then they could not be used to separate these two genera. Since some nephridial hypertrophy is associated with reproductive activity or with sexual maturity (Schroeder & Hermans 1975), nephridial development cannot be relied upon as a discriminating feature. An early description of enlarged nephridial pa- pillae was provided by Moore (1910:369; Pl. 31, Fig. 60) when he described Polynoe (?) renotubulata. This species was later moved to a new genus, Bathy- moorea, by Pettibone (1967) due in part to its extended nephridial papillae. This Research Note presents SEM and histological evidence that the ventrolateral struc- tures in Loandalia Monro are nephridial papillae and not branchiae. Posterior fragments of Loandalia riojai Salazar-Vallejo were prepared accord- ing to standard methods for SEM and for histological analysis; some modifications were employed (Sosa-Rodriguez, 1993) from standard Hematoxylin and Eosin techniques. SEM analysis was performed in the Electronic Microscopy Unit of the Instituto de Biologia, UNAM. The histological process was performed in the Laboratory of Invertebrates, Facultad de Ciencias, UNAM. All photographs were processed in the Photography Lab. of ECOSUR. In ventral view, nephridial papillae can be seen clearly set off from neuropo- dium (Fig. 1A); each neuropodium is larger than the papillae and clearly distin- guished by the presence of neurosetae which arise from setal bundles containing 2—3 setae each (Fig. 1B). If seen from their tip, nephridial papillae appear either 112 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 2. Cross sections of posterior segments of Loandalia riojai. A. Anterior section without nephridial papillae; B. More posterior setiger with enlarged nephridial papilla (indicated by an arrow); C. Close-up of sections with nephridial ciliated funnel; D. Same, in another section; E. Close-up of a section of the enlarged nephridial papilla; F Same, in another section (Scales: A 150 wm; F 25 wm; A and B are at the same scale; C—F are at the same scale). closed (Fig. 1C), or have a distal nephridiopore opening (Fig. 1D). Since the distal pore might be an artifact of the dehydration process, histological inspection of the internal structure was employed to reveal whether these were branchiae or nephridial papillae. The papillae are not seen in cross section (Fig. 2A) of anterior segments but in more posterior setigers, a clearly digitate process emerges separate from the neuropodium (Fig. 2B). Throughout the slide series, some tissue sections show a circular ciliated structure that corresponds with the nephridial funnel and is exactly NEPHRIDIAL PAPILLAE IN LOANDALIA Li3 at the end of the lateral muscular bundles (Figs. 2C, D). More posterior slides show that the enlarged papillae are hollow (Figs. 2E, F). Some major blood ves- sels can be seen in the tissue but there are no blood vessels associated with the papillae so they cannot be branchiae. Without true branchiae, the presence of enlarged nephridial papillae cannot be employed to separate Loandalia from Parandalia. Thus the genus Parandalia is a junior synonym of Loandalia. Since there is one species with gravid females that lacks enlarged papillae (P. vivianneae Salazar-Vallejo & Reyes-Barragan, 1990), these papillae might be associated only with mature males. If some mature males lack these papillae, it might represent an alternative to sperm release. Fur- ther study will be necessary on these worms’ reproductive biology. Acknowledgments SEM work was kindly performed by Sara Fuentes and allowed by Hilda Flores, both from Instituto de Biologia, UNAM. Histological preparations were made easily by the help of Teresa Sosa, Eva Mufioz and José Luis Bortolini and allowed by Maria Ana Fernandez, Facultad de Ciencias, UNAM. Photographs were pro- cessed by Humberto Bahena from ECOSUR. This note was triggered by the interaction with Thomas Parker, Marine Biology Lab, County Sanitation Districts, Los Angeles; he also provided a complete ms on his proposal to synonymize two species of Parandalia. Careful reviews by two referees and by the editor improved the clarity of this note. The original regard of the ventrolateral branchiae in Loan- dalia as peculiar structures comes directly from Marian H. Pettibone (1986 in a letter); I’m glad she is right, again. Partial funding for this publication was pro- vided by CONACYT (4120-P). Literature Cited de Leén-Gonzalez, J. A. 1991. Poliquetos de fondos blandos de la costa occidental de Baja California Sur, México, 1. Pilargidae. Cah. Biol. Mar. 32:311—321. Emerson, R. R., and K. Fauchald. 1971. A revision of the genus Loandalia Monro with description of a new genus and species of pilargiid polychaete. Bull. So. Cal. Acad. Sci. 70:18-—22. Fauchald, K. 1992. A review of the genus Eunice (Polychaeta: Eunicidae) based upon type material. Smithson. Contr. Zool. 523:1—422. Intes, A., and P. le Loeuff. 1975. Les annélides polychetes de Cote d’Ivoire, 1. Polychétes errantes— compte rendu systématique. Cah. ORSTOM, sér. Océanogr. 13:267—321. Monro, C. C. A. 1936. Polychaete worms, 2. Discovery Rep. 12:59-198. Moore, J. P. 1910. The polychaetous annelids dredged by the U.S.S. “‘Albatross”’ off the coast of Southern California in 1904: 2. Polynoidae, Aphroditidae and Segaleonidae (sic). Proc. Acad. Nat. Sci. Phila. 62:328—402. Orensanz, J. M. 1990. The eunicemorph polychaete annelids from Antarctic and Subantarctic seas, with addenda to the Eunicemorpha of Argentina, Chile, New Zealand, Australia and the South- ern Indian Ocean. Antarctic Res. Ser. 52:1—183. Pettibone, M. H. 1967. Some bathyal polynoids from Central and Northeastern Pacific (Polychaeta: Polynoids). Proc. U.S. Natl. Mus. 121(3575):1-15. Salazar-Vallejo, S. I. 1987. Pilargidae (Annelida: Polychaeta) de México: Lista de especies, nueva especie y bio(geo)grafia. Cah. Biol. Mar. 27:193—209. . 1990. Redescriptions of Sigambra grubii Miiller, 1858 and Hermundura tricuspis Miller, 1858 from Brazil and designation of neotypes (Polychaeta: Pilargidae). J. Nat. Hist. 24:507— S17. , and M. P. Reyes-Barragan. 1990. Parandalia vivianneae and P. tricuspis (Miller), two es- tuarine polychaetes (Polychaeta: Pilargidae) from Eastern Mexico. Rev. Biol. Trop. 38:87—90. Schroeder, P. C., and C. O. Hermans. 1975. Annelida: Polychaeta. Pp. 1-213 in Reproduction of marine 114 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES invertebrates, Vol. 3. Annelids and Echiurans. (A. C. Giese and J. S. Pearse, eds.), Academic, New York. Sosa-Rodriguez, T. 1993. Estudio histolégico del poliqueto pelagico Alciopina parasitica (Claparéde & Panceri, 1867). Tes. Prof., Fac. Ciencias, UNAM, México, 45 pp. Accepted for publication 29 December 1997. Bull. Southern California Acad. Sci. 97(3), 1998, pp. 115-120 © Southern California Academy of Sciences, 1998 Status of Beavers (Castor canadensis) in Valle de Mexicali, México Eric Mellink and Jaime Luévano Centro de Investigacion Cientifica y de Educaci6én Superior de Ensenada, B.C. Apartado Postal 2732, Ensenada, Baja California, México! Abstract.—To determine the current status of beavers (Castor canadensis) in Valle de Mexicali, Mexico, we surveyed the area in late 1995 and early 1996. We found evidence of current presence of beavers at 20 sites; 11 additional sites had evi- dence of past use. Most sites were along the Rio Colorado. Population levels of beavers in the area are highly variable and depend on extraordinary water releases through the Rio Colorado. Resumen.—Con el fin de determinar el estado del castor (Castor canadensis) en el Valle de Mexicali realizamos prospecciones a finales de 1995 y principios de 1996. Encontramos evidencias de presencia actual de castor en 20 sitios; otros 11 sitios tenfan evidencias de uso anterior. La mayoria de estos sitios se encontraban a lo largo del Rio Colorado. Las poblaciones de castores en esta regi6n son altamente variables y dependen de las aportaciones extraordinarias de agua del Rio Colorado. The Rio Colorado and tributaries historically had abundant water and main- tained large stands of willows (Salix gooddingii, S. exigua, and S. hindsiana) and cottonwoods (Populus macdougalii) along its banks and inundation flats (Wiggins 1980; Ezcurra et al. 1988). These areas supported abundant beavers (Castor can- adensis; Stone and Rhoads 1905; MacDougal 1906; Mearns 1907; Pattie 1831). The watercourse in the area was changed extensively early in the 20" century as a requirement for agriculture. This lead from time to time to the near dissapear- ance of beavers from most of the area (Sykes 1937a, b). Overall, however, beavers continued to be a typical component of the region (Burt 1938; Dixon 1922; Huey 1964; Leopold 1953). Indeed, some areas that once were unsuitable for beavers developed suitable habitat as a result of management of water for agriculture (Dixon 1922; Grinnell et al. 1937; Tappe 1942). Irrigation practices and agricul- tural development in the Valle de Mexicali intensified in the 1960s, reducing the extent of wetlands (Mellink 1995). Also, hunting periodically substantially re- duced beaver populations (Grinnell 1914; Pattie 1831). The current status of beavers in the Mexican portion of the drainage of the Rio Colorado was unknown, although it was supposed that they persisted in small numbers (Ceballos 1985; Ceballos and Navarro 1991). The purpose of this survey was to determine the extent of the presence of beavers in this area. Methods In October and November 1995 we visited all major water bodies in the Valle de Mexicali. As we had surveyed the Ciénega de Santa Clara in previous years, 'U.S. Mailing address: CICESE; P.O. Box 434844; San Diego, CA 92143. US) 116 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES we did not visit it at this time. We also interviewed colleagues with ample knowl- edge of the Ciénega. We inspected 80 sites, and five stretches of the Rio Hardy. The sites and river sections were surveyed from a car, on foot, or with a kayak. In all places, we searched for past and current evidence of use by beavers: felled trees, stumps, branches in the water, dikes, dens, and slides. On 7 February 1996, we searched for dams or dens, aboard a Cessna 182 airplane, flying over the central and northern sections of the mexican portion of the Rio Colorado, and over the Ciénega de Santa Clara. Results We found evidence of current occupation by beavers at 20 sites, some of which had also evidence of past use; 11 other sites had evidence only of past use (Figure 1). Beavers likely occur at some other, unsurveyed, locations in the area. Beavers were found from Presa Morelos, at the northern border with the United States, to the Vado de la Carranza, the intersection of the Rio Colorado with the road south out of Colonia Carranza; in one spot at the end of Canal El Caiman (formerly Canal Pescaderos), and in the Rio Hardy south of Campo Mosqueda. Evidences of past use followed the same pattern, and filled in current gaps. Evidence of former use was found also at two sites at the seldom-watered Canal Médanos, and local residents informed us about past presence of beavers in Canal Alamo. This channel had been cleaned a few months prior to our visit, and no evidence of its past occupation was left. The places that had only signs of past use by beavers reflect a current contrac- tion of the area occupied by them, resulting from the drying of the water bodies. Also, sites with beavers to which we made succesive visits 2—3 weeks apart were rapidly drying. Beavers were clearly associated with willows and cottonwoods, in addition to the water, as elsewhere in the Lower Colorado. On occasions beavers were present where water was limited to small stagnant pools, or to thin, shallow currents. The absence of beavers from the Rio Hardy north of Campo Mosqueda, which has abundant water and where beavers were once common, can be associated with the lack of willows and cottonwoods; the only trees now present are shruby tamarisks (mainly Tamarix pentandra). Current existence of beaver dams, as op- posed to their absence earlier in the century (Leopold 1959), is the result of the change in the type of watercourses, from a large river to small currents. Some of the sites occupied by beavers were depressions that resulted from the construction of a protection levee on the eastern side of the Rio Colorado during 1979-1981 (Sanchez-Ramirez 1990). These depressions, which can be several meters deep, were surrounded by willows and cottonwoods. In all such holes that we inspected, we found evidences of activity of beavers, either current or past. The Ciénega de Santa Clara is a large wetland created and maintained by brine water from the Wellton-Mohawk Irrigation District, Arizona (Glenn et al. 1992). Beavers can occupy this type of marsh, when they have willows, as in Mittry Lake, Arizona (R. Henry, pers. comm.; Todd 1986). However, the Ciénega de Santa Clara has few trees, mostly restricted to the edges. Moreover, these are not the beavers’ preferred trees, but western honey mesquites (Prosopis glandulosa var. torreyana), screwbean mesquites (P. pubescens), and tamarisks (Tamarix ra- BEAVERS IN MEXICALI 1k lig 115945) 115°30. 115915! 32°45) ERS MEXICALL ry ‘SS SAN LUIS RIO COLORADO Fig. 1. Sites with beaver activity in the Valle de Mexicali, México. Circles represent beaver pres- ence in October-November 1995. Plus signs, sites with evidence of past use by beavers. Numbers indicate waterbodies, and letters, specific locations: 1 = Canal Médanos, 2= Canal El Alamo, 3 = Rio Colorado, 4 = Rio hardy, 5 = Canal El Caiman (formerly Canal Pescaderos), 6 = Ciénega de Santa Clara, A = Presa Morelos, B = Campo Mosqueda, and C = Vado de la Carranza. mosissima) (Zengel et al. 1995), which explains why neither we nor colleagues who have worked in the area have seen any evidence of beavers. In two cases, beavers had cut stems of young tamarisks, and in two sites they seemed to be relying exclusively on tule roots (Scirpus americanus). Tappe (1942) considered that tules could be a more important food than commonly considered. We found them to be used rarely, and in one of the sites with heavy use of tules, beavers had been using willows until the drying of the steep-sided pool they lived in left such trees out of reach. Tules seemed, therefore, to be an emergency food. Some people in the area eat beavers on occasion, but this does not seem to happen often. In two places, beavers were a nuisance, as they were felling trees that had been planted or were being cared for. Rather than killing the beavers, the people in charge protected the trees with old barrels and salvaged metal. 118 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Discussion Once control of water for agriculture began, flow in natural watercourses was reduced and the courses themselves suffered major changes. Beavers followed these changes, and some locations where beavers have been found were suitable only as a result of watercourse management. The locality of the only museum specimen from the region of which we are aware, and the locations reported by Dixon (1922), Grinnell et al. (1937), Leopold (1953), and Tappe (1942), and three of our sites were all in such newly developed riparian habitat. The completion of Hoover Dam (in 1935) and Glenn Canyon Dam (in 1963) caused a severe reduction in water flow through the Rio Colorado, and an ac- cordingly severe reduction in populations of beavers in the Valle de Mexicali. Before 1960, beavers were locally abundant, but even then they suffered fluctu- ations due to dry periods, as in 1934 (Tappe 1942). Between 1960 and 1978, water in the Rio Colorado south of the border was extremely scarce, and the beaver population surely was reduced. Since 1978, there have been some important flow events in the mexican portion of the Rio Colorado. These flows, especially the one resulting from the 1982— 1983 El Ninio Southern Oscillation (ENSO) event, promoted the development of riparian vegetation and an increase in the beaver population along the Rio Col- orado. Elsewhere, waterways that are usually dry (the Alamos and the Médanos, for example) received plenty of water at this time and, also, beavers. Information on the Rio Hardy’s water history is difuse. During 1960-1978, it seems to have had rather stable, deep water, as the area north of Campo Mosqueda does today. Since 1947, a naturally formed dam near the entrance of the river to the Gulf of California caused the development of a large wetland, the Rio Hardy marsh (J.M. Payne, pers. comm.). At least the northern section of this marsh had abundant willows and cottonwoods. The 1982-1983 ENSO caused the flooding of a vast area below the protection levee and created a large wetland that joined the Rio Hardy marsh. A large community of willows and cottonwoods developed, and, according to riverside inhabitants, beavers were abundant. However, the in- tense flows of the mid-1980s also eroded the dam (J.M. Payne, pers. comm.), and when water flow ceased the marsh drained. Later on, the water level in the Rio Hardy dropped as well. Water flow was negligible during 1989-1992, but the 1992-1993 river dis- charges revived some beaver colonies. At the time of our survey, the pools in the river were drying once more; some colonies had disappeared, and others were drying rapidly. In November 1995, water levels in the Rio Hardy were lowering rapidly and only a few beavers remained. Photographs take by us in 1994 contrast with the 1995 condition. However, during the aerial reconnaissance of February 1996, the Rio Hardy seemed to have a higher water level than during the previous autumn. When large amounts of water are released into the Rio Colorado they can destroy existing beaver dams and carry animals away, sometimes for great dis- tances. During the 1982-1988 flows, beavers were seen by the fuel dock at the Estero de Santa Clara, and one was captured on the sandy seashore of Golfo de Santa Clara, in the Gulf of California (R. Pita and M. J. Sanchez, pers. comm.). BEAVERS IN MEXICALI je) These animals were a distance of about 70 Km from the closest colony at that time, across unsuitable habitat. Although numbers of beavers in the Valle de Mexicali fluctuate dramatically and often approach extirpation, it is difficult to give them a legal risk status. The local subspecies (C. c. frondator, after Hoffmeister 1986) is widespread and has healthy populations in adjacent areas in the United States, where beavers are often considered a nuisance, and are controlled accordingly. There is no management plan for beavers in the valle de Mexicali, and the conservation of the habitat, in this area is fortuitous and completely marginal to agricultural production. Cur- rently, beaver habitat is created mostly by rare extraordinary water releases through the Rio Colorado. Acknowledgments This project was financed by the Programa Ambiental Frontera Norte, Instituto Nacional de Ecologia. Rocio Esquivel and Rodrigo Medellin assisted the funding process. Robert Henry assisted during early stages of the project. Alberto Tapia- Landeros, Prisciliano Gonzalez, Enrique Galindo, Jesus Zazueta, Cecilio Lomeli- Lopez, and several inhabitants of the Valle de Mexicali provided information. Iriana Zuria, assisted during field work, and Lourdes Méndez and Martin Diaz provided logistical support. Sandy Lanham (Environmental Flying Services) kind- ly supported us with the aerial flight. Vicente Ferreira, Eduardo Palacios, Horacio de la Cueva, Dan Guthrie and several anonymous reviewers made important sug- gestions. To all of them, our deepest appreciation. Literature Cited Burt, W. H. 1938. Faunal relationships and geographic distribution of mammals in Sonora, Mexico. Misc. Publ. Mus. Zool., Univ. Michigan, 39:1—77. Ceballos-G., G. 1985. The importance of riparian habitats for the conservation of endangered mammals in Mexico. U. S. Dep. Agric., For. Serv. Gen. Tech. Rep., RM-120:96—100. Ceballos, G., and D. Navarro L. 1991. Diversity and conservation of Mexican mammals. Pp. 167— 198, in Latin American mammalogy: history, biodiversity and diversity (M. A. Mares and D. J. Schmidly, eds.). Univ. Oklahoma, 468 pp. Dixon, J. 1922. Rodents and reclamation in the Imperial Valley. J. Mamm., 3:136—146. Ezcurra, E., R., S. Felger, A., D. Russell, and M. Equihua. 1988. Freshwater islands in a desert sand sea: the hydrology, flora, and phytogeography of the Gran Desierto oases of northwestern Mexico. Desert Plants, 9:35—44, 55-63. Glenn, E., P, R. S. Felger, A. Burquez, and D. S. Turner. 1992. Ciénega de Santa Clara: endangered wetland in the Rio Colorado delta. Nat. Res. J., 32:817—824. Grinnell, J. 1914. An account of the mammals and birds of the Lower Colorado Valley. Univ. of Calif. Publ. in Zoology 12(4):51—294. Grinnell, J., J. S. Dixon, and J. M. Linsdale. 1937. Fur-bearing mammals of California. Univ. California ress, 777 pp: Hoffmeister, D. E 1986. Mammals of Arizona. Univ. Arizona Press, 602 pp. Huey, L. M. 1964. The mammals of Baja California, Mexico. Trans. San Diego Soc. Nat. Hist., 13: 85-168. Leopold, A. 1953. Round River; from the journals of Aldo Leopold (L. B. Leopold, ed.). Oxford Univ., 173. pp. Leopold, A. S. 1959. Wildlife of Mexico: the game birds and mammals. Univ. California Press, 568 pp. MacDougal, D. T. 1906. The delta of the Rio Colorado. Contrib. New York Bot. Garden, 77:1—16. Mearns, E. A. 1907. Mammals of the Mexican boundary of the United States; part 1 Didelphidae to Muridae. Bull. U. S. Nat. Mus., 56:1—530. 120 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Mellink, E. 1995. Status of the muskrat in the Valle de Mexicali and the Delta del Rio Colorado, México. Calif. Fish and Game, 8:33-38. Pattie, J. O. 1831. The personal narrative of James O. Pattie of Kentucky. John H. Wood, 269 pp. Sanchez-Ramirez, O. 1990. Crénica agricola del Valle de Mexicali. Univ. Aut. Baja California, 274 pp. Stone, W., and S. N. Rhoads. 1905. On a collection of birds and mammals from the Colorado Delta, Lower California. Proc. Acad. Nat. Sci. Philadelphia 1905:676—690. Sykes, G. 1937a. Delta, estuary, and lower portion of the channel of the Rio Colorado 1933 to 1935. Carnegie Inst. Washington Publ., 480: 1-70. Sykes, G. 1937b. The Colorado delta. Am. Geog. Soc. Publ. 19:1—193. Tappe, D. T: 1942. The status of beavers in California. Game Bull., Calif. Div. Game and Fish, 3:1—59. Todd, R. L. 1986. A saltwater marsh hen in Arizona. Ariz. Game and Fish Dep., 290 pp. Wiggins, I. L. 1980. Flora of Baja California. Stanford Univ., 1025 pp. Zengel, S. A., V. J. Meretsky, E. P. Glenn, R. S. Felger, and D. Ortiz. 1995. Cienega de Santa Clara, a remnant wetland in the Rio Colorado delta (Mexico): vegetation distribution and the effects of water flow reduction. Ecol. Engin. 4:19-—36. Accepted for publication 7 May 1998. Bull. Southern California Acad. Sci. 97(3), 1998, pp. 121-124 © Southern California Academy of Sciences, 1998 Re-occurrence of the Threebanded Butterflyfish, Chaetodon humeralis (Chaetodontidae), with Notes on its Distribution in Southern California Daniel J. Pondella, II,' Robert Snodgrass,* Matthew T. Craig,' and Hugh Khim? 'Vantuna Research Group, Moore Laboratory of Zoology, Occidental College, Los Angeles, California 90041 ?11671 Tierra del Sur, San Diego, California 92130-2614 3National Marine Fisheries Service 76 N. King St., Suite 200, Honolulu, Hawaii 96817 The threebanded butterflyfish, Chaetodon humeralis Ginther 1860, is an eastern Pacific endemic species whose range has been reported as northern Chile to San Diego, California, reaching the Galapagos and Cocos Islands (Miller and Lea 1972, Grove and Lavenberg 1997). It is easily distinguishable from the three other chaetodontid species, C. falcifer, Johnrandallia nigrirostris, and Forcipiger flav- issimus, found in the eastern Pacific and cannot be confused with any Indo-West Pacific species. Two juvenile specimens are herein reported from southern Cali- fornia. On 1 November 1997, Hugh Khim, a student at the University of California, San Diego, was free diving at 10 m along the wall of the underwater canyon in the La Jolla marine reserve. While observing other juvenile fish hiding along the one-meter high siltstone wall, he discovered a juvenile threebanded butterflyfish, approximately 4 cm total length (TL). This fish was found some three weeks later by Khim and Robert Snodgrass who videotaped it; it had not moved from its original location. Water temperatures in La Jolla at this time and during the pre- ceding weeks were above 20°C, even at depths exceeding 20 m (R. Mc- Connaughey, pers. comm.). In King Harbor, Redondo Beach, California, on 12 December 1997, while con- ducting routine ichthyotransects with Matthew Craig, Daniel Pondella of the Van- tuna Research Group (VRG) at Occidental College observed a solitary three- banded butterflyfish at the end of the west breakwater (latitude 33°50.5 N, lon- gitude 118°23.7 W). That day on a subsequent dive it was found at the same location and captured with a hand net. The depth of capture was 10 m and the water temperature was 17.3°C, the ambient temperature of Santa Monica Bay at the time. This specimen, 46.5 mm TL, was photographed (Fig. 1), preserved and given to the Marine Vertebrates Collection at the Scripps Institution of Ocean- ography in La Jolla (SIO 98-23). The last verified collection or known observation of C. humeralis in California was during a warm-water period approximately 140 years ago (Hubbs and Rech- nitzer 1958). During the Pacific Railroad Survey, Lt. W. P. Trowbridge collected two specimens from San Diego (USNM 3170); however, these fish were not catalogued until after Girard’s work on these collections (1854; 1858) and were not included in books of California fishes (Barnhart 1936, Roedel 1948, 1953). Hubbs and Rechnitzer (1958, p. 279) note: 121 122 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 1. Left lateral view of the threebanded butterflyfish, Chaetodon humeralis, (38.6 mm SL, S1098-23) captured in King Harbor, Redondo Beach, California on December 12, 1997. Photograph by Daniel J. Pondella, II. The low catalog number (3170) indicates that the specimens were in fact entered in the collection nearly 100 years ago. Lt. W. P. Trowbridge was one of the most effective of the West Coast collectors on the Pacific Railroad Surveys. Somehow the species escaped inclusion in Girard’s reports on the fishes collected by these surveys. Presumably the specimens came to light after the bulk of the collections had been studied and cataloged, for the number is higher than those recorded for the species reported by Girard. In this connection it may be noted that Girard (1858: 338) referred to other specimens from San Diego that had become ‘“‘mislaid in the moving of the Smithsonian collections from one end of the building to another a few months since.’’ Some slight doubt regarding the validity of the San Diego record can not be dispelled, but we believe that C. humeralis is to be added to the list of tropical fishes that occurred at San Diego during the warm period a century ago. A trip by Matthew Craig to the National Museum of Natural History found the specimens were catalogued as described by Hubbs and Rechnitzer (1958) with the locality listed simply as “‘San Diego’’. The specimens (68.9, 89.2 mm TL, Table 1) were in good condition. The two fish observed in 1997 are fairly small, allowing inferences into settlement processes. Two possibilities are that they either recruited from the ichthyoplankton or rafted into these reefs. The USNM speci- mens are much larger than the 1997 specimens. Although previously collected as far north as the San Benito Islands (SIO 84- RE-OCCURRENCE OF THREEBANDED BUTTERFLYFISH IN CALIFORNIA [25 Table 1. Counts and measurements for the three museum specimens of threebanded butterflyfish, Chaetodon humeralis, from southern California. Lengths given in millimeters. Specimens Counts and measurements SIO98-23 USNM3170 USNM3170 Standard length 38.6 59.9 75.9 Total length 46.5 68.9 89.2 Dorsal fin elements XIII, 20 XIII, 18 XII, 19 Anal fin elements a 17: Ill, 16 Ill, 16 Lateral line scales 32 34 35 Pectoral rays (1) 16 Ig) 16 227), the typical northern range does not extend past Magdalena Bay (eg. SIO 62-105, SIO 64-55 and R. N. Lea, pers. comm.). The specimen found in King Harbor, Redondo Beach, California represents the known northern limit of this species, an extension of some 150 kilometers from the historic San Diego locality. Much attention has been given to the warming trends of coastal waters along our coasts of the Americas beginning in the mid 1970’s (Hayward 1997) and recently exasperated by the 1997-98 major El Nino Southern Oscillation (ENSO) event. It is probable that the recruitment of these two juvenile individuals is related to this event. Various expatriated fishes to the Southern California Bight with the northern limits of their ranges normally at Magdalena Bay, or occasionally at the San Benito Islands, have been noted since the shift from Oregonian dominated fauna to a San Diegan fauna beginning with the 1978-79 ENSO (eg. Brooks 1987, Lea and Fukuhara 1991, Lea and Rosenblatt 1992, Lea and McAlary 1994, Lea and Walker 1995, Pondella 1997). The long-term success of these individuals and the continued presence of these species in the Southern California Bight are uncertain. However, as indicators of environmental change (Radovich 1961, Mearns 1988, Stephens et al. 1988) the recruitment of tropical and subtropical (Panamic) species in the temperate waters of southern California is strong evidence of the current ENSO strength. Acknowledgments We are indebted to the curatorial assistance of C. Klepadio and H. J. Walker at the Scripps Institution of Oceanography and S. Jewitt and J. T. Williams at the Smithsonian Institution National Museum of Natural History. We would like to acknowledge the critical review and assistance of R. N. Lea, H. J. Walker and J. S. Stephens, Jr. and information provided by R. McConnaughey. Literature Cited Barnhart, P. S. 1936. Marine Fishes of Southern California. University of California Press, Berkeley, California. iv + 209. Brooks, A. J. 1987. Two species of Kyphosidae seen in King Harbor, Redondo Beach, California. Calif. Fish and Game. 73:49-61. Grove, J. S. and R. J. Lavenberg. 1997. The Fishes of Galapagos Islands. Stanford University Press, Stanford, California. XLIV + 863 p. Hayward, T. L. 1997. Pacific Ocean climate change: atmospheric forcing, ocean circulation and eco- system response. TREE. 12(4):150—154. 124 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Hubbs, C. L. and A. B. Rechnitzer. 1958. A new fish, Chaetodon facifer, from Guadalupe Island, Baja California, with notes on related species. Sci., Proc. Calif. Acad. 29(8):273-313. Girard C. E 1854. Observation upon a Collection of Fishes Made on the Pacific Coast of the United States, by Lieut. W. P. Trowbridge, U.S.A., VII (no. 4). Proc. Acad. Nat. Sci. Phila. pp. 142— 156. . 1858. Fishes of the Pacific Railroad Survey. Reports of explorations and surveys, to ascertain the most practicable and economical route for a railroad from the Mississippi River to the Pacific Ocean. House of Rep. Ex. Doc. No. 91. 400 pp., 76 pls. Lea, R. N. and L. Fukuhara. 1991. The Gulf coney, Epinephelus acanthistius, from the marine waters of southern California. Bull. S. Calif. Acad. Sci. 90(2):80—82. Lea, R. N. and R. H. Rosenblatt. 1992. The Cortez grunt, Haemulon flaviguttatum, recorded from two embayments in southern California. Calif. Fish and Game, 78:163—165. Lea, R. N. and F McAlary. 1994. Occurrence of the swallow damselfish, Azurina hirundo, from islands off southern California. Bull. S. Calif. Acad. Sci. 93:43—44. Lea, R. N. and H. J. Walker, Jr. 1995. Record of the bigeye trevally, Caranx sexfasciatus, and Mexican lookdown, Selene brevoorti, with notes on other carangids from California. Calif. Fish and Game. 81:89-95. Mearns, A. J. 1988. The “‘odd fish’’: unusual occurrences of marine life as indicators of changing ocean conditions. Pp. 137-176, in Marine Organisms as Indicators. (D. E Soule and G. S. Kleppel, ed.), Springer-Verlag, New York. xii + 342 p. Miller, D. J. and R. N. Lea. 1972. Guide to the coastal marine fishes of California. Calif. Dept. Fish and Game, Fish Bull. (157):1—249. Pondella, D. J., Il. 1997. The first occurrence of the Panamic sergeant major, Abudefduf troschelii (Pomacentridae), in California. Calif. Fish and Game. 83(2):84—86. Radovich, J. 1961. Relationships of some marine organisms of the northeast Pacific to water temper- atures, particularly during 1957 through 1959. Calif. Dept. Fish and Game, Fish Bull. (112):1—62. Roedel, P. M. 1948. Common Marine Fishes of California. Calif. Dept. Fish and Game, Fish Bull. (68):1-153. . 1953. Common Ocean Fishes of the California Coast. Calif. Dept. Fish and Game, Fish Bull. (91):1-184. Stephens, J. S., Jr, J. Hose, and M. S. Love. 1988. Fish assemblages as indicators of environmental change in nearshore environments. Pp. 91—105, in Marine Organisms as Indicators. (D. E Soule and G. S. Kleppel, ed.), Springer-Verlag, New York. xii + 342 p. Accepted for publication 30 July 1998. Bull. Southern California Acad. Sci. 97(3), 1998, pp. 125-126 © Southern California Academy of Sciences, 1998 INDEX TO VOLUME 97 Araeoschizus agustinus, new species, 90 Araeoschizus blomi, new species, 93 Borgon-Gonzalez, Dora J., see Elena Solana-Arellano Brattstrom, Bayard H: Strategies of Predator Attacks on the Schooling Fish, Selar crumenophthalmus, in Academy Bay, Soccoro Island, Islas Revillagigedo, Mexico, 76 Bursey, Charles, R., see Stephen R. Goldberg Campos, Alma Rosa de, see Ernesto Campos Campos, Ernesto and Alma Rosa de Campos: Distribution and Taxonomic Re- marks for Five Crab Species of the Family Grapsidae (Crustacea: Sesarminae and Varuninae) of the Mexican Pacific, 96 Constantine, Denny G: Range Extensions of Ten Species of Bats in California, 49 Craig, Matthew T., see Daniel J. Pondella, I Diaz-Castaneda, V. and V. Rodriguez-Villanueva: Polychaete Fauna from San Quintin Bay, Baja California, Mexico, 9 Echavarria-Heras, Hector, see Elena Solana-Arellano Figueroa-Carranza, Ana-Luisa, see Juan-Pablo Gallo-Reynoso Gallo-Reynoso, Juan-Pablo and Ana-Luisa Figueroa-Carranza: Cetaceans of Isla De Guadalupe, Baja California, Mexico, 33 Goldberg, Stephen R., Charles R. Bursey amd Mei Q. Wu: Composition of the Helminth Community of a Montane Population of the Coastal Whiptail, Cnemidophorus tigris multiscutatus (Sauria: Teiidae) from Los Angeles County, California, 82 Hartney, Kristine Behrents and Lusine Tumyan: Temporal Changes in Diet and Foraging Habitat of California Killifish (Fundulus parvipinnis) in Marina del Rey, California, | Khim, Hugh, see Daniel J. Pondella, I Luevano, Jaime, see Eric Mellink Martinez-Mufnoz, Marco A., see Ricardo R. Murillo Mellink, Eric and Jaime Luevano; Status of Beavers (Castor canadensis) in Vaile de Mexicali, Mexico, 115 Murillo, Ricardo R., A. A. Ortega-Salas, and Marco A. Martinez-Munioz: A Di- etary Analysis of Hippoglossina stomata Eigenmann and Eigenmann, 1980 (Pisces: Bothidae) along the Western Coast of Baja California, Mexico, 104 15 126 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Ortega-Salas, A. A., see Ricardo R. Murillo Papp, Charles, S: Two New Stenosini Species in the Genus Araeoschizus LeConte from Baja California, Mexico (Coleoptera: Tenebrionidae), 89 Pondella, Daniel J. I], Robert Snodgrass, Matthew T. Craig, and Hugh Khim: Re- occurrence of the Threebanded Butterflyfish, Chaetodon humeralis (Chaeto- dontidae), with Notes on its Distribution in Southern California, 121 Rodriguez- Villanueva, V., see V. Diaz-Castafieda Salazar- Vallejo, Sergio I: SEM and Histological Evidence of Enlarged Nephridial Pappillae in Loandalia Monro (Polychaeta: Pilargidae), 110 Snodgrass, Robert, see Daniel J. Pondella, II Solana-Arellano, Elena, Dora J. Borbon-Gonzalez, and Hector Echavarria-Heras: A General Allometric Model for Blade Production in Zostera marina, 39 Tumyan, Lusine, see Kristine Behrents Hartney Wu, Mei Q., see Stephen R. Goldberg Yohe, Robert M. II: Notes on the Late Prehistoric Extension of the Range for the Muskrat (Ondatra zibethicus) Along the Ancient Shoreline of Lake Cahuilla, Coachella Valley, Riverside County, California, 86 a ai ss —— = = \ r / * —— \ Gt L 1 . \ } i i t ) \ \ & x = ‘ S : = i, , ‘ > : - \ et J Za ; 5 " a 2 7 . cf i Z s ‘ & i [ = } INSTRUCTIONS FOR AUTHORS The BULLETIN is published three times each year (April, August, and December) and inc!udes articles in English in any field of science with an emphasis on the southern California area. Manuscripts submitted for publication should contain results of original research, embrace sound principles of scientific investigation, and present data in a clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide for contributors. Consult also recent issues of the BULLETIN. 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The literature cited: Entries for books and articles should take these forms. MeWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp. Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54:452—458. Brattstrom, B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.), Univ. California Press, xii + 635 pp. Tables should not repeat data in figures (/ine drawings, graphs, or black and white photographs) or contained in the text. The author must provide numbers and short legends for tables and figures and place reference to each of them in the text. Each table with legend must be on a separate sheet of paper. All figure legends should be placed together on a separate sheet. 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Such illustrations along with a brief caption should be sent to the Editor for review. PROCEDURE All manuscripts should be submitted to the Editor, Daniel A. Guthrie, W. M. Keck Science Center, 925 North Mills Avenue, Claremont, CA 91711. Authors are requested to submit the names, addresses and specialities of three persons who are capable of reviewing the manuscript. Evaluation of a paper submitted to the BULLETIN begins with a critical reading by the Editor; several referees also check the paper for scientific content, originality, and clarity of presentation. Judgments as to the acceptability of the paper and suggestions for enhancing it are sent to the author at which time he or she may be requested to rework portions of the paper considering these recom- mendations. The paper then is resubmitted and may be re-evaluated before final acceptance. Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He or she should promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations, legends, and bibliographical references. He or she marks corrections on the galley (copy editing and proof procedures in Style Manual) and promptly returns both galley and manuscript to the Editor. Manuscripts and original illustra- tions will not be returned unless requested at this time. All changes in galley proof attributable to the author (misspellings, inconsistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with the printer, not the Editor. CONTENTS Two New Stenosini Species in the Genus Araeoschizus LeConte from Baja California, Mexico (Coleoptera: Tenebrionidae). Charles S. Papp —_ Distribution and Taxonomic Remarks for Five Crab Species of the Family Grapsidae (Crustacea: Sesarminae and Varuninae) of the Mexican Pa- cific. Ernesto Campos and Alma:Rosa de Campos — 3 A Dietary Analysis of Hippoglossina stomata Eigenmann and Eigenmann, 1980 (Pisces: Bothidae) along the Western Coast of Baja California, Mexico. Ricardo R. Murillo, A. A. Ortega-Salas, and Marco A. Miartineéz-MUNOZ 02.0.2 ee ee SEM and Histological Evidence of Enlarged Nephridial Papillae in Loan- dalia Monro (Polychaeta: Pilargidae). Sergio I. Salazar-Vallejo Status of Beavers (Castor canadensis) in Valle de Mexicali, México. Eric Mellink and Jaime Luévano: —.. 220. ee eee eee Re-occurrence of the Threebanded Butterflyfish, Chaetodon humeralis (Chaetodontidae), with Notes on its Distribution in Southern Califor- nia. Daniel J. Pondella, II, Robert Snodgrass, Matthew T. Craig, and Hugh: Kittin 22.202t.0 le ee COVER: Araeoschizus agustinus Papp, n. sp. 89 96 104 110 115