ISSN 0038-3872 fee ueERN CALIFORNIA ACADEMY OF. SCIENCES BULLET Volume 98 Number 1 oe ad : ~. ; as - a “ie an aii hie he es BCAS-A98(1) 1-44 (1999) APRIL 1999 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 © Southern California Academy of Sciences, 1999 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 Dayid Hyckgb Donn Gorsline ipbigitized bythe internet Archive. Gloria Takahashi witha Phillips Cheryl C. Swift in 2012 withefuading from California Academy of Sciences Library 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 29 March 1999 http://arenive:org/detaksfoulietre1t0seut Bull. Southern California Acad. Sci. 98(1), 1999, pp. 1-14 © Southern California Academy of Sciences, 1999 A Hemphillian (Late Miocene) Mammalian Fauna from the Desert Mountains, West Central Nevada Thomas S. Kelly! and Thomas P. Lugaski? 'Vertebrate Paleontology Section Natural History Museum of Los Angeles County 900 Exposition Blvd., Los Angeles, California 90007 -W. M. Keck Museum Mackay School of Mines University of Nevada Reno, Nevada 89557 LIZRARY CUSF TAN GF | SOE SANTIS IT a at Abstract.—The Churchill Butte Local Fauna, a new fossil mammalian assemblage of Hemphillian (late Miocene) age, is now recognized from the “‘Coal Valley Formation”’ of the western Desert Mountains, Lyon County, Nevada. It is the first biostratigraphically datable fauna from the formation and consists of the following taxa: Canidae, gen. indet.; Felidae, genera indet. (two spp.); Proboscidea, fam. indet.; Tayassuidae, gen. indet.; Camelidae, genera indet. (at least two spp.); Rhin- ocerotidae, gen. indet.; Equidae, gen. indet.; Leporidae, gen. indet.; Dipoides cf. D. wilsont; and Parapliosaccomys oregonensis. In 1993, Ross Secord, then a student at the University of Nevada, Reno, dis- covered a locality (W. M. Keck Museum locality number P-0101) in the western Desert Mountains that yielded a small sample of fossil mammals. The locality occurs along the western flank of the Desert Mountains about 7 km southeast of Churchill Butte in Lyon County, Nevada. A subsequent field investigation of the locality by the authors has resulted in the discovery of additional mammal fossils. The vertebrate fossil assemblage from this locality is herein referred to as the Churchill Butte Local Fauna. The purpose of this report is to: 1) document the mammalian taxa that comprise the new fauna; and 2) review the stratigraphic and geochronologic relations of the formation that yielded the fauna. Methods Measurements of larger mammal teeth and appendicular elements were taken with a vernier caliper to the nearest 0.1 mm, and those of smaller mammals were taken with an AO optical micrometer disc to the nearest 0.01 mm. All teeth were measured along their greatest anteroposterior and transverse dimen- sions. Metric abbreviation, dental terminology (except for those of Dipoides Jager 1835), and dental formulae follow standard usage. Dental terminology for Dipoides follows Stirton (1935). Upper teeth are indicated by uppercase letters and lower teeth by lowercase letters. All specimens are in the W. M. Keck Museum (WMK) at the Mackay School of Mines, University of Nevada, Reno. The locality section was measured using a Jacob’s staff and Brunton transit to the nearest 0.25 m. NW SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 119° 15’ = \__Lahontan Reservoir or \ CHURCHILL BUTTE Carson River” CHURCHILL VALLEY Gr mee oe ESERT CHURCHILL NARROWS 4460" MASON VALLEY —1460 m—Contour line — —:-—-—Watercourse Body of Water sx Fossil Locality PINE NUT MOUNTAINS Fig. 1. Generalized map showing location of vertebrate fossil locality P-0101 on western flank of Desert Mountains near Churchill Butte. Abbreviations are as follows: A-P, greatest anteroposterior dimension; d, deciduous; L, left; R, right; s. s., sensu stricto; TR, greatest transverse dimen- sion. Geology and Occurrence of Fossils Moore (1969) mapped outcrops of Tertiary sedimentary rocks in Lyon County that extend eastward from the northeast flanks of the Pine Nut Mountains across the Churchill Narrows area to the western flanks of the Desert Mountains. He also mapped sediments of similar lithology that outcrop further east along the north side of Mason Valley and in the Wildhorse Basin of the Desert Mountains. Kelly (1998) considered these rocks as correlatives of the type Coal Valley For- mation of Gilbert and Reynolds (1973), but not as strict lithocorrelatives because they were probably deposited in a separate depositional basin from that of the type Coal Valley Formation. For this reason, Kelly (1998) referred to these rocks as “Coal Valley Formation’’. The “Coal Valley Formation’”’ of the Pine Nut Mountains-Churchill Narrows- Desert Mountains area consists of lacustrine and fluvial deposits of diatomite, siliceous shale, siltstone, mudstone, sandstone, and pebble conglomerate. The HEMPHILLIAN FAUNA FROM NEVADA 3 80 COVERED GRAY SILTY CONGLOMERATE 70 LIGHT BUFF TUFFACEOUS SILTY SANDSTONE 60 BUFF TO REDDISH-BUFF SILTSTONE INTERBEDDED WITH LIGHT BUFF 50 TUFFACEOUS SANDSTONE AND CONGLOMERATE 40 BUFF, GREENISH-BUFF, BROWN, AND GREENISH- BROWN SILTSTONE WITH THIN BEDS OF CONGLOMERATE AND GREEN TO GRAY SILICIFIED SHALE i al MAMMALS 20° ORANGE TO DARK BROWN SILTSTONE AND MUDSTONE BUFF TO BROWN SANDSTONE + FISH GRAY SILTY CONGLOMERATE ra BLUISH-GRAY TO WHITE P| TUFFACEOUS SANDSTONE METERS COVERED Fig. 2. Stratigraphic section of W. M. Keck Museum locality P-0101 showing occurrences of fossils within section. sandstones and siltstones are commonly tuffaceous and often contain abundant fossilized rootlets. The formation unconformably overlies Miocene andesite in- terbedded with sedimentary deposits, probably a correlative of the Kate Peak Formation, and is unconformably overlain by late Pliocene to Pleistocene basalts and Quaternary alluvium (Moore 1969; Kelly 1995). Based on lithology, the for- mation can be roughly divided into a lower and upper portion. The lower portion is comprised primarily of sandstone interbedded with minor amounts of siltstone, 4 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES mudstone, shale, and conglomerate.. The upper portion is comprised primarily of diatomite interbedded with minor amounts of sandstone, siltstone, mudstone, evaporites, and conglomerate. The lower portion appears to be dominated by fluvial deposits, whereas the upper portion appears to be dominated by lacustrine deposits. The presence of diatomite and mudstones with evaporites in the upper lacustrine lithofacies indicates that it was probably deposited in shallow lakes that occasionally dried up. On the western flanks of the Desert Mountains, isolated outcrops of ‘“‘Coal Valley Formation” are exposed along the sides of several small westerly-drain- ing canyons. In this area, the beds within the “‘Coal Valley Formation”’ dip eastward, from about 25 to 33 degrees, and their line of strike is generally in a north-south direction. The fossil locality (P-0101) is located in a small west facing outcrop of “‘Coal Valley Formation”’ in the Desert Mountains, about seven km southeast of Churchill Butte (Figure 1; detailed locality data on file at WMK). The outcrop covers an area of 330 m by 210 m with 76.5 m of section exposed. A generalized stratigraphic section of the outcrop is presented in Figure 2. The actual thicknesses of the uppermost conglomerate and low- ermost sandstone (Figure 2) are uncertain because the section is covered by Quaternary alluvium at the top and bottom. A thick outcrop of diatomite in- terbedded with sandstone and shale occurs 0.8 km east of locality P-0101. Although this outcrop is clearly higher in the section than locality P-0101, its exact stratigraphic position is difficult to determine because it is separated from locality P-0101 by a small alluvium filled valley. In addition to the mammalian fossils, numerous disarticulated fossil fish bones were recovered from the lower buff to brown sandstone at locality P-0101 (Figure 2). The fish fossils are currently under study and will be the subject of a separate report. Age of Fauna The Churchill Butte Local Fauna consists of the following taxa: Canidae, gen. indet.; Felidae, genera indet. (two spp.); Proboscidea, fam. indet.; Tayassuidae, gen. indet.; Camelidae, genera indet. (at least two spp.); Rhinocerotidae, gen. indet.; Equidae, gen. indet.; Leporidae, gen. indet.; Dipoides cf. D. wilsoni Hib- bard (1949); and Parapliosaccomys oregonensis Shotwell (1967). The age of the Churchill Butte Local Fauna can be determined by the shared occurrences of certain taxa within the fauna. The beaver Dipoides first appears in the early Hemphillian and last occurs in the Blancan (Korth 1994). Most inves- tigators generally regard the extinction of the Rhinocerotidae as one of the events to mark the end of the Hemphillian (Tedford et al. 1987). However, Madden and Dalquest (1990) reported finding a single fragment of a rhino tooth during screen washing of matrix from the Blancan Yellow Quarry, Scurry County, Texas (Beck Ranch Local Fauna). With the exception of this record, no other occurrences of the Rhinocerotidae in the Blancan are known (Prothero 1998). The geomyid Par- apliosaccomys oregonensis was previously known only from the Hemphillian McKay Reservoir Fauna of Oregon. The combined presence of the Rhinoceroti- dae, Dipoides, and Parapliosaccomys oregonensis in the Churchill Butte Local fauna indicates the fauna is Hemphillian (late Miocene) in age. HEMPHILLIAN FAUNA FROM NEVADA 5 Systematic Paleontology Class Mammalia Linnaeus, 1758 Order Carnivora Bowdich, 1821 Canidae Gray, 1821 ?Canidae, gen. indet. Specimen.—Partial left dentary with root of m3, WMK 6504. Discussion.—The morphology of the partial dentary with its small, single-root- ed m3 is most similar to those of the Canidae, to which it is questionably assigned. Family Felidae Gray, 1821 Felidae, genera and spp. indet. Specimens.—Proximal portion of left fifth metacarpal, WMK 6569; first pha- lanx, WMK 6532; second phalanx, WMK 6542. Discussion.—A few isolated foot bones demonstrate the presence of the Feli- dae. These specimens are inadequate for generic identification, but they appear to represent two different species. For example, the partial fifth metacarpal (WMK 6569) and the second phalanx (WMK 6542) are robust and only slightly smaller than those of the large Pleistocene lion Panthera atrox (Leidy, 1853), whereas the first phalanx (WMK 6532) has slender, elongated proportions and appears to represent a smaller cheetah sized cat. Some measurements of the felid specimens are as follows: partial fifth meta- carpal (WMK 6569), proximal A-P = 20.1 mm, proximal TR = 21.8 mm; first phalanx (WMK 6532), length = 43.1 mm, proximal TR = 15.0 mm, distal TR = 11.4 mm; second phalanx (WMK 6542), length = 30.7, proximal TR = 14.5 mm, distal TR = 11.7 mm. Order Proboscidea Illiger, 1811 Proboscidea, family indet. Specimens.—Cheek tooth fragment, WMK 6579. Discussion.—The enamel morphology of the cheek tooth fragment indicates it represents either the Gomphotheriidae or Mammutidae. Order Artiodactyla Owen, 1848 Family Tayassuidae Hay, 1902 Tayassuidae, gen. indet. Specimen.—Lower cheek tooth fragment, WMK 6512. Discussion.—The lower cheek tooth fragment is inadequate for generic iden- tification, but indicates the presence of a peccary in the fauna. Family Camelidae Gray, 1821 Camelidae, genera and spp. indet. Specimens.—Associated broken LP4-M2 WMK 6552; associated partial dRP4 and partial dLP4, WMK 6514, WMK 6515; partial lower left incisor, WMK 6516; distal humerus, WMK 6501; partial distal radius, WMK 6507; scaphoid carpal, WMkK 6540; scaphoid carpal, WMK 6576; partial ?scaphoid carpal, WMK 6573; unciform carpal, WMK 6559; accessory carpal, WMK 6511; accessory carpal, WMkK 6571; partial accessory carpal, WMK 6557; astragalus, WMK 6506; cuboid 6 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES tarsal, WMK 6553; cuboid tarsal, WMK 6565; partial cuboid tarsal, WMK 6578; entocuneiform tarsal, WMK 6574; distal metapodial condyle, WMK 6503; distal metapodial condyle, WMK 6513; first phalanx, WMK 6505; partial first phalanx, WMkK 6510; partial first phalanx, WMK 6508; partial first phalanx, WMK 6572; partial first phalanx, WMK 6538; partial first phalanx, WMK 6563; partial first phalanx, WMK 6577; partial second phalanx, WMK 6541; Discussion.—The camel material consists of numerous isolated appendicular elements and a few fragmentary teeth that can be divided into three distinct size groups: a small-sized camel (WMK 6503, WMK 6505, WMK 6506, WMK 6510, WMK 6511, WMK 6514, WMK 6515, WMK 6516, WMK 6538, WMK 6540, WMK 6541, WMK 6557, WMK 6559, WMK 6563, WMK 6573, WMK 6574, WMK 6576, WMK 6577, WMK 6578), similar in size to Hemiauchemia vera (Matthew, 1909, in Matthew and Osborn 1909); a medium-sized camel (WMK 6501, WMK 6507, WMK 6508, WMK 6513, WMK 6552, WMK 6565, WMK 6571, WMK 6572), similar in size to Pliauchenia Cope (1875); and a very large camel (WMK 6553), similar in size to Megatylopus Matthew and Cook (1909). The broken teeth are of typical camelid morphology and not complete enough to allow generic identification. The three size groups appear to indicate that three camel species are present in the fauna. However, some species of camels are known to exhibit a fair degree of sexual dimorphism and, therefore, it is possible that the small and medium-sized camel elements could represent the same species. The large-sized camel specimens are significantly larger than the small and me- dium-sized camel specimens and can be confidently regarded as representing a different species. Thus, at least two, possibly three, species of camels are present in the fauna. Measurements of selected specimens demonstrate the three size groups. Mea- surements of the small-sized camel specimens are as follows: broken d(LP4 (WMK 6514), A-P = 17 mm estimated, TR = 14—15 mm estimated; astragalus (WMK 6506), height = 42.0 mm, greatest width = 24.3 mm; distal metapodial condyle (WMK 6503), A-P = 23.5 mm, TR = 18.5 mm; first phalanx (WMK 6505), length = 70.5 mm, proximal TR = 21.4 mm, distal TR = 17.0 mm. Measurements of the medium-sized camel specimens are as follows: damaged P4-M2 (WMK 6552), P4 A-P = 14.9 mm, M1 A-P (broken) = 27.9 mm, M2 (broken) A-P = 28.9 mm; distal humerus (WMK 6501), A-P across condyles = 53.3 mm, TR across condyles = 52.8 mm; distal radius (WMK 6507), greatest TR across distal articular surface = 53-54 mm estimated; distal metapodial condyle (WMK 6513), A-P = 30.9 mm, TR = 22.8 mm; proximal first phalanx (WMK 6572), proximal A-P = 28.3 mm, proximal TR = 29.3 mm. Measurements of the large-sized camel specimen are as follows: left cuboid tarsal (WMK 6553), height = 42.0 mm, A- P = 61.6 mm, TR = 42.0 mm. Order Perissodactyla Owen, 1848 Family Rhinocerotidae Owen, 1845 Rhinocerotidae, gen. indet. Specimens.—Lower cheek tooth fragment, WMK 6561; lower cheek tooth frag- ment, WMK 6562. . Discussion.—The cheek tooth fragments can only be identified as belonging to HEMPHILLIAN FAUNA FROM NEVADA 7 Fig. 3. Dipoides cf. D. wilsoni Hibbard (1949) of Churchill Butte Local Fauna; A, RP4, WMK 6528; B, LM1, WMK 6523; C, LM3, WMK 6529; D, Lp4, WMK 6527; E, Rm1 or 2, WMK 6526. All occlusal views with anterior up; A and D, lateral to left; B, C, and E, lateral to right. Scale = 1 mm. the Rhinocerotidae. The presence of a rhino in the fauna indicates that the fauna is probably no younger than Hemphillian. Family Equidae Gray, 1821 Equidae, gen. indet. Specimens.—Deciduous upper premolar fragment, WMK 6566; lower cheek tooth fragment, WMK 6567; lower cheek tooth fragment (preflexid), WMK 6568; first phalanx, WMK 6509. Discussion.—Although the equid specimens can only be identified to family, a few characters can be distinguished. The cement is moderately thick on the de- ciduous upper premolar fragment (WMK 6566). The preflexid of the lower cheek tooth fragment (WMK 6568) has a simple occlusal enamel morphology, similar to those of Dinohippus Quinn (1955) and Pliohippus Marsh (1874). As compared with other Hemphillian horses, the slightly crushed, first phalanx (WMK 6509) is moderate in size, measuring 55.0 mm in length, 30.5 mm proximal TR, and 31.1 mm distal TR. Order Lagomorpha Brandt, 1855 Family Leporidae Gray, 1821 Leporidae, gen. indet. Specimens.—Partial upper cheek tooth, WMK 6530; calcaneum, WMK 6517; partial calcaneum, WMK 6518; partial calcaneum, WMK 6519. Discussion.—The rabbit specimens were only identified to family. Order Rodentia Bowdich, 1821 Family Castoridae Gray, 1821 Genus Dipoides Jager, 1835 Dipoides cf. D. wilsoni Hibbard, 1949 Figure 3, Table 1 Specimens.—Partial maxilla with partial incisor and LP4-M1, WMK 6523; RP4, WMK 6528; LM3, WMK 6529; partial right dentary with base of incisor, 8 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1. Measurements (in mm) of teeth of Dipoides cf. D. wilsoni Hibbard (1949) from Desert Mountains. Specimen number Position A-P TR Crown height WMK 6523 P4 (unworn) 3:65 3.65 os WMK 6528 P4 4.63 4.50 14.4 WMK 6523 M1 3:52 3.60 10.4 WMK 6529 M3 395 3.35 10,2 WMK 6527 p4 5.10 3550 163 WMK 6526 ml son 3.80 3.97 14.3 WMK 6522 m2 3.95 eye) | os WMkK 6524; partial dentary with partial incisor, WMK 6525; Lp4, WMK 6527; partial dentary with Lm2, WMK 6522; Rm1 or 2, WMK 6526; associated cheek tooth fragments, WMK 6534. Discussion.—The small sample of Dipoides from the Desert Mountains consists of isolated teeth, partial dentaries with broken incisors and a m2, and a partial maxilla with P4-M1. All of the cheek teeth are rootless and hypsodont. The transverse widths are usually greater anteriorly in the upper molars, whereas in P4 and the lower cheek teeth the transverse widths are usually greater posteriorly. Cement is present in the flexi and flexids of the upper and lower cheek teeth, respectively. Two P4’s are represented in the sample; an unworn P4 (WMK 6523) in the process of erupting and a moderately worn P4 (WMK 6528). The unworn P4 (WMK 6523) exhibits the following characters: 1) a paraflexus, mesoflexus, and hypoflexus are present; 2) multiple small cusps are present on the mesostyle; and 3) several small cusps are present on the paracone. The morphology of the unworn P4 is not specifically diagnostic because the unworn P4 occlusal pattern is very similar in all species of Dipoides (Shotwell 1955). The moderately worn P4 (WMK 6528) exhibits the following characters: 1) a shallow, but persistent, par- aflexus is present that extends lingually across the occlusal surface to almost meet with the deep hypoflexus, resulting in a narrow isthmus connecting the anterior loph to the middle loph; 2) a metaflexus and metastria are lacking on the occlusal surface and labial side of the posterior loph, respectively; 3) the mesoflexus ex- tends lingually almost completely across the occlusal surface, almost isolating the posterior loph from the middle loph; and 4) the parastria, mesostria, and hypostria extend down the entire depth of the tooth. The M1 (WMK 6523) is in early wear and is characterized by having the following: 1) a general ‘‘S”’ shaped occlusal pattern; 2) a paraflexus and meta- flexus are lacking; 3) the hypoflexus extends labially across almost the entire occlusal surface, with its labial termination squared off, giving the hypoflexus an elongated rectangular shape; 3) the mesoflexus extends posterolingually across the occlusal surface, almost uniting with the lingual enamel wall, resulting in a narrow enamel connection of the middle loph to the posterior loph (this connec- tion probably widens with wear); and 4) the hypostria and mesostria extend down the entire depth of the tooth. The M3 (WMK 6529) is characterized by having the following: 1) a paraflexus and metaflexus are lacking; 2) the hypoflexus extends labially almost completely HEMPHILLIAN FAUNA FROM NEVADA S) across the occlusal surface, with its labial termination squared off and its sides nearly parallel; 3) the mesoflexus extends lingually across the occlusal surface, where it almost meets the lingual enamel wall, resulting in a narrow enamel connection between the middle and posterior lophs (however, at the base of the tooth the mesoflexus lingual termination is not so close to the lingual enamel wall and it is no longer squared off, but is bifurcated); and 4) the hypostria and me- sostria extend down the entire depth of the tooth. The p4 (WMK 6527) is characterized by having the following: 1) a paraflexid is absent, but a faint, vestigial parastriid is present, which extends down the entire depth of the tooth and results in a very minor indentation along the lingual aspect of the occlusal surface of the anterior lophid; 2) the mesoflexid extends antero- labially across the occlusal surface, almost abutting the labial enamel wall, re- sulting in anteroposteriorly oriented anterior and middle lophids; 3) the hypoflexid extends lingually across about three-quarters of the occlusal surface wherein it turns anterolingually and further extends to end in a rounded terminus close to the lingual enamel wall; and 4) the mesostriid and hypostriid extend down the entire depth of the tooth. The lower molars (WMK 6522, WMK 6526) are characterized by having the following: 1) simple “‘S”’ shaped occlusal patterns, with only mesoflexids and hypoflexids present; 2) the mesoflexids and hypoflexids have nearly parallel sides and rounded labial and lingual termini, respectively; and 3) the mesostriids and hypostriids extend down the entire depths of the teeth. Measurements of the Dipoides specimens are presented in Table 1. Species of Dipoides are differentiated primarily by size and cheek teeth occlusal morphologies (Wilson 1934; Stirton 1935; Shotwell 1955; Baskin 1979). A fair degree of intraspecific variation occurs in the cheek teeth occlusal patterns of Dipoides, so that species are often defined by frequencies of particular occlusal patterns rather than by the absolute presence or absence of a particular dental character (Shotwell 1955). The amount of intraspecific variation in the cheek teeth occlusal patterns of the Desert Mountains Dipoides cannot be determined because of the small sample size. The Desert Mountains specimens are regarded as be- longing to one species because they all appear to be about the same size and all were recovered from a single locality. In fact, based on the degree of wear, some of the isolated teeth may represent the same individual. Even though the number of Dipoides specimens from the Desert Mountains is small, they exhibit a distinctive suite of dental characters that can be compared with other recognized species of Dipoides. Six North American species of Di- poides are currently recognized: D. stirtoni Wilson (1934); D. williamsi Stirton (1936); D. rexroadensis Hibbard and Riggs (1949); D. wilsoni Hibbard (1949); D. smithi Shotwell (1955); and D. vallicula Shotwell (1970). One other species that was originally assigned to Dipoides, D. intermedius Zakrzewski (1969), has recently been referred to Procastoroides Barbour and Schultz (1937) by Repen- ning et al (1995). The Desert Mountains Dipoides differs from D. stirtoni by having the following characters: 1) significantly smaller in size; 2) a P4 metaflexus is lacking; and 3) a p4 paraflexid is lacking. It differs from D. williamsi by having the following characters: 1) slightly smaller in size; 2) a P4 paraflexus is present; and 3) a p4 paraflexid is lacking. It differs from D. rexroadensis, a poorly known species (Hibbard and Riggs 1949; Woodburne 1961; Zakrzewski 1969), by its 10 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES much smaller size. It differs from D. wilsoni by having the following characters: 1) a P4 paraflexus is present; and 2) the lower molar flexids have rounded rather than flattened terminations and are not as closely abutted to the opposing enamel walls. It differs from D. smithi by having the following characters: 1) significantly smaller in size; and 2) the P4 anterior loph is slightly less inflated and relatively narrower. It differs from D. vallicula by having the following characters: 1) the molar flexi and flexids are relatively narrower; 2) the P4 paraflexus and hypoflexus are slightly overlapping and do not abut, resulting in an oblique isthmus instead of an anteroposteriorly directed isthmus; and 3) a p4 paraflexid is lacking. It differs from Procastoroides intermedius by having the following characters: 1) much smaller size; 2) less hypsodont cheek teeth; and 3) the M3 lacks a meta- flexus and metastria. In addition to the six recognized species of Dipoides, pre- vious investigators have referred a number of specimens from various formations to Dipoides. Stirton (1935) referred specimens from the Rattlesnake Formation of Oregon and the Thousand Creek Beds of Nevada to Dipoides. The Desert Mountains form differs from the Rattlesnake and Thousand Creek forms by its much smaller size and lack of a p4 paraflexid. Macdonald (1959) referred spec- imens from the Coal Valley Formation of Smith Valley, Nevada, to Dipoides. The Desert Mountains form differs from the Smith Valley form by having the follow- ing characters: 1) slightly smaller in size; 2) a P4 paraflexus present 3) a P4 metaflexus is lacking; and 4) a p4 paraflexid is lacking. Although the differences noted above indicate the Desert Mountains Dipoides specimens may represent a new species, they also exhibit certain similarities to D. wilsoni and specimens referred by Shotwell (1955) to D. cf. D. wilsoni from the Malheur River area, Oregon. The Desert Mountain teeth are similar to those of D. wilsoni by having the following characters: 1) small size; 2) relatively hypsodont cheek teeth; and 3) a p4 with a weak or absent paraflexid. The size and overall occlusal patterns of the Desert Mountains Dipoides teeth are morpho- logically most similar to those of certain specimens referred to D. cf. D. wilsoni from the Malheur River area, including P4’s with paraflexi and parastriae that result in isthmuses between the anterior and posterior lophs, and p4’s that lack paraflexids. The Desert Mountains form differs from D. cf. D. wilsoni from the Malheur River area by having the lower molar flexid terminations not squared off, but more rounded. A much larger sample of Dipoides from the Desert Mountains is needed before this form can be fully characterized. Until such time, the Desert Mountain spec- imens are herein referred to Dipoides cf. D. wilsoni. In North America, Dipoides has its first occurrence in the early Hemphillian (late Miocene) and its last occurrence in the Blancan (Pliocene) (Tedford et al. 1987; Korth 1994). The presence of Dipoides in the Churchill Butte Local Fauna indicates the fauna is no older than Hemphillian. Family Geomyidae Gill, 1872 Genus Parapliosaccomys Shotwell, 1967 Parapliosaccomys oregonensis Shotwell, 1967 Figure 4 Specimen.—Rp4, WMK 6531. Discussion.—The slightly worn Rp4 is subhypsodont with a basic two col- HEMPHILLIAN FAUNA FROM NEVADA it & A B Fig. 4. Parapliosaccomys oregonensis Shotwell (1967) of Churchill Butte Local Fauna: Rp4, WMK 6531, A—occlusal view, anterior up, lateral to right, B—medial view, anterior left. Scales for A and B = | mm. umned structure (anterior and posterior lophids). Two distinct roots are present that are still partially open at their bases. The anterior lophid has a distinctive trefoil occlusal outline. This occlusal pattern would rapidly disappear with wear because it only extends down the anterior lophid about 1.3 mm from the crown, after which the anterior lophid would assume a simple ovoid occlusal pattern with a strong central union to the posterior lophid. Dentine tracts are present on the labial and lingual aspects of the anterior and posterior columns. The tracts extend up from the base of the crown, ranging in length from 1.5 mm to 3.3 mm, but do not reach the occlusal surface during early wear. In later wear, about half way down the tooth, the dentine tracts would interrupt the lateral occlusal enamel outlines of the anterior and posterior lophids. Measurements of WMK 6531 are as follows: occlusal A-P = 1.27 mm, occlusal anterior TR = 1.35 mm, occlusal posterior TR = 1.87 mm, crown height = 5.3 mm. The p4 is indistinguishable from those of Parapliosaccomys oregonensis from McKay Reservoir, Oregon, and it is tentatively assigned to this species. It differs from those of the closely related Pliosaccomys Wilson (1936) by its relatively greater hypsodonty and the presence of dentine tracts. Three species have been previously assigned to Parapliosaccomys: P. orego- nensis of the Hemphillian McKay Reservoir Fauna of Oregon; P. hibbardi (Storer, 1973) from the Clarendonian WaKeeney Local Fauna of Kansas; and P. annae Korth (1987) from the Barstovian Valentine Formation of Nebraska. Recently, Akersten (1988) and Korth (1994) questioned the assignment of P. hibbardi and P. annae to Parapliosaccomys because of differences between the lower premo- lars of these species and those of the type species P. oregonensis. It appears that P. hibbardi and P. annae are generically distinct from P. oregonensis and should not be included in Parapliosaccomys s. s. As such, Parapliosaccomys is a mono- typic genus restricted to the Hemphillian. The presence of P. oregonensis in the fauna from the Desert Mountains indicates the fauna is Hemphillian in age. Summary The ‘“‘Coal Valley Formation” of the western Desert Mountains, Lyon Country, Nevada, has yielded a new fossil mammalian assemblage, the Churchill Butte 12 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Local Fauna. The fauna consists of the following taxa: Canidae, gen. indet.; Fe- lidae, genera indet. (two spp.); Proboscidea, fam. indet.; Tayassuidae, gen. indet.; Camelidae, genera indet. (at least two spp.); Rhinocerotidae, gen. indet.; Equidae, gen. indet.; Leporidae, gen. indet.; Dipoides cf. D. wilsoni; and Parapliosaccomys oregonensis. The shared occurrences of Dipoides, Parapliosaccomys oregonensis, and Rhinocerotidae indicate that the fauna is Hemphillian (late Miocene) in age. The Churchill Butte Local Fauna is the first biostratigraphically datable fauna from the ‘“‘Coal Valley Formation” of the western Desert Mountains. Exposures of “Coal Valley Formation”’ can be traced westward from the west- ern flanks of the Desert Mountains across the Churchill Narrows area to the northeastern flanks of the Pine Nut Mountains in Lyon County, Nevada (Moore 1969; Kelly 1998). The discovery of the Churchill Butte Local Fauna indicates that the ““Coal Valley Formation” of the Pine Nut Mountains-Churchill Narrows- Desert Mountains area was deposited, at least in part, during the Hemphillian (late Miocene). Additional exposures of “Coal Valley Formation” occur north and northeast of the Pine Nut Mountains-Churchill Narrows-Desert Mountains area as follows: 1) along the southeastern flanks of the Virginia Range, Lyon Country; 2) in the central Virginia Range, Storey Country; and 3) in the vicinity of the communities of Mogul and Verdi, Washoe County (Kelly 1998). In the Mogul- Verdi area, the “‘Coal Valley Formation” is about 900 m thick and has been radiometrically and biostratigraphically dated from the Clarendonian (late Mio- cene) to the latest Hemphillian or earliest Blancan (early Pliocene) (Axelrod 1958, 1962; Bonham 1969; Kelly 1998). In the Chalk Hills area of the central Virginia Range, the “‘Coal Valley Formation” is about 850 m thick and, based on bio- stratigraphic data and radiometric dates on underlying and overlying strata (Bon- ham 1969; Kelly 1998), was deposited from the Clarendonian to the late Hem- phillian or early Blancan. Along the southeastern flanks of the Virginia Range, the “Coal Valley Formation” is about 228 m thick and has yielded the Churchill Valley Local Fauna of probable Clarendonian age from the lower 92 m of the formation (Kelly 1998). In the Churchill Narrows area, the Churchill Narrows Site has yielded a mammalian assemblage of undetermined age from near the base of the “Coal Valley Formation”’ (Kelly 1998). The thickness of the “Coal Valley Formation’’ in the Pine Nut Mountains-Churchill Narrows-Desert Moun- tains area has not been determined, but appears to be thickest along the north- eastern flanks of the Pine Nut Mountains, where at least several hundred meters are exposed. From the east side of the Churchill Narrows to the western flanks of the Desert Mountains, the formation thins, so that in the vicinity of locality P- 0101, the formation is only about 150 m thick. The ‘“‘Coal Valley Formation” of west central Nevada may have been deposited in two or possibly three different depositional basins. The thick deposits in Mogul- Verdi area and the central Virginia Range probably represent two separate de- positional basins. A third depositional basin may have been centered in the area now exposed along the northeastern flanks of the Pine Nut Mountains. However, because the “‘Coal Valley Formation”’ exhibits a general eastward thinning from the central Virginia Range across the southern flanks of the Virginia Range and southeastward to the Pine Nut Mountains-Churchill Narrows-Desert Mountains area, it is also possible that these areas represent a single depositional basin. The composition of the Churchill Butte Local Fauna provides evidence of the HEMPHILLIAN FAUNA FROM NEVADA L3 paleoenvironmental setting during the time of deposition at locality P-0101. The fauna includes browsers (a bunodont proboscidean and Rhinocerotidae), grazers (Equidae, Camelidae, and Leporidae), and aquatic animals (fish and the beaver Dipcides). This faunal combination indicates the paleoenvironment was probably characterized by grasslands or open woodlands with thicker woodlands occurring along perennial streams or small lakes. The streams or lakes probably supported riparian vegetation suitable for browsing animals, whereas, further from the wa- tercourses, grasslands or open woodlands supported grazing animals. The presence of tuffaceous deposits at locality P-0101 and throughout the ‘‘Coal Valley For- mation” of the Pine Nut Mountains-Churchill Narrows-Desert Mountains area indicates that the area was periodically supplied with volcaniclastic debris of epiclastic and airfall origin. Acknowledgments We are indebted to James R. Firby of the MacKay School of Mines, University of Nevada, Reno (UNR), Everett Lindsay of the University of Arizona, Donald R. Prothero of Occidental College, Charles A. Repenning of the United States Geological Survey, David S. Webb of the University of Florida, and David P. Whistler of the Natural History Museum of Los Angeles County for their com- ments and helpful suggestions on the original draft of this report. Special thanks is given to Ross Secord, a former student at UNR, for his untiring field work that led to the discovery of the fossil locality reported herein. Literature Cited Akersten, W. A. 1988. Affinities of “‘“Pliosaccomys” and ‘*Parapliosaccomys” from the Great Plains. J. Vert. Paleont., Sup. 3:8A. Axelrod, D. I. 1958. The Pliocene Verdi flora of western Nevada. Univ. Calif., Pub. Geol. Sci., 34: 91-160. . 1962. A Pliocene Sequoiadendron forest from western Nevada. Univ. Calif., Pub. Geol. Sci., 39:195—268. Barbour, E. H. and C. B. Schultz. 1937. An early Pleistocene fauna from Nebraska. Amer. Mus. Novitates, 942:1—10. Baskin, J. A. 1979. Small mammals of the Hemphillian age White Cone Local Fauna, northeastern Arizona. J. Paleont., 53:695—708. Bonham, H. E 1969. Geology and mineral deposits of Washoe and Storey counties, Nevada. Nevada Bur. Mines Geol. Bull. 70:1—140. Cope, E. D. 1875. On some new fossil Ungulata. Proc. Acad. Nat. Sci., Philadelphia, 19:258—263. Gilbert, C. M. and M. W. Reynolds. 1973. Character and chronology of basin development, western margin of the Basin and Range Province. Geol. Soc. Amer. Bull., 84:2489—2510. Hibbard, C. W. 1949. Pliocene Saw Rock Canyon Fauna in Kansas. Univ. Michigan, Contr. Mus. Paleont., 7:91—105. Hibbard, C. W. and E. S. Riggs. 1949. Upper Pliocene vertebrates from Keefe Canyon, Meade County, Kansas. Geol. Soc. Amer. Bull. 60:829—860. Jager, G. EF 1835. Ueber die Fossilen Saugethiere, welche in Wurttemberg gefunden worden sind, pp. 17-18. Kelly, T. S. 1995. A Pleistocene mammalian fauna from Adrian Valley, Lyon Country, west central Nevada. Cur. Res. in the Pleistocene, 12:99—102. . 1998. New Miocene mammalian faunas from west central Nevada. J. Paleont., 72:137—149. Korth, W. W. 1987. New rodents (Mammalia) from the late Barstovian (Miocene) Valentine Formation, Nebraska. J. Paleont., 61:1058—1064. . 1994. The Tertiary record of rodents in North America. Topics in Geobiology 12, Plenum IETess, x1 “319 pp: 14 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Leidy, J. 1853. Description of an extinct species of American lion: Felis atrox. Trans. Amer. Philos. Soc., 10:319=321- Macdonald, J. R. 1959. The middle Pliocene mammalian fauna from Smiths Valley, Nevada. J. Pa- leont., 33:872—887. Madden, C. T. and W. W. Dalquest. 1990. The last Rhinoceros in North America. J. Vert. Paleont., 10:266—267. Marsh, O. C. 1874. Notice of new equine mammals from the Tertiary formation. Amer. J. Sci., 7: 247-258. Matthew, W. D. and H. J. Cook. 1909. A Pliocene fauna from western Nebraska. Bull. Amer. Mus. Nat. Hist., 26:361—414. Matthew, W. D. and H. E Osborn. 1909. Faunal lists of the Tertiary Mammalia of the West. U. S. Geol. Sur. Bull., 361:91—138. Moore, J. G. 1969. Geology and mineral deposits of Lyon, Douglas, and Ormsby Counties, Nevada. Nevada Bur. Mines Geol. Bull., 75:1—45. Prothero, D. R. 1998. Rhinocerotidae. Pp. 595—605 in Evolution of Tertiary Mammals of North Amer- ica (C. M. Janis, K. M. Scott, and L. L. Jacobs, eds.), Cambridge Univ. Press, x + 691 pp. Quinn, J. H. 1955. Miocene Equidae of the Texas Gulf Coastal Plain. Univ. Texas, Bur. Econ. Geol. Pub. 5516:1—102. Repenning, C. A., T. R. Weasma, and G. R. Scott. 1995. The early Pleistocene (latest Blancan-earliest Irvingtonian) Froman Ferry Fauna and history of the Glenns Ferry Formation, southwestern Idaho. U. S. Geol. Sur. Bull., 2105:1—86. Shotwell, J. A. 1955. Review of the Pliocene beaver Dipoides. J. Paleont., 29:129—-144. . 1967. Late Tertiary geomyoid rodents of Oregon. Univ. Oregon, Bull. Mus. Nat. Hist., 9:1—51. . 1970. Pliocene mammals of southeast Oregon and adjacent Idaho. Univ. Oregon, Bull. Mus. Nat. Hist., 17:1—103. Stirton, R. A. 1935. A review of the Tertiary beavers. Univ. California, Pub. Geol. Sci., 23:391—458. . 1936. A new beaver from the Pliocene of Arizona with notes on the species of Dipoides. J. Mamm., 17:279-—281. Storer, J. E. 1973. The entoptychne geomyid Lignimus (Mammalia: Rodentia) from Kansas and Ne- braska. Can. J. Earth Sci., 10:72—83. Tedford, R. H., T. Galusha, M. FE Skinner, B. E. Taylor, R. W. Fields, J. R. Macdonald, J. M. Rens- berger, S. D. Webb, and D. P. Whistler. 1987. Faunal succession and biochronology of the Arikareean through Hemphillian interval (late Oligocene through earliest Pliocene), North America. Pp. 153-210 in Cenozoic Mammals of North America (M. O. Woodburne, ed.), Univ. California Press, xv + 336 pp. Wilson, R. W. 1934. A new species of Dipoides from the Pliocene of eastern Oregon. Carnegie Inst. Washington, Contr. Paleont., 453:19—29. . 1936. A Pliocene rodent fauna from Smiths Valley, Nevada. Carnegie Inst. Washington, Contr. Paleont., 473:15—34. Woodburne, M. O. 1961. Upper Pliocene geology and vertebrate paleontology of part of the Meade Basin, Kansas. Pap. Michigan Acad. Sci., 46:61—101. Zakrzewski, R. J. 1969. The rodents from the Hagerman Local Fauna, upper Pliocene of Idaho. Univ. Michigan, Contr. Mus. Paleont., 23:1—36. Accepted for publication 30 July 1998. Bull. Southern California Acad. Sci. 98(1), 1999, pp. 15-25 © Southern California Academy of Sciences, 1999 Fossil Wood from the Middle Miocene Conejo Volcanics, Santa Monica Mountains, California Carol J. Stadum and Peter W. Weigand RMW Paleo Associates, Inc., 23392 Madero, Suite L, Mission Viejo, California 92691] Department of Geological Sciences, California State University, Northridge, California 91330-8266 Abstract.—Evidence that the upper Conejo Volcanics erupted subaerially is con- firmed by the discovery of fossil wood preserved in a ~13.5 Ma tuff breccia exposed on the northern flank of the Santa Monica Mountains, California. This middle Miocene wood represents a low montane geoflora of hardwoods and co- nifers that lived at elevations greater than 1,300 m. Ponderosa Pine, Douglas Fir, Incense Cedar, and specimens from the dicotyledonous families Fagaceae (ever- green and live oaks), Rosaceae (mountain mahogany), and Rhamnaceae (mountain lilac) have been identified by thin section analysis. Preservation of this wood varies from amorphous clayey limb casts to splintered surfaces and detailed cell features. This is the first record of fossil trees in a tuff breccia from the Los Angeles basin region. In 1995, development of a 70-acre construction project in the Thousand Oaks area northwest of Los Angeles exposed the middle Miocene Upper Topanga For- mation and the underlying Conejo Volcanics. The volcanic units form ridges in the Santa Monica Mountains and represent a series of eruptions that commenced as submarine basalt flows about 16 Ma and concluded as subaerial lava flows and explosive lahars about 13.5 Ma (Yerkes and Campbell 1979). Site mitigation required paleontologists to monitor the excavation of the Upper Topanga marine sandstones exposed along the northern portion of the project. As the monitors were crossing volcanic material to reach the sandstone, they discov- ered petrified wood fragments and two logs in tuff breccia. Grading operations were temporarily diverted while paleontologists determined the extent of the fossil wood and recovered specimens. Thin sections of these specimens show that pres- ervation varies from detailed cell features to no identifiable features. Site The Conejo wood was recovered 65 km northwest of Los Angeles in the City of Westlake Village, north of California State Freeway 101, east of Lindero Can- yon Road, and south of Thousand Oaks Boulevard (Fig. 1A). The 70-acre com- mercial/residential development is in a valley bounded on the north by hills of the Monterey Formation and on the south by the Conejo Volcanics. Extending east/west across the center of the valley floor is a small ridge that separates the volcanic rocks exposed along the south flank from the sandstone of the Upper Topanga Formation exposed to the north. As construction equipment removed colluvium, sandstone, and tuff breccia from the site, fossil wood fragments were 15 16 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Thousand Oaks Bivd U.S. Westlake Blvd Highway 4 Ventura Fwy WESTLAKE XK VILLAGE Ladyface Mtn. Qa eRe) Quaternary alluvium Tm ISSN Monterey Formation Tlvc Boooo| Detrital Sediments of Lindero Canyon iPtue ee Upper Topanga Formation VVVVV ° . Tcva Conejo Volcanics 0.5 mi | 0.5 km | Fig. |. Maps of the study area in the City of Westlake Village, Los Angeles County. A. Location map. Position of Fig. 1B outlined. B. Geologic map after Dibblee and Ehrenspeck (1993a). The “X” indicates the fossil wood site. exposed across a 100-m? area along the southern flank of the ridge (Fig. 1B). Two broken logs were found encased in the tuff unit southwest of the ridge. This concentration of middle Miocene fossil wood may not be unique to the area. Local residents have reported that reworked fossil wood occurs throughout the Thousand Oaks region in the upper Conejo Volcanics and overlying Upper To- panga and Monterey Formations. [Note: The stratigraphic nomenclature of Dib- blee and Ehrenspeck (1993a) is used here; Yerkes and Campbell (1979) refer to the latter two formations as the Calabasas and Modelo Formations respectively. ] Geologic Environment The Conejo Volcanics lie between the Lower Topanga and Upper Topanga Formations (Dibblee and Ehrenspeck 1993a; Fig. 2) and include up to 2 km of extrusive basaltic flow breccias, flow and pillow lavas, andesite and tuff breccias, dacite breccias, and a variety of intrusive forms that have been described by Dibblee and Ehrenspeck (1993b). The lower portions of the volcanic sequence were formed in a marine environment, indicated by an abundance of pillow lavas, pillow breccias, and occasional layers of fossil oyster debris. Higher in the se- quence, the rocks were deposited in a subaerial environment and include laharic breccias, fluvial and alluvial fan breccia-conglomerates, volcanic bombs (Dibblee and Ehrenspeck 1993b), and the fossil wood described in this paper. K-Ar dates, FOSSIL WOOD FROM CONEJO VOLCANICS 17 FORMATION [ital] Epoch DETRITAL MONTEREY SEDIMENTS 10 FORMATION OF LINDERO CONEJO V JO VOLCANICS FORMATION MIOCENE LOWER TOPANGA FORMATION SESPE FORMATION OLIGOCENE Fig. 2. Chronostratigraphic diagram of the rock units in the western Santa Monica Mountains based on the nomenclature of Dibblee and Ehrenspeck (1993a). “‘U” represents an unconformable contact. determined by Turner (1970) and Yerkes and Campbell (1979) on plagioclase grains separated from samples collected throughout the volcanic section, range from 16.0 + 0.6 to 13.5 = 0.9 Ma (recalculated to new IUGS constants: Dalrym- ple 1979). Although the wood-containing tuff has not been dated, its occurrence at the top of the volcanic sequence suggests that it was deposited approximately 13.5 Ma. In the project area, the volcaniclastic rocks containing the fossil wood have been mapped by Dibblee and Ehrenspeck (1993a) as part of an andesite flow and tuff breccia unit. This gently north-dipping unit occurs at the top of the volcanic sequence and is overlain conformably by marine clastic rocks of the Upper To- panga Formation. An anticline has been mapped east of the wood site; however, no attitudes could be measured in the immediate sample area. Dibblee and Ehrenspeck (1993a) described these volcaniclastic units as ranging ‘from tan to light brown, massive, coherent, fine-grained feldspathic rock to less coherent, somewhat darker andesite breccia...composed of unsorted angular fragments in coherent matrix of same rock; deposited subaerially (?) as laharic flows.”’ A laharic flow or lahar is a volcanic debris flow in which air-borne vol- canic material that accumulates on the slopes of an active volcano becomes sat- urated with water and is mobilized, similar to lahars observed during the 1980 eruption of Mount St. Helens. Lahars can be mobilized during volcanic eruptions or long afterward and thus can be either warm or cold. In the case of the Conejo 18 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES lahars, the mobilizing event was probably heavy rainfall. The source of the lahars may occur 10 to 12 km to the west where possible dacite and andesite volcanic centers crop out near Lake Sherwood. Tuff deposits from these centers have been traced eastward beyond Lindero Canyon and Ladyface Mountain (Dibblee and Ehrenspeck 1993a). The tuff breccia, exposed during development, is composed of rounded lapilli and larger flow clasts in a fine-grained ash matrix. It is unsorted and poorly bedded. Thin sections of the matrix rock show that the material consists of eu- hedral phenocrysts of abundant, concentrically-zoned plagioclase and subordinate dipyramidal quartz and biotite set in an altered, possibly devitrified, matrix. The phenocrysts exhibit pervasive jigsaw fractures, and some occur in aggregates. The fracturing, observed in virtually all the thin section crystals, is a typical feature of hydroclastic deposits produced by lahars. At least a third, and possibly as many as half, of the phenocrysts are complete. This, combined with their angular shape and the unsorted and unbedded nature of the deposit, suggests that the matrix is a primary, not reworked, pyroclastic deposit. The tuff samples contain oblong inclusions 1 to 5 cm in length that are com- posed of quartz and sanidine (?) crystals set in smectite and possibly zeolite. These areas originally may have been pumice lapilli. Also present are spherical voids into which large zeolite laths have grown. A uniform isotropic substance, observed microscopically, may be kaolinitic clay with lesser amounts of small, possibly collapsed, pumice fragments, crystals of plagioclase and quartz, and la- pilli-sized clasts of lava, possibly rhyodacite. These microscopic characteristics and field analyses of the volcanic matrix encasing the fossil wood confirm that the rock was deposited as a lahar. Field Collection The resistance of the tuff breccia limited the amount of wood that could be salvaged, and the collection represents an incomplete geoflora. Fossil coniferous and dicotyledonous woods were concentrated along the southern flank of an an- desite ridge. The wood fragments appear to have been tumbled and worn. The surfaces of two logs found west of the fragments are splintered and have retained twig nodes. A 170 cm long section of a Ponderosa pine log was removed before recovery operations were halted. The greatest diameter of the fossil pine is 36 cm (Fig. 3A). Depending upon the quality of soil, Ponderosa pines today can attain di- ameters greater than 45 cm in 75 to 150 years (Hughes and Dunning 1949). This suggests that the tree was less than 75 years in age when it was caught in the lahar. The branch nodes and the splintered edges on the pine are unusually well preserved (Fig. 3B). A reddish sandy sediment, not occurring elsewhere in the tuff deposits, was plastered on one side of the log. R. B. Waitt (pers. comm. 1997) noted that during the 1980 eruption of Mount St. Helens, Washington, and the 1986 eruption of Augustine Volcano, Alaska, trunks and limbs of trees caught in lahars and cool pyroclastic flows show little or no charring and are commonly bruised, splintered, and sometimes shredded. None of the Conejo wood shows evidence of charring. A portion of an oak log, 31 cm in diameter and 175 cm long, was recovered FOSSIL WOOD FROM CONEJO VOLCANICS Ig Fig. 3. Fossil wood. A. Splintered and abraded section of fossil pine log, DMW 615. Width is 150 mm. B. Fossil pine limb with broken branch nodes, DMW 615. Width is 85 mm. 20 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 4 m north of the pine. Both logs were inclined relative to the horizon. Unlike the pine that was abraded and had no bark, the broken oak log has retained its fur- rowed bark. The complete dimensions of the oak and pine logs could not be measured before they were buried by construction activities. Sixteen m east of the two logs, three broken oak branches and numerous wood fragments were found. The branches are 28 to 30 cm in length and 5.5 cm in width. A polished cross section of branch CS 971 shows the trace of a stem and faint growth rings, rays, and bark. A cluster of wood fragments found south of the oak branches contained an unusual oak branch, 13 cm in length with 4-mm- thick bark, with five limb nodes that extend in different directions and range from 3 to 4 cm in width. Thin Section Analysis Selected samples were coated with Epo-Tek 301 resin prior to cutting to prevent splintering. The wood was then cut at 90° angles (cross section, radial, and tan- gential) into billets from which the thin sections were prepared. To achieve max- imum cell definition, the thin sections were polished to 35 to 45 pm in thickness. No cover slips were applied to the slides. With this technique and the remarkably high degree of preservation in the selected wood specimens, minor wood struc- tures, e.g. circular-bordered pits on tracheid cell walls, could be distinguished and used for identification (Fig. 4A). Based on thin section analysis, the fossil conifers have been identified as Doug- las Fir (Pseudotsuga menziesii), Incense Cedar (Calocedrus decurrens), and Pon- derosa Pine (Pinus ponderosa). The dicotyledonous taxa include the families Fa- gaceae (evergreen and deciduous oaks), Rosaceae (mountain mahogany and haw- thorn), and Rhamnaceae (mountain lilac or buck brush). The representative fossil woods selected for study are listed in Table 1. These include three dicots and two conifers that had cellular structures distinctive enough to be described (S. Carlquist pers. comm. 1996). Thin sections of a pine limb, Pinus ponderosa (RMW 5), contain resin canals, uniseriate rays, and uniseriate circular-bordered pits with tracheids that are squarish in outline (Fig. 4B). Conejo conifer RMW 12, possibly an Incense Cedar (Calocedrus decurrens) or a Douglas Fir (Pseudotsuga menziesii), has more rounded tracheids, as seen in thin section, uniseriate rays, resin canals, and circular-bordered pits. Conifers lack a diversity of water-conducting cells and the homogeneity of their cells produces discrete annual growth rings. Cross-section cuts of this fossil wood show annual rings as the contacts between the small cells of late wood and the large cells of early wood (Fig. 4C). The more complex cellular organization of the dicot wood shows distinct uni- seriate and multiseriate (aggregate) rays, diffuse and aggregate axial parenchyma with mostly solitary vessels, and fiber tracheids (Fig. 4D). Thin sections of an oak branch (RMW 2) have clearly defined tracheids and fiber tracheids, large vessels, and aggregate rays. The diversity of cellular organization in the oak thin sections suggests more than one species. Other dicots collected from the Conejo tuff breccia include a rosaceous wood specimen (RMW 4) that resembles, but does not belong to, the genus Heteromeles or Adenostoma. Thin sections of this unidentified dicot show diffuse axial paren- chyma, simple perforation plates, tracheids, possible fibers, short vertical uniser- FOSSIL WOOD FROM CONEJO VOLCANICS ps S22 s * 2 ele” - se ov. --;'-‘e™, 5s~* = 2a A St '@ #408? 0 aie | oe ere e Ot rv eeslee gies Se] Sse ase jetesees © ‘ = “.e } lo ¢ es Ce 2 a's jeg © = = Se ep Ny 5 pars 0 e > Re Sg: Orage Uepe ce ote se ‘ >". © - @ Ye e 2 fee ote ‘ an = -@ S&S. a OO a : x - ae 4 € ; 3 oe 108, Maggs! = BP Be i oth bes, 2) aA jbe‘e ‘409-4 2 Peed hestey @,25 24 § e. 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WO a me ~—* « ~~ y 2 Sx 3 ae ; “s ‘ { € he: fe > AVA vee Ls OR * . 4 ae Se « 4, > ee Ls Sao SS Lee Fig. 4. Photomicrographs of fossil wood from the Conejo Volcanics. A. Ponderosa pine (Pinus ponderosa); specimen RMW SC, radial section. Note circular-bordered pits on tracheid cell walls. Bar represents 85 wm. B. Ponderosa pine (Pinus ponderosa); specimen RMW 5A, cross section. Note the well defined, squarish tracheids with thin secondary walls. Two uniseriate rays extend vertically across the image. Bar represents 55 wm. C. Conifer (possibly Pseudotsuga or Calocedrus); specimen RMW 12 A, cross section. Two resin canals occur among the tracheids of early growth wood (E). Smaller tracheids represent late wood (L). (R) represents a ray. Bar represents 135 wm. D. Oak (Quercus sp.); specimen RMW 3C. Radial section shows xylem ray (R) and vessels (V) of axial system. Bar rep- resents 90 wm. iate rays, and multiseriate rays. The vessels are mostly solitary and growth rings were not observed. Specimen RMW 11 is identified as belonging to the family Rhamnaceae, possibly the genus Rhamnus or Ceanothus. Its thin sections show unusually narrow multiseriate rays with wings, uniseriate rays, vessels, and tra- cheids. Karowe and Jefferson (1987, p. 198) report that fossil woods, associated with tuffs from numerous localities and ages, can undergo silica replications of detailed cellular structures as small as bordered pits on tracheid walls. They noted that SOUTHERN CALIFORNIA ACADEMY OF SCIENCES N N [erpel pvsosapuod snuid OS MINY jeurIpnysuoy] ‘ds snuigd S19 MNWG jeuosur} psosapuod snuid as MN SNAPIIO[DI jenuosur} IO Dénsjopnasd OZI MN UOT]IOS SSOIO vsosapuod snuld VS MIN SNAPAIO[DI jerpes IO Dsnsjopnasd AzZI MN UOT]OOS SSOID SNODdvSOY Or MANY SNAPIIOIVI UOT]OAS SSOID IO Dsnsjopnasd V7l MN erpel SsnosdRsoYy dr MN [erpel snyjouvay Io snuwupyy OL MING jenuasur} SnOddeSOY Vr MN jenuosur) snyjouvay JO snuwpyy all MN jenuosur} DIOfIASD “CG “JO SNIAANO OF MIN onbrjgo snyjouvay Io snuupyy VII MN [erpel DILOfIASD °C “JO SNIAANO qe MN UOT]OAS SSOID IOJIUODZ 8 MN UOT]DOS SSOIO DIJOfIASD “CG “JO SNIAANO VE MANY UOT}DAS SSOID ‘ds sno1anO L MWY jenuosur} ‘ds sno1san@O ZT MN UOT]OAS SSOIO ‘ds snosanO IL6 SO UOT]DOS SSOIO JOTI Z, I MWY MOT A UOXeL # PS MNOTA uOxeL # SPS ‘sasATeUR UOT}DAS UI} 1OJ payooyas sopdures T S1QeL FOSSIL WOOD FROM CONEJO VOLCANICS 28} **... lahars provide conditions which are conducive to the excellent preservation of wood: burial is generally rapid and complete, leading to the isolation of organic matter within an anaerobic environment for thousands of years’? (Karowe and Jefferson 1987, p.203). The exceptional quality of preservation seen in the Conejo wood thin sections appears typical for flora preserved in lahars. Discussion Deposits of fossil wood from eastern Los Angeles Basin formations have been observed by the senior author in the middle to upper Miocene Monterey and Capistrano Formations of the San Joaquin Hills and the foothills of the Santa Ana Mountains, and in the upper Miocene Puente Formation in the Puente Hills and San Jose Hills. Remains of wood and leaves collected from these formations and the Modelo Formation have been studied extensively by Axelrod (1956). He noted that they belong to a subtropical south coastal province and represent a distinctive assemblage that he termed the Madro-Tertiary Geoflora. Included in this flora are willows (Salix), buck brush (Ceanothus), avocado (Persea), palms (Sabal), syc- amore (Platanus), live oak (Quercus), cottonwood (Populus), mountain mahogany (Cercocarpus), juniper (Juniperus), and magnolia (Magnolia). This assemblage from the Miocene Los Angeles basin region appears *‘... to have survived in mild coastal valleys adjacent to the coast ...’’; similar flora live today “*... in tropical and subtropical regions extending from Mexico southward”’ (Axelrod 1956, p. 262). In attempting to compare the Conejo wood assemblage with Ax- elrod’s Madro-Tertiary Geoflora, it should be noted that fossil Miocene trees (syc- amore, palm, and avocado), common in other Los Angeles Basin formations, are notably absent. The subtropical paleoenvironment that characterized southern California in the middle Miocene has become more temperate as temperatures have declined. It is estimated that present-day oaks and conifers live ~600 m lower in elevation than during the warm middle Miocene (J. R. Haller pers. comm. 1997). On the western slopes of the central Sierra Nevada, Ponderosa Pine and Incense Cedar, species represented in the Conejo geoflora, live today from ~700 to 1,500 m elevation (Hughes and Dunning 1949). This implies that the 13.5-Ma Conejo geoflora lived at elevations between 1,300 and 2,100 m. Axelrod’s (1956) Arcto-Tertiary Geoflora from cooler paleoclimates includes maples (Acer), mountain lilac or buck brush (Ceanothus), Oregon grape (Maho- nia), pines (Pinus), alder (Alnus), aspens and cottonwoods (Populus), ash (Frax- inus), Willows (Salix), redwood (Sequoiadendron), Douglas Fir (Pseudotsuga), spruce (Picea), and sumac (Rhus). Apparently missing are oaks, which are com- mon in the Conejo flora, and this absence of oaks makes it impossible to assign the Conejo geoflora to the Arcto-Tertiary Geoflora. Unlike the Tertiary geofloras described by Raven and Axelrod (1995), the Co- nejo flora appears closely related to oak and pine forests, found today in the west- central Sierra Nevada Mountains and northern California. These transitional for- ests contain Valley Oak, Live Oak, Coast Live Oak, Pacific Madrone, California Laurel, Incense Cedar, Douglas Fir, and Ponderosa Pine (Kricher and Morrison 1993, p. 328). The life zone of this conifer/hardwood forest belongs to the Low Montane Zone of 730 to 1,850 m elevation with an annual precipitation of 62.5 cm (Kricher and Morrison 1993, p. 282—287). 24 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES The geoflora preserved in the Conejo Volcanics tuff appears to represent a low montane forest that grew on an active volcanic highland ~13.5 Ma when coastal temperatures were subtropical. The gradual decline in temperatures since the me- dial Miocene and the composition of the Conejo geoflora suggest that the ele- vation of this volcano was greater than 1,300 m. It is plausible that a lahar carried conifers from a higher elevation down slope where the pines, firs, and cedars were buried with oaks and other dicots. However, the absence of genera common in other Los Angeles Basin Miocene deposits (e.g. palms, sycamores, and avocados) suggests that the Conejo fossil wood may represent a low montane geoflora pre- viously undescribed from the geologic record of the Los Angeles Basin region. Acknowledgments We thank Dr. David Whistler for early encouragement on this project. The authors wish to acknowledge the construction site personnel from J. D. Diffen- baugh, Inc. Their cooperation and the work of the RMW Paleo Associates, Inc. field monitors expedited the fossil wood recovery. Our gratitude is also extended to the following botanists at the Santa Barbara Botanic Garden: Dr. Edward L. Schneider, who reviewed the thin sections; Dr. Sherwin Carlquist, who identified the wood; and Dr. J. Robert Haller, who discussed the Neogene paleoenviron- mental conditions of southern California. Jeffrey A. Myers, Department of Geo- logical Sciences, University of California, Santa Barbara, made the thin section analyses of the tuff. Dr. Tanya M. Atwater, Department of Geological Sciences, University of California, Santa Barbara, has provided valuable direction. Richard C. Meyers, Scripps Institution of Oceanography, prepared the thin sections and supervised the photomicrography. Figures were drafted by Karen L. Savage and Kathryn van Roosendaal. Laboratory work, including thin-section preparation, was funded by a grant from RMW Paleo Associates, Inc. We appreciate comments on early versions of this manuscript from Marilyn W. Morgan, RMW Paleo As- sociates, Inc., Helmut E. Ehrenspeck, Dibblee Geological Foundation, and Ray- mond Prouty, Historian, City of Westlake Village. The fossil wood specimens have been deposited at the Santa Barbara Botanic Garden, the Geology Department of Pomona College, the Santa Monica National Recreation Area Visitor’s Center, and the City of Westlake Village, California. Literature Cited Axelrod, D. I. 1956. Mio-Pliocene Floras from West Central Nevada: Univ. of California Publ. in Geol. Sci., 33:1-—322. Dalrymple, G. B. 1979. Critical tables for conversion of K-Ar ages from old to new constants: Ge- ology, 7:558-—560. Dibblee, T. W., Jr., and H. E. Ehrenspeck. 1993a. Geologic map of the Thousand Oaks quadrangle, Ventura and Los Angeles Counties, California: Santa Barbara, California, Dibblee Geological Foundation, Map #DF-49, scale 1:24,000. Dibblee, T. W., Jr., and H. E. Ehrenspeck., 1993b. Field relations of Miocene volcanic and sedimentary rocks of the western Santa Monica Mountains, California. Pp. 75—92 in Weigand, P. W., A. E. Fritsche, and G. E. Davis, eds., Depositional and volcanic environments of middle Tertiary rocks, in the Santa Monica Mountains, southern California: SEPM (Society for Sedimentary Geology), Pacific Section, Book 72. Hughes, B. O., and D. Dunning. 1949. Pine forests of California. Pp. 352-358 in Trees—The yearbook of agriculture. Washington, D. C., U.S. Department of Agriculture, 353 pp. FOSSIL WOOD FROM CONEJO VOLCANICS 25 Karowe, A. L., and T. H. Jefferson, 1987, Burial of trees by eruptions of Mount St. Helens, Washing- ton: Implications for the interpretation of fossil forests: Geological Magazine, 24:191—302. Kricher, J. C., and G. Morrison. 1993. Ecology of Western Forests: The Peterson Field Guide Series, Boston—New York, Houghton Mifflin Company: 281-290. Raven, P. H., and D. I. Axelrod. 1995. Origin and relationships of the California Flora. Univ. of Calif. Publ. in Botany, 72:9—43. Turner, D. L. 1970. Potassium-argon dating of Pacific Coast Foraminiferal Stages: Geol. Soc. of Amer. Special Paper, 124:91—129. Yerkes, R. E., and R. H. Campbell. 1979. Stratigraphic nomenclature of the central Santa Monica Mountains, Los Angeles County, California: U. S. Geol. Surv. Bulletin 1457-E: 31 pp. Accepted for publication 7 May 1998. Bull. Southern California Acad. Sci. 98(1), 1999, pp. 26-38 © Southern California Academy of Sciences, 1999 First Record of [socyamus kogiae Sedlak-Weinstein, 1992 (Crustacea, Amphipoda, Cyamidae) from the Eastern Pacific, with Comments on Morphological Characters, a Key to the Genera of the Cyamidae, and a Checklist of Gyamids and their Hosts Joel W. Martin’ and John E. Heyning Research and Collections Branch, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007 Abstract.—The cyamid amphipod species [socyamus kogiae Sedlak-Weinstein, 1992, is reported for the first time from southern California, extending the known range of the species from Moreton Island, Queensland, Australia, to the north- eastern Pacific. Additional descriptive notes are provided based on a single adult male taken from a pygmy sperm whale, Kogia breviceps (de Blainville, 1838), stranded near San Diego, California. Morphological differences between this spec- imen and the description of the type series are discussed. A revised key to the six currently recognized genera of the family Cyamidae is provided, as well as a checklist of all described species of the family, their cetacean hosts, and the ranges of those hosts. The amphipod crustacean family Cyamidae Rafinesque, 1815, is a relatively species-poor taxon, all members of which are ectoparasites on cetaceans (Laubitz 1982). Subsequent to Leung’s (1967) review, where only 16 species in 5 genera were treated, there have been only one new genus (Scutocyamus Lincoln and Hurley, 1974) and relatively few other species described (e.g. Leung 1970b, Lin- coln and Hurley 1974, 1980, 1981, Berzin and Vlasova 1982, Waller 1989, Sed- lak-Weinstein 1992a, b). Currently, the family contains six genera and approxi- mately 27 species (see Table 1). Prior to 1992, there were no descriptions of cyamids associated with pygmy sperm whales, Kogia breviceps (de Blainville, 1838). Although Caldwell et al. (1971) reported a cyamid attached to one of seven pygmy sperm whales from the western Atlantic examined by them, they did not describe the cyamid, referring to it only as ‘“‘Cyamidae, form D, genus and species undetermined” (Caldwell et al. 1971: 4). All specimens of cetacean parasites mentioned in that paper as being in the personal collection of Stephen Zam were subsequently lost in an office move (personal communication, Stephen Zam, 29 June 1993). In 1992, a new species of cyamid was described by Sedlak-Weinstein (1992a) from a pygmy sperm whale that stranded on Moreton Island, Queensland, Aus- tralia, marking the first record of cyamids taken from the genus Kogia (with the exception of the western Atlantic record above). Sedlak-Weinstein’s species proved to belong to the genus Jsocyamus, but differed sufficiently from J. del- phinii, the sole previous member of the genus, to necessitate the erection of a second species of the genus, Isocyamus kogiae. A third species of Isocyamus, I. ‘ Corresponding author: Joel W. Martin, Phone 213-763-3440, Fax 213-746-2999 26 FIRST RECORD OF JISOCYAMUS KOGIAE FROM EASTERN PACIFIC Poe | Table 1. Known cyamid amphipods of the world and their cetacean hosts and distributions, com- piled mostly from Leung (1965, 1967), Gruner (1975), Berzin and Vlasova (1982), and Sedlak-Wein- stein (1991, 1992a, b). Additional earlier specific names now considered junior synonyms are listed in Gruner (1975). Selected host references are not meant to be exhaustive; numerous additional ac- counts exist in the cetacean and crustacean literature. Taxonomy of the host species has been updated based on Mead and Brownell (1993). Family Cyamidae Rafinesque, 1815 Selected Host References Genus Cyamus Latreille, 1796 Cyamus antarcticensis Vlasova, 1982, in Berzin and Vlasova Orcinus orca (Linnaeus, 1758) (killer whale; worldwide) Cyamus bahamondei Buzeta, 1963 Physeter catodon Linnaeus, 1758 (sperm whale; non-polar worldwide) Cyamus balaenopterae Barnard, 1931 Balaenoptera musculus (Linnaeus, 1758) (blue whale; worldwide) Balaenoptera physalus (Linnaeus, 1758) (fin whale; worldwide) Balaenoptera acutorostrata Lacépede, 1804 (minke whale; worldwide) Cyamus boopis Lutken, 1870 Megaptera novaeangliae (Borowski, 1781) (humpback whale; worldwide) Physeter catodon Linnaeus, 1758 (sperm whale; non-polar worldwide) unidentified New Zealand whale unidentified south Australian whale Cyamus catodontis Margolis, 1954 Physeter catodon Linnaeus, 1758 (sperm whale; non-polar worldwide) Cyamus ceti (Linnaeus, 1758) Balaena mysticetus Linnaeus, 1758 (bowhead whale; Arctic) Eschrichtius robustus Lilljeborg, 1861 (gray whale; North Pacific; also North Atlantic in historic times) Cyamus erraticus Roussel de Vauzeme, 1834 Eubalaena australis (Desmoulins, 1822) (southern right whale; southern hemisphere) Eubalaena glacialis (Muller, 1776) (northern right whale; northern hemisphere) Megaptera novaeangliae (Borowski, 1781) (humpback whale; worldwide) Cyamus gracilis Roussel de Vauzeme, 1834 Eubalaena australis (Desmoulins, 1822) (southern right whale; southern hemisphere) Eubalaena glacialis (Muller, 1776) (northern right whale; northern hemisphere) Berzin and Vlasova, 1982 Leung, 1965; Gruner, 1975 Berzin & Vlasova, 1982 Gruner 1975 Leung 1965; Gruner, 1975 Gruner, 1975 Leung, 1965; Gruner, 1975 Gruner, 1975 Berzin & Vlasova, 1982 Sedlak-Weinstein, 1991 Sedlak-Weinstein, 1991 Keune, 1965-"Gruner, 1975 Berzin & Vlasova, 1982: Fransen and Smeenk, 1991 Gruner, 1975 Leung, 1965; 1976 Lueng, 1965 Gruner, 1975 Leung, 1965: Gruner, 1975 euns; 1965: (Gruner, 1975 Gruner, 1975 Leung, 1965; Gruner, 1975 Leung, 1965: (Gruner, 1975 28 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1. Continued. Selected Host References Family Cyamidae Rafinesque, 1815 Cyamus kessleri A. Brandt, 1873 Eschrichtius robustus (Lilljeborg, 1861) (gray whale; North Pacific; also North Atlantic in historic times) Cyamus monodontis Lutken, 1870 Monodon monoceros Linnaeus, 1758 (narwhal; Arctic) Delphinapterus leucas (Pallas, 1776) (beluga; Arctic) Cyamus nodosus Lutken, 1861 Monodon monoceros Linnaeus, 1758 (narwhal; Arctic) Delphinapterus leucas (Pallas, 1776) (beluga; Arctic) Cyamus orcini Leung, 1970b Orcinus orca (Linnaeus, 1758) (killer whale; worldwide) Cyamus orubraedon Waller, 1989 Berardius bairdii Stejneger, 1883 (Baird’s beaked whale; North Pacific) Cyamus ovalis Roussel de Vauzeme, 1834 Eubalaena australis (Desmoulins, 1822) (southern right whale; southern hemisphere) Eubalaena glacialis (Muller, 1776) (northern right whale; northern hemisphere) Physeter catodon Linnaeus, 1758 (sperm whale; non-polar worldwide) Cyamus rhytinae (J. EK Brandt, 1846) Hurley and Mohr, 1957 Leung, 1965; Gruner, 1975 Leung, 1965; Gruner, 1975 Gruner, 1975 Leung, 1965; Gruner, 1975 Gruner, 1975 Leung, 1970b; Gruner, 1975 Waller 1989 Gruner, 1975; Leung, 1965, 1970a, b Leung, 1965; Gruner, 1975 Leung, 1965; Gruner, 1975 Leung, 1965; Gruner, 1975 Berzin & Vlasova, 1982 {dubious species, supposedly found on Steller’s sea cow; See Gruner, 1975: 88} Cyamus scammoni Dall, 1872 Eschrichtius robustus Lilljeborg, 1861 (gray whale, North Pacific; also North Atlantic in historic times) Genus /socyamus Gervais & Van Beneden, 1859 Isocyamus delphinii Guerin-Meneville, 1837 Globicephala melas (Traill, 1809) (long-finned pilot whale; temperature waters) Globicephala macrorhynchus Gray, 1846 (short-finned pilot whale; temperate and tropical, worldwide) Pseudorca crassidens (Owen, 1846) (false killer whale; temperate and tropical, worldwide) Gruner, 1975 Leung, 1965, 1976 Sedlak-Weinstein, 1991 Gruner, 1975 Berzin and Vlasova, 1982 Sedlak-Weinstein, 1992a Leung, 1965 Raga et al., 1983 Hiro, 1938 Sedlak-Weinstein, 1992a Sedlak-Weinstein, 1991 Gruner, 1975 Sedlak-Weinstein, 1992a Bowman, 1955 FIRST RECORD OF JSOCYAMUS KOGIAE FROM EASTERN PACIFIC 29 Table 1. Continued. Family Cyamidae Rafinesque, 1815 Steno bredanensis (Lesson, 1828) (rough toothed dolphin; temperate and tropical worldwide) Delphinus delphis* Linnaeus, 1758 (common dolphin; temperate and tropical, worldwide) Grampus griseus (G. Cuvier, 1812) (Risso’s dolphin; temperate and tropical, worldwide) Phocoena phocoena (Linnaeus, 1758) (harbor porpoise; northern hemisphere, temperate ) Tursiops truncatus (Montagu, 1821) (bottlenose dolphin; tropical and temperate) Lagenorhyncus albirostris Gray, 1846 (white-beaked dolphin; temperate North Atlantic) Mesoplodon europeaus (Antillean beaked whale; North Atlantic) Orcinus orca (killer whale; worldwide) TIsocyamus deltobranchium Sedlak-Weinstein, 1992b Globicephala macrorhynchus Gray, 1946 (short-finned pilot whale; temperate and tropical waters) Globicephala melas (Triall, 1809) (long-finned pilot whale; temperate) Isocyamus kogiae Sedlak-Weinstein, 1992a Kogia breviceps (de Blainville, 1838) (pygmy sperm whale; tropical to warm temperate) Genus Neocyamus Margolis, 1955 Neocyamis physeteris (Pouchet, 1888) Physeter catodon Linnaeus, 1758 (sperm whale; non-polar worldwide) Phocoenoides dalli (True, 1885) (Dall’s porpoise; North Pacific, temperate) Genus Platycyamus Lutken, 1870 Platycyamus thompsoni (Gosse, 1855) Mesoplodon grayi Von Harst, 1876 (Gray’s beaked whale; southern hemisphere) Hyperoodon planifrons Flower, 1882 (southern bottlenose whale; southern hemisphere) Hyperoodon ampullatus (Forster, 1770) (northern bottlenose whale; northern hemisphere) Selected Host References Gruner, 1975 Lincoln and Hurley, 1974 Sedlak-Weinstein, 1992b Gruner, 1975 Berzin and Vlasova, 1982 Sedlak-Weinstein, 1992a Gruner, 1975 Berzin and Vlasova, 1982 Sedlak-Weinstein, 1992a Gruner, 1975 Stock, 11977. Berzin and Vlasova, 1982 Sedlak-Weinstein, 1992a Fransen and Smeenk, 1991 Balbuena and Raga, 1991 Sedlak-Weinstein, 1992a Stock, 1977 Sedlak-Weinstein, 1992a Fransen and Smeenk, 1991 Balbuena and Raga, 1991 Sedlak-Weinstein, 1992a Best, 1969 Sedlak-Weinstein, 1992a Sedlak-Weinstein, 1992b Sedlak-Weinstein, 1992b Sedlak-Weinstein, study 1992a; This Leung, 1965; Gruner, 1975 Berzin and Vlasova, 1982 Gruner, 1975 Berzin and Vlasova, 1982 Sedlak-Weinstein, 1991 Berzin and Vlasova,1982 Gruner, 1975 Berzin and Vlasova, 1982 Fransen and Smeenk, 1991 30 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table |. Continued. Selected Host References Family Cyamidae Rafinesque, 1815 Platycyamus flaviscutatus Waller, 1989 Berardius bairdii Stejneger, 1883 (Baird’s beaked whale; North Pacific) Genus Scutocyamus Lincoln and Hurley, 1974 Scutocyamus parvus Lincoln and Hurley, 1974 Lagenorhynchus albirostris (Gray, 1846) (white-beaked dolphin; temperate North Atlantic) Scutocyamus antipodensis Lincoln and Hurley, 1980 Cephalorhynchus hectori (Van Beneden, 1881) (Hector’s dolphin; New Zealand) Genus Syncyamus Bowman, 1955 Syncyamus chelipes (Costa, 1866) Delphinus delphis' Linnaeus, 1758 (common dolphin; temperate and tropical waters) Syncyamus pseudorcae Bowman, 1955 Pseudorca crassidens (Owen, 1846) (false killer whale; temperate and tropical waters) Syncyamus aequus Lincoln and Hurley, 1981 Tursiops truncatus (Montagu, 1821) (bottlenose dolphin; tropical and temperate) Stenella longirostris* (Gray, 1828) (Spinner dolphin; tropical) Stenella coeruleoalba (Meyen, 1833) (striped dolphin; tropical and temperate) Waller, 1989 Leung, 1967, 1970a; Berzin and Vlasova, 1982 Lincoln & Hurley, 1974 Gruner, 1975 Fransen and Smeenk, 1991 Lincoln & Hurley, 1980; Sedlak-Weinstein, 1991 Gruner, 1975 Bowman, 1955 Gruner, 1975 Sedlak-Weinstein, 1991 Sedlak-Weinstein, 1991 Sedlak-Weinstein, 1991 Raga and Raduan, 1982 Raga, 1988 Syncyamus sp. Bowman, 1958 (close to S. pseudorcae Bowman, 1955) Stenella attenuata* (Gray, 1846) (spotted dolphin; pan-tropical) Bowman, 1958 Gruner, 1975 Syncyamus sp. Leung, 1970 (close to S. pseudorcae Bowman, 1955) Delphinus delphis' Linnaeus, 1758 (common dolphin; temperate and tropical) Stenella coeruleoalba (Meyen, 1833) (striped dolphin; tropical and temperate) Stenella longirostris? (Gray, 1828) (spinner dolphin; tropical) Tursiops truncatus (Montagu, 1821) (bottlenose dolphin; tropical and temperate) Gruner (1975: 93) lists as “‘doubtful species”’ the following: Cyamus latreilleii A. Smith, 1831 Cyamus leachii A. Smith, 1831 ' Delphinus delphis has been divided into D. delphis and D. capensis Gray, 1828; see Heyning and Perrin (1994). * Stenella longirostris has been divided into S. longirostris and S. clymene (Gray, 1846); see Perrin et al. (1981). * Stenella attenuata has been divided into S. attenuata and S. frontalis (G. Cuvier, 1829); see Perrin et al. (1987). Leung, 1965; Gruner, 1975 Leung, 1965; Gruner, 1975 Leung, 1965; Gruner, 1975 Leung, 1965; Gruner, 1975 FIRST RECORD OF ISOCYAMUS KOGIAE FROM EASTERN PACIFIC 31 deltobranchium, has now been described from Japanese and Australian pilot whales (see Sedlak-Weinstein 1992b). On 21 May 1993, a pygmy sperm whale stranded alive in San Diego County, California. The whale was taken to Sea World™ in San Diego where it died the same day. The whale was frozen intact and then transported to the Natural History Museum of Los Angeles County (LACM). The cetacean was subsequently found to have a single attached cyamid amphipod, which we have attributed to Isocy- amus kogiae Sedlak-Weinstein, 1992a. However, sufficient morphological differ- ences exist between this specimen and the original description of the type of J. kogiae, and between other members of the genus Jsocyamus, that we felt it ap- propriate to describe and discuss our specimen below. Additionally, we provide a more detailed diagnosis of the genus Jsocyamus, a key to the six known cyamid genera, and an updated checklist of species of the Cyamidae and their hosts. Materials and Methods The stranded pygmy sperm whale was a 3.00 m male, the skeleton and selected tissues of which are catalogued in the Natural History Museum of Los Angeles County’s collection of mammals as LACM 88938. The cyamid was found on the dorsolateral flank of the host, approximately two-thirds the body length posterior to the head. The dactyli of the pereiopods were firmly embedded in the epidermis of the whale. The whale’s body at the site of the cyamid was free of wounds. However, prior to the discovery of the cyamid, the whale was transported by truck to Sea World, subsequently died, and was frozen. Therefore, the relative position of the cyamid upon the whale’s body may be a post-stranding artifact. Illustrations were made with a Wild MSAPO stereo microscope and a Nikon Labophot™, both equipped with a drawing tube. The cyamid specimen is cata- logued as LACM 93-34.1 in the Crustacea collections of the Natural History Museum of Los Angeles County, a collection that contains the most extensive holdings of cyamids in the world (see Leung 1965). Results Isocyamus Gervais and van Beneden, 1859 Emended diagnosis.—Body thin and elongate, not ovate. Pigment lacking, im- parting whitish color. Head widest at base, tapering slightly anteriorly. Antenna 1 long, exceeding length of either first or second pereiopod, composed of 4 ar- ticles. Antenna 2 with 3 articles (possibly 4 articles; distinction between distal most articles difficult to discern). Maxillipeds extremely reduced, flap-like. Max- illipedal palps lacking. First pereon fused with head. All subsequent pereonal somites fused dorsally at midline. Pereiopod (gnathopod) | slightly shorter and much thinner than pereiopod 2, composed of 6 articles. Dactylus of pereiopod 1 with comb row along cutting border; comb row either single (. delphinii) or double, extending from tip proximally to at least midway point along ventral border. Dactylus with or without distinct unguis. Pereiopod (gnathopod) 2 robust, composed of 4 articles; dactylus smooth. Somite 2 with epaulet-like infoldings of cuticle along anterodorsal border. Somites 3 and 4 bearing single (unpaired), elon- gate and tubular primary gill and well-developed accessory gill equal in size to (I. delphinii) or smaller than primary gills. Sternal area just posterior to and lateral to each primary gill bearing laterally directed acute process. Oostegites rounded, 32 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES not triangular or wedge-shaped. Pereiopods 5, 6, and 7 similar, each terminating in smooth, sickle-shaped dactylus (as per family), each lacking ventral spines on any article, each with dorsal ridge-like process extending proximally on basis; this process most obvious on pereiopod 5. Adjacent sternal region with or without (I. kogiae) ventrally-directed spines. Species: Isocyamus delphinii (Guerin-Meneville, 1836); Isocyamus kogiae Sed- lak-Weinstein, 1992a; Isocyamus deltobranchium Sedlak-Weinstein, 1992b (see Table 1 for host species). Comparison of Jsocyamus kogiae specimens from California and Australia Our specimen (Figs. 1, 2), a mature male, agrees with Sedlak-Weinstein’s (1992a) original description of J. kogiae in most major morphological features, such as the relative size of the gnathopods, size and shape of the head, fusion of all pereonites dorsally, presence of epaulet-like processes on the second pereon, size and shape of primary and accessory gills, absence of sternal spines adjacent to pereiopods 5—7, and presence of an acute tooth on the propodus of gnathopod 2. Size of the specimens is also in agreement; our specimen (measured following Sedlak-Weinstein 1992a, and with corresponding measurements of the holotype from Australia given in brackets) measured: total length 4.7 mm [4.75]; width of body 1.4 mm [1.5]; length of primary gill 1.0 mm [1.0]; length of propodus of first gnathopod 0.7 mm [0.5]. Because our sole specimen is a mature male, we can not offer comparisons with Sedlak-Weinstein’s allotype. Our specimen differs from the Australian specimens in having a more strongly curved dactylus on pereiopods 5-7, a slightly different shape to the propodal distal expansion of the second gnathopod (which in the type specimens also bears a second, more proximal, blunt expansion not seen on our specimen), and a slightly different shape to the dactylus and propodus of gnathopod 1. In our specimen, the rounded ventral border of the propodus of gnathopod 1 bears a row of minute spines or teeth, corresponding to what is found on the dactylus; these spines were not mentioned or illustrated by Sedlak-Weinstein (1992a), but may have been overlooked. Another difference is that the accessory gills of our specimen are distally rounded (Figs. la, c), whereas they appear more tapered in Sedlak-Wein- stein’s illustration (1992a: fig. 2), but are well rounded in the paratypes (T. Haney, pers:comy); Because we had only one specimen, we did not remove mouthparts for separate illustration or subject the specimen to SEM; mouthparts shown here (Fig. 2c) were illustrated in situ, so details are more difficult to see and illustrate. Still, some differences in the mouthparts between our specimen and the Australian series are evident. The most obvious is that in our specimen the second maxillae are fused basally, whereas in Sedlak-Weinstein’s illustration (1992a: 4, fig. 11) these are shown as being separate basally. Additionally, her figure of the maxil- lipeds shows two distinct processes, each with three terminal setae (4—5 in text), whereas we detected only a small, crescent-shaped flap of cuticle, and did not see any setation. Finally, our specimen differs from the Australian specimens of J. kogiae, and indeed from all other described cyamids, in having the last pair of legs displaced anteriorly so that they fall anterior to the penultimate pair; this is unique in the family. Although perhaps appearing as an artifact, the amount of dislocation of FIRST RECORD OF JSOCYAMUS KOGIAE FROM EASTERN PACIFIC 35 Fig. 1. Jsocyamus kogiae Sedlak-Weinstein, 1992a, from pygmy sperm whale (Kogia breviceps) stranded in San Diego, California. a, Entire animal, ventral view. b, same, dorsal view. c, Primary and accessory gills and ventrally projecting processes of somites 3 and 4. Note ventrolateral process of somite 3 directed more laterally than that of somite 4. d, Dorsal view of somite 2. Note epaulet- like infoldings of cuticle at anterodorsal margins. e, First gnathopod with tip of dactylus enlarged at lower right showing comb row. f, Second gnathopod. g, Ventral view of proximal two articles of gnathopod 2 and part of propodus. the legs that would be necessary to return the posterior most leg to its “‘usual’”’ position is substantial, and would involve bilaterally pulling the back legs a con- siderable distance upward in order for the long claw-like dactylus to clear the other legs. This very unusual feature is immediately diagnostic, but could be overlooked in a disarticulated or damaged specimen, or even one that has been manipulated to the point that the pereiopods are in the “‘normal’’ arrangement. 34 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 2. Isocyamus kogiae Sedlak-Weinstein, 1992a, from pygmy sperm whale (Kogia breviceps) stranded in San Diego, California. a, First and second antenna. b, Close up of second antenna showing indistinct suture at base of distal most article. c, Entire mouthfield, ventral view. d, Pereiopod 7. e, Abdominal region and male genitalia, ventral view. This condition does not occur in the collection of 9 females and 5 males of /. kogiae from Australia. Based on known variability of some cyamid features (e.g. Raga 1988), and because of the similarities of the southern California specimen to Sedlak-Wein- stein’s (1992a) description of Isocyamus kogiae from the same host (Kogia brev- iceps) off Australia, we do not believe the above differences warrant erection of another species of the genus. Currently Recognized Genera of the Cyamidae As currently recognized, the family Cyamidae contains only the genera Cyamus Latreille, 1796; Isocyamus Gervais and van Beneden, 1859; Platycyamus Lutken, 1870; Syncyamus Bowman, 1955; Neocyamus Margolis, 1955; and Scutocyamus Lincoln and Hurley, 1974. Most species in the family are members of the genus Cyamus, currently containing 16 nominal species (Table 1; see also Gruner 1975, FIRST RECORD OF JSOCYAMUS KOGIAE FROM EASTERN PACIFIC 35 Berzin and Vlasova 1982). Species within the genus Cyamus have a second max- illa that bears an outer lobe and possess a maxillipedal palp (variously developed, and absent in early stages of some species, and lacking in all stages of Cyamus nodosus [T. Haney, personal communication]), features lacking in members of the other genera (Bowman 1955, Leung 1967, Lincoln and Hurley 1974). Other genera of the Cyamidae contain few species; Neocyamus is monotypic, and the genera Platycyamus, Syncyamus, and Scutocyamus have but two species each. Isocyamus was erected by Gervais and van Beneden (1859), and later de- scribed in more detail by Barnard (1932), to accommodate J. delphinii (often incorrectly spelled delphini), originally described (as Cyamus) by Guerin-Mene- ville (1836) from the common dolphin, Delphinus delphis. Isocyamus dephinii has since been reported from a number of small odontocetes (listed in table 1 of Sedlak-Weinstein 1992a; see also Table 1). Leung (1967), in his key to the genera, employed as a character separating Isocyamus from Cyamus the fact that Jsocyamus bears laterally directed spines at the bases of the gills. Unfortunately, these laterally directed spines are not unique to Isocyamus; Leung himself illustrated these quite clearly for Cyamus kessleri (see Leung 1967: fig. 4a), and they are also evident in the most recently described member of that genus, Cyamus antarcticensis Berzin and Vlasova (1982: 152, figs. LA, 2G), as well as in Jsocyamus. Thus, Leung’s (1967) key to cyamid genera was seriously flawed even at the time of publication, and included only 5 genera and 16 species. There exists today no valid key for the separation and identifi- cation of cyamid genera. An additional problem is that some morphological char- acters employed in cyamid taxonomy were poorly or incompletely described in the original species accounts. Mouthparts, details of which may prove to be the best or the only indicators of phylogenetic affinity, remain undescribed—or are incorrectly described—for several genera and most species. For example, although Bowman (1955) stated that Jsocyamus has a 2-segmented palp on the first maxilla, this is incorrect as all three species, J. delphinii, I. deltobranchium, and I. kogiae, have a l-segmented palp on this appendage (Sedlak-Weinstein 1992a, b). Details of the male abdomen and reproductive structures are sketchy at best in the liter- ature, and are not illustrated for most species of Cyamus; even the terminology and homologies of these structures are unclear. Clearly, although the Cyamidae contains relatively few genera and species, there remains a considerable amount of taxonomic confusion, and a systematic revision of the group, including rede- scriptions of all species, is badly needed. To facilitate more accurate identification of cyamids, at least to the level of genus, we have constructed a revised key that is admittedly artificial, employing what appear to us (from the literature and from examination of select taxa in the LACM Crustacea Collection) to be fairly con- servative characters. Revised Key to the Genera of the Cyamidae 1. Gills fasciculate (highly branched, appearing as bundles). Males lacking CRESS OGY ONS: Petes crete ess ee Nee meee ce een otc cotta as a Jere Me an Neocyamus — Gills not fasciculate. Males with or without accessory gills ............. 2 2. Gnathopods | and 2 approximately equal in size. Pereon | extending lat- erally beyond lateral margins of head, clearly separated from head by amet imdentatony ease et oak aches ) maids Bot tahe ee SE Platycyamus 36 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES — Gnathopod 1 distinctly smaller (shorter and thinner) than gnathopod 2. Pereon | not extending laterally beyond lateral margins of head, or if so then fused to head, not separated from it by distinct indentation ........ 3 3. Pereon 3 and 4 greatly reduced in male, fused dorsally in both sexes. Gnathopod 2 composed of 3 articles. Pereon 6—7 fused. Males without accessory GillsiG aR i Sit oe Oe PR sR ees, DE OO 4 Scutocyamus — Pereon 3 and 4 not reduced or reduced in male, fused or unfused dorsally in both sexes. Gnathopod 2 composed of 5 articles. Pereon 6—7 fused or unfused, Males withiaccessory eilisst. 0. ree RI, Se ee 4 4. Antenna 2 with 4 articles. Maxilliped occasionally with palp in adult. Pereon 2 without infoldings of cuticle along anterodorsal region ... Cyamus — Antenna 2 composed of 2 or 3 articles. Maxilliped greatly reduced, never with palp in adult. Pereon 2 with epaulet-like cuticular infoldings on an- terodorsalresiomns Suc ee Os ee OR, EO) A Aa 5 5. Combined head and pereon | arising from recessed area along front of pereon 2. Antenna | shorter than either gnathopod and shorter than fused head + pereon 1. Pereiopods 5—7 short, heavy. Adults not greater than 3 ministotal Meme chat ae on ls ee , PO A SY ol ae Syncyamus — Combined head and pereon | not arising from recessed area in pereon 2. Antenna | elongate, clearly exceeding length of head + pereon 1. Pereio- pods 5-7 elongate, delicate. Adults greater than 4 mm total length (usually G2 Fumi): eee ten, ISS, Ee AU 2 ER Re Te a ese Isocyamus Discussion In the first report of any cyamid from a pygmy sperm whale, Caldwell et al. (1971) reported that the undescribed cyamid (now lost) was taken from “infected tissue in [an] unhealed open lesion penetrating the skin into the underlying adipose tissue on the side of the body.’ The Kogia breviceps from which that cyamid was taken was a 3.2 m male (Caldwell and Caldwell 1989: 255) from northeastern Florida in the vicinity of St. Augustine (Caldwell et al. 1971: 4). Similarly, Sed- lak-Weinstein’s (1992a) pygmy sperm whale cyamids were collected “from among 20 ’golf ball-sized’ wounds” on the stranded host. These described wounds are characteristic of those inflicted by cookie-cutter sharks. In contrast, the southern California cyamid specimen from Kogia breviceps was found firmly attached to healthy tissue just posterior to midlength. Interestingly, only species of the genus Cyamus have been taken from baleen whales (suborder Mysticeti), whereas species of all six genera, including Cyamus, have been found on various members of the toothed whales (suborder Odontoceti) (see Leung 1967 and Table 1). Unfortunately, all systematic investigations of cyamids to date have been of a phenetic rather than cladistic nature, precluding for the moment comparisons of cyamid and cetacean phylogenies. Acknowledgments We are grateful to Dr. Stephen Zam for providing information on the eventual fate of the collection of cetacean parasites described in Caldwell et al. (1971), to the staff of Sea World™ of San Diego, California, for their assistance, to Mr. Grant Graves for help in locating literature, and to Don Cadien, Tim Stebbins, FIRST RECORD OF ISOCYAMUS KOGIAE FROM EASTERN PACIFIC oY and especially Todd Haney for their useful comments on this manuscript. We further thank Todd Haney for examining paratypes of Queensland Museum spec- imens of Jsocyamus kogiae (Reg. No. W17108) for comparison with our descrip- tion. Literature Cited Balbuena, J. A., and J. A. Raga. 1991. Ecology and host relationships of the whale-louse /socyamus delphini (Amphipoda: Cyamidae) parasitizing long-finned pilot whales (Globicephala melas) off the Faroe Islands (Northeast Atlantic). Canad. J. Zool. 69:141—-145. Barnard, K. H. 1932. Amphipoda: Cyamidae. Discovery Rept. 5:307—315. Berzin, A. A. and L. P. Vlasova. 1982. Fauna of the Cetacea Cyamidae (Amphipoda) of the world ocean. Investigations on Cetacea 13:149—164. Best, P. B. 1969. The sperm whale (Physeter catodon) off the west coast of South Africa. 3. Repro- duction in the male. Invest. Rept., Div. Sea Fisheries, Republic of South Africa 72:1—20. Bowman, T. E. 1955. A new genus and species of whale-louse (Amphipoda: Cyamidae) from the False Killer Whale. Bull. Mar. Sci. Gulf Carib. 5:315—320. Bowman, T. E. 1958. First Pacific record of the whale-louse genus Syncyamus (Amphipoda: Cyami- dae). Pac. Sci. 12:181—182. Caldwell, D. K. and M. C. Caldwell. 1989, Pygmy sperm whale Kogia breviceps (de Blainville, 1838): dwarf sperm whale Kogia simus Owen, 1866. In: Ridgway, S. H. and Sir. R. Harrison, eds., Handbook of Marine Mammals, volume 4. River dolphins and the larger toothed whales. Pp. 235—260. Academic Press, London. Caldwell, D. K., M. C. Caldwell, and S. G. Zam. 1971. A preliminary report on some ectoparasites and nasal-sac parasites from small odontocete cetaceans from Florida and Georgia.—ZJn: S. G. Zam, D. K. Caldwell, and M. C. Caldwell, eds., Studies on Cetacean Parasites, Technical Report Number 5, Marineland Research Laboratory, St. Augustine, Florida. Fransen, C. H. J. M., and C. Smeenk. 1991. Whale-lice (Amphipoda: Cyamidae) recorded from The Netherlands. Zoologische Mededelingen 65:393—405. Gervais, P., and H. van Beneden. 1859. Zoologie Medicale: expose methodoique du Regne Animal (Paris). [not seen] Gruner, H-E. 1975. Amphipoda II, Caprellidea Il, Cyamidae.—/n: H-E. Gruner and L. B. Holthuis, eds., Crustaceorum Catalogus 5:79—104. Guerin-Meneville, F E. 1836. Iconographie du regne animal de G. Cuvier. Tome II. Plances des animaux invertebres (Paris). [not seen] Heyning, J. E., and W. EF Perrin. 1994. Evidence for two species of common dolphins (genus Del- phinus) from the eastern North Pacific. Contributions in Science, Natural History Museum of Los Angeles County, No. 442:1—35. Hiro, E 1938. Cyamus elogatus n. sp., a new whale lice from Japan. Annotat. Zool. Japon. 17:71—77. (not seen) Hurley, D. E., and J. L. Mohr. 1957. On whale-lice (Amphipoda: Cyamidae) from the California gray whale E. glaucus. J. Parasit. 43(3):352—357. Laubitz, D. R. 1982. Caprellidea. In: S. P. Parker, ed., Synopsis and classification of living organisms, Pp. 292-293. McGraw-Hill. Leung, Y-M. 1965. A collection of whale-lice (Cyamidae: Amphipoda). Bull. South. Calif. Acad. Sci. 64(3):132—-143. Leung, Y-M. 1967. An illustrated key to the species of whale-lice (Amphipoda, Cyamidae), ectopar- asites of Cetacea, with a guide to the literature. Crustaceana 12(5):279-291. Leung, Y-M. 1970a. First record of the whale-louse genus Syncyamus (Cyamidae: Amphipoda) from the western Mediterranean, with notes on the biology of odontocete cyamids. In: G. Pilleri, ed., Investigations on Cetacea 2:243—247. Leung, Y-M. 1970b. Cyamus orcini, a new species of whale-louse [Cyamidae, Amphipoda] from a killer whale. Bulletin de |’ Institue Francais d’ Afrique Noire (A) 32:669—675. Leung, Y-M. 1976. Life cycle of Cyamus scammoni (Amphipoda: Cyamidae), ectoparasite of Gray Whale, with a remark on the associated species. Sci. Rept. Whales Res. Inst. 28:153—160. Lincoln, R. J. and D. E. Hurley. 1974. Scutocyamus parvus, a new genus and species of whale-louse 38 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES (Amphipoda: Cyamidae) ectoparasitic on the North Atlantic white-beaked dolphin. Bull. British Mus. (Nat. Hist.), Zoology 27(2):59—64. Lincoln, R. J. and D. E. Hurley. 1980. Scutocyamus antipodensis n. sp. (Amphipoda: Cyamidae) on Hector’s dolphin (Cephalorhynchus hectori) from New Zealand. New Zealand J. Mar. Freshw. Res. 14(3):295-301. Lincoln, R. J. and D. E. Hurley. 1981. A new species of the whale-louse Syncyamus (Crustacea: Amphipoda: Cyamidae) ectoparasite on dolphins from South Africa. Annals of the Cape Pro- vincial Museum (Nat. Hist.) 13(13):187—-194. Mead, J. G., and R. L. Brownell, Jr. 1993. Order Cetacea. Pp. 349-364 In: D. E. Wilson and D. M. Reeder (eds.), Mammal Species of the World, a taxonomic and geographic reference, second edition. Smithsonian Institution Press, Washington, D.C. Perrin, W. FE, E. D. Mitchell, J. G. Mead, D. K. Caldwell, and P. J. H. Van Bree. 1981. Stenella clymene, a rediscovered tropical dolphin of the Atlantic. J. Mammalogy 62:583—598. Perrin, W. F, E. D. Mitchell, J. G. Mead, D. K. Caldwell, M. C. Caldwell, P. J. H. Van Bree, and W. H. Dawbin. 1987. Revision of the spotted dolphins, Stenella spp. Mar. Mammal Sci. 3:99—170. Raga, J. A. 1988. On some morphological variations of Syncyamus aequus Lincoln & Hurley, 1981 (Amphipoda, Cyamidae) from the Mediterranean Sea. Crustaceana 54:149—152. Raga, J. A., and M. A. Raduan. 1982. First record of Syncyamus aequus Lincoln and Hurley, 1981 (Amphipoda, Cyamidea) in the Mediterranean Sea. Raga, J. A., A. Raduan, and A. Blanco. 1983. Sobre la presencia de /socyamus delphini (Guerin- Meneville, 1836) (Amphipoda: Cyamidae) en aguas del Mediterraneo Espanol. Actas del Con- gresso Iberico de Entomologica 1:627—630. Sedlak-Weinstein, E. 1991. Three new records of cyamids (Amphipoda) from Australian cetaceans. Crustaceana 60(1):90—104. Sedlak-Weinstein, E. 1992a. A new species of /socyamus (Amphipoda: Cyamidae) from Kogia brev- iceps (De Blainville, 1838) in Australian waters. Syst. Parasit. 23:1—6 Sedlak-Weinstein, E. 1992b. The occurrence of a new species of /socyamus (Crustacea, Amphipoda) from Australian and Japanese pilot whales, with a key to species of /socyamus. J. Nat. Hist. 26:937—-946. Stock, J. H. 1977 Whale-lice (Amphipoda, Cyamidae) on Lagenorhynchus albirostris in Dutch waters. Crustaceana 32:206—207. Waller, G. N. H. 1989. Two new species of whale lice (Cyamidae) from the ziphioid whale Berardius bairdii. Investigations on Cetacea 22:292—297. Accepted for publication 13 November 1997. Bull. Southern California Acad. Sci. 98(1), 1999, pp. 39-44 © Southern California Academy of Sciences, 1999 Helminths of the Western Toad, Bufo boreas (Bufonidae) from Southern California Stephen R. Goldberg,' Charles R. Bursey,? and Sonia Hernandez! 'Department of Biology, Whittier College, Whittier, California 90608, U.S.A. *Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, U.S.A. Abstract.—Museum specimens of Bufo boreas from southern California were ex- amined for helminths. The helminth fauna consisted of one trematode species, Haematoloechus kernensis, one cestode species, Distoichometra bufonis, six nem- atode species, Aplectana itzocanensis, Cosmocercoides variabilis, Falcaustra in- glist, Oswaldocruzia pipiens, Rhabdias americanus, Physaloptera sp. (third stage larvae only). Helminths found in B. boreas are generalists occurring in other anuran species. Bufo boreas represents a new host record for H. kernensis, A. itzocanensis, F. inglisi, and Physaloptera sp. (larvae). The western toad, Bufo boreas Baird and Girard 1852, occurs from southern Alaska to northern Baja California, from the Pacific Coast to the Rocky Mountains and from sea level to over 3600 m (Stebbins 1985). There are five previous reports of helminths in B. boreas: California (Ingles 1936; Walton 1941; Koller and Gaudin 1977), Idaho (Waitz 1961), and Utah (Frandsen and Grundmann 1960). The purpose of this study is to present additional helminth records for B. boreas from southern California. Sixty-nine B. boreas from the herpetology collection of the Natural History Museum of Los Angeles County (LACM) were examined: Los Angeles County (N = 30, collected 1958, 1972, mean snout-vent length [SVL] = 73.4 mm + 14.8 SD, range 37-95 mm, LACM 115185-—115208, 144207—144208, 144210— 144213); Orange County (N = 3, collected 1972, SVL = 84.7 mm = 7.2 SD, range 80—93 mm, LACM 144209, 144214—144215); Riverside County (N = 11, collected 1958-1959, 1964, 1966-1967, SVL = 82.0 mm = 10.1 SD, range 64— bOZ smm, LACM. 11218, 11220-11221, 11223, 87313-87316; 87323, 87325, 87327); San Bernadino County (N = 25, collected 1955, SVL = 67.2 mm = 9.3 SD, range 47-81 mm, LACM 11297-11300, 11302, 11305-11307, 11309-11315, 11318, 11343-11344, 11351-11352, 11355-11357, 11359, 11362). Toads had been fixed in 10% formalin and preserved in 70% ethanol. The body cavity was opened by a longitudinal incision from throat to vent and the digestive tract (esophagus, stomach, small and large intestine), lungs and urinary bladder were removed and examined for helminths with a dissecting microscope. The liver and body cavity were also searched for helminths. Helminths were intitially examined using the standard glycerol wet-mount procedure. Nematodes were identified from these wet mounts. Trematodes and cestodes were stained in he- matoxylin, dehydrated in increasing concentrations of ethanol, cleared in xylene and mounted in balsam for identification. Terminology is in accordance with Bush et al. (1997). Sy) 40 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES The helminth fauna of B. boreas examined in this study consisted of one species of Trematoda, Haematoloechus kernensis Ingles 1932; one species of Cestoda, Distoichometra bufonis Dickey 1921; and six species of Nematoda, Aplectana itzocanensis Bravo Hollis 1943, Cosmocercoides variabilis (Harwood 1930), Fal- caustra inglisi (Anderson 1964), Oswaldocruzia pipiens Walton 1929, Rhabdias americanus Baker 1978 and Physaloptera sp (third-stage larvae only). Selected helminths were placed in vials of 70% ethanol and deposited in the U.S. National Parasite Collection (USNPC), Beltsville, Maryland as: Haematoloechus kernensis (88065); Distoichometra bufonis (88066); Aplectana itzocanensis (88067); Cos- mocercoides variabilis (88068); Falcaustra inglisi (88069); Oswaldocruzia pi- piens (88070); Rhabdias americanus (88071); Physaloptera sp., third stage larvae (88072). Prevalence and mean abundance + SD by county are presented in Table 1. Aplectana itzocanensis occurred in the small and large intestines; O. pipiens was found in the stomach and small and large intestines. Other helminths were site specific: D. bufonis, small intestine; H. kernensis and R. americanus, lungs; C. variabilis and F. inglisi, large intestines; Physaloptera sp. stomach. None of the helminths found in this study is unique to B. boreas. These hel- minths are generalists which occur in other anurans. Haematoloechus kernensis is known previously only from its type host, Rana aurora, from California (Ingles 1932). Two of the six North American species of Haematoloechus exhibit short extracaecal uterine loops but can be separated on the shape and location of the testes: H. kernensis with round almost parallel testes; H. varioplexis with elongate oblique testes (Kennedy 1981). Bufo boreas represents a new host record for H. kernensis. The monotypic D. bufonis is known throughout western North America from species of Hyla, Bufo, Scaphiopus and Spea (see Douglas 1958; Goldberg and Bursey 1991a,b; Goldberg et al. 1995). Koller and Gaudin (1977) reported Dis- toichometra sp. from B. boreas collected in Los Angeles County. Thus, this is the second report of this cestode from southern California from B. boreas. Aplectana itzocanensis is known from species of Bufo, Gastrophryne, Scaphio- pus and Spea from Costa Rica, México and the southwestern United States (Gold- berg et al. 1998). The two North American species, A. incerta and A. itzocanensis are separated on the basis of the number of eggs in gravid females: A. incerta with approximately 50 eggs; A. itzocanensis with several hundred eggs (Baker 1985). Bufo boreas represents a new host record for A. itzocanensis. Cosmocercoides variabilis has been reported from North American salaman- ders, frogs, lizards, snakes and turtles (see Baker 1987). Some uncertainty exists for its hosts because of confusion between C. variabilis and Cosmocercoides dukae, a molluscan parasite. Vanderburgh and Anderson (1987) demonstrated that the two species are distinct. The major difference in morphology for the two species is the number of rosette papillae in the male: C. dukae with 12 pairs, C. variabilis with 14 to 20 pairs. Ingles (1936) reported C. dukae from Taricha torosa, Rana aurora and B. boreas from California but illustrated 16 pairs of papillae and for this reason we refer his specimens to C. variabilis. We also refer the Cosmocercoides sp. of Koller and Gaudin (1977) to C. variabilis. Thus, this is the third report of C. variabilis from B. boreas. Falcaustra inglisi has been reported from Rana catesbeiana, Rana clamitans 41 ‘p1OSOI SOY MOU = ‘PoUurWIeXS SpeO} JO Joquinu ][eI0} sy Aq poprArp satoads oytsesed v JO syenprlArpur Jo Joquinu [RIO], - ‘pourulexe Speo} JO Jaquinu oy} Aq paprArp satoads oyIsesed & JO S[eNPIAIPUT dIOU JO DUO YIM poJdoFUT speo} Jo JoquINYY ; SS —— a Se a se SO ee eee eee EGO) = GEO Gl = = = = EO» SE 100) € IG 6) se Gey 8c = — = =a — a Se ee ee ee ee ee ae ed ds (%) ds= (%) (dSF) (%) Gig = (%) souepunge JOUD[RADI gouRrpunqe JOUD[RBAIIg souerpunqge JOUD[BADI | zoouepunge ;VOUD|[BADId ues ues] uevofJ uel] (Gc = N) Cae N) (c= ND (OG —SN)) Ajuno, OUIpIeUIog ues AJUNOD ApIsIOARY AWUNOD ssurIO AyunoZ sojasuy soy HELMINTHS OF THE WESTERN TOAD “RIUIOJI[E) UJYINOS WO, svas0g Ofng 69 WOIZ SYIUTUTAFY “[ 2]qQey, —6—$“~—S=_€—_€_86—0—0$0$”9090©”0. SSoaoaoaoaoaoaoamamamamamnmnmwsxoayeoqyoeqoejéaé*sos=~=~=~— oS 42 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 2. Helminth parasites of Bufo boreas. Prevalence Helminth Locality (%) Reference Trematoda Glypthelmins shastai California not given Ingles 1936 California 2/115 (2) Koller and Gaudin 1977 Gorgoderina sp. California not given Ingles 1936 Gorgoderina translucida Idaho 1/8 (13) Waitz 1961 Haematoloechus kernensis California 7/69 (10) this study Megalodiscus microphagus California not given Ingles 1936 Phyllodistomum bufonis Utah not given Frandsen and Grundmann 1960 Cestoda Distoichometra bufonis California 19/140 (14) Koller and Gaudin 1977 California 4/69 (6) this study Nematoda Aplectana itzocanensis California 7/69 (10) this study Cosmocercoides variabilis California not given Ingles 1936 California 40/255 (16) Koller and Gaudin 1977 California 2/69 (3) this study Utah not given Frandsen and Grundmann 1960 Falcaustra inglisi California 1/69 (1) this study Falcustra pretiosa California not given Walton 1941 Megalobatrachonema gigantica Utah not given Frandsen and Grundmann 1960 Oswaldocruzia pipiens California not given Ingles 1936 California 119/255 (47) Koller and Gaudin 1977 California 11/69 (16) this study Rhabdias americanus California not given Ingles 1936 California 13/255 (5) Koller and Gaudin 1977 California 8/69 (12) this study Physaloptera sp. (larvae) California 1/69 (1) this study and Rana septentrionalis from Ontario, Canada (Baker 1987). The nine North American species of Falcaustra (see Baker 1987) are separated by size of spicule and position of caudal papillae. Falcaustra inglisi is the only species having 2 pairs of large and 2 pairs of small precloacal papillae. Bufo boreas represents a new host record and California a new location record for F. inglisi. All North American specimens of Oswaldocruzia have been referred to O. pipiens by Baker (1977). This species is widely distributed in North America and has been reported from frogs, toads, salamanders, lizards and tortoises (see Baker 1987). Oswaldocruzia sp. was reported from B. boreas in California by Koller and Gaudin (1977). This is the second report of O. pipiens from B. boreas. Rhabdias americanus was first described from Bufo americanus collected in Canada (Baker 1978) and has been reported from species of Bufo from the south- western United States (see Goldberg et al. 1996). In addition, Baker (1978) re- ferred reports of Rhabdias bufonis in Bufo woodhousii and Bufo americanus of eastern North America to R. americanus. We refer the reports of Rhabdias sp. in B. boreas from California (Ingles 1936; Koller and Gaudin 1977) to R. ameri- canus because Ingles (1936) in his survey of amphibian parasites reported this Rhabdias to be a species other than Rhabdias joaquinensis or Rhabdias ranae; HELMINTHS OF THE WESTERN TOAD 43 R. americanus was not described until 1978 (Baker 1978). This is the third report of R. americanus from B. boreas. Third stage larvae of Physaloptera sp. are common in toads, but no parasitism by adult physalopterans in toads has been reported (Goldberg et al. 1995). Bufo boreas represents a new host record for larvae of Physaloptera sp. Reasons for the patchy distribution of helminths found in this study (Table 1) are unknown. No helminth species occurred in all locations; yet, given general distribution ranges, each helminth species might be expected to be present. Koller and Gaudin (1977) also report similar differences in helminth species distribution for two populations of B. boreas from southern California. Likewise, other hel- minths have been reported from populations of B. boreas in California (Table 2), but were not found in this study. Of the parasites listed in Table 2, the trematodes require an aquatic intermediate host (Smyth and Smyth 1980); the life cycle of the cestode D. bufonis is unknown (Hardin and Janovy 1988). With the exception of Physaloptera which has an indirect life cycle requiring an insect intermediate host, the nematodes are ac- quired orally or by skin penetration (direct life cycles) and have moisture sensitive larval stages (Anderson 1992). In either case, indirect or direct life cycle, envi- ronmental conditions have a deciding role in patchy distribution patterns by caus- ing local changes in the distribution of intermediate hosts or the hydrologic cycle while not affecting the historical pattern of host distribution. More work will be required to determine the factors responsible for the patchy distribution patterns of parasites within this host; specifically, annual prevalence studies with environ- mental monitoring for several populations of B. boreas should be initiated. We thank Robert L. Bezy (Natural History Museum of Los Angeles County) for permission to examine B. boreas for helminths. Literature Cited Anderson, R. C. 1992. Nematode parasites of vertebrates. Their development and transmission. C.A.B. International, Wallingford, Oxon, U.K. x11 + 578 pp. Baker, M. R. 1977. Redescription of Oswaldocruzia pipiens Walton, 1929 (Nematoda: Trichostron- gylidae) from amphibians of eastern North America. Can. J. Zool., 55:104—109. . 1978. Morphology and taxonomy of Rhabdias spp. (Nematoda: Rhabdiasidae) from reptiles and amphibians of southern Ontario. Can. J. Zool., 56:2127-2141. . 1985. Redescription of Aplectana itzocanensis and A. incerta (Nematoda: Cosmocercidae) from Amphibians. Trans. Amer. Microsc. Soc., 104:272—277. . 1987. Synopsis of the Nematoda parasitic in amphibians and reptiles. Mem. Univ. Newfound- land, Occas. Pap. Biol., 11:1—325. 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. Douglas, L. T. 1958. The taxonomy of nematotaeniid cestodes. J. Parasitol., 44:261—273. Frandsen, J. C., and A. W. Grundmann. 1960. The parasites of some amphibians of Utah. J. Parasit., 46:678. Goldberg, S. R., and C. R. Bursey. 1991la. Helminths of three toads, Bufo alvarius, Bufo cognatus (Bufonidae), and Scaphiopus couchii (Pelobatidae), from southern Arizona. J. Helm. Soc. Wash., 58:142—146. , and C. R. Bursey. 1991b. Helminths of the red-spotted toad, Bufo punctatus (Anura: Bufon- idae), from southern Arizona. J. Helm. Soc. Wash., 58:267—269. , C. R. Bursey, and H. Cheam. 1998. Nematodes of the Great Plains narrow-mouthed toad, Gastrophryne olivacea (Microhylidae), from southern Arizona. J. Helm. Soc. Wash., 65:102— 104. 44 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES , C. R. Bursey, K. B. Malmos, B. K. Sullivan, and H. Cheam. 1996. Helminths of the south- western toad, Bufo microscaphus, Woodhouse’s toad, Bufo woodhousii (Bufonidae), and their hybrids from central Arizona. Great Basin Nat., 56:369-—374. , C. R. Bursey, and I. Ramos. 1995. The component parasite community of three sympatric toad species, Bufo cognatus, Bufo debilis, (Bufonidae), and Spea multiplicata (Pelobatidae) from New Mexico. J. Helm. Soc. Wash., 62:57—61. Hardin, E. L., and J. Janovy, Jr. 1988. Population dynamics of Distoichometra bufonis (Cestoda: Nematotaentidae) in Bufo woodhousii. J. Parasitol., 74:360—365. Ingles, L. G. 1932. Four new species of Haematoloechus (Trematoda) from Rana aurora draytoni from California. Univ. Cal. Publ. Zool., 37:189—202. . 1936. Worm parasites of California amphibia. Trans. Am. Micro. Soc., 55:73—92. Kennedy, M. J. 1981. A revision of species of the genus Haematoloechus Looss, 1899 (Trematoda: Haematoloechidae) from Canada and the United States. Can. J. Zool., 59:1836—1846. Koller, R. L., and A. J. Gaudin. 1977. An analysis of helminth infections in Bufo boreas (Amphibia: Bufonidae) and Hyla regilla (Amphibia: Hylidae) in southern California. Southwest. Nat., 21: 503-509. Smyth, J. D., and M. M. Smyth. 1980. Frogs as host-parasite systems I. The Macmillan Press Ltd., London, ix + 112 pp. Stebbins, R. C. 1985. A field guide to western reptiles and amphibians. Houghton Mifflin Company, Boston, xiv + 336 pp. Vanderburgh, D. J., and R. C. Anderson. 1987. Seasonal changes in prevalence and intensity of Cosmocercoides dukae (Nematoda: Cosmocercoidea) in Deroceras laeve (Mollusca). Can. J. Zool., 65:1662—1665. Waitz, J. A. 1961. Parasites of Idaho amphibians. J. Parasitol., 47:89. Walton, A. C. 1941. Amphibian nematodes from the Gaspé Peninsula and vicinity. J. Parasitol., 27: 59-61. Accepted for publication 29 September 1998. INSTRUCTIONS FOR AUTHORS | a “The BULLETIN is published three times each year (April, August, and December) and includes articles in English in any field of science with an emphasis on the southern California area. Manuscripts submitted for publication ould 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 Bielogical Journals is recommended as a guide for contributors. Consult also recent issues of the BULLETIN. 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Héeyning. 2 ESTE Helminths of ‘the Westeiii Toad, Bufo boreas (Bufonidae) from Southern California. Stephen R. Goldberg, Charles R. Bursey, and Sonia Hernandez | oe COVER: Quercus sp. (Oak) Radial section shows large vessels of axial system. From Conejo Volcanics Tuft (13 million years old). RMW Paleo Associates. 15 26 OR Sele NiGEss aOAD TE Mey See ERN CALIFORNIA LLETIN BU Number 2 Volume 98 AUGUST 1999 BCAS-A98(2) 45-90 (1999) Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 © Southern California Academy of Sciences, 1999 OFFICERS David Huckaby, President Robert S. Grove, Vice-President Susan E. Yoder, Secretary Robert Lavenberg, Treasurer Daniel A. Guthrie, Editor Hans Bozler, Past President David Soltz, Past President BOARD OF DIRECTORS 1997-2000 1998-2001 1999-2002 Robert S. Grove Kathryn A. Dickson Ralph G. Appy David Huckaby Donn Gorsline Jonathan N. Baskin Robert Lavenberg Robert F Phalen John W. Rcberts Kenneth E. Phillips Cheryl C. 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All other communications should be addressed to the Southern California Academy of Sciences in care of the Natural History Museum of Los Angeles County, Exposition Park, Los Angeles, California 90007-4000. Date of this issue 2 August 1999 © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Bull. Southern California Acad. Sci. 98(2), 1999, pp. 45-56 © Southern California Academy of Sciences, 1999 | CALIFORNIA ; 534 ; | ACADEMY OF SCIENCES Checklist of Amphibians and Reptiles on Islands in the Gulf of California, México AUG 2 4 1999 L. Lee Grismer | IBRARY Department of Biology, La Sierra University, Riverside, California 92515-8247 Abstract.—A cross-referenced checklist of the herpetofauna on islands in the Gulf of California is provided which contains 58 additional records and 28 additional islands not reported in the most recent checklist. The taxonomy of the herpeto- fauna is based on an evolutionary species concept. Resumen.—Una lista de la herpetofauna de las islas del Golfo de California, México esta presentada y contiene 58 registros adicional y 28 islas adicional que no habian reportados en la lista mas reciente. La taxonomia de la herpetofauna esta basada en un concepto de especie evolutiva. The islands in the Gulf of California have been heralded to be among North America’s most distinctive natural laboratories for broad-based evolutionary stud- les (e.g., Case and Cody 1983). As such, these islands have received considerable attention from evolutionary biologists of widely varying disciplines, most notably herpetologists (e.g. Case 1975, 1983; Cliff 1954a,b; Dixon 1966; Grismer 1994a,b,c, 1999a,b; Hews 1990a,b; Hollingsworth 1998; Murphy 1983a,b; Petren and Case 1997; Radtkey et al., 1997; Robinson 1972, 1974; Savage 1960; Soulé 1964, 1966; Upton and Murphy 1997; Wilcox 1978). Necessary prerequisites for many evolutionary studies concerned with insular lineages or ecosystems is an understanding of which species are present on the island(s) of concern as well as an appropriate taxonomy that is consistent with their evolutionary history. Gris- mer (1999b) has revised the taxonomy of the insular herpetofauna of the Gulf of California using the criterion of the evolutionary species concept (Wiley 1978; Frost and Hillis 1990) in an attempt to make it consistent with each taxon’s evolutionary history. This paper provides a working, up-to-date checklist of the herpetofauna of the all the islands in the Gulf of California based on the taxonomy of Grismer (1999b). Since the turn of the century, at least four herpetological checklists concerning the gulf island herpetofauna have been published (Schmidt 1922; Cliff 1954b; Soulé and Sloan 1966; Murphy and Ottley 1984). Although the most recent check- list by Murphy and Ottley (1984) has been extremely useful and is considerably more comprehensive than its predecessors, it is problematic in that it includes numerous taxonomic changes lacking supportive data (see Flores-Villela 1993; Grismer, 1999b), it contains misidentified islands resulting in erroneous records, and it omits many published insular accounts and museum records that were in existence prior to its publication. In this paper, there are 58 additional insular records which were not reported in Murphy and Ottley (1984). These include records reported here for the first ttme (Appendix I), records from accounts pub- lished subsequent to Murphy and Ottley (1984) (.e., Cryder et al. 1997; Felger 45 46 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES . El Muerto . Coloradito . Encantada . Blancos . San Luis . Willard . Mejia . Granito . Angel de La Guarda . Pond . Cardonosa Este and Partida Norte . La Rasa . Salsipuedes . San Lorenzo Norte . San Lorenzo Sur . Tiburén . Cholludo . Alcatraz . Danzante . San Esteban . Monserrate . San Pedro Martir . Santa Catalina . Datil . Santa Cruz . San Pedro Nolasco . San Diego . Tortuga . Las Animas . San Marcos . San José . Santa Inez . San Francisco . San Ildefonso . Partida Sur . Coronados . Espiritu Santo . Carmen . Cerralvo 1h 2 3 ~ 5 6 7 8 9 Fig. |. Location of islands in the Gulf of California, México. and Moser, 1985; Grismer 1989a,b, 1996a,b; Grismer et al. 1996; Ramirez 1989; Reynoso 1989; Hollingsworth and Grismer 1996; Hollingsworth et al. 1996, 1997; Lava-G. et al. 1993; Mahrdt and Grismer 1995; Wong, 1997; Wong et al. 1995, 1996), and previously existing records (Lowe 1955; Soulé and Sloan 1966) over- looked by Murphy and Ottley (1984; Appendix I). Twenty-eight additional gulf islands are included that were not listed in Murphy and Ottley (1984). Figure 1 illustrates the location of the major islands. Descriptive locations of islands not AMPHIBIANS AND REPTILES ON ISLANDS IN THE GULF OF CALIFORNIA 47 figured are provided in Appendix II. Table 1 and Appendix II were compiled from new records from Appendix I, the checklist of Murphy and Ottley (1984), and from literature records subsequent to Murphy and Ottley (1984). Erroneous or Questionable Insular Records Malkin (1962) reported Bufo alvarius from Isla Tibur6n which was followed by Foquette (1970). However, R. Crombie (pers. comm. 1996) informed me that surveys for amphibians on Isla Tiburé6n revealed only B. punctatus and that there were no documented records of B. alvarius from Isla Tiburon. It is presumed here that Malkin (1962) mistook B. punctatus for B. alvarius. Therefore, this record is not included in the checklist. Duellman (1970) reported a specimen of Ayla regilla from Isla Coronados off the east coast of Loreto in the Gulf of California. Isla Coronados is a landbridge, volcanic island that has never been known to have any source of fresh water. Therefore, although a specimen does exist with locality data, the locality is con- sidered here to be dubious. Felger (1966) reported Gopherus agassizzii as occurring on Isla Datil. However, this sighting has yet to be confirmed with a specimen or voucher photograph. Therefore, its occurrence on Isla Datil remains in question. Murphy and Ottley (1984) provided an undocumented report of Uta stansbu- riana from Isla Cholludo. A search of the holdings of five museums (CAS, LACM, MVZ, SDSNH, USNM; museum acronyms follow Leviton et al. 1985) revealed no specimens from Isla Cholludo. Additionally, no U. stansburiana were observed on two surveys (6 October 1995 and 25 March 1997) of Isla Cholludo totaling 18 person-hours. Therefore, U. stansburiana is presumed to be absent from Isla Cholludo. Crotalus atrox was first reported from Isla Santa Cruz by Soulé and Sloan (1966). Since that time there has been no indication that it was anything other than C. atrox. Stewart et al. (1990), without comment, considered this population to be C. ruber. Specimens I have observed in the field (LACM PC 1303) and in museums are all C. atrox. Therefore, the report of Stewart et al. (1990) is con- sidered here to be an oversight. Grismer et al. (1994) placed Crotalus ruber in the synonymy of C. exsul. A petition was submitted to the International Commission on Zoological Nomen- clature to give precedence to C. ruber over C. exsul (Smith et al. 1998). According to Article 80 of the International Code of Zoological Nomenclature, the most commonly used classification should be employed until a decision has been made by the commission. Because the earlier classification has a wider usage, it is adopted here. Acknowledgments For comments on the manuscript I express my thanks to E. Gergus, R. Ether- idge, B. Hollingsworth, J. McGuire, and H. Wong. Field work was conducted under Mexican research permits issued to L. Grismer in 1990 (permit 03445), E. Mellink in 1993 and 1994 (permits 01303 end 01678, respectively), and E. Gergus in 1995 (permits 00944) by Instituto Nacional de Ecologia, Dirreci6n General de Aprovechamiento, Ecol6gico de los Recursos Naturales, Direcci6n de Flora y Fauna Silvestres in Méxcio City and La Paz. I thank the curators and collection 48 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES managers of the following institutions; C. Myers (AMNH), J. W. Sites, Jr. and W. W. Tanner (BYU), J. Gauthier and J. Vindum (CAS), H. Marx (FMNBH), R. L. Bezy and J. W. Wright (LACM), P. Alberch and J. Rosado (MCZ), D. B. Wake (MVZ), R. W. Murphy (ROM), D. C. Cannatella (TNHC), G. Pregill (SDSNH), C. H. Lowe and G. Bradly (UA), EF Reynoso (UABCS), and G. Zug (USNM) for allowing me access to specimens. Literature Cited Case, T. J. 1975. Species numbers, density compensation, and colonizing ability of lizards on islands in the Gulf of California. Ecology 56:3-18. . 1983. The Reptiles: Ecology. Pp. 159-209. In T. J. Case and M. L. Cody (Eds.), Island Biogeography in the Sea of Cortéz. University of California Press, Los Angeles, California, U.S.A. Case, T. J., and M. L. Cody. 1983 (eds.). Island Biogeography in the Sea of Cortéz. University of California Press, Los Angeles, California. Cliff, E S. 1954a. Snakes of the islands in the Gulf of California, Mexico. Trans. San Diego Soc. Nat. Hist. 12:67—98. . 1954b. Variation and evolution of the reptiles inhabiting the islands in the Gulf of California, Mexico. Ph.D. Dissertation, Stanford University, Stanford, California, U.S.A. Dixon, J. R. 1966. Speciation and systematics of the gekkonid lizard genus Phyllodactylus of the islands in the Gulf of California. Proc. Calif. Acad. Sci., 4th series 33:415—452. Cryder, M. R., H. Wong, L. L. Grismer, and B. D. 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The Evolutionary and Ecological Biogeography of the Herpetofauna of Baja California and the Sea of Cortés, México. Ph.D. Dissertation, Loma Linda University, Loma Linda, Cal- ifornia, U.S.A. . 1994b. Geographic origins for the reptiles on islands in the Gulf of California, México. Herpetol. Nat. Hist. 2:17—40. . 1994c. The origin and evolution of the peninsular herpetofauna of Baja California, México. Herpetol. Nat. Hist. 2:51—106. . 1996a. Phyllodactylus xanti. Geographic distribution. Herpetol. Rev. 27:32—33. . 1996b. Lampropeltis getula nigrita. Geographic distribution. Herpetol. Rev. 27:34. . 1999a. Phylogeny, taxonomy, and biogeography of the Cnemidophorus hyperythrus and C. ceralbensis (Squamata: Teiidae) in Baja California, México. Herpetologica 55:28—42. . 1999b. An evolutionary classification of the reptiles on islands in the Gulf of California, México. Herpetologica 55: Grismer, L. L., B. D. Hollingsworth, M. R. Cryder, and H. Wong. 1996. Crotalus enyo. 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Evol. 8:104—113. 50 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Wilcox, B. A. 1978. Supersaturated island faunas: a species-age relationship for lizard faunas on post- Pleistocene land-bridge islands. Science 199:996—998. Wiley, E. O. 1978. The evolutionary species concept reconsidered. Syst. Zool. 29:76—-80. Wong, H. 1997. Chilomeniscus cinctus. Geographic Distribution. Herpetol. Rev. 28:51. Wong, H., E. Mellink, and B. D. Hollingsworth. 1995. Proposed recent overwater dispersal by Calli- saurus draconoides to Isla Danzante, Gulf of California, México. Herptol. Nat. Hist. 3:179—-182. Wong, H., L. L. Grismer, B. D. Hollingsworth and M. R. Cryder. 1996. Urosaurus nigricaudus. Geographic Distribution. Herpetol. Rev. 28:51. Accepted for publication 27 August 1998. Table 1. Checklist of the herpetofauna on islands in the Gulf of California, México. Asterisked taxa are insular endemics. Taxon Insular locale ANURA Bufonidae Bufo punctatus Cerralvo, Espiritu Santo, Partida Sur, Tibur6n Pelobatidae Scaphiopus couchii Cerralvo, Espiritu Santo, Partida Sur, Tibur6n CHELONIA Testudinidae Gopherus agassizii Datil (?), Tibur6n SQUAMATA (Lizards) Crotaphytidae Crotaphytus dickersonae Tibur6n Crotaphytus insularis* Angel de La Guarda Gambelia wislizenii Tibur6n Iguanidae Ctenosaura conspicuosa* Cholludo, San Esteban Ctenosaura hemilopha Cerralvo Ctenosaura nolascensis* San Pedro Nolasco Dipsosaurus dorsalis Angel de La Guarda, Carmen, Cerralvo, Coronados, Es- piritu Santo, Monserrate, Partida Sur, Santiago, San José, San Luis, San Marcos Dipsosaurus catalinensis* Santa Catalina Sauromalus ater Ballena, Danzante, El Coyote, Espiritu Santo, Gallo, Pardo, Partida Sur, San Cosme, San Diego, San Fran- cisco, San José, San Marcos, Santa Cruz, Tiburon, Willard Sauromalus hispidus* Angel de La Guarda, Cabeza de Caballo, Flecha, Grani- to, La Ventana, Mejia, Mitlan, Piojo, Pond, San Lor- enzo Norte, San Lorenzo Sur, Smith Sauromalus klauberi* Santa Catalina Sauromalus slevini* Carmen, Coronados, Monserrate Sauromalus varius* Roca Lobos, San Esteban Sauromalus hispidus X ater X varius Alcatraz Phrynosomatidae Callisaurus draconoides Angel de La Guarda, Carmen, Cerralvo, Coronados, Danzante, Espirirtu Santo, Partida Sur, Patos, San Francisco, San‘ José, San Luis, San Marcos, Santa Inez, Smith, Tibur6n AMPHIBIANS AND REPTILES ON ISLANDS IN THE GULF OF CALIFORNIA 5] Taxon Table 1. Continued. Insular locale Petrosaurus mearnsi Petrosaurus repens Petrosaurus slevini* Petrosaurus thalassinus Phrynosoma solare Sceloporus angustus* Sceloporus clarkii Sceloporus grandaevus* Sceloporus hunsakeri Sceloporus lineatulus* Sceloporus magister Sceloporus orcutti Sceloporus zosteromus Urosaurus nigricaudus Urosaurus ornatus Uta encantadae* Uta lowei* Uta nolascensis* Uta palmeri* Uta squamata* Uta stansburiana Uta tumidarostra* Eublepharidae Coleonyx gypsicolus* Coleonyx variegatus Gekkonidae Phyllodactylus bugastrolepis* Phyllodactylus partitus* Phyllodactylus homolepidurus Phyllodactylus unctus Phyllodactylus xanti El Muerto Danzante Angel de La Guarda, Mejia Espiritu Santo, Partida Sur Tiburon San Diego, Santa Cruz San Pedro Nolasco, Tiburon Cerralvo Ballena, Espiritu Santo, Gallo, Partida Sur Santa Catalina Tibur6n Carmen, Coronados, San Francisco, San Ildefonso, San José, San Marcos, Tortuga Carmen, Coronados, Espiritu Santo, Monserrate, Partida Sur, San José Ballena, Carmen, Cayo, Coronados, Danzante, El Coy- ote, E] Requeson, Espiritu Santo, Gallina, Gallo, Gav- iota, Islitas, Las Animas, Pardo, Partida Sur, San Cos- me, San Damian, San Francisco, San José, San Marcos, Tijeras Tiburon Encantada, Islotes Blancos El Muerto San Pedro Nolasco San Pedro Martir Santa Catalina Alcatraz, Angel de La Guarda, Ballena, Bota, Cabeza de Caballo, Cardonosa Este, Carmen, Cerraja, Coronados, Danzante, Datil, El Pardito, Espiritu Santo, Flecha, Gallo, Granito, Lagartija, La Raza, La Ventana, Las Galeras, Mitlan, Mejia, Monserrate, Partida Norte, Partida Sur, Patos, Piojo, Pond, Roca Lobos, Salsi- puedes, San Esteban, San Francisco, San Ildefonso, San José, San Lorenzo Norte, San Lorenzo Sur, San Luis, San Marcos, Smith, Tiburé6n, Tortuga, Willard Coloradito San Marcos Angel de La Guarda, Coronados, Danzante, Espiritu Santo, Partida Sur, San José, San Marcos, Santa Inez, Tibur6n Santa Catalina Cardonosa Este, Partida Norte San Pedro Nolasco Ballena, Cerralvo, Espiritu Santo, Gallina, Gallo, Partida Sur Alcatraz, Angel de La Guarda, Carmen, Cayo, Cholludo, Coronados, Danzante, Datil, El Coyote, El Muerto, El Pardito, La Raza, La Ventana, Las Animas, Mejia, Monserrate, Mosca, Pardo, Piojo, Pond, Salsipuedes, San Diego, San Esteban, San Francisco, San Ildefon- so, San José, San Lorenzo Norte, San Lorenzo Sur, San Marcos, Santa Cruz, Smith, Tiburon Taxon SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1. Continued. Insular locale Teiidae Cnemidophorus bacatus* Cnemidophorus canus* Cnemidophorus carmenensis* Cnemidophorus catalinensis* Cnemidophorus celeripes* Cnemidophorus ceralbensis* Cnemidophorus danheimae* Cnemidophorus espiritensis* Cnemidophorus franciscensis* Cnemidophorus hyperythrus Cnemidophorus martyris* Cnemidophorus pictus* Cnemidophorus tigris © SQUAMATA (Snakes) Leptotyphlopidae Leptotyphlops humilis Boiidae Lichanura trivirgata Colubridae Bogertophis rosaliae Chilomeniscus cinctus Chilomeniscus punctatissimus* Chilomeniscus savagei* Chilomeniscus stramineus Eridiphas marcosensis* Eridiphas slevini Hypsiglena gularis* Hypsiglena torquata Lampropeltis catalinensis* Lampropeltis getula Masticophis barbouri* Masticophis bilineatus Masticophis flagellum Masticophis slevini* Phyllorhynchus decurtatus Pituophis melanoleucus Pituophis vertebralis Rhinocheilus etheridgei* San Pedro Nolasco Salsipuedes, San Lorenzo Norte, San Lorenzo Sur Carmen Santa Catalina San Francisco, San José Cerralvo San José Espiritu Santo, Partida Sur San Francisco Coronados, San Marcos San Pedro Martir Monserrate Angel de La Guarda, Cardonosa Este, Carmen, Corona- dos, Danzante, Espiritu Santo, Partida Norte, Partida Sur, Pond, San Cosme, San Esteban, San Marcos, Smith, Tibur6n Carmen, Cerralvo, Danzante, San Marcos, Santa Catalina Angel de La Guarda, Cerralvo Mejia, San Marcos, Tibu- ron Danzante Danzante, Monserrate, San José, San Marcos, Tiburén Espiritu Santo, Partida Sur Cerralvo Tibur6én San Marcos Cerralvo, Coronados, Danzante Partida Norte Angel de la Guarda, Carmen, Cerralvo, Coronados, Dan- zante, Mejia, Monserrate, Partida Sur, San Esteban, San Francisco, San José, San Lorenzo Sur, San Mar- cos, Santa Catalina, Smith, Tiburon, Tortuga Santa Catalina Angel de La Guarda, Cerralvo, Monserrate, Salsipuedes, San Esteban, San Lorenzo Norte, San Lorenzo Sur, San Pedro Martir, San Pedro Nolasco, Tortuga Espiritu Santo, Partida Sur Tibur6n Carmen, Cerralvo, Coronados, Danzante, Datil, Espiritu Santo, Monserrate, Partida Sur, San Ildefonso, San José, San Marcos, Tiburon San Esteban Angel de La Guarda, Cerralvo, Monserrate, San José, San Marcos Tibur6n San José Cerralvo AMPHIBIANS AND REPTILES ON ISLANDS IN THE GULF OF CALIFORNIA Taxon Salvadora hexalepis Sonora semiannulata Tantilla planiceps Trimorphodon biscutatus Nn eS) Table 1. Continued. Insular locale Espiritu Santo, San José, Tibur6n San José, San Marcos Carmen Cerralvo, Danzante, El Muerto, San José, San Marcos, Tibur6n Elapidae Micruroides euryxanthus Tiburon Viperidae Crotalus angelensis* Angel de La Guarda Crotalus atrox Datil, San Pedro Martir, Santa Cruz, Tibur6n Crotalus catalinensis* Santa Catalina Crotalus cerastes Tiburon Crotalus enyo Carmen, Cerralvo, Coronados, Espiritu Santo, Pardo, Partida Sur, San Francisco, San José, San Marcos Crotalus estebanensis* San Esteban Crotalus lorenzoensis* San Lorenzo Sur Crotalus mitchellii Carmen, Cerralvo, Espiritu Santo, Monserrate, Partida Sur, Piojo, Salsipuedes, San José, Smith Crotalus muertensis* El Muerto Crotalus molossus Tibur6n Crotalus ruber Angel de La Guarda, Danzante, Monserrate, Pond, San José, San Marcos Crotalus tigris Tibur6én Crotalus tortugensis* Tortuga Appendix I New insular records of amphibians and reptiles on islands in the Gulf of California, México are listed below. The records are followed by voucher photograph numbers (LACM PC) or museum catalogue numbers and, when possible, the date the specimens were observed or collected. Literature records Or museum specimens present prior to 1984 not reported by Murphy and Ottley (1984) are followed by a literature citation or museum catalogue number, respectively. Museum acronyms follow Leviton et al. (1985). ANURA Pelobatidae. Scaphiopus couchii.—Isla Tibur6n, Sonora; Felger and Moser (1985). CHELONIA Testudinidae. Gopherus agassizii.—Isla Datil, Sonora; Felger (1966); questionable. SQUAMATA (Lizards) Iguanidae. Dipsosaurus dorsalis.—Isla Santiago (1 km north of Loreto). Sauromalus ater.—Isla Gallo, BCS, LACM PC 1275, 31 August 1987; Isla San Cosme, BCS, LACM PC 1276, 4 April 1993; Isla El Coyote (in Bahia Concepcion), BCS, LACM PC 1277, 5 June 1987; Isla Willard, BC, fecal material, 18 December 1986. Sauromalus hispidus.—Islas Flecha and Mitlan, BC (scat recorded by G. A. Polis; pers. comm. 30 June 1991). Phrynosomatidae. Callisaurus draconoides.—Isla Patos, Sonora, SDSNH 46427—28. Sceloporus hunsakeri.—Isla Gallo, BCS, LACM PC 1278, 31 August 1987. Urosaurus nigricaudus.—Isla Cayo, BCS, LACM PC 1279, 23 June 1992; Isla El Coyote, BCS, LACM PC 1280, 5 June 1987; Isla Gallo, BCS, LACM PC 1281, 31 August 1987; Isla Reques6n, BCS, LACM PC 1282, 10 August 1988; Islas San Cosme and San Damian, BCS, LACM PC 1283 and 1284, respectively, 4 April 1993. Uta stansburiana.—Isla Bota, LACM PC 1285, 30 June 1991; Isla Cabeza de Caballo, BC, LACM PC 1286, 26 April 1989; Isla Cardonosa Este, LACM PC 1287; Isla Cerraja, BC, LACM PC 1288, 30 54 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES June 1991; Isla Datil, Sonora, Felger (1966); Isla Flecha, BC, LACM PC 1290, 30 June 1991; Isla Lagartija, BC, LACM PC 1291, 7 October 1994; Isla Mitlan, BC, G. Polis (pers. comm., 5 June 1992); Isla Pata, BC, LACM PC 1292, 30 June 1991; Isla Roca Lobos, BC, LACM PC 1293, 7 October 1994; Isla Willard, BC, LACM PC 1294, 18 December 1986. Gekkonidae. Phyllodactylus partitus.—Isla Cardonosa Este, BC, LACM PC 1295, 7 October 1994. Phyllodactylus unctus.—Isla Gallo, BCS, LACM PC 1296, 31 August 1987. Phyllodactylus xanti.— Isla Cholludo, Sonora, Felger (1966); Isla El Coyote, BCS, Dixon (1966); Isla Moscas (in Bahia Concepciéon) BCS, Dixon (1966); Isla Smith, BC, LACM 131406-12. Teiidae. Cnemidophorus tigris.—Isla Cardonosa Este, BC; LACM PC 1297, 7 October 1994; Isla San Cosme, BCS, LACM PC 1298, 8 April 1993. SQUAMATA (Snakes) Colubridae. Chilomeniscus stramenius.—Isla Tibur6n, Sonora, Felger (1966). Masticophis flagel- lum.—Isla Datil, Sonora, Felger (1966). Salvadora hexalepis.—Isla Espiritu Santo, BCS, CAS 146563—64. Isla El Muerto, BC, LSUHC 4000, 26 March, 1998, shed skin. Viperidae. Crotalus atrox.—Isla Datil, Sonora, Lowe (1955). Crotalus mitchellii.—Isla Cabeza de Caballo, BC, G. A. Polis (pers. comm., 1993). Appendix II Checklists of the herpetofauna of the islands in the Gulf of California, México. Less commonly used island names are listed in parentheses. Order of species follows Table 1. Asterisked taxa are insular endemics. Isla Alcatraz. (Isla Pelicano). Sauromalus hispidus * ater X varius, Uta stansburiana, Phyllodac- tylus xanti. Isla Angel de La Guarda. Crotaphytus insularis*, Dipsosaurus dorsalis, Sauromalus hispidus*, Cal- lisaurus draconoides, Petrosaurus slevini*, Uta stansburiana, Coleonyx variegatus, Phyllodactylus xanti, Cnemidophorus tigris, Lichanura trivirgata, Hypsiglena torquata, Lampropeltis getula, Phyl- lorhynchus decurtatus, Crotalus angelensis*, Crotalus ruber. Isla Ballena. (0.5 km W of Isla Espiritu Santo). Sauromalus ater, Sceloporus hunsakeri, Urosaurus nigricaudus, Uta stansburiana, Phyllodactylus unctus. Isla Bota. (within Bahia de Los Angeles, BC). Uta stansburiana. Isla Cabeza de Caballo. (within Bahia de Los Angeles, BC). Sauromalus hispidus*, Uta stansburi- ana, crotalus mitchellii. Isla Cardonosa Este. (Isla Cardonosa; 0.5 km NE of Isla Partida Norte). Uta stansburiana, Phyllo- dactylus partitus*, Cnemidophorus tigris. Isla Carmen. Dipsosaurus dorsalis, Sauromalus slevini*, Callisaurus draconoides, Sceloporus or- cutti, Sceloporus zosteromus, Urosaurus nigricaudus, Uta stansburiana, Phyllodactylus xanti, Cnem- idophorus carmenensis*, Cnemidophorus tigris, Leptotyphlops humilis, Hypsiglena torquata, Masti- cophis flagellum, Tantilla planiceps, Crotalus enyo, Crotalus mitchellii. Isla Cayo. (within Bahia Concepcion, BCS). Urosaurus nigricaudus, Phyllodactylus xanti. Isla Cerraja. (within Bahia de Los Angeles, BC). Uta stansburiana. Isla Cerralvo. Bufo punctatus, Scaphiopus couchii, Ctenosaura hemilopha, Dipsosaurus dorsalis, Callisaurus draconoides, Sceloporus grandaevus*, Phyllodactylus unctus; Cnemidophorus ceralben- sis*, Leptotyphlops humilis, Lichanura trivirgata, Chilomeniscus savagei*, Eridiphas slevini, Hypsi- glena torquata, Lampropeltis getula, Masticophis flagellum, Phyllorhynchus decurtatus, Rhinocheilus etheridgei*, Trimorphodon biscutatus, Crotalus enyo, Crotalus mitchellii. Isla Cholludo. (Isla Lobos, Son.; Isla Roca Foca; between islas Turners and Tibur6n). Ctenosaura conspicuosa*, Phyllodactylus xanti. Isla Coloradito. (Isla Lobos, BC; Lobera). Uta tumidarostra*. Isla Coronados. Dipsosaurus dorsalis, Sauromalus slevini*, Callisaurus draconoides, Sceloporus orcutti, Sceloporus zosteromus, Urosaurus nigricaudus, Uta stansburiana, Coleonyx variegatus, Phyl- lodactylus xanti, Cnemidophorus hyperythrus, Cnemidophorus tigris, Eridiphas slevini, Hypsiglena torquata, Masticophis flagellum, Crotalus enyo. Isla Danzante. Sauromalus ater, Callisaurus draconoides, Petrosaurus repens, Urosaurus nigricau- dus, Uta stansburiana, Coleonyx variegatus, Phyllodactylus xanti, Cnemidophorus tigris, Leptotyph- lops humilis, Bogertophis rosaliae, Chilomeniscus cinctus, Eridiphas slevini, Hypsiglena torquata, Masticophis flagellum, Trimorphodon biscutatus, Crotalus ruber. AMPHIBIANS AND REPTILES ON ISLANDS IN THE GULF OF CALIFORNIA 35) Isla Datil. (Isla Turners). Gopherus agassizii (?), Uta stansburiana, Phyllodactylus xanti, Masticophis flagellum, Crotalus atrox. Isla El Coyote. (within Bahia Concepcion, BCS). Sauromalus ater, Urosaurus nigricaudus, Phyllo- dactylus xanti. Isla El Muerto. (Isla Miramar; Link). Petrosaurus mearnsi, Phyllodactylus xanti, Uta lowei*, Tri- morphodon biscutatus, Crotalus muertensis*. Isla El Pardito. (Isla Coyote). Uta stansburiana, Phyllodactylus xanti. Isla El Requeson. (within Bahia Concepcion, BCS). Urosaurus nigricaudus. Isla Espiritu Santo. Bufo punctatus, Scaphiopus couchii, Dipsosaurus dorsalis, Sauromalus ater, Callisaurus draconoides, Petrosaurus thalassinus, Sceloporus hunsakeri, Sceloporus zosteromus, Uro- saurus nigricaudus, Uta stansburiana, Coleonyx variegatus, Phyllodactylus unctus, Cnemidophorus espiritensis*, Cnemidophorus tigris, Chilomeniscus punctatissimus*, Masticophis barbouri*, Masti- cophis flagellum, Salvadora hexalepis, Crotalus enyo, Crotalus mitchellii. Isla Encantada. Uta encantadae*. Isla Flecha. (within Bahia de Los Angeles, BC). Sauromalus hispidus*, Uta stansburiana. Isla Gallina. (0.5 km W of Isla Espiritu Santo, BCS). Urosaurus nigricaudus, Phyllodactylus unctus. Isla Gallo. (0.5 km W of Isla Espiritu Santo). Sauromalus ater, Sceloporus hunsakeri, Urosaurus nigricaudus, Uta stansburiana, Phyllodactylus unctus. Isla Gaviota. (0.5 km N of Pichilingue, BCS). Urosaurus nigricaudus. Isla Granito. Sauromalus hispidus*, Uta stansburiana. Isla Islitas. (11.5 km S of Puerto Escondido, BCS). Urosaurus nigricaudus. Isla Lagartija. (0.25 km N of Isla Salsipuedes). Uta stansburiana. Isla La Raza. Uta stansburiana, Phyllodactylus xanti. Isla La Ventana. (Isla Nuevo Amor; within Bahia de Los Angeles, BC). Sauromalus hispidus*, Uta stansburiana, Phyllodactylus xanti. Isla Las Animas. (BCS). Urosaurus nigricaudus, Phyllodactylus xanti. Isla Las Galeras. (there are two islands, east and west; 3 km N of Isla Monserrate). Uta stansburiana. Isla Mejia. Sauromalus hispidus*, Petrosaurus slevini*, Uta stansburiana, Phyllodactylus xanti, Li- chanura trivirgata, Hypsiglena torquata. Isla Mitlan. (within Bahia de Los Angeles, BC). Sauromalus hispidus*, Uta stansburiana. Isla Monserrate. (Isla Monserrat; Monserrato). Dipsosaurus dorsalis, Sauromalus slevini*, Scelo- porus zosteromus, Uta stansburiana, Phyllodactylus xanti, Cnemidophorus pictus*, Chilomeniscus cinctus, Hypsiglena torquata, Lampropeltis getula, Masticophis flagellum, Phyllorhynchus decurtatus, Crotalus mitchellii, Crotalus ruber. Isla Moscas. (within Bahia Concepcion, BCS; Dixon, 1966). Phyllodactylus xanti. Isla Pardo. (13.5 km S of Puerto Escondido, BCS). Sauromalus ater, Urosaurus nigricaudus, Phyl- lodactylus xanti, Crotalus enyo. Isla Partida Norte. (Isla Partida; Cardonosa). Uta stansburiana, Phyllodactylus partitus, Cnemido- Phorus tigris, Hypsiglena gularis*. Isla Partida Sur. Bufo punctatus, Scaphiopus couchii, Dipsosaurus dorsalis, Sauromalus ater, Cal- lisaurus draconoides, Petrosaurus thalassinus, Sceloporus hunsakeri, Scelporus zosteromus, Urosau- rus nigricaudus, Uta stansburiana, Coleonyx variegatus, Phyllodactylus unctus, Cnemidophorus es- piritensis*, Cnemidophorus tigris, Chilomeniscus punctatissimus*, Hypsiglena torquata, Masticophis barbouri*, Masticophis flagellum, Crotalus enyo, Crotalus mitchellii. Isla Pata. (within Bahia de Los Angeles, BC). Uta stansburiana. Isla Patos. (16 km north of Isla Tibur6n, SON). Callisaurus draconoides, Uta stansburiana. Isla Piojo. (within Bahia de Los Angeles, BC). Sauromalus hispidus*, Uta stansburiana, Phyllo- dactylus xanti, Crotalus mitchellii. Isla Pond. (Isla Estanque; La Vibora). Sauromalus hispidus*, Uta stansburiana, Phyllodactylus xanti, Cnemidophorus tigris, Crotalus ruber. Isla Roca Lobos. (0.25 km SW of north end of Isla Salsipeudes). Sauromalus varius*, Uta stans- buriana. Isla Salsipuedes. Uta stansburiana, Phyllodactylus xanti, Cnemidophorus canus*, Lampropeltis ge- tula, Crotalus mitchellii. Isla Santiago. (0.25 km W of Isla Coronados, BCS). Dipsosaurus dorsalis. Isla San Cosme. (13 km W of Isla Monserrate). Sauromalus ater, Urosaurus nigricaudus, Cnemi- dophorus tigris. 56 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Isla San Damian. (12 km W of Isla Monserrate). Urosaurus nigricaudus. Isla San Diego. Sauromalus ater, Sceloporus angustus*, Phyllodactylus xanti. Isla San Esteban. Crenosaura conspicuosa*, Sauromalus varius, Uta stansburiana, Phyllodactylus xanti, Cnemidophorus tigris, Hypsiglena torquata, Lampropeltis getula, Masticophis slevini*, Crotalus estebanensis*. Isla San Francisco. Sauromalus ater, Callisaurus draconoides, Sceloporus orcutti, Urosaurus nigri- caudus, Uta stansburiana, Phyllodactylus xanti, Cnemidophorus celeripes*, Cnemidophorus francis- censis*, Hypsiglena torquata, Crotalus enyo. Isla San Ildefonso. Sceloporus orcutti, Uta stansburiana, Phyllodactylus xanti, Masticophis flagellum. Isla San Jose. Dipsosaurus dorsalis, Sauromalus ater, Callisaurus draconoides, Sceloporus orcutti, Sceloporus zosteromus, Urosaurus nigricaudus, Uta stansburiana, Coleonyx variegatus, Phyllodac- tylus xanti, Cnemidophorus celeripes*, Cnemidophorus danheimae*, Chilomeniscus cinctus, Hypsi- glena torquata, Masticophis flagellum, Phyllorhynchus decurtatus, Pituophis vertebralis, Salvadora hexalepis, Sonora semiannulata, Trimorphodon biscutatus, Crotalus enyo, Crotalus mitchellii, Cro- talus ruber. Isla San Lorenzo Norte. (Isla Las Animas, BC). Sauromalus hispidus*, Uta stansburiana, Phyllo- dactylus xanti, Cnemidophorus canus*, Lampropeltis getula. Isla San Lorenzo Sur. (Isla San Lorenzo). Sauromalus hispidus*, Uta stansburiana, Phyllodactylus xanti, Cnemidophorus canus*, Hypsiglena torquata, Lampropeltis getula, Crotalus lorenzoensis*. Isla San Luis. (Isla Encantada Grande; Salvatierra; San Luis Gonzaga). Dipsosaurus dorsalis, Cal- lisaurus draconoides, Uta stansburiana. Isla San Marcos. Dipsosaurus dorsalis, Sauromalus ater, Callisaurus draconoides, Sceloporus or- cutti, Urosaurus nigricaudus, Uta stansburiana, Coleonyx gypsicolus*, Coleonyx variegatus, Phyllo- dactylus xanti, Cnemidophorus hyperythrus, Cnemidophorus tigris, Leptotyphlops humilis, Lichanura trivirgata, Chilomeniscus cinctus, Eridiphas marcosensis*, Hypsiglena torquata, Masticophis flagel- lum, Phyllorhynchus decurtatus, Sonora semiannulata, Trimorphodon biscutatus, Crotalus enyo, Cro- talus ruber. Isla San Pedro Martir. Uta palmeri*, Cnemidophorus martyris*, Lampropeltis getula, Crotalus atrox. Isla San Pedro Nolasco. Ctenosaura nolascensis*, Sceloporus clarkii, Uta nolascensis*, Phyllodac- tylus homolepidurus, Cnemidophorus bacatus*, Lampropeltis getula. Isla Santa Catalina (Isla Catlan; Catalana; Catalano). Dipsosaurus catalinensis*, Sauromalus klaub- eri*, Sceloporus lineatulus*, Uta squamata*, Phyllodactylus bugastrolepis*, Cnemidophorus catali- nensis*, Leptotyphlops humilis, Hypsiglena torquata, Lampropeltis catalinensis*, Crotalus catalinen- SUS: Isla Santa Cruz. Sauromalus ater, Sceloporus angustus, Phyllodactylus xanti, Crotalus atrox. Isla Smith. (Isla Coronado; within Bahia de Los Angeles, BC). Sauromalus hispidus*, Callisaurus draconoides, Uta stansburiana, Phyllodactylus xanti, Cnemidophorus tigris, Hypsiglena torquata, Cro- talus mitchellii. Isla Tijeras. (13 km S of Puerto Escondido, BCS). Urosaurus nigricaudus. Isla Tiburon. Bufo punctatus, Scaphiopus couchii, Gopherus agassizii, Crotaphytus dickersonae, Gambelia wislizenii, Sauromalus ater, Callisaurus draconoides, Phrynosoma solare, Sceloporus clar- kii, Sceloporus magister, Urosaurus ornatus, Uta stansburiana, Coleonyx variegatus, Phyllodactylus xanti, Cnemidophorus tigris, Lichanura trivirgata, Chilomeniscus cinctus, Chilomeniscus stramineus, Hypsiglena torquata, Masticophis bilineatus, Masticophis flagellum, Pituophis melanoleucus, Salva- dora hexalepis, Trimorphodon biscutatus, Micruroides euryxanthus, Crotalus atrox, Crotalus cerastes, Crotalus molossus, Crotalus tigris. Isla Tortuga. Sceloporus orcutti, Uta stansburiana, Hypsiglena torquata, Lampropeltis getula, Cro- talus tortugensis*. Isla Willard. (within Bahia de San Luis Gonzaga, BC). Sauromalus ater, Uta stansburiana. Islas Santa Inez. (three islands; records from southernmost island only). Callisaurus draconoides, Coleonyx variegatus. Islotes Blancos. (0.25 km SE of Isla Encantada). Uta encantadae*. Bull. Southern California Acad. Sci. 98(2), 1999, pp. 57—65 © Southern California Academy of Sciences, 1999 Identification and Distribution of Spiny Pocket Mice (Chaetodtpus) in Cismontane Southern California Richard A. Erickson! and Michael A. Patten? 'LSA Associates, One Park Plaza, Suite 500, Irvine, California 92614 e-mail: richard.erickson@ lsa-assoc.com 2Department of Biology, University of California, Riverside, California 92521 e-mail: patten@ citrus.ucr.edu Abstract.—The San Diego Pocket Mouse (Chaetodipus fallax) and California Pocket Mouse (C. californicus) frequently are confused with each other. The shorter, more rounded ears of San Diego Pocket Mice, and to a lesser extent their smaller hind feet, are the best external means of distinguishing them from Cali- fornia Pocket Mice. The two species are largely allopatric, californicus to the north and fallax to the south, but their ranges overlap somewhat in cismontane southern California. California Pocket Mice occupy higher elevations in this area and farther south, whereas San Diego Pocket Mice are generally found at lower elevations and extend no farther north than southeastern Los Angeles County. Two species of spiny pocket mouse (Chaetodipus; formerly considered a sub- genus of Perognathus; Hafner and Hafner 1983) coexist in cismontane southern California. The California Pocket Mouse (C. californicus) occurs from central California to the mountains of northern Baja California, and the San Diego Pocket Mouse (C. fallax) occurs from southwestern California to northwestern Baja Cal- ifornia Sur (Hall 1981, Schmidly et al. 1993, Williams et al. 1993). Based upon our Own experiences in the field, and our examination of museum specimens and the scientific literature, it is clear that these species often have been confused and misidentified. This note is intended to draw attention to this issue, and was pre- pared with biological consultants primarily in mind. More than any other group, they are charged with documenting the status of wildlife in this rapidly developing region. Moreover, they are among the least likely to produce specimen material in their work. Although we are aware of no evidence suggesting these species are threatened at this time, two subspecies under review here (C. c. femoralis and C. f. fallax) were Category-2 candidates for federal listing as threatened or en- dangered (Federal Register 59:58982—59028) before that category was eliminated in 1995. Given the propensity for heteromyids to populate such lists, these species may once again receive attention by the regulatory wildlife agencies. If so, reliable data for their occurrence and distribution will be vital to management planning. Methods We obtained ear (to the notch) and hind foot measurements for all specimens of these two species taken from Pacific coastal drainages in southwestern Cali- fornia housed in the Natural History Museum of Los Angeles County (LACM; n = 413), San Diego Natural History Museum (SDNHM; n = 252), California Academy of Sciences (CAS; n = 37), and the San Bernardino County Museum 37 58 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1. Comparisons of ear length (at the notch) and hind foot length between Chaetodipus californicus and Chaetodipus fallax fallax. C. californicus C.. fi fallax Cal notch sample size n = 1,083 he = 367, mean + SD 11.44 + 0.88 mm 8.45 + 0.79 mm (t = 59.01, P < 0.0001, d.f. = 1,468) range 8—14 mm 6-10 mm coefficient of variation 0.08395 0.09436 hind foot sample size n = 347 ni-t3is3 mean + SD 25.16 = 1.42 mm 23.36 + 1.18 mm (t = 18.63, P < 0.0001, d.f. = 718) range 21-30 mm 20-27 mm coefficient of variation - 0.05641 0.05057 (SBCM; n = 17). We recorded measurements from the original specimen tags unless we had reason to believe they were in error (e.g., ear measurements from the crown), in which case we remeasured the ear if possible, but generally dis- regarded foot measurements due to the difficulty of remeasuring them on dried specimens. Measurements of pocket mice live-trapped by Erickson and others in the course of consulting work from 1994—1997, primarily in southern Orange and northern San Diego counties (LSA unpublished data; n = 751) also were used in our analysis. We compared mean lengths of ear and hind foot between the two species using standard t-tests. We performed a canonical discriminant function analysis to ex- amine the degree to which we could distinguish the species based only on these two mensural characters. Prior probabilities for groups classification were set to one-half. Correct placement of individuals to species was determined using both jacknifed classification percentages and Mahalanobis D? values with their asso- ciated posterior probabilities of correct classification. Statistical analyses were performed using BMDP Statistical Software ver. PC-90. In addition to standard reference works (i.e., Grinnell 1933, McLaughlin 1959, Huey 1964, Bond 1977, Hall 1981) and localities listed on the specimens ex- amined, we used our own trapping records and those maintained by the Orange County Vector Control District (OCVCD), data from the Museum of Vertebrate Zoology, University of California, Berkeley (MVZ) and Field Museum of Natural History (Chicago) collections, and the reports of fellow investigators to formulate general distributional patterns. Results Identification.—Cismontane southern California examples of these species dif- fer significantly in length of ear (to the notch) and in length of hind foot (Table 1). These two mensural characters were combined to yield two discriminant func- tions: DISTRIBUTION OF SPINY POCKET MICE IN SOUTHERN CALIFORNIA 59 Table 2. Jackknifed classification resulting from a canonical discriminant function analysis of lengths of ear and hind foot of Chaetodipus californicus and Chaetodipus fallax fallax. Number of cases classified into group Group % correct californicus fallax californicus 92.7 319 25 fallax 97.9 8 365 Total 95.4 327 390 californicus = 13.4712(hind foot) + 11.09899(ear,,,..,) — 234.0003 fallax 12.87714(hind foot) + 7.33506(ear,,,.,) — 182.0511 These functions correctly classified 95% of the individuals we examined (Table 2). Only one-third as many individual fallax were misclassified compared to mis- classifications of californicus. Based on an examination of posterior probabilities of correct classification, the majority of misclassified californicus were immatures, suggesting that these animals had not yet attained full adult size in ear length and hind foot length. In any case, scores for these species on the first canonical dis- criminant function axis formed a distinctly bimodal distribution, and virtually all of the misclassified individuals were near the valley between the two peaks (Fig- ure 1). Thus, the above formulas can be used to classify reliably virtually all individuals of these two species of Chaetodipus based only on their ear length at notch and their hind foot length. Even without statistical procedures, California Pocket Mice and San Diego Pocket Mice are readily distinguished by the length and shape of their ears. The greater ear length of californicus is not merely the result of the species’ slightly larger size. The width of the ears do not appear to be appreciably different, so the length of californicus imparts a nearly parallel-edged, elliptical shape com- 90 ME Chaetodipus californicus == Chaetodipus fallax Frequency Canonical Variate | Fig. 1. Mensural statistics; see text. 60 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig. 2. Ears of San Diego (Cbaetodipus fallax; a) and California (C. californicus; b) pocket mice. Note the nearly parallel-sided elliptical shape of the latter compared to the nearly round shape of the former. Drawing from Osgood (1900). pared to the nearly round ears of fallax (Figure 2). Even very young californicus, with ear lengths more typical of fallax, can be identified by ear shape. Hind foot measurements are also significantly different in these two species, but the dif- ference is more proportional to their difference in overall size and there is con- siderable overlap. In addition to these external characteristics, and beyond the scope of this paper, these species may be distinguished by the shape of their skulls (Hall 1981, Williams et al. 1993) and apparently the presence (califor- nicus) or absence (fallax) of white spinous hairs on the shoulders (S. J. Mont- gomery in litt.). California Pocket Mouse distribution.—This species is primarily found in the Upper Sonoran and Transition life zones (Grinnell 1933, Bond 1977). It occurs at low to moderate elevations in central California, occurs primarily at moderate elevations in southern California, and is restricted to high elevations at the south- ern end of its range in the Sierra San Pedro Martir. Three subspecies of California Pocket Mouse have been described from south- ern California: C. c. dispar, C. c. bernardinus, and C. c. femoralis (Figure 3). C. c. bernardinus is found primarily in chaparral at moderate to high elevations (750—2100 m) in the San Bernardino, San Jacinto, and eastern San Gabriel moun- tains of Los Angeles, San Bernardino, and Riverside counties (Grinnell 1933); note that Williams et al. (1993) erroneously excluded the San Jacinto Mountains from the range of this subspecies. A specimen labeled bernardinus from a low elevation site at the eastern edge of the Chino Hills in extreme northwestern Riverside County (MVZ 132520) is from outside the published range of the taxon, but was apparently examined by Seth Benson (J. L. Patton pers. comm.), who first described the subspecies (Benson 1930). The locality fits better with the southeastern range of dispar described below. The southern limits of dispar and the northern limits of femoralis are unclear. DISTRIBUTION OF SPINY POCKET MICE IN SOUTHERN CALIFORNIA 61 Thy, Adhk _SAN JOAQUIN, HILLS; pe i California Pocket Mouse Chaetodipus californicus San Diego Pocket Mouse Chaetodipus fallax Crosses Indicate Areas of Presumed Intergradation Fig. 3. Approximate distribution of two species of pocket mice in southwestern California. Grinnell (1933) and Williams et al. (1993) described the range of dispar as ex- tending south only to Los Angeles County, and the California range of femoralis as limited to San Diego County, thus ignoring all of Orange County. Specimens in MVZ labeled dispar come from the southern foothills of the San Gabriel Moun- tains (Vaughan 1954) and the Palos Verdes Peninsula in Los Angeles County, and near Laguna Beach, Orange County. MVZ specimens labeled femoralis extend north to the vicinity of Las Pulgas Creek, San Diego County and Aguanga in southwestern Riverside County. Pequegnat (1951) considered the Santa Ana Mountains to represent an intergradation zone between these two subspecies; he 62 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES identified most specimens from there as dispar, but considered three specimens representative of femoralis. California Pocket Mice have been trapped throughout the undeveloped low- lying portions of Orange County (OCVCD, LSA, and Patten unpubl. data) up- wards to Santiago Peak (Pequegnat 1951), but are essentially unknown in the valleys of western Riverside and southwestern San Bernardino counties (SBCM, pers. obs.). In San Diego County, californicus has been collected on the coast south to Oceanside (Hall 1981; MVZ and LSA unpubl. data), but farther south has been found only at interior sites east to the desert slopes of the mountains. San Diego Pocket Mouse distribution.—The more southerly distributed San Diego Pocket Mouse is primarily resident in the Lower Sonoran Life Zone, and thus is generally restricted to relatively low elevations (exceptionally to 1800 m on the north side of the San Bernardino Mountains) throughout its range (Grinnell 1933, Pequegnat 1951, Vaughan 1954, Huey 1964, Bond 1977). San Diego Pocket Mice are apparently widespread throughout western Baja California, south to ex- treme northwestern Baja California Sur (Huey 1964, Hall 1981). Two subspecies are found in southern California: C. f pallidus is restricted to the western deserts, north to northeastern Los Angeles County and nominate fallax is found south and west of the mountains (Grinnell 1933, Hall 1981). A skeletal specimen labeled pallidus from Devore, San Bernardino County (MVZ 158943) is from an area outside the biogeographical limits of the subspecies and here is considered dubious. San Diego Pocket Mice have been collected at scattered localities throughout the lowlands of western San Diego County all the way to San Onofre (Grinnell 1933) and the mouth of San Mateo Creek (LSA unpubl. data) on the coast. To the north, fallax has been found in Orange County only locally in the foothills of the Santa Ana Mountains (Pequegnat 1951, Bontrager 1975; MVZ, OCVCD and Patten unpubl. data) north to the Santa Ana River Canyon (MVZ) and Chino Hills (Patten unpubl. data). We examined two purported fallax specimens from the San Joaquin Hills, Orange County (University of California, Irvine Museum of Sys- tematic Biology) and found them to be typical californicus; one had been rela- beled as such. The specimens have since been deposited at MVZ (uncataloged at this time), where their identities were confirmed by J. L. Patton (pers. comm.). Two additional specimens from the same locality (UCIMSB 274 and 282; now in the Museum of Wildlife and Fisheries Biology at the University of California, Davis) have ear measurements of 9 and 10 mm. Only californicus had been found previously in the San Joaquin Hills (Pequegnat 1951; MVZ) or subsequently during extensive trapping since at least 1990 (LSA and OCVCD unpubl. data). C. f. fallax is widespread in the valleys of western Riverside and southwestern San Bernardino counties (pers. obs.) and extends northwest to the vicinity of Claremont, Los Angeles County (Hall 1981, Williams et al. 1993). At the southern base of the San Gabriel Mountains, Vaughan (1954) reported fallax below 570 m, and californicus had replaced fallax by ca. 610 m at a site north of Etiwanda (LSA unpublished data). In the Santa Ana Mountains, Pequegnat (1951) found fallax only below 610 m (‘‘much more abundant on the interior slopes’’), and Bontrager (1973) found fallax and californicus coexisting at three of his study plots (elevation 400—670 m) on the Santa Rosa Plateau, Riverside County. DISTRIBUTION OF SPINY POCKET MICE IN SOUTHERN CALIFORNIA 63 Discussion The different ear proportions of these species have been known for at least 100 years (Figure 2 first appeared in 1900) and fallax was known formerly as the Short-eared Pocket Mouse. Nevertheless, we believe this review is warranted in light of the confusion between these species that we have witnessed in southern California. We admit to misidentifying numerous animals ourselves before gaining sufficient experience and accepting the sample technique of measuring the ears. Spurred on by popular field guides, and even more technical works, that may (e.g., Ingles 1965, Whitaker 1980) or may not (e.g. Palmer 1954, Burt and Gros- senheider 1976, Hall 1981, Jameson and Peeters 1988) include ear measurements for both species, observers often attempt to identify these species by such unre- liable means as habitat, overall pelage coloration, or the extent and color of spines on the rump, without resorting to measurements. This identification problem has not been limited to the unsophisticated. We found misidentified animals in all of the museums where we examined specimens, and the same was true of the OCVCD collection (S. G. Bennett pers. comm.). Claims of either species outside of the general ranges described here should be carefully documented, preferably by specimen. Not surprisingly, the misidentification problem has crept into the scientific lit- erature as well. M’Closkey (1972, 1976) and Meserve (1976) reported on a pop- ulation of San Diego Pocket Mice in the San Joaquin Hills of Orange County that we have identified as californicus (see Results). Their findings on the eco- logical characteristics of this population have begun to be incorporated in the species specific literature on the San Diego Pocket Mouse (e.g., Zeiner et al. 1990). By calling attention to the potential problem of misidentifying Chaetodipus in southern California, we hope to spare others the misfortune we have gone through, and to prevent any further muddying of the literature on these species. This is but one of several mammal identification challenges confronting biol- ogists in southern California. These problems are especially acute for those, such as consultants, who rarely collect and/or handle specimens. Price et al. (1992) clarified the situation concerning Stephens’ Kangaroo Rat (Dipodomys stephensi), but other taxa in need of attention include the Brush Rabbit (Sylvilagus bach- mani), the “‘Pacific’”’ kangaroo rats (D. agilis and D. simulans; Sullivan and Best 1997), and the white-footed mice (Peromyscus, especially the Brush Mouse P. boylii). Acknowledgments Specimen examination at LACM, SDNHM, SBCM, CAS, and the University of California, Irvine (Museum of Systematic Biology) was facilitated by Kimball L. Garrett, Philip Unitt, Robert L. McKernan, Andrea Jesse, and Peter A. Bowler, respectively. Barbara R. Stein sent us a printout of specimen data from MVZ, Thomas S. Schulenberg and David E. Willard did the same for the Field Museum of Natural History, and Ronald E. Cole provided measurements of specimens in his care at the Museum of Wildlife and Fisheries Biology, University of Califor- nia, Davis. Peter Bowler arranged the transfer of important specimens to MVZ, where James L. Patton kindly commented on them for us. Patton also alerted us to Osgood’s publication and gave us other welcome advice. Stephen G. Bennett 64 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES of the Orange County Vector Control District shared collection data from Orange County, as well as portions of Los Angeles, Riverside, and San Diego counties. Kris Walden prepared the map. Peter H. Bloom, Scott D. Cameron, J. R. Easton, Karen Kirtland, John N. Ko, Spencer Langdon, and Ruben S. Ramirez, Jr. assisted with data collection in the field. We discussed the status of these species in south- ern California with Shana C. Dodd, Stephen J. Montgomery, and Mark A. Pav- elka; Montgomery also shared Chaetodipus photographs with us. David R. Bon- trager provided unpublished materials and discussed habitat preferences, much of which could not be incorporated here. Peter L. Meserve’s refreshingly honest correspondence was especially appreciated. Bontrager, Daniel A. Guthrie, Mont- gomery, and an anonymous reviewer made helpful comments on the manuscript. Literature Cited Benson, S. B. 1930. Two. new pocket mice, genus Perognathus, from the Californias. Univ. Calif. Publ. in Zool. 32:449—454. Bond, S. I. 1977. An annotated list of the mammals of San Diego County, California. Trans. of the San Diego Soc. Nat. Hist. 18:229—248. Bontrager, D. R. 1973. Rodent ecology of the Santa Rosa Plateau, Riverside County, California. Unpublished M.A. Thesis, California State University, Long Beach. Bontrager, D. R. 1975. An ecological survey of the mammals of the Starr Ranch Audubon Sanctuary, Orange County, California. Unpublished report. 69 pp. Burt, W. H. and R. P. Grossenheider. 1976. A Field Guide to the Mammals, 3rd ed. Houghton Mifflin Company, Boston. Grinnell, J. 1933. Review of the recent mammal fauna of California. Univ. Calif. Publ. in Zool. 40: 71-234. Hafner, J. C. and M. S. Hafner. 1983. Evolutionary relationships of the heteromyid rodents. 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North American Fauna 18:1—65. Palmer, R. S. 1954. The Mammal Guide, Mammals of North America North of Mexico. Doubleday, Garden City, New York. Pequegnat, W. E. 1951. The biota of the Santa Ana Mountains. J. Entomology and Zoology vol. 42, nos. 3&4. Price, M. V., P. A. Kelly, and R. L. Goldingay. 1992. Distinguishing the endangered Stephens’ Kan- garoo Rat (Dipodomys stepbensi) from the Pacific Kangaroo Rat (Dipodomys agilis). Bull. So. Calif. Acad. of Sci. 91:126—136. Schmidly, D. J., K. T. Wilkins, and J. N. Derr. 1993. Biogeography. pp. 319-356 in H. H. Genoways and J. H. Brown (eds.). Biology of the Heteromyidae. Spec. Publ. No. 10, Amer. Soc. of Mammalogists. Sullivan, R. M. and T. L. Best. 1997. Systematics and morphologic variation in two chromosomal forms of the agile kangaroo rat (Dipodomys agilis). J. Mammalogy 78:775—797. DISTRIBUTION OF SPINY POCKET MICE IN SOUTHERN CALIFORNIA 65 Vaughan, T. A. 1954. Mammals of the San Gabriel Mountains of California. Univ. Kansas Publ., Mus. of Nat. Hist. 7:513—582. Whitaker, J. O., Jr. 1980. The Audubon Society Field Guide to North American Mammals. Alfred A. Knopf, New York. Williams, D. FE, H. H. Genoways, and J. K. Braun. 1993. Taxonomy. pp. 38—196 in H. H. Genoways and J. H. Brown (eds.). Biology of the Heteromyidae. Spec. Publ. No. 10, Amer. Soc. of Mammalogists. Zeiner, D. C., W. EK Laudenslayer, Jr., K. E. Mayer, and M. White (eds.). 1990. California’s Wildlife, vol. 3, Mammals. California Department of Fish and Game, Sacramento. Accepted for publication 29 September 1998. Bull. Southern California Acad. Sci. 98(2), 1999, pp. 66-74 © Southern California Academy of Sciences, 1999 Growth and Mortality of the Fantail Sole, Xystreurys holepis (Jordan and Gilbert 1881) off the Western Coast of Baja California, Mexico Marco A. Martinez-Mufioz and A. A. Ortega-Salas Instituto de Ciencias del Mar y Limnologia, UNAM, México 04510, D.F. Ap. Post. 70-305 e-mail: marcoa@mar.icmyl.unam mx; ortsal@mar.icmyl.unam.mx Abstract.—Data on growth and mortality were obtained for fantail sole, Xystreu- rys liolepis collected with otter trawls during 11 cruises off the western coast of Baja California, Mexico, from April 1988 to December 1990. The Bottom tem- perature varied between 11 to 19°C. In all, 712 X. liolepis were caught and over the sampling period the sex ratio of males to females was 1:1.2. Males were smaller than females. The relationship between weight and length is described for males by, W = 1.1013 X 10° SL?!?!5# and for females by, W = 7.6274 X 10~° SL?!9?. Von Bertalanffy growth parameters were determined to be: L,, = 443 mm SL, k = 0.16, t) = —0.098 total sample, for males by, L,, = 354.85 mm SL, k = 0.151, t) = —0.918, and for females by, L., = 444.68 mm SL, k = 0.1636, t, = —0.423. The total mortality (Z) rate was 0.62 and the estimated fishing mor- tality (F) was 0.46. Although fantail sole is not a target species of commercial fisheries, it suffers high mortality as part of the bycatch in the shrimp fishery. Resumen.—Se obtuvieron datos de crecimiento y mortalidad del lenguado de dos manchas, Xystreurys liolepis, éstos fueron colectados con redes de arrastre de fondo durante 11 cruceros, que se llevaron a cabo en la costa occidental de Baja California, México, de abril de 1988 a diciembre de 1990. La temperatura de fondo vari6 de 11 a 19°C. Se capturaron 712 individuos de X. liolepis, y durante el periodo de muestreo, la proporciédn de hembras y machos fue de 1:1.2. Los machos presentaron tallas menores comparadas con las de las hembras. La rela- ci6n peso-longitud esta descrita con la siguiente ecuaci6n para los machos W = 1.1013 SL*?"5* y para las hembras fue de W = 7.6274 X 1.1013 ie SL2-25% 7.6274 X 10°-° SL?-"°?. Los parametros de crecimiento de Von Berar lanffy fueron los siguientes L,, = 443 mm SL, k = 0.16, t, = —0.098 para la muestra total, para los machos fue de L,, = 354.85 mm SL, k = 0.151, to = —0.918, y para las hembras por L,, = 444.68 mm SL, k = 0.1636, t) = —0.423. La tasa de mortalidad total (Z) estimada fue de 0.62 y la mortalidad por pesca fue de 0.46. Aunque al lenguado de dos manchas, no se le considera como una pesqueria directa, éste pez sufre una alta mortalidad, debido a la captura incidental como fauna de acompafiamiento, en la pesca del camaron. The fantail sole ranges from Monterey Bay to the Gulf of California at depths from 4 to 79 m (Miller and Lea, 1972). A number of studies on the distribution and abundance of the fantail sole, Xystreurys liolepis, have been conducted. Love et al (1986) reported X. liolepis in soft sediments off southern California between 66 GROWTH AND MORTALITY OF THE FANTAIL SOLE OFF BAJA CALIFORNIA 67 6 to 18 m depth. In this area it is most commonly found in shallow offshore waters and its biomass is relatively constant across all open coast depth strata, ranging from 14 to 16 kg (Kramer 1991). However Allen and Herbinson (1991) reported that its standing crop may vary inversely with temperature. Along the western coast of Baja California, Martinez-Mufnoz and Ramirez-Cruz (1992) stat- ed that fantail sole is an abundant species, reaching up to 50 cm in length between 24° to 28° north latitude at depths from 13 to 150 m. It was also observed that in summer these fish move into shallow waters to reproduce. Moser and Watson (1990) using CALCOFI samples between 1951 and 1981 mentioned that X. lio- lepis larvae were captured from Punta Concepcion to Magdalena Bay, B.C.S and were relatively abundant between the coast of Sebastian Vizcaino Bay to Mag- dalena Bay from July to October, reaching peak abundance in August (10 larvae/ m°). Despite these frequent references to its occurrence, there have been very few studies on the biology of X. liolepis. Off California adults of fantail sole feed on polychaetes, shrimp, crabs, and euphasiids. They spawn at the end of winter and the beginning of spring (Frey, 1971). Although it is not considered a commercially desirable fish, fantail sole has often been found as part of the bycatch in the trawl fishery for shrimp off Baja California and in the Gulf of California (Ramirez-Hernandez and Arvizi Matinez 1965). Off the coast of Sinaloa and Baja California X. liolepis was caught with shrimp trawls Ramirez-Hernandez et al (1965). Castro-Aguirre et al (1970) and van der Heiden (1985) found that fantail sole is frequently caught by shrimp trawlers in the Gulf of California. Pérez-Mellado and Findley (1985) reported 2% of shrimp catches along the Sonora and Sinaloa coasts consisted of X. liolepis. Many reach sizes > 30 cm long off the coast of Orange County, California (Mearns 1979). In Mexico there are no regulations to monitor catching of flattish (Balart 1996), so the numbers reported are reliable. However it is known there is a great fishery for them associated with shrimp trawling with most of the catches sold to Cali- fornia markets. As Martinez-Mufioz and Ramirez-Cruz (1992) consider this species a potential exploitable flatfish, the purpose of this paper is to provide information on the growth and mortality of X. liolepis populations in the Mexican Pacific Ocean, which is needed for the regulation of X. liolepis as a fishery resource. Materials and Methods Fish were collected from April 1988 to December 1990 from 160 otter trawls conducted during the course of eleven cruises aboard the B/O ““EL PUMA” and B/I MARSEP XVI. The 34 sampling stations were located off the western coast of Baja California, from Boca del Carrizal (23°00’ N) to Sebastian Vizcaino Bay (28°51' N) in depths of 10 and 250 m (Fig. 1). All fish were obtained by an otter trawl that measured 20 m wide and 9 m high at the mouth, 24 m in length, and has a stretched mesh size of 3 cm. Trawling time and speed were recorded to estimate the area swept by net. The shrimp boat speed was standardized to 2.2 knots while for the ““EL PUMA” this was 3 knots. Trawls were towed for 30 minutes along isobaths. Depth was re- corded with the Simrad sounder aboard the ““EL PUMA’, and Furuno sounder 68 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 115° 114° 113° 1125 111° 110° Xystreurys ae Fig. 1. Stations sampled off Baja California, México, 1988-1990. on the ““MARSEP’’. Temperature at bottom was recorded with Van Dorn bottles and substrate temperature was directly taken from grab samples. Specimens of X. liolepis were measured (standard length, SL), to the nearest mm, weighed (g) and sexed by eye immediately on recovery. Gonads were ex- amined macroscopically according Nikolskii (1963). These data were used to calculate length-weight relationships varying the logarithmic form of the equation W = aL? for each sex, where W is weight in grams, a, is the y-intercept, SL is standard length in mm, b: slope using FISAT software (The FAO-ICLARM Stock Assessment Tools; Gayanilo et al. 1994). Age groups based in length frequency data of X. liolepis were estimated using the modal progression method of Petersen (1939). Mean lengths for age groups were fitted with the von Bertalanffy (1938) equation L, = L,, (1 — exp *“*%) using the Ford (1933) and Walford (1949) method by FISAT software to describe theoretical growth and growth parameters L, = standard length of fish at t years, L.. = theoretical asymptotic length, k = growth coefficient rate of approach to L.., t) = theoretical age at which L, = 0. LFDA (Length Frequency Distribution Analysis v. 3.1). Holden and Bravington (1992) and Ortega-Salas (1981, 1988a) basic programs were also used. Estimation of the instantaneous total mortality rate (Z) was calculated by the GROWTH AND MORTALITY OF THE FANTAIL SOLE OFF BAJA CALIFORNIA 69 100 40 FREQUENCY (%) 20 125 170 215 260 305 350 308 STANDARD LENGTH (mm) E53 Undetermined Hi Males [_]Females 0 = 80 Fig. 2. Length frequency distributions showing population structure of Xystreurys liolepis during 1988-1990. Beverton and Holt (1959) catch curve method described in Ricker (1975) which was based on all fish = 3 years of age. The difference between the instantaneous total mortality coefficient (Z) which includes migration and the natural mortality coefficient (M) gave an estimate of fishing mortality (F): F = Z — M. Although X. liolepis is considered an exploited species, the natural mortality (M) was es- timated as an index from the rate of growth (K) as described by Beverton and Holt (1959). Results Fish from all trawls were combined such that 712 fantail sole, ranging in size from 80 to 390 mm and weights from 1.45 to 9 kg were analysed. Of these 338 (47.5%) were females, 331 (46.5%) were males, and the sex of 43 (6.0%) were undetermined. As with most flatfish, females were considerably larger in length and weight relative to males (Fig. 2). The largest female measured 390 mm SL and 1,450 g weighed, compared with 320 mm and 500 g, respectively, for the largest male. The mean values for females were 222.4 mm SL + 46.44 SD and ge = 2013)SD' weight. For males’ values were 191.6,mm SL = 30.26 SD and 158.6 g + 67.74 SD respectively (Table 1). The ratio of males to females throughout the year was generally 1:1.2 but it varied in winter when the ratio of females was slightly higher. A chi square test showed there was not a significant difference in numbers of females relative to males (x? = 0.0732; d.f. = 1; P < 0.01). Length and weight were closely correlated with the regression coefficients rang- ing from 0.972 in females to 0.982 in males. The length-weight relationship was determined by non-linear least squares methods (Fig. 3). Use of a t-test and mean differences showed there was not a significant differ- ence between sexes according to the mean length with 31.19; confidence interval was 95%; d.f. = 669; t = 10.29; significance level = 3.30 X 10°? at P < 0.05, 70 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1. Sex n Minimum Maximum Mean St. dev. Males 331 Standard length (mm) 93.0 320.0 191.64 30.26 Weight (g) 155 500.0 158.64 67.74 Females 338 Standard length (mm) 100.0 390.0 2227598 46.44 Weight (g) 12 1450.0 Zi Tea 201.3 Undetermined 43 Standard length (mm) 80.0 197.0 127.26 24.59 Weight (g) 9.1 210.0 AaS 38.35 even with mean weight there was not significant differences between sexes with 115.87; 95%; d.f. = 669; t = 10.43; sig. level = 3.97 X 10°°. The von Bertalanffy equations describing theoretical growth of males and fe- males of the fantail sole are shown in Fig. 4. After age 3 (210 mm), females were consistently longer and heavier than males due to differences in reproductive condition or general robustness of fish, although the rate of growth (K) for the two sexes was similar (Fig. 5). However females attained an older age and longer length than males. For the estimation of mortality rates the best results were obtained by the Beverton and Holt (1959) method (Z = 0.62; M = 0.16; F = 0.46) (Fig. 6). Although these populations of X. liolepis are considered exploited, the natural mortality coefficient (M) was taken as the (K) value, as an index, obtained from the von Bertalanffy growth equation (Beverton and Holt 1959). These results suggest that even though fantail sole is not a target species in the commercial fishery, the rate of fishing mortality is higher than natural mortality, possibly due to the large bycatch of flatfish, including fantail sole, in the shrimp fishery. Chavez and Ramos-Padilla (1974) mentioned that along the western coast of Baja Cali- fornia between 25 and 100 m depth that of 75.5 tons caught 3.5 tons were brown —e Females -* Males FW 7.6274 x 10°SL3-1932 7 n=375 ; r = 0.972 M:W= 1.1013x 10° SL3-1215 ; n= 380 ; r = 0.982 50 100 =150 200 ©6250 300 @6©350 8=400 Standard length (mm) Fig. 3. Length-weight relationships of male and female Xystreurys liolepis from Baja California. GROWTH AND MORTALITY OF THE FANTAIL SOLE OFF BAJA CALIFORNIA Tl 500 F SL; = 444.68[ 1 - e 0-1696 (t + 0.423 )] 400 300 200 : MSL = 354.85 [ 1 -e-0-151 (1+ 0.918) ] Standard length (mm) Age (years) —=—— Males —@— Females Fig. 4. Growth curve of female and male Xystreurys liolepis in standard length, showing standard dispersion error at 95% confidence. shrimp, Penaeus californiensis, shark, flatfish, an other fish and 72 tons were considered rubbish such as red crab (Pleuroncodes planipes). Discussion There have been few previous studies concerned with the biology of X. liolepis so the growth and mortality parameters estimated here can only be compared with those obtained in studies of other species of flatfish (Table 2 and 3). In comparison with many other flatfish species for which growth data are avail- 2500 2000 | FLW = 7.6274 x 10*SL3.1%2 ~~ oN 71500 oie = ot = 1000 | 500 M'W = 1.1013 x 10 $S1,32215 0 1 3 5 1 9 1] iis Ni) 17 19 Age (years) —=— Males —e— Females Fig. 5. Weight curve of female and male Xystreurys liolepis in weight, showing the standard dispersion error at 95% confidence. 12 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 0 3 6 9 12 AGE (years) Fig. 6. Catch curve for fantail sole (Xystreurys liolepis) caught off Baja California, Mexico, (N = 712). The curve is slightly non-linear but regression analysis suggests a value of Z = 0.62. Open symbols indicate values not included in the regression. able, the rate of growth of X. liolepis is comparatively high and the species attains a size large enough for commercial exploitation at 3 years age. The ratio of the sexes in all species was generally 1:1. The females are also larger than males and have a higher rate of growth. According to Dagang et al. (1992) as cited in Liu (1990), flatfish generally have a life span of up to 5—6 years, while the oldest fish are occasionally more than 10 years old. At most X. liolepis attains an age of 13 years. The age com- positions for both sexes of the same stock are clearly different and males generally have a shorter life span than females. Thus, while males dominate the younger age-groups, females are more abundant in the older groups. Off the western coast of Baja California, a high fishing mortality of flatfish, including X. liolepis, occurs resulting from intensive trawl fishing for shrimp according to Ramirez-Hernandez and Arvizi Matinez (1965); Ramos-Padilla (1974); Ramirez-Hernandez et al. (1965); Castro-Aguirre et al. (1970) and van der Heiden (1985). Table 2. Specie Location Sex i. (mm) k i Source Hippoglossina — Baja California 208.09 SL 0.1843 0.2096 Ramirez Murillo stomata México. (1995) Xystreurys Mar del Plata, Ar- S27 0.43 —0.169 Fabre and Cous- rasile gentina. seau (1988) Eopsetta Sea Yelow, China. Male 342.47 O23 —0.184 Dagang et al., grigorijewi Fem. 34257 0.3 OZ (1992) as cited by Liu (1990) Limanda Isle of Man, UK. Male. 221 52 0.604 0.4254 Ortega-Salas (1981, limanda Fem. ©329:73 O.SL7S 0.2394 1988a, b) Xystreurys Baja California, Mates 354.85. SL~ (O928 =O)151 Present Paper liolepis México. Fem. 444.68 SL 0.163 —0.918 GROWTH AND MORTALITY OF THE FANTAIL SOLE OFF BAJA CALIFORNIA 73 Fable: Specie Location 7b; Source Hypsopsetta guttulata Anaheim Bay, USA 2.6 Lane (1975) Limanda limanda Isle of Man, UK Ortega-Salas (1981, 1988 a and b) Female 1.06 Male 1.39 Xystreurys liolepis Baja California, México 0.62 Present paper This paper shows that females (L., = 444.68 mm SL) grow bigger than males (L.. = 354.85 mm SL) and the total mortality rate was 0.62 of which fishing mortality was 0.46. Although X. liolepis is not a target species of commercial fisheries, the rate of fishing mortality is higher than natural mortality, possibly due to the large bycatch of flatfish, including fantail sole, in the shrimp fishery. It is considered a potential exploitable flatfish but needs to have regulations as a fishery resource. Acknowledgements This study was partially supported by the Consejo Nacional de Ciencia y Tec- nologia de México (CONACYT), grant P22O0CCOR880518, and also by the Uni- versidad Nacional Aut6noma de México, which provided the B/O *“‘EL PUMA” from 1988 to 1991. We also thank the Instituto de Ciencias del Mar y Limnologia (UNAM) for analysed the information and the Centro de Investigaciones Biol6- gicas del Noroeste for processed the raw material. Literature Cited Allen, M. J., and K. T. Herbinson. 1991. Beam-Trawl survey of bay and nearshore fishes of the soft- bottom habitat of southern California in 1989. Calif. Coop. Oceanic Fish. Invest. Rep. 32:112— W7le Balart, E. FE 1996. Pesqueria de lenguados. Eds. Casas Valdez, M. y G. Ponce Diaz. Estudio del potencial pesquero y acuicola de Baja California Sur. 285 p. Beverton, R. J. H. and S. J. Holt. 1959. 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Taxonomia, biologia y evaluacion de la ictiofauna demersal del Golfo de California, cap. 4; 149-200. I: Yafiez-Arancibia, A. (Ed.). Recursos pesqueros potenciales de México: la pesca acompajfiante del camar6n. Univ. Nal. Auton. México/Secr. Pesca, México 748 p. von Bertalanffy, L. 1938. A quantitative theory of organic growth of animals. Human Biology 10(2): 181-213. Walford, L. A. 1949. A new graphic method of discribing the growth of animals. Biol. Bull. 90(2): 141-147. Accepted for publication 4 August 1998. Bull. Southern California Acad. Sci. 98(2), 1999, pp. 75-79 © Southern California Academy of Sciences, 1999 The Euryhaline Gobiid Fish, Gillichthys mirabilis Cooper 1864, Second Intermediate Host of the Trematode, Pygiopsoides spindalis Martin 1951 Mark H. Armitage Azusa Pacific University, Mary Hill Center, P.O. Box 7000, Azusa, California 91702-7000 Abstract.—Pygidiopsoides spindalis Martin 1951 is reported from the gills of a new second intermediate host, Gillichthys mirabilis Cooper 1864 (the longjaw mudsucker), collected at Point Mugu Lagoon, (Naval Air Weapons Station), Point Mugu, California. Ascocotyle (Phagicola) diminuta Stunkard and Haviland 1924 were also found. Metacercariae were enzymatically excysted from gills and ex- amined by scanning electron microscopy. Fundulus parvipinnis Girard were ex- amined and yielded P. spindalis and Ascocotyle sexidigita Martin and Steele 1970. Parasites reported from the longjaw mudsucker were reviewed. The presence of these trematodes in spite of heavy contamination may be indicative of the health of the Mugu lagoon ecosystem. Martin (1951) described Pygidiopsoides spindalis, a heterophyid trematode, from naturally infected Fundulus parvipinnis Girard collected at Newport Bay, CA, but did not describe the infection site in the killifish. Martin further elucidated the life cycle of P. spindalis (Martin 1964) by exposing F. parvipinnis to cercariae shed from naturally infected Cerithidea californica Haldeman, and determined that the cercaria “‘works its way to the bony support of the [fish] gill before encysting’’. The trematode was not described from other killifish organs nor was the naturally occuring definitive host for P. spindalis described. Earlier, Martin described Euhaplorchis californiensis (1950a) from the brain of F. parvipinnis collected at Playa del Rey, CA, indicating that “Another fish, Gillichthys mirabilis Cooper 1864, commonly called the ‘mudsucker,’ having sim- ilar opportunities for infestation, was not parasitized by this trematode ...’’. No other parasites of G. mirabilis were mentioned. Gillichthys mirabilis was reported, however, as a successful experimental second intermediate host for another new trematode, Stictodora hancocki, described by Martin (1950b, as Parastictodora) which encysted in “the tissues of the lower jaw, around the eyes ...”’ and else- where. I have observed many such naturally occuring metacercarial cysts on mud- suckers collected in the course of this study. Gillichthys mirabilis has been the subject of many studies, several of which have focused on the gill arches and gill filaments, but litthe mention of gill par- asites has been made. Barlow (1963) briefly discussed the parasites of G. mirabilis noting that, ‘““The Salton Sea [variety of these] fish at times are heavily infested with a monogenetic trematode, but so are coastal populations.”’ Todd and Ebeling (1966) studied aerial respiration in the mudsucker, and even sectioned the buccal roof of that and the killifish, but did not discuss gill parasites, although this may have significantly impacted their study. Martin and Multani (1970) reported mono- 1S 76 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES genetic trematode metacercaria in mud suckers from Baja, California, Seal Beach, and Newport Bay. Love and Moser (1983) presented a listing of heterophyids for G. mirabilis on the West Coast, but did not include P. spindalis. Armitage (1997) looked at several G. mirabilis collected at the Twelfth Street bridge at Mugu, but no metacercariae were found. No other report has been made for other second intermediate hosts for P. spindalis, natural or experimental in California. In this paper, a new, naturally occuring second intermediate host, the longjaw mudsucker, is reported for P. spindalis. Materials and Methods Twenty G. mirabilis were collected by trap from the Laguna Road bridge over the western Mugu lagoon area of the Point Mugu Naval Air Weapons Station, CA, from January 1998 through May 1998. Twenty seven F. parvipinnis were also collected. The gills, intestines and livers of these fish were separately har- vested and examined under a dissecting microscope for the presence of metacer- carial cysts. Hearts were not examined. Experimental definitive hosts were not employed. Over 200 of each of the trematodes in this study were successfully excysted. Excysted live worms were fixed in hot formalin and infected gills were fixed in glutaraldehyde. Dehydrated worms were sputter coated at 30mA for 4 minutes and plastic gill sections were stained in Fuchsin and coverslipped. Pho- tomicrographs were taken of whole live material under cover slip pressure, of sections and of fixed worms on a compound microscope and on a scanning elec- tron microscope. Anatomical measurements were not made. Results and Discussion Twenty G. mirabilis were collected and only 4 lacked cysts in the gills, which averaged 8 cysts per filament when infected. Most had cysts of varying sizes, indicating that infections possibly continued occuring over time. Additionally, Ascocotyle (Phagicola) diminuta Stunkard and Haviland 1924 metacercariae were excysted from the mudsucker gills at the same time as P. spindalis, representing a new host record for this trematode as well. These worms exhibited the same anterior row of 16 oral spines with a 2nd row of 2 accessory dorsal spines (Fig. 1A) as described from Leptocottus armatus Girard 1854 collected at Mugu (Ar- mitage 1997). Love and Moser (1983) reported the presence of A. angrense Tra- vassos 1916 from G. mirabilis in Southern California, based on an unpublished personal communication with Baker, however, none of the ascocotylid specimens excysted in this or the previous study (Armitage 1997) exhibited over 16 spines in the anterior row, therefore, it is quite possible that Baker, as well as Yoshino (1972) were looking at A. (P.) diminuta, and not A. angrense. All F. parvipinnis specimens examined contained P. spindalis in their gills (Fig. 1, B, E, F). Ascocotyle sexidigita Martin and Steele 1970 (Fig. 1C—D) were ex- cysted from the intestines of F. parvipinnis as reported by Martin and Steele (1970). No cysts were found in or along the intestines or livers of the mudsuckers collected at the same time as the killifish, although A. sexidigita was reported from the mudsucker intestine by Martin and Multani (1970). The area chosen for collection, along Laguna Road, bisecting the lagoon, is marked as contaminated by pesticides. The California Toxic Substances Monitor- ing program has shown that the fish in Mugu lagoon contain “‘among the highest ARMITAGE: PYGIDIOPSOIDES IN GILLICHTHYS 77 Fig. 1. A.A. (P.) diminuta excysted metacercaria from G. mirabilis scale bar = 10 micrometers. B. P. spindalis excysted metacercaria from G. mirabilis scale bar = 30 micrometers. C. A. sexidigita excysted metacercaria from F. parvipinnis scale bar = 70 micrometers. D. A. sexidigita excysted metacercaria from F. parvipinnis scale bar = 20 micrometers. E. W.M., P. spindalis excysted meta- cercaria from G. mirabilis, scale bar = 35 micrometers. EK W.M., P. spindalis excysted metacercaria from G. mirabilis, scale bar = 35 micrometers. 78 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Fig 2. A. P. spindalis in G. mirabilis gill W.M. scale bar = 100 micrometers. B. Sectioned G. mirabilis gill, uninfected. scale bar = 70 micrometers. C. Sectioned G. mirabilis gill, infected scale bar = 40 micrometers. concentrations of arsenic, silver, DDT, and methoxychlor measured in California”’ (Saiki 1994), yet the abundant presence of this and other trematode parasites indicates that all of the requirements for it to complete its complex life cycle are in place and well established. This could mean that in spite of the contamination, the health of the ecosystem is such that large numbers of cercaria are being ARMITAGE: PYGI/IDIOPSOIDES IN GILLICHTHYS 79 produced by snails, and that piscivorous birds and mammals frequent the lagoon to feed. Like A. (P.) diminuta, P. spindalis requires a snail (probably C. califor- nica (Martin 1964)) as the first intermediate host, and a bird or mammal as the definitive host. Further work must be done to identify naturally occuring definitive hosts of these trematodes at the Mugu lagoon. Acknowledgements The author wishes to thank the reviewers, Les Eddington of Azusa Pacific University for use of the electron microscopy facilities as well as Thomas Keeney, Ecologist/Natural Resources Manager and Capt. Stephen Beal, Commanding Of- ficer Naval Air Station Point Mugu, for access to the Mugu Lagoon. Literature Cited Armitage, M. 1997. The euryhaline cottid fish Leptocottus armatus Girard 1854, second intermediate host of the trematode, Ascocotyle (Phagicola) diminuta Stunkard and Havilland 1924. Bull. So. Calif. Acad. Sci. 96(3):112—116. Barlow, G. W. 1963. Species structure of the gobiid fish Gillichthys mirabilis from coastal sloughs of the eastern Pacific. Pacific Sci. 17:47—72. Love, M. S. and M. Moser. 1983. A checklist of parasites of California, Oregon and Washington marine and estuarine fishes. NOAA Tech. Rept. NMFS SSRF-777. 572 pp. Martin, W. E. 1950a. Euhaplorchis californiensis N. G. n.sp. Heterophyidae:Trematoda, with notes on its life-cycle. Trans. Am. Micro. Soc. 59(2):194—209. 1950b. Parastictodora hancocki N. GEN., n.sp. (Trematoda:Heterophyidae), with observations on its life cycle. J. Parasit. 36(4):360—370. 1951. Pygidiopsoides spindalis N. GEN., n.sp., (Heterophyidae:Trematoda), and its second intermediate host. J. Parasit. 37(3):297—299. 1964. Life cycle of Pygidiopsoides spindalis Martin 1951 (Heterophyidae: Trematoda). Trans. Am. Micro. Soc. 83:270—272. and S. Multani. 1970. Some helminths of the mudsucker fish Gillichthys mirabilis Cooper. Bull. So. Calif. Acad. Sci. 69:161—168. and D. E Steele. 1970. Ascocotyle sexidigita sp. n. (Trematoda:Heterophyidae) with notes on its life cycle. Proc. Helm. Soc. Wash. 37(1):101—104. Saiki, M. K. 1994. Survey of fishes and selected physiochemical variables in Mugu Lagoon and its tributaries, Sept.-Nov. 1993. US Department of the Navy Report, Natural Resources Manage- ment, Environmental Division, Naval Air Station, Point Mugu, CA 93042-5000. 134 pp. Todd, E. S. and A. W. Ebeling. 1966. Aerial respiration in the longjaw mudsucker Gillichthys mirabilis (Teleostei: Gobiidae). Biol. Bull. 130(2):265—288. Yoshino, T. P. 1972. Helminth parasitism in the Pacific killifish, Fundulus parvipinnus from Southern California. J. Parasit. 58:635—636. Accepted for publication 30 July 1998. Bull. Southern California Acad. Sci. 98(2), 1999, pp. 80-89 © Southern California Academy of Sciences, 1999 First Fossil Record of the Pteropod Limacina from the Pacific Coast of North America Richard L. Squires,' James L. Goedert,” and Steven R. Benham? ‘Department of Geological Sciences, California State University, Northridge, California 91330-8266 °15207 84th Ave. Ct. NW. Gig Harbor, Washington 98329-8765, and Section of Vertebrate Paleontology, Natural History Museum of Los Angeles County 3Department of Geosciences, Pacific Lutheran University, Tacoma, Washington 98447 Abstract.—Pteropods from upper Eocene to lower Miocene deep-marine rocks in western Washington represent the first fossil record of genus Limacina from the Pacific coast of North America and the northernmost fossil occurrence in North America for this genus. Two species were found. Both occur in concretions in siltstone, and one was also found in limestone formed by chemosynthetic pro- cesses. Both species closely resemble other late Eocene to early Miocene Lima- cina spp. from Europe, Australia, and New Zealand, but poor preservation of the Washington specimens prevents their identification to the species level. Pteropods are a group of rather poorly known free-swimming, holoplanktonic gastropods. Generally, they are found only in very restricted numbers. Their shells are thin, very fragile (Janssen 1990a), and consist of aragonite, which is easily subject to dissolution (Hodgkinson et al. 1992; Janssen 1991). Limacinids are euthecosomatous pteropods with a small (1-5 mm in height), sinistrally Cleft- handed) coiled shell. Living species of Limacina are found in all the world’s oceans (Tesch 1946, 1948; McGowan 1968; Bé and Gilmer 1977). Most of these species have a free- swimming veliger stage that hatches from a free-floating egg mass, although some species have developed brood protection through the early or entire larval stage (Bandel et al. 1984). The planktonic mode of life of pteropods can allow for wide geographic distribution, and there are indications that fossil species can be used successfully for long-distance correlations (Janssen 1990Qa). The oldest known pteropods are Paleocene in age, but only two species are known. One is Limacina mercinensis (Watelet and Lefévre 1885), and its earliest occurrence is latest Paleocene (Janssen and King 1988; Janssen 1991). It has been reported from Paleocene rocks of Alabama (Tracey et al. 1993), England, and Denmark (Janssen and King 1988). The other species is Limacina advenulata (Darragh 1997), from Victoria, Australia. Whether this species is of early or late Paleocene age has not been determined. Other fossil species of Limacina have been reported from Eocene rocks of the southeastern United States (see Hodgkinson et al. 1992 for a modern summary), England and France (Curry 1965, 1981), the Ukraine (Korobkov 1966; Bielokrys 1997), and New Zealand (Maxwell 1992); Eocene/Oligocene rocks of Mississippi (MacNeil and Dockery 1984; Dockery and Zumwalt 1986), Germany (Koenen 80 LIMACINA FROM THE PACIFIC COAST OF NORTH AMERICA 81 1892), and Australia (Janssen 1989a); Oligocene rocks of Denmark (Janssen 1990b); Oligocene/Miocene rocks of the North Sea (Janssen 1989b) and Australia (Janssen 1989a); Miocene of Poland (Janssen and Zorn 1993), other areas in Europe (see Janssen 1984 for a modern summary), Mexico, and the Dominican Republic (Collins 1934); Pliocene rocks of Japan (Ujihara 1996); and Jamaica (Janssen 1998). The fossil record indicates that Limacina was confined to warm waters and had widespread geographic dispersal during the Eocene and early to middle Miocene. The dispersal of this genus to present-day, worldwide distribution (tropical to arctic and antarctic waters) took place relatively recently, probably during the Pleistocene and possibly during the Pliocene. Several modern species of Limacina are cosmopolitan (Bernasconi and Robba 1982). There are few reports of Tertiary pteropods from the Pacific coast of North America (Collins 1934). The only detailed account is by Squires (1989), who reported three species in two genera, Praehyalocylis and Clio, from upper Eocene to middle Miocene rocks in Oregon and Washington. Specimens representing a third genus, Limacina, are the subject of the present investigation. They are of latest Eocene to early Miocene in age and represent the first known fossil record of genus Limacina from the Pacific coast of North America and the northernmost fossil occurrence in North America for this genus. Today, there are five species of Limacina found in the northeastern Pacific Ocean, and their geographic distri- butions are given by McGowan (1968). Only one of these species, Limacina helicina (Phipps), is found primarily north of Point Conception, California. The other species are found in southern California and/or Baja California, Mexico. The pteropod specimens described here, as well as the associated fauna, are reposited in the Natural History Museum of Los Angeles County, Invertebrate Paleontology Section (abbreviated LACMIP). Systematic Paleontology Class Gastropoda Cuvier, 1797 Order Thecosomata Blainville, 1824 Suborder Euthecosomata Meisenheimer, 1905 Family Limacinidae Gray, 1847 [=Spiratellidae Dall, 1921] Genus Limacina Bosc, 1817 Type species.—Clio helicina Phipps, 1774, by monotypy; Recent, polar seas, North Atlantic, and Pacific coast of North America. Remarks.—There has been considerable inconsistency and confusion in the literature regarding the proper generic name to use for this group of pteropods. As reported by Janssen (1989a) and Janssen and Zorn (1993), biologists usually use the name Limacina, but paleontologists commonly use the name Spiratella Blainville 1817. Both genera were named in the same month (December) of 1817, and both have the same type species. Curry (1981) reported that the name Spir- atella was registered in the archives of the “‘Bibliothéque Nationale de Paris” before the name Limacina was registered, and some workers (e.g., Curry 1981; Maxwell 1992) have argued that the name Spiratella should be used in preference to the name Limacina. There has not been uniform acceptance of this informal conclusion (e.g., Janssen 1989a; Ujihara 1996), and it is apparent that an appli- cation to the International Commission on Zoological Nomenclature (abbreviated 82 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES \n/ Seattle WASHINGTON 50 km 122° Fig. 1. Index map to LACMIP localities where fossil Limacina spp. have been found in Wash- ington. ICZN) is necessary for a formal decision of name priority. To date, no application has been made, but one will be made in the near future by A. W. Janssen (pers. comm.). Pending the ICZN decision, we have subjectively decided, like Bé and Gilmer (1977) and Janssen (1989a), to use the name Limacina because it is more frequently used than Spiratella. In addition, usage of the name Limacina will help bring the work of paleontologists and biologists in line. Adding to the confusion around the synonymous taxa Limacina and Spiratella, in older papers (e.g., Kittl 1886) the name Spirialis Eydoux and Souleyet 1840 has been used (at least, in part) for this same group of pteropods. For a full synonymy of genus Limacina, see Spoel (1967:36). Traditionally, Limacina is the only genus recognized within this family. At- tempts to split the genus into three subgenera have not received wide acceptance because the boundaries between the subgenera are unclear (Bielokrys 1997) and the fossil forms were not fully treated (Janssen 1989a). Limacina is characterized by a sinistrally coiled shell (anatomically dextral) that can be conispiral (the term ‘“‘trochoid”’ is used by some authors) or more flattened and involute. Characters such as relative whorl height, aperture outline or aperture elongation, and development of the umbilicus are quite variable and can coincide in different species (Bielokrys 1997). In order to differentiate spe- cies, it is critical to have the entire adult teleoconch preserved. LIMACINA FROM THE PACIFIC COAST OF NORTH AMERICA 83 Limacina sp. 1 Figures 2—5 Description.—Shell very small (height 0.7 to 0.83 mm), smooth, sinistral, with up to five whorls, rather quickly increasing in diameter; shell is 1.7 times wider than high. Apical side of the body whorl flat, with the spire barely protruding. Protoconch smooth. Aperture oval, slightly opisthocline, periphery seemingly evenly rounded; outer lip broken but apparently with a slight flare. Umbilicus about 20 percent of shell diameter. Large specimens with faint axial and spiral threads on body whorl. Remarks.—More complete specimens of Limacina sp. 1 are needed before a species identification can be made. In terms of the nearly flattened spire, shape of the body whorl, and relative width of the umbilicus, Limacina sp. 1 is very similar to Limacina atypica (Laws 1944; Janssen 1989a:7-8, pl. 1, figs. 1—2; pl. 10, figs. 1-3) from ?upper Oligocene to lower Miocene rocks in Australia and from Miocene rocks in New Zealand. Limacina sp. 1 differs from L. atypica by having a slightly wider umbilicus (20 percent of the shell diameter rather than 15 percent). In addition, although the apertural periphery is not completely preserved on Limacina sp. 1, the specimens seem to differ from L. atypica by having a rounded anterior part of the aperture rather than a projected anterior part. Material.—Thirty-five specimens, including hypotypes LACMIP 12722 and 12723. Age.—Latest Eocene to early Miocene. Stratigraphic occurrence.—Three specimens of Limacina sp. 1 were found in limestone at LACMIP loc. 5802 (Bear River deposit). The limestone, which is late Eocene in age and temporally equivalent to the lower part of the Lincoln Creek Formation (Squires and Goedert 1991), is rich in mollusks and formed in a deep-marine chemosynthetic environment (Goedert and Squires 1990). Limacina sp. 1 was found in two concretions at LACMIP loc. 17102. One of the concretions contained the following: 24 specimens of Limacina sp. 1, a few internal molds of an unidentifiable triangular-shaped pteropod, some specimens of minute gastropods (including the opisthobranchs Tornatellaea? sp. and Sca- phander? sp.), the bivalve Delectopecten sp., minute bivalves, a cheliped of the crab Portunites triangulum Rathbun, fish scales, and carbonized-wood fragments. The other concretion contained a single specimen of Limacina sp. 1 and a spec- imen of the nautiloid Aturia angustata (Conrad). In the gray siltstone surrounding the concretions, specimens of the gastropod Turritella oregonensis (Conrad), the turrid gastropod Spirotropsis? sp., Delectopecten sp., the bivalve Cyclocardia sp.., Aturia angustata, and Portunites triangulum were moderately common. Turritella oregonensis ranges from early to middle Miocene (Addicott 1976). Portunites triangulum ranges from latest Eocene to latest Oligocene or early Miocene, but it is most common in rocks of late Eocene to early Oligocene age (R. Berglund person. commun.). Aturia angustata ranges from latest Eocene to early Miocene (Armentrout 1973). Based on the overlapping ranges of these three species, the rocks at locality 17102 are of early Miocene age. Locality 17102 plots in the Lincoln Creek Formation on the geologic map of Pease and Hoover (1957). This formation ranges in age from late Eocene to early Miocene (Moore 1984), and the rocks at locality 17102 are assignable to the upper part of the formation. 84 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES LIMACINA FROM THE PACIFIC COAST OF NORTH AMERICA 85 The association at locality 17102 of pteropods, other small gastropods, small bivalves (including the “‘mud pecten’’ Delectopecten), crab remains, fish scales, and carbonized wood is very similar to other pteropod-bearing bathyal (greater than 200 m depth) assemblages found elsewhere in Tertiary formations of Wash- ington and Oregon. These other bathyal assemblages commonly are a mixture of faunal remains derived from pelagic communities (pteropods, fish scales) and from nearby nonmarine communities (wood) (Squires 1989). The nautiloid at locality 17102 is a pelagic component, and the Turritella specimens probably represent a shallow-marine component that underwent post-mortem transport by turbidity currents. Five specimens of Limacina sp. 1 were found in a concretion in the Lincoln Creek Formation at LACMIP locality 17105, which is stratigraphically and pa- leoenvironmentally equal to locality 17102. Also in the concretion were some specimens of the scaphopod Fustiaria? sp., a few unidentified turrids, Delecto- pecten sp., and a crab fragment. A single specimen of Limacina sp. 1 was found in a concretion at LACMIP locality 8232, and the concretion also contained a specimen of the scaphopod Dentalium sp., minute bivalves, and wood fragments. This locality is in the lower Oligocene undifferentiated part of the Makah Formation, and this part of the formation, like the rest of the Makah Formation, was deposited in a predominantly lower to middle bathyal environment (Snavely et al. 1980). A single specimen of Limacina sp. 1 was found in deep-marine rocks in the lower Oligocene Jansen Creek Member of the Makah Formation at LACMIP loc. 17101, in association with Limacina sp. 2. Limacina sp. 2 Figures 6—8 Description.—Shell minute (up to 5 mm in height), conispiral (naticiform), slightly higher than wide, sinistral, 4 to 5 whorls, spire low to moderately low; shell smooth except for regularly spaced sinuous, shallow axial grooves (growth lines?); suture moderately impressed, protoconch smooth. Aperture large, outer lip mostly incomplete, columella missing. Umbilicus area mostly missing, appar- ently narrow. Remarks.—More complete specimens of Limacina sp. 2 are needed before a species identification can be made. In terms of the relatively low spire, subglobose body whorl, relative height of the aperture versus the height and width of the body whorl, and the apparently very narrow umbilicus, Limacina sp. 2 has the most similarity to some specimens of Limacina pygmaea (Lamarck 1804:30; Wa- telet and Lefévre 1885:101, pl. 5, figs. 3a—c; Curry 1965:362, figs. 18a, 18b, 19; o Figs. 2-8. SEM micrographs of fossil Limacina spp. from Washington; 2—5, Limacina sp. 1, LACMIP loc. 17102, lower Miocene part of the Lincoln Creek Formation, Washington; 2—4, hypotype LACMIP 12722, X50, height 0.83 mm; 2, apertural view; 3, abapertural view; 4, apical view; 5, hypotype LACMIP 12723, 500, maximum diagonal distance 0.14 mm, apical view showing proto- conch; 6-8, Limacina sp. 2, LACMIP loc. 17101, lower Oligocene Jansen Creek Member of the Makah Formation, Washington; 6—7, hypotype LACMIP 12724, 35, height 1.9 mm; 6, apertural view; 7, apical view; 8, hypotype LACMIP 12725, abapertural view, X34, height 1.6 mm. 86 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Curry 1981:37, pl. 1, figs. 3a, 3b; Hodgkinson et al. 1992:19, pl. 3, figs. 14, 15) from middle Eocene (Lutetian Stage) rocks of Paris Basin, France, England, and Texas. Limacina pygmaea has considerable morphologic variation in the relative spire height, from flush to nearly half of the height of the shell. Because of this variation, some authors have recognized subspecies of L. pygmaea. Limacina pygmaea bernayi (Laubriére 1881:377, pl. 8, fig. 5; Cossmann and Pissarro 1910— 1913:pl. 60, figs. 1-2 of the pteropods) from middle Eocene (Lutetian Stage) rocks of France is one of these subspecies, and it has a relatively low spire and can have axial grooves. In these two features, it is closely similar to the specimens of Limacina sp. 2. Limacina sp. 2 also has some close similarity to Limacina nemoris (Curry 1965: 362, figs. 16a—b; Curry 1981:37, pl. 1, figs. 5a, b; Hodgkinson et al. 1991:18, pl. 3, figs. 9, 10) from the middle Eocene upper Bracklesham Beds of southern England, the upper Eocene “‘marnes bleues”’ of France, the upper Eocene Stone City and Cook Mountain formations of eastern Texas, and the upper Eocene Lisbon Formation of Alabama. Limacina sp. 2 does not have the inclined suture that can be present on some specimens of L. nemoris. Material.—Twelve specimens, including hypotypes LACMIP 12724 and 12725, Age.—Early Oligocene. Stratigraphic occurrence.—The specimens were found in a concretion at LAC- MIP locality 17101, and they were in the matrix that surrounded fragments of a small squat crab (Munida? sp., R. Berglund pers. comm.). This locality is in the lower Oligocene Jansen Creek Member of the Makah Formation, and this member represents a transported olistostromal rock unit containing mostly shallow-water marine conglomerate and fossiliferous sandstone enclosed in deep-water (1,000 to 2,000 m) marine siltstone and sandstone (Snavely et al. 1980; Squires and Goedert 1994). Acknowledgments We thank Ross E. Berglund (Bainbridge Island, Washington) for sharing his considerable knowledge of fossil crabs. We thank A. W. Janssen (Gozo, Malta) and R. R. Seapy (Department of Biology, California State University, Fullerton) for their opinions on the status of Limacina versus Spiratella. Arie W. Janssen also provided many, very useful reprints, as well as comparative specimens of representative European Paleogene pteropods. Lindsey T. Groves (LACMIP) ob- tained some important literature. Literature Cited Addicott, W. O. 1976. Neogene molluscan stages of Oregon and Washington. Pp. 95-115 in The Neogene Symposium (eds. A. E. Fritsche, H. TerBest Jr., and W. W. Wornardt). Pac. Sec. Soc. Econ. Mineral. Paleont., Los Angeles. Armentrout, J. M. 1973. Molluscan paleontology and biostratigraphy of the Lincoln Creek Formation, late Eocene-Oligocene, southwestern Washington. Ph.D. dissertation, University of Washington, 479 pp. Bandel, K., A. A.-Labin, C. Hemleben, and W. G. Deuser. 1984. The conch of Limacina and Peraclis (Pteropoda) and a model for the evolution of planktonic gastropods. Neues Jahr. Geol. Palaont. Abhand., 168(1):87—107. Bé, A. W. H., and R. W. Gilmer. 1977. A zoogeographic and taxonomic review of euthecosomatous LIMACINA FROM THE PACIFIC COAST OF NORTH AMERICA 87 Pteropoda. Pp. 733-808 in Oceanic Micropaleontology. (A. T. S. Ramsay, ed.). Academic Press, Vol? 1, Chpt.6: Bernasconi, M. P., and E. Robba. 1982. The thecosomatous pteropods: a contribution toward the Cenozoic Tethyan paleobiogeography. Boll. Soc. Paleont. Ital., 21(2—3):211—222. Bielokrys, L. S. 1997. Pteropod gastropods from the Eocene of the Ukraine. Paleont. J., 31(4):356— BGS: Blainville, M. H. 1817. Dictionnaire des Sciences Naturelles. Vol. 7. Levrault and Strasbourg: Paris, 534 pp. Bosc, L. A. G. 1816-1817. Nouveau dictionnaire d’ Histoire nauturelle. Deterville, Paris, Vol. 7, 586 pp. Collins, R. L. 1934. A monograph of the American Tertiary pteropod mollusks. Johns Hopkins Univ. Stud. Geol., 11:137—234. Cossmann, A. E. M., and G. Pissarro. 1910-1913. Iconographie completé des coquilles fossiles de l’Eocéne des environs de Paris. Société Géologique de France. Vol. 2 (Gastropodes, etc.). Paris, 65 pls. Curry, D. 1965. The English Palaeogene pteropods. Proc. Malacol. Soc. London, 36:357—371. . 1981 [1982]. Ptéropodes Edceénes de la Tuilerie de Gan (Pyrénées-Atlantiques) et de quelques autres localités du SW de la France. Cahiers Micropaléont., 4(1981):35—44. [English summary]. Dall, W. H. 1921. Summary of the marine shell-bearing mollusks of the north west coast of America, from San Diego, California to the polar sea, mostly contained in the collection of the United States National Museum, with illustrations of hitherto unfigured species. Smithson. Inst., U.S. Natl. Mus. Bull., 112:1—217. Darragh, T. A. 1997. Gastropoda, Scaphopoda, Cephalopoda and new Bivalvia of the Paleocene Pebble Point Formation, Victoria, Australia. Proc. Roy. Soc. Victoria, 109(1):57—108. Dockery, D. T., HI, and G. S. Zumwalt. 1986. Pteropods (Mollusca: Gastropoda) from the upper Yazoo Formation (Eocene) at Cynthia, Mississippi. Miss. Geol., 6(4):9-12. Eydoux, F, and FL. A. Souleyet. 1840. Description sommaire de plusieurs ptéropodes nouveaux ou imparfaitement connus, destinés a étre publiés dans le voyage de la Bonite. Rev. Zool. Soc. Cuvier, Paris 3:235—239. Goedert, J. L., and R. L. Squires. 1990. Eocene deep-sea communities in localized limestones formed by subduction-related methane seeps, southwestern Washington. Geology, 18:1182—1185. Gray, J. E. 1847. A list of the genera of Recent Mollusca, their synonyma and types. Proc. Zool. Soc. London, pt. 15:129—219. Hodgkinson, K. A., C. L. Garvie, and A. W. H. Bé. 1992. Eocene Euthecosomatous Pteropoda (Gas- tropoda) of the Gulf and eastern coasts of North America. Bull. Amer. Paleont., 103(341):1—62. Janssen, A. W. 1984. Type specimens of pteropod species (Mollusca, Gastropoda) described by Rolle (1861), Reuss (1867) and Kittl (1886), kept in the collection of the Naturhistorisches Museum at Vienna. Meded. Werkgr. Tert. Kwartaire Geol., 21(2):61—91. . 1989a. Pteropoda (Gasteropoda, Euthecosomata) from the Australian Cenozoic. Scripta Geo- logica, 91:1—76, pls. 1-13. . 1989b. Some new pteropod species from the North Sea basin Cainozoic (Mollusca: Gastro- poda, Euthecosomata). Meded. Werkgr. Tert. Kwartaire Geol., 26(3):91—133. . 1990a. Long distance correlation of Cainozoic deposits by means of planktonic gastropods (“‘pteropods’’); some examples of future possibilities. Tert. Res., 11(2—4):65—72. . 1990b. Pteropod species (Mollusca, Gastropoda, Euthecomsomata) from the late Oligocene of Mogenstrup, Jylland, Denmark. Contrib. Tert. Quat. Geol., 27(2—3):83-91. . 1991. Biostratigraphic application of ‘Pteropoda’ (Gastropoda, Euthecosomata) in Cainozoic deposits of the North Sea basin and interregional correlations. Proc. Tenth Internatl. Malacol. Congr. (Tiibingen, 1989):489—492. . 1998. Holoplanktonic Mollusca (Gastropoda: Heteropoda and Thecosomata) from the Plio- cene Bowden beds, Jamaica. Contr. Tert. Quatern. Geol. 35(1—4):95—-111. , and C. King. 1988. Planktonic molluscs (pteropods). Pp. 356—368 in The northwest European Tertiary basin. Results of the International Geological Correlation Programme Project No. 124 (eds. R. Vinken et al.). Geolog. Jahr., Reihe A, Heft 100. , and I. Zorn. 1993. Revision of middle Miocene holoplanktonic gastropods from Poland, published by the late Wilhelm Krach. Scripta Geologica, Special Issue, 2:155—236. Kittl, E. 1886. Ueber die Miocenen Pteropoden von Oesterreich-Ungarn. Ann. K. K. Naturhist. Hof- mus., 1:47—74. 88 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Koenen, A. Von. 1892. Das Norddeutsche Unter-Oligocin und seine Mollusken-Fauna. IV. Abhand. Geolog. Special. Preus. Thiiring. Staaten, 6:819—1004. Korobkoy, I. A. 1966. Pteropoda from Paleogene deposits in the south of the USSR. Voprosy Paleont., 5:71—92. [In Russian. ] Lamarck, J. B. 1804. Mémoires sur les fossiles des environs de Paris. Ann. Mus. Natl. Hist. Nat., Paris, Vol. 5, variously paged. (Reprinted 1978 by Paleont. Res. Inst., Ithaca, New York). Laubriere, L. P. de. 1881. Description d’espéces nouvelles du Bassin de Paris. Bull. Soc. France, Paris, 3:377-384. Laws, C. R. 1944. The molluscan faunule at Pakaurangi Point, Kaipara, 3. Trans. Proc. Roy. Soc. New Zealand, 73(4):297-312. MacNeil, F S., and D. T. Dockery, III. 1984. Lower Oligocene Gastropoda, Scaphopoda, and Ceph- alopoda of the Vicksburg Group in Mississippi. Miss. Dept. Nat. Res., Bur. Geol., Bull., 124: 1-415. McGowan, J. A. 1968. Thecosomata and Gymnosomata. The Veliger, 3 (Supplement): 103-135. Maxwell, P. I. 1992. Eocene Mollusca from the vicinity of McCulloch’s Bridge, Waihao River, South Canterbury, New Zealand: paleoecology and systematics. New Zeal. Geol. Surv. Paleontol. Bull., 65:1—280. Moore, E. J. 1984. Molluscan paleontology and biostratigraphy of the lower Miocene upper part of the Lincoln Creek Formation in southwestern Washington. Nat. Hist. Mus. Los Angeles Co., Contrib. in Sci., 351:1—42. Pease, M. H., Jr., and L. Hoover. 1957. Geology of the Doty-Minot Peak area, Washington. U.S. Geol. Surv., Map OM 188. Phipps, C. J. 1774. A voyage towards the North Pole undertaken by his Majesty’s Command 1773. W. Bowyer and J. Nichols, London, 275 pp. Snavely, P. D., Jr, A. R. Niem, N. S. Macleod, J. E. Pearl, and W. W. Rau. 1980. Makah Formation— a deep-marginal-basin sequence of late Eocene and Oligocene age in the northwestern Olympic Peninsula, Washington. U.S. Geol. Surv. Prof. Pap., 1162—B:1—28. Spoel, S. Van Der. 1967. Euthecosomata, a group with remarkable developmental stages (Gastropoda, Pteropoda). J. Noorduijn Zoon, Gorinchem, 375 pp. Squires, R. L. 1989. Pteropods (Mollusca: Gastropoda) from Tertiary formations of Washington and Oregon. J. Paleont., 63:443—448. , and J. L. Goedert. 1991. New late Eocene mollusks from localized limestone deposits formed by subduction-related methane seeps, southwestern Washington. Jour. Paleo., 65(3):412—416. , and J. L. Goedert. 1994. A new species of the volutid gastropod Fulgoraria (Musashia) from the Oligocene of Washington. The Veliger 37(4):400—409. Tesch, J. J. 1946. The thecosomatous pteropods. I. The Atlantic. Dana Report 5(28):1—82. . 1948. The thecosomatous pteropods. II. The Indo-Pacific. Dana Report 5(30):1—45. Tracey, S., J. A. Todd, and D. H. Erwin. 1993. Mollusca: Gastropoda. PP. 131—167 in The Fossil Record 2. (M. J. Benton, ed.). Chapman and Hall, London, 845 pp. Ujihara, A. 1996. Pteropods (Mollusca, Gastropoda) from the Pliocene Miyazaki Group, Miyazaki Prefecture, Japan. J. Paleont., 70:771-788. Watelet, A., and T. Lefévre. 1885. Note sur les pteropodes du genre Spirialis decouverts dans le Bassin de Paris. Soc. Royal Malacol. Belg., 15:100—103. Accepted for publication 29 September 1998. Appendix Localities LACMIP 5802. “Bear River deposit”. In an abandoned quarry on the south side of Bear River in the SE %, SE % of section 20, T. 10 N, R. 10 W, U.S. Geological Survey 15-minute Chinook Quad- rangle, 1949 (photorevised 1984), Pacific County, Washington. Strata temporally equivalent to the lower part of the Lincoln Creek Formation. Collectors: J. L. and G. H. Goedert, May 30, 1998. LACMIP 8232. Float from beach terrace exposures approximately 2000 m northwest of the mouth of the Sekiu River, near center of SW % NW % sec. 5, T. 32 N, R. 13 W, U.S. Geological Survey 7.5-minute Sekiu River Quadrangle, 1984 (provisional edition), south shore of Strait of Juan de Fuca, Clallam County, Washington. Lower Oligocene part of the Makah Formation. Collector: J. L. Goedert, April 30, 1998. LACMIP 17101. Approximately 320 m northwest of the mouth of Jansen Creek, near center of sec. LIMACINA FROM THE PACIFIC COAST OF NORTH AMERICA 89 26, T. 33 N, R. 14 W, U.S. Geological Survey 7.5-minute Sekiu River Quadrangle, 1984 (provisional edition), south shore of Strait of Juan de Fuca, Clallam County, Washington. Lower Oligocene, Jansen Creek Member of the Makah Formation. Collector: J. L. Goedert, March, 1997. LACMIP 17102. Siltstone 2 m below a hardground with reworked concretions, on the east side of a logging road, approximately 360 m north and 230 m west of the southwest corner of sec. 31, T. 17 N, R. 6 W, U.S. Geological Survey 7.5-minute South Elma Quadrangle, 1986 (provisional edition, minor revisions 1993), Grays Harbor County, Washington. Lower Miocene part of the Lincoln Creek Formation. Collectors: J. L. Goedert, G. H. Goedert, and E. Z. Nordlander, Spring, 1998. LACMIP 17105. Concretion 1 m below a 4 cm-thick hardground (with reworked crab fossils) on steep slope on north side of ridge, approximately 720 m east and 480 m north of the southwest corner of sec. 28, T. 17 N, R. 6 W, U.S. Geological Survey 7.5-minute South Elma Quadrangle, 1986 (provisional edition, minor revisions 1993), Grays Harbor County, Washington. Lower Miocene part of the Lincoln Creek Formation. Collectors: J. L. and G. H. Goedert, May 23, 1998. SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 1999 ANNUAL MEETING APRIL 30-MAY 1, 1999 CALIFORNIA STATE UNIVERSITY DOMINGUEZ HILLS The Annual Meeting of the Southern California Academy of Sciences was held April 30—May 1 at California State University, Dominguez Hills. Student Award winners were as follows. Margaret Barber Award for Best Paper Steve I. Lonhart, Dept. of Biology, University of California, Santa Cruz Feeding Preferences of Invasive and Native Turban Snail Predators Jules Crane Award for Best Paper Chugey Sepulveda, Dept. of Biological Science, California State University, Fullerton Are Tunas Faster or more Efficient Swimmers as a Result of Endothermay? Best Poster Award Gloria Jean Baca, Dept. of Biological Science, California State University, Fullerton The Effects of Lateral Line Development on the Escape Response (C-Start) of the California Halibut, Paralichthyes californicus. (Coauthors A. C. Gibb and K. A. Dickson) Best Poster Award Kristina D. Louis, Dept. of Organismic Biology, Ecology and Evolution, U.C.L.A. An Investigation of the Subspecies of the Staghnorn Sculpin, Leptocottus armatus (Scor- paeniformes: Cottidae) (coauthors, A. Y. Han and D. K. Jacobs) Honorable Mention Kristin M. Ward, Pacific Estuarine Research Laboratory of San Diego State Univ. Episodic Colonization of an Intertidal Mudfiat by Cordgrass at Tijuana Estuary (Coauthors, J. C. Callaway and J. B. Zedler) NEXT MEETINGS MAY 5-6, 2000 at the University of Southern California ° MAY 5-6, 2001 at California State University, Los Angeles The Southern California Junior Academy of Sciences met concurrently with the Academy. Twenty-six students presented oral presentations and twenty-one papers were turned in for judging. On the basis of their oral presentations and written papers, the following students were selected to attend the Na- tional Meeting of the American Junior Academys of Science, to be held in Washington D.C. Feb. 16— 20, 2000 in conjunction with the annual meeting of A.A.A.S. Daniel Chen, Villa Park High School. Mentor: Ping Wang, U.C. Irvine Insulin-Like Growth Factor I Retards Apoptotic Signaling induced by Ethanol in Cardiomy- ocytes. Nathan Fleischaker, La Costa Canyon High School. Mentor: David Emmerson, La Costa Development of an Anodic Stripping Voltammetry Electrode for Measurement of Heavy Metal Ions and its use in Monitoring Clean-up of Contaminated Soils Sophia Tran, Alhambra High School. Mentor: Donn Gorsline, U.S.C. Clay Mineral Groups in the Gulf of California Michael Wang, Troy High School. Mentor: Christian Lytle, U.C. Riverside Immunolocalization of NA-K-2Cl Cotransporter and Cl/HCO, Exchange Proteins in Human Stomach. Elizabeth Williams, Palos Verdes High School, Mentor: Irving Biederman, U.S.C. Perceived Lightness as a Measure of Perceptual Grouping Terry Yen, West High School, Torrance. Mentor: Elisheva Goldstein, Calif. Poly. Univ. Pomona. 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Ortega-Salas The Euryhaline Gobiid Fish, Gillichthys mirabilis Cooper 1864, Second Intermediate Host of the Trematode, Pygiopsoides spindalis Martin 1951. Mark H. Armitage First Fossil Record of the Pteropod Limacina from the Pacific Coast of North America. Richard L. Squires, James L. Goedert and Steven R. Benham COVER: Detail of Ascocotyle sexidigita excysted metacercaria from Fundulus par- vipinnis. Scale bar = 20 micrometers. ISSN 0038-3872 meee RN CALIFORNIA ACADEMY OF SCIENCES BOLLETIN Volume 98 Number 3 BCAS-A98(3) 91-138 (1999) DECEMBER 1999 Southern California Academy of Sciences Founded 6 November 1891, incorporated 17 May 1907 © Southern California Academy of Sciences, 1999 OFFICERS David Huckaby, President Robert S. Grove, Vice-President Susan E. Yoder, Secretary Robert Lavenberg, Treasurer Daniel A. Guthrie, Editor Hans Bozler, Past President David Soltz, Past President BOARD OF DIRECTORS 1997-2000 1998-2001 1999-2002 Robert S. Grove Kathryn A. Dickson Ralph G. 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For further information on posters, abstracts, registration and deadlines, see the Southern California Academy of Science web page at: www.lam.mus.ca.us/~scas/ or contact gorsline @earth.usc.edu — 5 c - = 55a = ‘ . - i‘ ’ a wy - thes 4 yt adernry ot ni Sw LE ee Py WIAD Vaan L726 AAD ito 4 Hrs, OS —-@l valy | y : ‘ear we aries A ve : - oe ‘ge VROULEAD WASH Tie ; J Pa) = , ; wo) etanorert. iia at OmODIe- Mi asSNeouls MM ‘uarineetam ajo8 ‘wwsleal e< wet Sy. 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Sci. 98(3), 1999, pp. 91-102 © Southern California Academy of Sciences, 1999 Rare Species as Bioindicators in Marine Monitoring Don Maurer', Tom Gerlinger? and George Robertson? 'Department of Biological Sciences, California State University Long Beach, Long Beach, California 90840 ?County Sanitation Districts of Orange County, California, Fountain Valley, California 92708-7081 Abstract.—Soft-bottom invertebrates commonly form the core of marine moni- toring studies. Accordingly, those species which occur frequently and abundantly are favored in analyses, and rare species are normally excluded from consider- ation. Since rare species commonly occur at their geographic limits under sub- optimal conditions, it has been postulated that they would be vulnerable to natural and anthropogenic stresses. We examined the potential of rare species as bioin- dicators in monitoring a major marine outfall on the San Pedro Shelf, California. A rare species was operationally defined as occurring only once per sample (0.1m? Van Veen Grab). Based on 780 quantitative benthic samples, distributed over thirteen 60 m (outfall depth) stations, and 12 years, we concluded that the number of rare species generally declined towards outfall stations. We suggested that a more conventional definition of rarity based on actual geographic range might provide an increased degree of sensitivity to identify bioindicators. Soft-bottom macroinvertebrates commonly represent the basis of many marine monitoring programs (Bilyard 1987). In turn, the enumeration of macrobenthos depends on accurate identification (Brinkhurst 1980) prior to being quantitatively analyzed for measures of community structure (number of species, abundance, biomass, diversity, species composition) (Diener et al. 1995). Species richness is a focus of many ecological theories and applications (Cao et al. 1998). In addition, identification and enumeration of various species frequently reveal taxa as can- didate bioindicators. The latter species are sensitive to a contaminant gradient by increasing or decreasing in abundance (Pearson and Rosenberg 1978). Bioindi- cators have been commonly used as major tools in monitoring programs (Wilson 1994). We have found that using groups or suites of species is preferable to relying on a single or a few species as bioindicators. During the course of benthic surveys with a diverse fauna, a common practice among benthic ecologists is to exclude rare species from data analysis (Cao et al. 1998). There are several reasons for this. Species that occur frequently and abun- dantly dominate most of the functional processes on which an ecological com- munity depends (Main 1982). As a result, they are preferred candidates as dom- inant species and bioindicators. According to Cao et al. (1998) rare species are routinely deleted from data sets because many workers believe they contribute little to community analysis and add noise to statistical solutions. Moreover, ex- clusion of rare species from quantitative analyses normally simplifies data pro- 91 92 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES cessing, thereby reducing problems frequently encountered in the analysis of data matrices containing large numbers of zeros (Harvey et al. 1998). During its monitoring program the Districts has produced over 3000 quantita- tive samples yielding 2 million individuals and over 1200 species. Monitoring in Santa Monica Bay, California by the City of Los Angeles has produced about 2400 infaunal species (Dorsey et al. 1995). It is estimated that there are over 5000 marine benthic invertebrates in the Southern California Bight (SCB) (Thompson et al. 1993). Assuming the accuracy of this order of magnitude the SCB supports one of the richer bottom faunas in the world. This presents a huge task and challenge to practitioners analyzing data from the SCB. From the body of work on rare species (Rabinowitz 1981; Groves and Ride 1982) we have adopted the construct that species are rare at the limit of their geographical and/or physiological range, and as such they occur under suboptimal conditions (Hummel et al. 1995). Cao et al. (1998) cite literature asserting that small populations are more likely to go extinct, and therefore rare species should be more sensitive to disturbance than common ones. If the premise is valid, then rare species might be particularly vulnerable to stresses imposed by natural pro- cesses and anthropogenic activities. In this account we focus on frequency of occurrence in samples from the study area as a measure of rarity rather than actual geographic ranges of incumbent species. In any faunistic survey it is a common observation for a collection of species that some occur in relatively few samples and/or occur at relatively low abun- dances (Schoener 1987). The growth of a species or a population in its habitat is subjected to constraints (Grieg-Smith and Sagar 1981). If those constraints are relaxed population size may increase, if they are exercised, populations may de- crease or grow and develop more slowly. Causes of rarity are found by identifying the constraints on the potential rate at which the population size of the selected species can increase. Anthropogenic activities can provide constraints unrelated to natural processes. There is a considerable literature on rare species, rarity or rarities (Preston 1948, 1962, 1980; Harper 1981; Groves and Ride 1982; Schoener 1987; Prendergast et al. 1993; Boero 1994; Gaston 1995). Criteria for defining rarity may involve uniqueness, relict nature, frequency of occurrence, functional role in community dynamics, geographic range, habitat specificity, and local abundance (Rabinowitz 1981; Main 1982). For the present account rare species are operationally defined as those species occurring once per sample (0.1 m? Van Veen Grab). A practical purpose for monitoring is to ascertain the status of particular pop- ulations or species (Davy and Jeffries 1981). Successful management of a rare population, making it less rare or preventing its local extinction, implies the ability to manipulate the size and structure of that population. If monitoring is to con- tribute significantly to this ability, it must yield a predictive understanding of population structure and function. Accordingly, monitoring based on enumeration of species is most relevant to problems of rarity (Davy and Jeffries 1981; Cao et al. 1998). This argument provides support for the definition of rarity used herein. Boero’s (1994) discussion of fluctuations and variations in coastal marine en- vironments provides a useful point of departure. According to him the higher the number of rare species available to replace the dominant species, the higher the possibility for a community to successfully face important disturbances. Gray RARE SPECIES IN MARINE MONITORING 93 33.6° Districts' Outfall 40 3355" Los Angeles \* County Sanitation Districts LS of Orange County g,, Dio 118.1° 117.9° Fig. 1. Benthic sampling stations from the San Pedro Shelf, California. (1989) expressed the view that in most unstressed communities there are many rare species contributing to high species richness. Recalling our earlier character- ization of the relative vulnerability of rare species to natural and anthropogenic stresses compared to common species, the number of rare species might be ex- pected to decline towards an ocean outfall (Hummel et al. 1995; Cao et al. 1998). Moreover, stress may also be imposed by increased abundance of organisms due to organic enrichment. It occurred to one of the authors (TG), that if rare species are more responsive to stress than common ones, the former should be included on a more regular basis as a tool in marine monitoring. This would greatly elevate their ecological significance, and serve as a stimulus fostering studies of natural history and population dynamics of species generally ignored for this purpose. Accordingly, we examined the proposition that the number of rare species should be lower in the vicinity of an ocean outfall. Materials and Methods Field.—From 1985 to the present the Districts has collected benthic samples of sediment and macrobenthic invertebrates with a double yoked 0.1 m* Van Veen grab from an array of stations on the San Pedro Shelf, California (Fig. 1). Sam- pling stations were located by Loran C and/or GPS (+ or — 15 m) and samples were sieved through a 1.0 mm mesh screen which is the accepted mesh size in the SCB. Specimens were fixed in 10% buffered formalin and subsequently trans- ferred to 70% ethanol in preparation for laboratory analyses. From 1985 to the present the Districts has sampled annually (summer) 40 Stations (N = 1) extending from 30 m to 324 m, and thirteen 60 m stations (N 94 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES = 5) quarterly (winter, spring, summer, fall), totaling over 3000 quantitative ben- thic samples for this monitoring program. In the summer of 1994 the Districts participated in the SCB Pilot Project (Bergen et al. 1998) which precluded regular sampling, and so this year is deleted from quantitative analyses in the present account. Based on this extensive sampling effort, collections from the summer characteristically yield the highest number of macrobenthic species (Districts 1995). Moreover, the thirteen 60 m stations were established to permit quarterly sampling on the same isobath as the ocean outfall, thus reducing the effect of depth on variation of estimates of community measures. For purposes of this account we have focused on the thirteen 60 m stations sampled during the summer from 1985 to 1997. Analyses are based on 780 benthic samples. Taxonomic analyses.—From the 780 sample database a rare species was defined as one individual per 0.1 m?’ grab. Single occurrences were extracted from the database. To provide conservative estimates of rarity for this analysis we applied some degree of filtering for inclusion. Mysids, euphasusiids, pteropods, calanoid copepods, larvae and other plankton collected during sampling were excluded from counts together with fishes and their parasites. Since the analysis focuses on benthic species, inclusion of planktonic and nektonic species would be inappro- priate. Collected nematodes, harpacticoid copepods, and halacarids were also ex- cluded because the sampling gear and sieving protocols preclude accurate sam- pling of these small organisms. In addition, taxa with plural notations in the taxonomic listings (Macoma spp.) were not included because of uncertainty about the actual number of species represented by such a designation. The same convention was applied to a genus identified as a juvenile (Amphiodia sp. juvenile). Finally, higher taxonomic cat- egories (phylum, class, order, family, genus) were excluded from rare species designation unless listed with a species designation (Phoronid species B). Statistical analyses.—After the number of rare species was tabulated, they were ranked according to station. Since this account focused on the presence or absence of rare species, we used the non parametric Kruskal-Wallis test with ties, an analysis of variance by ranks to test whether there were any statistically significant (x = 0.05) differences in rarity across stations (Zar 1984). For the Kruskal-Wallis test Hy was that the number of ranked rare species did not differ across stations. When H, was rejected, a nonparametric multiple comparison was applied to de- termine between which of the stations significant differences occurred. For the latter test we assumed a one-tailed distribution. When the nonparametric multiple comparison indicated significant differences in ranked rare species among stations, we used the nonparametric Mann-Whitney (U) distribution (Zar 1984). For the Mann-Whitney distribution we compared the number of ranked rare species at a reference station (CON) with those from the Zone of Initial Dilution (ZID) stations (0, ZB, ZB2, 4). We also assumed a one- tailed distribution for these analyses. Results Number of rare species.—The average number of rare species per thirteen 60 m stations (1985—1997) was computed (Table 1). Examination of these data re- vealed the following trends. First, the average number of rare species increased from the 1980s (37-38) into the 1990s (47—48) (Fig. 2). Second, rare species 95 RARE SPECIES IN MARINE MONITORING WeIBO01g 10[Iq GOS r66l x S =N = uonejs ‘uoneiAsg plepurig = Cs ‘urs = X :puosoT LV C 8V OV v OV Cc OV Oise STE VLE OV 8e Ease CBE x ESM GCCe IC BCiCOm OC OSL = CL OL) ee oP SP = NL Cin One le AZ A vato)I| KO Gs. wey Cole WSS SOS es: See Sis ace Ore OCS “Gr 1859 . Ge B9ICC™ wo "Sco MGIIVS Sy LG “cr. way Ol BS 99 OS Set yiS: 9%" “Give 10'GN Sil ve E Ve Cree es Geer 9rCo™ Ui SiS OC AV CS Be Se aco i, Ore. VOI9Ge 9's PHEGe “OI9y UOTCE, OC Si SiLE | Ge SCV, 39 IE SIC OB VIC Vee? Cl OVE DE MOLPSIS 88 8S @ OV WSC es Ge 9G SOV av 9. Cee BG WeCe., 266), G9. “8 6y VOW “GO Cue NEC OMY, 6 Gi Re CL OOS NL Be BONG” Wy NS cr BG 2oUr 0h, COS SCC. 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OV OS Gye “Soe CayINOS ds x ds x ds x ds x ds x ds x ds x ds x ds x ds x ds x aS = su0n L66l 9661 S66! c661 C66! 1661 0661 686l 8861 L86l 986l C86l an ‘L661 01 SQ6I BIUIOJTTLD ‘JJOUS OIpog URS oY) WOIZ sUONRIs (JOUTUUNS) JenuUUR WE Q9 €] Wor sordads ory}UNg aIe1 JO JoquINU aseIOAY “T IQRL 96 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 60 ” o Oo 50 oO Qa ” o oe = re Oo © 30/ ig | = =) Says w fey) | [4] | = 2 10- < | 0 4 T; = = T i. = T = = CONS Ac 13 5 1 0 ZB2. ZB 4 9 12 37 C2 Stations (sequence not actual distance) -— = —— /—*'85 —#-'90 _+'97| l 2 = x J Fig. 2. Average number of rare benthic species from 13 60m stations on the San Pedro Shelf, CA 1985, 1990, and 1997. were always lowest at Station C2 located in the Newport Submarine Canyon (Table 1, Fig. 1—2). Third, rare species were generally higher at farfield stations (CON. C, 13, 37). Fourth, the average number of rare species was generally lower at ZID stations compared to other 60 m stations (Fig. 2). The average number of rare species from 1987 were plotted as representative of the data base (Fig. 3). The marked decline of rare species at C2 in the canyon and the saddle posed by the ZID stations is very clear. However, at various times as many as 57—58 rare 60 | 50 40 - 30 | 20 10 | Average number of rare species CON. S 13 5 1 O00. 0282-2 426 4 9 12 37 C2 Stations (sequence not actual distance) Fig. 3. Average number of rare benthic species from 13 60m stations on the San Pedro Shelf, CA 1987. *Error bar equals | standard deviation. RARE SPECIES IN MARINE MONITORING i) /) species were recorded from ZID stations. This attests to the richness and status of the macrobenthos from this portion of the San Pedro Shelf. Rank number of rare species.—The rank number of rare species was computed (Table 2). These data reflected the trends noted for the average number of rare species (Table 1). Station C2 always received the lowest ranks. Moreover, ranks were usually lower at the ZID stations (0, ZB, ZB2, 4) compared to the nearfield (1, 5, 9, 12) and farfield stations (CON, C, 13, 37) (Table 2, Figure 1). Kruskal-Wallis tests were performed on the ranks of rare species. Ranks were always significantly (« = 0.05) lower at Station C2 (Figure 1) compared to other 60 m stations. As a result, data from Station C2 were deleted from remaining analyses (Table 3). Based on the computed H, value compared to the tabled H, (Zar 1984), the null hypothesis (H, = there were no differences in the number of ranked rare species across stations) was rejected eight of twelve years. When Hy was accepted, critical values were very close for significance for three years cigs; 1987, 1996). For those years when H, was rejected a nonparametric comparison was applied to distinguish significant ranking of rare species among stations (Table 3). A major trend was that ranks of rare species from ZID stations were commonly grouped and were significantly (« = 0.05) lower than those at other 60 m stations (Table 3). To examine further the Hj, the Mann-Whitney (U) distribution was employed comparing ranks of rare species at reference Station CON with those at ZID stations (Table 4). For these comparisons the null hypothesis (H, = the number of ranked rare species is not higher at CON compared to ZID stations) was rejected 23/48 (47.9%), accepted 17/48 (35.4%), and received 8/48 (16.7%) ties (accept/reject) where computed U was equal to the tabled critical U (Table 4, Zar 1984). The average number of rare species and the rank number of rare species was generally lower at ZID stations than other 60 m stations (Table 1 and 2). Kruskal- Wallis analyses indicated that ranked rare species differed significantly across sta- tions (Table 3). Moreover, the nonparametric multiple comparisons indicated that ranked rare species at ZID stations commonly grouped together and were signifi- cantly lower than those from nearfield and farfield stations. Although not without exceptions, the Mann-Whitney distribution further indicated significant separation between CON and the ZID stations. The weight of evidence supports the propo- sition that the number of rare species was lower in proximity to the ocean outfall. Discussion Several trends emerged from this account. Earlier it was noted that the average number of rare species increased from 1985 through 1997 (Fig. 2). This might be interpreted as increased immigration and successful colonization of benthic species to the study area due to improved environmental conditions. Increased source control and upstream recycling together with reduced emission of total suspended solids to the ocean has markedly reduced concentrations of many trace metals and organic contaminants (Districts 1995). Thus, there is solid evidence to support the position that environmental conditions have improved around the outfall from 1985 through 1997. Regardless, the increased number of rare species with time is probably due to increased refinement in taxonomy as the monitoring program has matured. Since this pattern is distributed across all stations including the 40 annual ones (Fig. 1), it does not affect the results of the study. SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 98 ak Jag uonrig Jog Ssuryuey Jo wing = surly puosas Joaquin uoT}eIg = UIT ISIL4 :puasayq S'O09TC SVC S°9CC CSIC C°COC S681 Tat CLIT CPrg9l OOT S6L L9 SI 3) 6 EC él el NOO ¢ v I 0 cdZ aZ (G@) L66I 6LC CLVC GEC SCL Sél CCc8I C 1st C081 OSI C’'S6 S'6L CLV CLI Le e) cl NOO I ¢ 6 4 el 0) aZ CUaZ CO 9661 COC HEE S'OCC NG AIG S'10C C86! CL8I Ocl 98 C8 EE Sl 6 V cl Le J NOO ¢ I el 0) CcaZ aZ CO S66! C 16C O€T SIT LOC SCOT ¢ S6l Cc rel SM VCl EAL C'V6 S’6S cl jig ¢ el e) NODO 6 V aZ CaZ (4! I 0) CO 66! 697 19C ec C LCC SLCC 8LI vLI ELI 19] Ceri 1Ol g°S9 cl NOO e) 4 ¢ Le 6 (| I el CaZ aZ 0 (6@) C66! COL S9C C6CC OCC 961 (6S) 9SI CPrsl Sor! C'S8 C8 S°€8 SI NOO ¢ LEE 6 2) cl (Gi I V cdZ aZ 0 CO 1661 O87 CG LSC LEC LOC vol L81 SOI Ceol S'6SI vol cOl Sele C91 NOO [BS e) 0) el ¢ 6 I CaZ v aZ (Gs! (69) 0661 L8C S'O8T VLC C'6CC CSIC col O¢l CIll las S801 901 8L cle LE el O ¢ I NOD 0) cl V cdZ 6 aZ CO 6861 CSc 8VC Ive C'6CC SKE 881 S6LI esl oSel S'6cl Sccl 8e tc 2) LE NOD el 6 V g I cl 0) CaZ aZ CO 886l [Sc VET 6CC CCle C'CO0C c'r6l ¢'991 991 ccl C'8cl col S8 91 J NOO 6 cl ¢ Le V et I 0) CHAZ aZ CO L86l COC 8ETC 8ITC 90C Lol SsI Cesl SOI COSI 61 LOI v6 SI LC I ¢ (Giles NOO e) 6 0 4 cl aZ cdZ (6@) 9861 G°99TC GEE LCC £07 C'cél 881 Coil rot COSI 6C1 col SLO 61 eh cl ¢ NOO Le 6 CcdZ e) aZ 4 0) I CO S86l ‘L661 01 SQ6T BIUIOJITeD ‘J[aYS O1pag ue dy} WO (¢ = N) suONEIs (J9WIUINS) [enuUR WI QO ¢] Wor satoads oryyuaq ore Jo Joquinu yueY “7 IquL RARE SPECIES IN MARINE MONITORING 95 Table 3. Summary of Hy results (N = 12 stations) from Kruskal-Wallis analysis and nonparametric multiple comparison 1985-1997. Kruskal-Wallis = Se eee eee Nonparametric Multiple Comparison Year Estimated H, Tabled H, Decision Station Separation 1985 19:3 19.6 Accept 1986 15.6 Accept 1987 N9ES Accept 1988 Did Reject CMTS CON 1379 4°95 Pal2202ZB2 ZB 1989 42.7 Reject 37 13 Ce AICONTOmMI2 4 ZED 9 ZB 1990 26.1 Reject CON 37-C0MI355 9 ZB2 40 ZB 2 199] 81.1 Reject SI CONS *C 37) 13 IDA 2B? eZ BO E992 58.4 Reject CONG e439 oI IS OE Z Be Zoe 1993 S129 Reject 37% 5) 187.6. CON, 9 4028 >ZB2 1201 0 1995 22.0 Reject ANZ CC CON 5 els 0SZB25Z258 1296 93 Accept 997 20.4 Reject Cro 372 1ST CON D4 I 0eZB2Z5 Another trend cited earlier involved Station C2 (Table 1 and 2). Extensive sam- pling of benthic organisms, sediment properties, and sediment geochemistry has demonstrated that the physical oceanography and associated hydrodynamic regime of the Newport Canyon (Fig. 1—3) is dramatically different from other 60 m sta- tions. The hydrodynamic regime produced different conditions for sediment prop- erties and benthos alike (Districts 1995). Accordingly, it was not surprising that the average number and rank of rare species was significantly lower at Station C2 in the canyon (Fig. 2—3) and so its exclusion from further analyses was justified. Although not without exceptions, the principal trend from this study revealed a reduction of rare species in proximity to the outfall (Table 1—4, Fig. 2—3). This trend agrees with the work of other researchers (Gray 1989; Boero 1994; Cao et al. 1998) which suggests that the distribution and abundance of rare species might be useful in marine monitoring. In the present account a rare species was operationally defined as being col- lected once per 0.1 m* rather than adopting a conventional definition based on geographic distribution. The operational definition was used to entertain the con- Struct and to initiate discussion emphasizing the importance of including rare Species in analyses of marine monitoring. This approach also maximizes infor- mation rather than characteristically discarding presumably useless data. Exclu- 100 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 4. Summary of Mann-Whitney (U) distribution comparing rank order of rare species at Station CON with ZID Stations (0, ZB2, ZB, 4) 1985 to 1997. Year 0 ZB2 ZB 4 1985 21=21 L922 16<21 PRS Accept/Reject Accept Accept Accept 1986 15<21 Vs) Woo say || 21=21 [si5=271 Accept Reject Accept/Reject Accept 1987 20221 DED) 232i 18<21 Accept Reject Reject Accept 1988 21=21 21=21 2521 19<21 Accept/Reject Accept/Reject Reject Accept 1989 [ese 21=21 D289) 21=21 Accept Accept/Reject Reject Accept/Reject 1990 21=2 24>21 Desi 2527 Accept/Reject Reject Reject Reject 1991 DIDI 25221 752k 2521 Reject Reject Reject Reject 1992 2574 235-21 221 23. 2N Reject Reject Reject Reject 1993 Paar V ae 2021 18<21 16.7<21 Reject Accept Accept Accept 1995 21=21 BN S| 20521 Ph2 ik Accept/Reject Reject Accept Accept 1996 24.5221 De IA| 2A aN 18.4<21 Reject Reject Reject Accept 1997 205<2 1 PoE S| eI | 14.5<21 Accept Reject Reject Accept Ist number = Estimated Mann-Whitney U. 2nd number = Critical value of Mann-Whitney U distribution. Underline = statistically significant x = 0.005 (1), n = 5, n = 5. sion of rare species seriously violates general ecological observations and theory leading to unacceptable losses of ecological information (Cao et al. 1998). Al- though some of the species included in the analyses may be rare according to geographic convention, that is, they occur near or at the limit of their geographic range, we presently do not know how many of the species used herein fall into this category. The operational definition may be overly simplistic in framing the physiological condition of rare species, thus reducing their sensitivity to respond to the potential stress (abiotic or biotic) posed by the outfall. Cao et al. (1998) examined the response of rare species as defined by occur- rence frequency to an aquatic gradient of contaminants. They concluded that ex- clusion of rare species coupled with small sample size can influence comparison of species richness compromising the reliability and sensitivity of community analysis and bioassessment. Rare species are critical for accurate community stud- ies and bioassessment (Cao et al. 1998). An anonymous reviewer noted that this view contrasts with the practice of taxonomic sufficiency advocated by researchers from the U. S. EPA (Ferraro and Cole 1990, 1995). Taxonomic sufficiency involves identifying organisms only to the taxonomic level necessary to meet study objectives assuming no loss of sta- tistical rigor. According to its proponents the practice minimizes the time, cost, RARE SPECIES IN MARINE MONITORING 101 and error of taxonomic identification. Taxonomic sufficiency is a cost-benefit ex- ercise. Clearly an analysis focusing on rare species places a demand on accurate identification and is not consistent with the savings attributed to taxonomic suf- ficiency. While recognizing that opportunities for cost savings associated with long-term monitoring deserve serious consideration, there are a number of other implications to taxonomic sufficiency which have not been addressed, and which should be considered prior to implementing this as a regular monitoring protocol. We have prepared a fuller rebuttal in a manuscript in preparation. Harper (1981) placed emphasis on the role of pathogens controlling natural plant populations of rare species. By analogy effluent from the outfall could serve as a medium for invertebrate pathogens potentially exerting negative pressure on benthic populations. However, there is no evidence to support that the outfall serves as the basis of an epizootic condition for invertebrates (Districts 1995). We conclude that the definition of rarity used herein provides evidence for using rare species as bioindicators in analyses of marine monitoring. This assertion derives independent support from the work of Cao et al. (1998). Alternatively, rare species defined through geographic conventions may provide an increased degree of sensitivity for this purpose. Since polychaetes and amphipods provide diverse taxa for SCB soft-bottom communities, they would seem to be likely candidates for identifying rare species based on their geographic range. Notwith- standing the extensive taxonomic work on polychaetes and amphipods by Hart- man (1968, 1968) and Barnard (1969), respectively, new species of both taxa are still being identified by the Southern California Association of Marine Invertebrate Taxonomists. Thus, the geographic range of new species of polychaetes and am- phipods and existing ones remains to be determined. As a first start we suggest that molluscs with their extensively documented geographic ranges (Morris 1966; Abbott 1974) and ecologic importance to benthic communities would be a good candidate taxon to initiate refined analyses of rare species in marine monitoring. 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Sci. 98(3), 1999, pp. 103-118 © Southern California Academy of Sciences, 1999 Morphologic and Genetic Variation Among Six Populations of the Spotted Sand Bass, Paralabrax maculatofasciatus, from Southern California to the Upper Sea of Cortez Gregory J. Tranah' and Larry G. Allen Department of Biology, California State University, Northridge, Northridge, California 91330 Abstract.—Spotted sand bass, Paralabrax maculatofasciatus, were examined for 19 morphometric and 7 meristic characters to determine the extent of morphologic and genetic variation among six Pacific and Gulf of California populations. Poly- merase Chain Reaction (PCR) and Restriction Fragment Length Polymorphism (RFLP) techniques were used to amplify and cleave the ITS region of the rDNA in order to detect differences among these populations. The morphological and genetic lines of evidence demonstrate that significant differentiation has occurred between the most distant Gulf and Pacific regions. The three Baja California populations share significant morphological and genetic affinity and appear to be different from the three Pacific populations. There is no significant morphologic differentiation between the three Pacific populations although genetically they form two different groups. The northernmost Pacific population of San Diego is significantly different from the two southern-Pacific groups as well as the entire gulf sample. The results of this study indicate that geographically isolated pop- ulations of nearshore marine fishes, under the influence of strong selection pres- sures, do not require long periods of time for divergence. The last 15,000 years may have been sufficient to allow significant divergence to occur between upper Gulf and Pacific groups of the spotted sand bass. The spotted sand bass, Paralabrax maculatofasciatus, is a serranid native to the eastern Pacific Ocean, having a historical range of San Francisco Bay south to Mazatlan, Mexico. Spotted sand bass rarely occur north of Santa Monica Bay (Love 1991) and are exceedingly rare in the warmer tropical waters of the south- erm cape region at Cabo San Lucas (Thomson and McKibbin 1976). Dense pop- ulations occur in the northern Gulf of California and also in the southern Cali- fornia region where they are restricted to shallow, warm-water back bays and harbors and the protected outer coast which provide warm-water refuges for this largely subtropical species (Fitch and Lavenberg 1975; Allen et al. 1995a). Tag- ging studies conducted by Allen (pers. obs.) throughout southern California have shown that spotted sand bass are nonmigratory and remain in their distinct region of settlement. This species leads a relatively sedentary lifestyle in association with a particular bay, harbor, or open coast rock relief structure. Gene flow and dis- persal are facilitated through the long distance movement of eggs and larvae by ocean currents, and settlement from the plankton occurs at approximately 21 days ' Present Address: Department of Animal Science, University of California, Davis. Davis, California 95616. 103 104 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES (Smith 1996). Spotted sand bass live to a maximum of 14 years although the average age is 6 to 10 years old. Sexual maturity is reached within the first year. Sizes up to 450 mm. standard length (SL) and 2.6 kg can be reached but indi- viduals larger than 330 mmSL are rare (Allen et al. 1995a). Many of the fish species which occur in the upper Gulf of California are also found on the Pacific side of southern and Baja California and are absent or rare at the southern cape region of Baja California. This disjunct distribution is shared by 42 ecologically diverse fish species belonging to 28 families (Hubbs 1960; Walker 1960; Present 1987). Highly mobile and sedentary species (Hubbs 1960; Walker 1960) as well as viviparous (Miller and Lea 1972) and planktonic spawn- ing species are represented. Fish species composition in the Gulf places it as part of the Panamic faunal province (Walker 1960). Approximately 75% of the shore- fish have a principal range south of the Gulf. About 8% range north of the gulf and are mostly disjuncts from southern California while 17% are endemic to the Gulf (Walker 1960). Ocean floor spreading separated southern Baja California from mainland Mex- icO approximately 5.5 million years ago and has subsequently opened the Gulf 260 km to the northwest (Larson et al. 1968; Spencer et al. 1989). It is widely held that the distinct Pacific and Gulf faunae were apparently established during the most recent of nine Pleistocene glaciation events. The maximum southerly extension of the most recent glaciation occurred approximately 18,000 years ago. Moore et al. (1980) estimate that cooler (by approximately 2—4 °C) sea surface isotherms were shifted sufficiently south of the cape to allow free movement of species otherwise limited by higher sea surface temperatures in this region. The colder waters in the north forced the temperate and subtropical species to the warmer southern waters in order to maximize feeding and reproduction rates (Webb 1992). Walker (1960) discusses the possibility of a late Pleistocene seaway in the La Paz region which would have allowed for movement between Pacific and Gulf regions although the occurrence of such a waterway has not yet been established (Durham and Allison 1960). As the ice sheet began to recede to the north, approximately 15,000 years ago, the current day fauna returned and the Pacific and Gulf distribution was probably established well before the disappear- ance of the ice sheet. The magnitude of isolation provided by the Baja peninsula is unknown and most likely varies according to each species’ ability to move long distances or disperse eggs and larvae. In addition to the Baja peninsula, oceanographic barriers such as currents and isotherms (Cowen 1985) as well as chemical cues may affect larval dispersal routes. The spotted sand bass occupies a 3200-km range, which is unusually wide for a nearshore marine fish. However, in southern California, adults are restricted to definable populations and large scale movement is unlikely due to discontinuities of suitable habitat (Allen, unpubl. data). The extent and direction of larval dispersal in this species is unknown. Plankton studies by Butler et al. (1982) found that Paralabrax eggs and larvae were con- fined mostly shoreward of the 36-mile contour off the mainland. California Co- operative Oceanic Fisheries Investigations (CalCOFI) cruise data from 1951—1984 report that serranid larvae occur as far as 250 km off of the southern and Baja California Pacific shoreline (Moser et al. 1993). The abundance of serranid eggs and larvae relative to all species sampled (Moser et al. 1993) was approximately VARIATION IN PARALABRAX MACULATOFASCIATUS 105 0.2%. At these latitudes the serranid larvae are most likely to be Paralabrax spp. One aggregation of serranid larvae was reported throughout the Channel Islands within the Southern California Bight. Another region of high larval density oc- curred off the central Baja, Pacific coast, stretching from Bahia de San Quintin beyond the tip of the peninsula. Within the Gulf, considerable mixing occurs in the northern regions (van Andel 1964). The relative abundance of serranid eggs and larvae were approximately 1.5% relative to all species sampled (Moser et al. 1986). Investigators have used both morphological and molecular techniques to deter- mine if gene flow is restricted between Gulf and Pacific populations. Morpholog- ical analyses can provide insight into the degree of adaptive differentiation which has occurred between disjunct populations. Although this may be reflected ge- netically and imply limited gene flow between regions, it is imperative to inves- tigate the amount of genetic differentiation through more direct methods (Patter- son 1987). Morphological differentiation has been reported in Gulf and Pacific populations of Hypsoblennius jenkinsi (Losey 1968), Sebastes macdonaldi (Chen 1975), and Zalembius rosaceus (Congleton 1968). Morphological differentiation has also been reported among Girella, Sebastes, and Leuresthes, all of which are recognized as having closely related Gulf and Pacific species. Protein electropho- retic variability indicates that gene flow is limited between disjunct Pacific and Gulf populations of H. jenkinsi (Present 1987) and between Girella nigricans and Girella simplicidens (Orton and Buth 1984) with no fixed allelic differences. Fixed allelic differences were found between Leuresthes populations (Crabtree 1983). The use of ribosomal DNA (rDNA) in restriction fragment length polymor- phism (RFLP) studies is commonly used in population biology (Zhuo et al. 1994; Jensen et al. 1993; Hall 1992; Phillips et al. 1992; Hillis and Moritz 1990). The rDNA unit occurs as clusters of tandem repeats located in the nucleolar organizing region (Long and Dawid 1980; Learn and Schaal 1987). In most vertebrates, this repeat unit consists of two nonconserved internal transcribed spacers, ITS-1 and ITS-2, flanked by three highly conserved gene coding regions, 18S, 5.8S, and 28S. Two external transcribed spacers (3'-ETS and 5’-ETS) flank the 18S and 28S genes and each repeat unit is separated by an intergenic spacer region (IGS). The internal and external transcribed spacers (ITS-1, ITS-2, 3’-ETS, and 5’-ETS) as well as the IGS region evolve more rapidly and can be used for comparisons of closely related species and geographically distant populations. The gene coding regions (18S, 5.8S, and 28S) evolve more slowly and are suitable for use in phylogenetic studies (Fain et al. 1992; Phillips et al. 1992; Learn and Schaal 1987). Furthermore, the number and location of ITS repeat units on the chro- mosomes varies among species (Allard et al. 1990). Mutations occurring within these repeat units can spread rapidly and homogenize within an interbreeding population (Silberman and Walsh 1992). Differences of restriction sites within the ITS regions due to the accumulation of mutations will result in restriction frag- ments that are unique to a particular population. Those populations that are most genetically similar will have more restriction sites in common and this can be determined through the comparison of RFLP banding patterns. The purpose of this investigation was to determine the extent of morphological and genetic variation among Gulf and Pacific populations of the spotted sand 106 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Guerrero Negro 5°N Pachiic Ocean Magdalena Bay Fig. |. Map showing six collection sites from southern California and Baja, California, Mexico. bass. If gene flow is limited between these regions, this system would provide an excellent opportunity to investigate the effects of isolation on the process and rate of speciation since a Pleistocene establishment has been estimated for the time of separation (Hubbs 1960; Walker 1960). Materials and Methods Collection and preservation of specimens.—Spotted sand bass were collected between March 1993 and January 1995 with hook and line angling. Fish were collected from six sites along the entire range of this species (Fig. 1). The three Pacific sites sampled were San Diego, Guerrero Negro, and Magdalena Bay and the three Gulf sites included Los Pulpos, Bahia de Los Angeles, and La Paz. Whole live specimens were immediately placed in individually labeled plastic bags, stored on ice and transported to California State University, Northridge. Gill filament tissue was clipped in the field with forceps and tweezers and placed into 1.5 ml. microfuge tubes containing 5X NET buffer (2.5M NaCl, 0.25M EDTA, 0.25M Tris Base, pH 8.0). Tubes were labeled according to location and time of capture and stored on ice until transfer to freezers at the laboratory. Morphological analysis.—A total of 30 P. maculatofasciatus individuals were collected from each site (n=180). Hook and line sampling limited the range of specimens to 110—307 mm. standard length. In total, seven meristic counts and nineteen morphometric measurements were made on each individual according to Cailliet et al. (1986). Measurements were made to the nearest 0.01mm with Mi- tutoyo CD-6B digital calipers. Genetic analysis.—A total of 15 specimens from each of the six sites were analyzed for variation within the ribosomal DNA repeat unit. Total nuclear ge- nomic DNA was extracted using the G-NOME Kit protocol from BIO 101, Inc. VARIATION IN PARALABRAX MACULATOFASCIATUS 107 DNA concentration and purity (absorbance at 260/280 nm) were estimated using a Beckman DU-64 spectrophotometer. DNA concentrations of sample stock were adjusted to 0.15 wg/pl by the addition of TE buffer (pH 8.0). The two internal transcribed spacer regions (ITS 1 and ITS 2) and the 5.8S subunit were amplified by polymerase chain reaction (PCR). Primers complimentary to the 28S and 18S flanking regions (Hillis and Dixon 1992) were purchased from Integrated DNA Technologies, Inc. Primer 28u has a 21 base sequence (5’-CGTTA- CTGGGGGAATCCTGGT-3’) and primer 18s has a 27 base sequence (5'-CA- CACCGCCCGTCGCTACTACCGATTG-3’). The following PCR reaction param- eters were used: 0.75 wg of template DNA, 0.75 wM of each primer (18D and 28U), 0.15 mM dNTP’s, 3.0 mM MgCl, 1.2 pl of 10 buffer, and 1 U Taq polymerase in a 20 pl reaction. Taq polymerase was pipetted into the mixture after an initial 8-minute heat shock phase. The following thermocycle conditions were used: 30 cycles consisting of a 95 °C denaturing step for 45 seconds, a 60 °C annealing phase of 45 seconds, and a 72 °C elongation period of 1 minute. These cycles were followed by a 5-minute elongation phase at 72 °C and then a 5 °C storage until removal from the thermocycler. PCR products were subjected to digestion in a 37 °C or 60 °C waterbath (depending on the reaction require- ments) for 16 hours with 1U of a restriction endonuclease. The following restric- tion enzymes were used: BstUI, Dpnil, Hhal, Haelll, MspI, Rsal, Taqal, Hindill, Stul, Ncil, and BamHI (purchased from New England BioLabs). Electrophoresis was conducted on a 65 ml 2.5% TBE agarose gel in 1X TBE buffer at 90 volts for 2 hours. New England Biolab’s ®X174 HaelIlII size marker was loaded onto each gel in order to determine fragment size. RFLP’s produced by endonuclease restriction were examined on an ultraviolet transilluminator and captured with the UVP ImageStore 5000 (Beaumont, CA) gel documentation system. The captured gel images were analyzed with UVP’s SW 5000 GelBase™ Windows software program and all fragment sizes were estimated in base pair lengths. Data analysis.—Statistica for Windows was used for data analysis and graph- ical representation. The morphological data were log-transformed in order to re- duce the correlation of measurement variances and means (Sokal and Rohlf 1981). Body counts in most fishes do not usually covary with growth after a threshold body size is reached (Strauss 1985). Meristic data were not transformed and were treated separately from measurement data. A stepwise multiple discriminate func- tion analysis was used in order to calculate the linear combinations of the original morphometric and meristic variables which maximally discriminated among the six known groups (Atchley and Bryant 1975). This analysis computed weights for each character and combined the weighted characters into a single score to best discriminate among groups. Graphical displays of these discriminant func- tions are useful for demonstrating both differences and similarities among groups. Codominant RFLP genotypes were recorded for each individual and analyzed with the ‘Genes in Populations’ analysis program (designed by B. May and C. C. Krueger; written by W. Eng and E. Paul, UC Davis). Roger’s genetic distances (Roger 1972) were calculated between all population pairs and a phenogram gen- erated using the unweighted pair-group method using arithmetic mean (UPGMA). Significance was tested for all population pairs using likelihood ratio (G7) tests for homogeneity of gene frequencies across populations. 108 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 1. Classification results of meristic data for six populations of spotted sand bass (Paralabrax maculatofasciatus). Rows are observed classifications and columns are predicted classifications. Bahia Percent San Guerrero Magda- de Los Los correct Diego Negro lena Bay La Paz Angeles’ Pulpos San Diego a7 7 1 8 0) 3 1 Guerrero Negro 30 3 2) 10 0) 6 2 Magdalena Bay 43 S) 1 1S) O 6 1 La Paz 17, 6 + 6 0 9 5 Bahia de Los Angeles 67 0) 4 4 0) 20 Z Los Pulpos O 8 3 i 0) 7 1 Total 36 43 22 Si 0) ail 12 Results Morphological analysis.—No variation occurred in four of the seven meristic counts (dorsal fin spines, anal fin spines, anal soft rays, and branchiostegal rays). Of the three discriminant function (DF) roots computed for the three remaining characters, only the first was statistically significant (F = 3.5, Wilk’s lambda = 0.75, p < 0.001). The correct classification results ranged from 0.00 to 66.6% with an overall rate of 35.6% (Table 1). No populations were grouped with La Paz, which also failed to group with itself. The plot of DF I (Fig. 2) with 95% confidence ellipses shows complete overlap among all sites with no discernible patterns. The morphometric discriminant function analysis computed five functions to account for 100% of the variation. DF I, II and III described a total of 85.1% of the variation (Table 2) and were highly significant (Table 3). The correct classi- San Diego Guerrero Negro Magdalena Bay Los Pulpos Bahia de Los Angeles La Paz Discriminant function I 85.1% Fig. 2. Meristic scatterplot showing Discriminant Function I. Ellipsoids indicate 95% confidence limits. VARIATION IN PARALABRAX MACULATOFASCIATUS 109 Table 2. Loadings of 19 log-transformed morphometric characters on three disciminant function roots. Character DFE I DF II DF III Standard length = 997, OOH —=():831 Body depth 1.476 =2 IAD 0.781 Caudal peduncle length = 2313 0.584 OWS Predorsal length 0:393 —0.616 —0.807 Dorsal base length = 0/618 —0.043 0.087 Anal base length = de. deh 8 —0.614 0.099 Dorsal fin height 0.826 0.583 0.645 Anal fin height =Qn25 0.688 0.692 Pectoral fin length = OS —0.300 0.683 Pelvic fin length 0.094 0.633 = Deas Longest dorsal spine =0:659 —0.414 1087 Head length —0.070 0.017 0.455 Head width OL Sai —0.894 =1.995 Snout length 16935 =0:265 —0.088 Suborbital width 0.247 1.218 0.677 Orbit to preopercle Dal 0.539 1.508 Eye diameter =0:202 = OL252 0.641 Upper jaw length 0.001 0.847 0:28) Gape width 0.785 1.595 1.134 Percent Variance Individual 50.40% 23.80% 10.80% Cumulative 50.40% 74.20% 85.10% fication rates ranged from 73.0 to 100.0% with an overall rate of 86.1% (Table 4). Samples were distinctive along DF I (50.4%) and DF II (23.8%) axes shown with 95% confidence ellipses (Fig. 3). Spotted sand bass from the uppermost Pacific population of San Diego were clearly distinct from the uppermost Gulf, Los Pulpos population. These are the most distant regions of the present range of spotted sand bass. Considerable overlap occurs among the four remaining Baja sites. The Guerrero Negro and Bahia de Los Angeles samples show overlap which follows their geographic positioning as southern adjacents between San Diego and Los Pulpos. The southernmost cape populations, Magdalena Bay and La Paz, overlap with one another and include Bahia de Los Angeles with most of Guerrero Negro. The characters most useful for discriminating between the upper Gulf and upper Pacific regions along the DF I axis are increasing orbit to preopercle and body depths, and shorter caudal peduncle and standard length, respectively. The plot of DF II groups San Diego and Los Pulpos away from Guerrero Negro and Table 3. Summary of stepwise discriminant analysis of morphometric data. Results for first three calculated roots shown. Discriminant Function F value Wilk’s lambda p level I 8.8 0.0267 <0.001 II 6.4 0.1138 <0.001 Ill 4.9 0.2818 <0.001 110 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 4. Classification results of morphometric data for six populations of spotted sand bass (Paral- abrax maculatofasciatus). Rows are observed classifications and columns are predicted classifications. Bahia Percent San Guerrero Magda- de Los Los correct Diego Negro lena Bay La Paz Angeles’ Pulpos San Diego 100 30 0) 0) 0) 0) 0) Guerrero Negro 80 24 1 3 1 0) Magdalena Bay Ue 1 l 9) 6 0) O La Paz 87 0) 2 1 26 1 O Bahia de Los Angeles 90 0) 0 1 2 27 0) Los Pulpos 87 0) 0) 4 O 0 26 Total 86 3 27 29 57, 29 26 Bahia de Los Angeles and reflects their north to south clinal relation to one another. Magdalena’ Bay and La Paz again overlap and include all sites. The characters most responsible for the variation are increasing gape width and sub- orbital width describing the southern group while smaller body depth and standard length characterize the northern sites. Results of DF I and DF II combined show that the cape populations are not easily discerned from the remaining sites and are more inclusive. The remaining upper Gulf and upper Pacific sites tend to have tighter clusters which allow for the discernment of geographic patterns. The upper Pacific samples are generally described by smaller fin and body sizes while the upper Gulf groups have larger facial measurements. The plot of DF III (10.8%) shows an overlap of San Diego, Guerrero Negro and Los Pulpos which are the range extremes (Fig. 4). La Paz and Bahia de Los Angeles are separated along the zero axis and Magdalena Bay overlaps the both of them reflecting its clinical San Diege Guerrero Negro Magdalena Bay Los Pulpos 23.8% Bahia de Los Angeles La Paz Discriminant function II 6 4 9 0 2 4 6 Discriminant function] 50.4% Fig. 3. Morphometric scatterplot showing Discriminant Functions I and II. Ellipsoids indicate 95% confidence limits. VARIATION IN PARALABRAX MACULATOFASCIATUS halt San Diego Guerrero Negro Magdalena Bay Los Pulpos Bahia de Los Angeles La Paz 10.8% Discriminant function III -6 -4 -2 0 2 4 6 Discriminant function I 50.4% Fig. 4. Morphometric scatterplot showing Discriminant Function III. Ellipsoids indicate 95% con- fidence limits. geographic position to these sites. The variation from La Paz to Magdalena Bay and then Bahia de Los Angeles is described by smaller pelvic fin and longest dorsal spine and increasing orbit to preopercle and gape width. Larger facial dimensions with DF III again describe the upper gulf region. Genetic analysis.—Frequencies of RFLP patterns were calculated for seven of the eleven restriction endonucleases that showed variation (Table 5). A phenogram obtained using Roger’s (1972) genetic distance and UPGMA (Sneath and Sokal 1973) clusters the six populations (Fig. 5). G-test results indicate that the gulf populations are genetically similar to one another and are significantly different from the three Pacific populations. The three Pacific populations form two sig- nificantly different groups. The northernmost Pacific population of San Diego is significantly different from the two southern-Pacific groups of Guerrero Negro and Magdalena Bay which are the most genetically similar of the six populations. Discussion The morphological and genetic results indicate that populations of P. macula- tofasciatus from the Baja California peninsula form a complex system which cannot simply be divided into disjunct Gulf and Pacific distributions. The morphometric results indicate that significant differentiation has occurred among populations from the range extremes. Morphometric DF I and DF II de- scribe these as discrete groups according to varying body shapes between these regions. The San Diego and Guerrero Negro populations are distinguished by smaller caudal peduncle, standard length, body depth, and fin characters. This indicates that the upper Pacific spotted sand bass are smaller-bodied and have smaller fins than the upper Gulf group. The Los Pulpos and Bahia de Los Angeles spotted sand bass have larger preopercle, body depth, gape width, and suborbital bi? SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Table 5. Arbitrary designation of restriction fragment patterns shown as frequencies. w o) so} es] bg] Enzyme Population A G H — BSTU I San Diego 0 Guerrero Negro OZ Magdalena Bay 0.4 La Paz Oat Bahia de LA 0.6 0.2 Los Pulpos 0) 0.1 DPN II San Diego 0.4 0.6 Guerrero Negro 0.6 1 Magdalena Bay 0) La Paz 0) Bahia de LA 0 Los Pulpos 0) O O ee oe ~ MN on NN OW —_ N N ~ Ww eceecos eeqoo eoeo ooe > fon HAE III San Diego Guerrero Negro Magdalena Bay 0.3 0.8 La Paz 0.1 0.6 Bahia de LA 0.5 0.5 Los Pulpos HhA I San Diego Guerrero Negro Magdalena Bay La Paz Bahia de LA Los Pulpos MSP I San Diego Guerrero Negro Magdalena Bay La Paz Bahia de LA Los Pulpos RSA I San Diego Guerrero Negro Magdalena Bay La Paz Bahia de LA Los Pulpos Taq al San Diego Guerrero Negro Magdalena Bay La Paz Bahia de LA Los Pulpos ON £N N TS cooooso Nn — me OW AN £ O oe) = nN Mn W eS) N N QI Nowa See oococoees = = KNN N i) N — o N Coo = Ww ~ \O No — Glioma ceocooeceacooceoeoeoooooooceocoeocosoooeoooooos NN Seeeccomeceococoecooormcocoone Nn © Gre Grote Go SOO oS oO SCletOTO OO OCoO SCS S erclemoemeieo cro eoocecomoeoocoecomo ooo ee oe Come Ccooo So N width characters. These characters describe a group of larger-headed fish with more robust facial characters. DF III overlaps San Diego, Guerrero Negro, and La Paz but shows some distinction among the cape sites and Bahia de Los An- geles. La Paz and Magdalena Bay are described by smaller pelvic fin and longest dorsal spine fin characters. Bahia de Los Angeles is characterized by larger orbit to preopercle and gape width measurements, again demonstrating the large head dimensions of the upper Gulf group. Magdalena Bay and La Paz samples are very similar morphologically. These share more similarity with the adjacent groups to VARIATION IN PARALABRAX MACULATOFASCIATUS 113 0.198 0.395 0.593 0.791 Ri ees Sh. She ee San Diego Guerrero Negro = P< 0.001 Magdalena Ba P < 0.001 La Paz Bahia de Los Angeles ns Los Pulpos ne ae ee | 0.198 0.395 0.593 0.791 Roger’s genetic distance Fig. 5. Phenogram obtained using Roger’s genetic distance and UPGMA. Non-significant G-test results are shown as (ns) and P-values for significant tests are shown on the graph. The gulf populations are genetically similar to one another and are significantly different from the three Pacific populations. The three Pacific populations form two significantly different groups. The northernmost Pacific pop- ulation of San Diego is significantly different from the two southern-Pacific groups of Guerrero Negro and Magdalena Bay that are the most genetically similar of the six populations. the north, Guerrero Negro and Bahia de Los Angeles, with little overlap with the most distant sites, San Diego and Los Pulpos. These trends describe two broadly inclusive southern populations that are less easily classified. The most distant northern groups become more distinguished from this intermediate, southern re- gion and have the highest classification rates. Determining the causes of morphological variation among groups has been a difficult task for researchers. The variation of one character or groups of characters may be caused by phenotypic plasticity or by genetic differences among popu- lations. The environment in which a fish develops modifies its shape by accel- erating or slowing the rate of development (Hubbs 1926). The northern represen- tatives of a population are usually larger than those to the south. Northern groups grow more slowly and frequently display smaller head and fin proportions than their southern counterparts (Martin 1949). The colder water temperatures of the north alter the timing of transition from one growth stanza to another and produce a new relative growth relationship (Huxley 1932; Martin 1949). Meristic variation is also greatly influenced by the environment. A correlation between cooler en- vironmental temperatures and higher meristic numbers usually result in northern groups having greater meristic counts (Hubbs 1926; Vladykov 1934; Taning 1952). Cooler waters slow growth rates which cause longer developmental periods and allow tissues to develop into a greater number of elements (Gabriel 1944). Southern fishes from warmer waters generally grow faster and display lower me- ristic numbers. Other influences such as high salinities and low oxygen concen- trations parallel the effects of lower temperatures by retarding growth rates (Tan- ing 1952). The genetic determination of morphology has been demonstrated with- in and between populations (Hubbs 1955). New Zoarces individuals transplanted among different environments continued to produce forms that were recognizable from the endemic population (Ege 1942). Individuals from different fish popu- lations reared under comparable conditions continued to produce dissimilar me- 114 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES ristic characters (Heuts 1947 1949; Seymour 1956). Determining the source of morphological variation among widely distributed spotted sand bass populations provides similar difficulties. The upper Gulf spotted sand bass are subjected to annual temperature fluctuations of 9-32 °C (Thomson et al. 1987) while San Diego fish experience 14—27°C extremes (Allen 1995b). The salinity in the upper Gulf is stable at 36—37 ppt annually (Brusca 1973) while in San Diego the salinity fluctuates between 32-36 ppt (Allen 1995). These environmental influences may be affecting the development of P. maculatofasciatus larvae and juveniles suffi- ciently to produce significantly different morphologies. Lindsey (1962) found that fin ray numbers are more strongly influenced by environmental temperature than by parental genotype in Gasterosteus aculeatus. If this is the case with P. maculatofasciatus, then the physical environment may be a less important influence on morphology than selection due to the complete lack of meristic variation found in this study. Significant morphometric variation could then be the result of natural selection rather than differential development induced by varying physical environments. Spotted sand bass within the southern California bight feed mainly on small fishes, crustaceans, and infaunal bivalves (Allen et al. 1995a). The three Pacific sampling sites and La Paz are soft sediment microhabitats which provide infaunal invertebrate prey. The upper Gulf habitat is dominated by harder, rocky reef substrate upon which spotted sand bass are highly abundant. Ferry and Clark (1997) have demonstrated that the Bahia de Los An- geles spotted sand bass feed on a wider variety of prey and take advantage of the annually fluctuating food resources due to periodic winterkills (Thomson and Lehner 1976). In some cases these fish are dislodging prey items from the reef substrate and are crushing hard-shelled invertebrates. The variable diet of upper Gulf spotted sand bass may require a more robust facial morphology as demon- strated in DF I, II, and HI. The higher temperature fluctuation in this region may be selecting for species that acclimate to these environmentally influenced changes in diet. Assuming that morphometric and meristic characters are equally labile, the genetic determination of meristics could weaken the argument for selection versus influence of physical environment on development. Genotype may be con- trolling meristics regardless of the possible influence of environment on the de- velopment of morphometric measures. If counts are more highly conserved than body proportions, then the lack of meristic variability may simply be reinforcing P. maculatofasciatus as a single species. Presumably, genetic differences among geographically widespread species arise from isolation by ecological, physical, and distance barriers. Natural selection, favoring different genotypes, may produce genetic differentiation among isolated regions (Prakash et al. 1969). Differences have been shown to occur among ma- rine organisms with widespread planktonic dispersal capabilities (Present 1987; Waples and Rosenblatt 1987; Grothues 1994). Genetic differentiation has been demonstrated among geographic groups of a larger panmictic population of An- guilla rostrata (Williams et al. 1973; Koehn and Williams 1978). In these studies, differential selection pressures were sufficient to change gene frequencies on the order of 10% per generation (Williams et al. 1973). Genetic differentiation of annual A. rostrata cohorts was directed in similar patterns by natural selection (Koehn and Williams 1978). The population affinities described by RFLP cluster analysis suggest that the VARIATION IN PARALABRAX MACULATOFASCIATUS 115 Gulf and Pacific populations are genetically distinct. Although spotted sand bass eggs and larvae stay in the plankton for nearly a month, direct gene flow between Gulf and Pacific regions seems to be limited. The California Current and Gulf surface waters converge to form a persistent oceanic front in the cape region (Roden and Groves 1959; Griffiths 1968). This may restrict the movement of adults as well as drifting eggs and larvae around Cabo San Lucas. The absence of most adults from this region may also be due to a lack of suitable habitat and unfavorably high water temperatures. In the Gulf of California, prevailing northerly (winter) and southerly (summer) winds cause considerable upwelling and produce year-round plankton blooms (van Andel 1964). Considerable amounts of gene flow are likely with such widespread mixing and strong tidal currents around the larger islands of the upper Gulf. The lack of significant genetic differentiation among the three gulf populations sug- gests that sufficient mixing may occur throughout this region. The southerly flowing California Current delivers eggs and larvae from the north along the Pacific coast (Schwartzlose 1964). The clustering of Guerrero Negro and Magdalena Bay in the Pacific indicates that sufficient gene flow is occurring throughout Baja California’s southern Pacific region. San Diego is the most genetically distant population in this study, indicating that gene flow between this region and the southern populations is limited. A paucity of serranid larvae between the Southern California Bight and Guerrero Negro may result from a region of upwelling at San Quintin which may limit consistent gene flow between these regions (Moser et al. 1993; Grothues 1994). A study of ten marine shore fishes from southern California and central Baja demonstrated that more rare alleles were delivered from the south and collected in the north (Waples and Rosenblatt 1987). Episodic northward movements of water from the south, in- cluding El Nino events, occasionally transport larvae with southern Baja affinities into southern and central California waters (Radovich 1961; Brinton 1981). The results herein indicate that these oceanic phenomena may not be facilitating sig- nificant gene flow between the southern Pacific and San Diego populations. The morphological and genetic evidence demonstrates that significant differ- entiation has occurred between the Gulf and Pacific regions. Agreement between both lines of evidence lends powerful support to the idea that these areas are restricted from sharing direct gene flow. A clear morphological distinction be- tween San Diego and Los Pulpos is evident with overlap of the southern inter- mediates, Guerrero Negro and Bahia de Los Angeles. As these groups become more distant, their divergent morphologies may reflect differential selection pres- sures between different habitats. The molecular evidence indicates that the Pacific and Gulf populations are different, most likely due to a significant reduction of gene flow. Additionally, the significant genetic differences between San Diego and the southern-Pacific populations of Baja indicate that gene flow is also re- stricted between these regions. The results of this study suggest that geographi- cally isolated populations, under the influence of strong selection pressures, do not require long periods of time for divergence. With a species that matures rapidly, many generations are subjected to selection through evolutionary time. The last 15,000 years may have been sufficient for allowing significant divergence to occur between upper Gulf and Pacific groups of the spotted sand bass. 116 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Acknowledgements We express our gratitude to Kenneth Jones for his assistance with the molecular aspects of this study. Our appreciation is also extended to Donald Buth, Robert Carpenter, and Shawn Nordell for providing advice on numerous aspects of this research and comments on the manuscript. Special thanks go to Tim Hovey, Tom Grothues, Carrie Wolfe, Jon Smith, Mike Franklin, Mara Morgan, Craig Camp- bell, Holly Harpham, Cheryl Baca, Andy Barbarena, Ron Klaver, Jorge Rosales- Casian, and the crew of the R/V Yellowfin: Jim Cvitanovitch and Dan Warren for their extensive field support. Greg Cailliet, Lara Ferry, and Alan Andrews pro- vided a number of the Bahia de Los Angeles samples. 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Sci. 98(3), 1999, pp. 119-126 © Southern California Academy of Sciences, 1999 Diet and Seed Dispersal Efficiency of the Gray Fox (Urocyon cinereoargenteus) in Chaparral James A. Wilson and Barry Thomas Department of Biological Science, California State University, Fullerton, CA, 92834-6828 Abstract.—The frequency of occurrence of prey items in the diet of gray foxes (Urocyon cinereoargenteus) in the chaparral of Southern California was deter- mined from scat analysis. Fruit occurred in 70% of scats (n = 106) analyzed and consisted of coffeeberry (Rhamnus californica), eastwood manzanita (Arctostaph- ylos glandulosa), toyon (Heteromeles arbutifolia), and hollyleaf redberry (Rham- nus illicifolia). Mammalian prey included white-footed mice (Peromyscus sp., 7%), wood rats (Neotoma fuscipes, 4%), and meadow voles (Microtus californi- cus, 1%). Insects, including grasshoppers (Orthoptera, 66%) and Jerusalem crick- ets (Stenopelmatus sp., 54%) composed 14% of prey occurrences. There was only one occurrence of reptilian prey (0.94%), a western fence lizard (Sceloporus oc- cidentalis). Seeds passing through the fox’s gut were tested to determine germi- nation period. Coffeeberry and redberry ingested by foxes germinated signifi- cantly sooner than seeds from fresh fruit (p = 0.0005 and 0.03), while toyon germination rates did not differ between fresh and ingested seeds. Days to ger- mination for fresh seeds and seeds from scat averaged 94.6 + 37.2 and 48.2 + 25.4 for coffeeberry, 68.9 + 15.2 and 52.0 + 8.2 for redberry, and 14.0 + 2.3 and 22.2 + 9.0 for toyon. Seed dispersal occurs in many fruit-eating animals including fish (Horn 1997), primates (Lieberman et al. 1979; Zhang 1995), bears (Applegate et al. 1979; Rogers and Applegate 1983), and most notably birds (Krefting and Roe 1949). Studies of frugivory and seed dispersal have focused on tropical species, and have not investigated temperate fruit and frugivores (Willson 1992). Frugivores act as seed dispersal agents by consuming fruit and defecating or regurgitating the seeds some distance from the parent plant (Traveset and Willson 1997). In addition to transporting seeds, frugivores may have an affect on seed morphology and/or physiology (Schupp 1993). The effects of frugivores on seeds includes releasing the seed from chemicals that inhibit germination (Barnea et al. 1991), scarification of the seed coat (Schopmeyer 1974), and deposition at suitable germination sites (Bustamonte and Canals 1995). In addition, the fruit may have some effect on the frugivore, including increased digestive rates from laxative chemicals in the fruit pulp (Murray et al. 1994; Putz 1993). Willson (1992) reports that nine families of mammals in North America con- sume fruit and are potential seed-dispersers. Of these mammals, carnivores are considered the best potential seed dispersal agents due to their large home ranges (Willson 1992). One such carnivore that consumes fruit throughout its range is the gray fox (Urocyon cinereoargenteus). The gray fox has morphological spe- cializations for frugivory, including shortened jaws that increase bite speed and ty) 120 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES higher digestive efficiency for fruit compared to the more carnivorous red fox (Vulpes vulpes; Jaslow 1987). Both the red and gray fox are found throughout the United States, and these specializations would allow the gray fox to partition available food resources, thereby reducing competition with the red fox. The gray fox has a wide distribution, extending from the Canada and United States border to northern South America (Fritzell and Haroldson 1982). Although gray foxes consume fruit throughout their range, they are more carnivorous in the eastern and southern United States than in the west (Bennett and English 1942; Chaddock 1939; Grinnell and Storer 1924; Hatfield 1939; Korschegen 1957; Nel- son 1933; Wood 1954; and Wood et al. 1958). In the Sierra Madre Mountains of Mexico, the diet of gray foxes may be as much as 96% fruit (Delibes et al. 1989). The difference in the amount of fruit consumed by gray foxes may be the result of a difference in climatic conditions between east and west. It has been hypoth- esized that foxes living in xeric environments may use berries as a reliable source of water (Ball and Golightly 1992). This study was designed to explore the relationship between gray foxes and the food plants occurring in the chaparral community. If the water use hypothesis is correct, foxes living in chaparral should show an increase in frugivory. In addition, if seeds ingested by foxes are viable after passage through the gut, the fox is a potential seed disperer. We quantified the use of fruit by gray foxes in chaparral habitats, and determined whether seeds were viable after passing through the fox digestive system. Materials and Methods Study site.—This study was conducted in the Cleveland National Forest in Orange County, California, at an elevation of 1460 m. The plant community is chaparral (Munz and Keck 1965) dominated by chamise (Adenostoma fascicula- tum), Ceanothus cuneata, C. facicuatum, C. spinosa, manzanita (Arctostaphylos glandulosa and A. glauca), scrub oak (Quercus berberidifolia), and buckthorn (Rhamnus spp). The climate is Mediterranean, with hot, dry summers and cold, wet winters. Water sources located in the study site included a perennial spring and two man-made quail guzzlers. Diet analysis.—A 0.5-km section of road was cleared of scat at the beginning of the study and then scat was collected approximately biweekly. The wet weight of each scat was taken, scats were washed for 3—5 minutes in a 0.5-mm metal strainer placed over a 500-ml beaker to catch small bone fragments that might pass through the strainer. Particles remaining in the sieve were saved for identi- fication. Bone fragments, seeds, carapaces, feathers, skin, hair, and unknown par- ticles were collected. Identification of prey remains was made using dichotomous keys and reference collections. Incisors and cuticular scale patterns on hair were used to identify rodents. Incisors were identified using Weintraub and Shockley (1980). Hair sam- ples were photographed using a scanning electron microscope at 1000 and cu- ticular scale patterns were referenced with photographs taken of hair from mu- seum specimens. Seeds were identified using museum specimens and by refer- encing Martin and Barkley (1961). Total and seasonal diet was analyzed inde- pendently. A Chi-square test was performed against the hypothesis that prey would be evenly distributed across the seasons. DIET AND SEED DISPERSAL IN GRAY FOX 124 The water content of fruit found in the diet was measured using the mean amount of weight lost after the fruit was dried for one week in an oven at 35°C. Fifty individual fruits were used to obtain the mean water content for each species. Germination Study.—Coffeeberry (Rhamnus californica), manzanita (Arcto- staphylos glandulosa), hollyleaf redberry (Rhamnus ilicifolia), and toyon (Het- eromeles arbutifolia) seeds were collected from scat, and by picking ripe fruit growing in the study site. Fruit pulp was mashed in a bowl of water and seeds were extracted and air-dried. Seeds were then put into a 5% bleach solution for 30 sec to kill fungi. One hundred seeds from fox scats and 100 seeds from fresh fruit were planted in compartmentalized plastic growing trays containing a soil mixture (Applegate et al. 1979; Krefting and Roe 1949; Rogers and Applegate 1983). Germination trays were left on benches outside the California State Uni- versity, Fullerton greenhouse and were watered daily. Trays were checked daily to determine if seed had germinated, and the date of germination was recorded. Trays were checked until no germination occurred for three weeks. Coffeeberry, hollyleaf redberry, and toyon did not require any pretreatment, such as cold stratification or scarifiction, prior to planting (Schopmeyer 1974). The mean time to germination for fresh and ingested seeds was compared using a two-tailed t-test. The hypothesis tested was that no difference would exist be- tween the two seed samples. In addition, the number of seeds that germinated between each sample was compared using a pooled probability comparison with a hypothesis of no difference. Results Diet.—One hundred and six gray fox scats were collected from the study site. The mean wet weight of scats was 5.15g (+ 2.28) with a range of 0.94g to 10.29. Fruit constituted the most frequently consumed food item and was found in 94.3% of scats. Insects comprised the second most frequent item (26.4%), followed by mammals (22.6%), reptiles (0.94%), and birds (0.94%). Many fox scats contained only one of the major food items. Scat containing only fruit, mammals, or insects occurred 59.4%, 3.8%, and 0.9% of the time respectively. Fruit from coffeeberry, toyon, eastwood manzanita, and hollyleaf redberry were consumed by foxes. Excepting manzanita, all plants possessed fleshy fruits high in water content. Manzanita fruit was dry and grainy with only 10.1% water. Coffeeberry had the highest water content at 68.8%, hollyleaf redberry had 56.2%, and toyon contained 55.1%. In addition, clumps of undigested grass were found in scats on five separate occasions. Incidental plant material found in scat included the fruit tips and leaf blades of lemonadeberry (Rhus integrifolia) and occasional whole dried leaves from assorted plants. Gray foxes in the study area consumed fruit throughout the year, and specific fruits used reflected the fruiting patterns of the plants (Figure 1). Fruiting times of species consumed were staggered. Manzanita, hollyleaf redberry, and coffee- berry fruited from summer through early winter while toyon fruited during winter and spring. Manzanita had the longest fruiting period and consequently was con- sumed more than other plants (68% of scat containing fruit had manzanita). Cof- feeberry (37%), toyon (14%), and hollyleaf redberry (11%) follow in frequency. Foxes consumed more insects in spring, perhaps reflecting the decreased avail- ability of fruit. Insect remains primarily consisted of Jerusalem crickets (54% of 122 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Month Plant JF MAM: J Jo Aas O07. a Coffeeberry Redberry Toyon Manzanita Flower [J Fruit Found in Diet * Fig. 1. Flowering and fruiting times of the four plants consumed by chaparral gray foxes. Flow- ering and fruiting times were determined using observations in the study site over the course of the 1995/1996 season. insect occurrences), but leg fragments from other orthopterans also were common. A spider, a fragment of a beetle carapace, a bee head, and a small caterpillar were also recorded. Mammal prey included white-footed mice (Peromyscus spp., 63%), wood rats (Neotoma, 29%), and meadow voles (Microtus, 8%). One scat con- tained a western fence lizard (Sceloporus occidentalis), and another scat contained an unidentified bird. The diet of the gray fox changed seasonally (Figure 2). Fall (September—No- vember) and winter (December—February) diets contained mostly fruit, which occurred in 97.8% and 100% of the scat respectively. Insects (24.4% fall, 16.0% winter) were the second most important prey types followed by mammals (15.6% fall, 12.0% winter). Spring (March—May) diet was significantly different (X? anal- ysis, p < 0.001) from fall and winter. The use of fruit in spring (50% occurrence) was about half of that used in fall and winter. In addition, the occurrence of insects (44% occurrence) in scats during spring was more than double that of other seasons. Mammal use declined during the spring months to 4.0%. Frequency (%) Fall Winter Spring Fig. 2. Seasonal difference in the diet of the chaparral gray fox. Summer data were not available due to an outbreak of canine distemper and the subsequent death of all foxes in the study population. Foxes had not returned to the study site as of April 1997. DIET AND SEED DISPERSAL IN GRAY FOX 12 eS) Rhamnus ilicifolia Number Germinated 20 40 60 80 100 120 140 160 30 40 50 60 70 80 90 100 Heteromeles arbutifolia Days —o— Fresh —@— Ingested Number Germinated Days Fig. 3. Germination rate for seeds from (a) coffeeberry (Rhamnus californica), (b) hollyleaf Redberry (Rhamnus illicifolia), and (c) toyon (Heteromeles arbutifolia). For each species, 100 seeds were collected from fresh fruit in the study site and 100 seeds were collected from fox scat. Fresh samples of redberry seeds were obtained from the seed bank at Rancho Santa Ana Botanic Gardens, Claremont, CA. Germination.—Seeds that passed through the fox’s digestive system showed different rates of germination (Figure 3). Coffeeberry seeds from scats germinated significantly earlier (Two tailed T-test, p = 0.0005) compared to seeds from fresh fruit (Figure 3a). The mean number of days to germination was 94.6 + 37.2 and 48.2 + 25.4 for the freshly picked and ingested seeds respectively. Ingested seeds began growing eight days earlier than the freshly picked seed and followed similar germination rate thereafter. Seventeen seeds germinated in the freshly picked sam- ple and 13 germinated from the ingested sample; however, this difference was not statistically significant (Pooled probability comparison, p = 0.22). Hollyleaf redberry also exhibited a significantly earlier germination time (Fig- ure 3b) for seeds from scats compared to fresh seeds (Two tailed T-test, p = 0.03). The mean days to germination for the fresh and scat samples was 68.9 + 15.2 and 52.0 + 8.2 respectively. Nine fresh and 4 scat seeds germinated; however, the difference was not significant (Pooled probability comparison, p = 0.08). The mean germination time for fresh and ingested toyon seeds was 14.0 + 2.3 and 22.2 + 9.0 days (Figure 3c). The depressed germination rate for the scat sample was not significant (Two tailed T-test, p = 0.08). Fresh seeds began ger- minating one day before the fox sample. The number of seeds that germinated in 124 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES the two samples was 10 fresh and 6 ingested. The difference in the number of seeds that germinated is not significant (Pooled probability comparison, p = 0.14). Discussion Diet.—A mostly frugivorous diet was observed in foxes from this study. Sim- ilar observations were made in oak-pine woodlands in the Sierra Madre of Mexico where 96% of the scat contained manzanita (Delibes et al. 1989). Grinnell and Storer (1924) state that scat of gray fox found in early winter and fall often contained manzanita seeds and hulls. Ball and Golightly (1992) suggest that gray foxes may consume fruit more for water than as a source of energy. In the hot, dry environment of the chaparral water could be a limiting factor; however, on our study site there are several sources of water. Laurel Spring, located within the fox’s home range, runs as a trickle during summer months, but flows steadily following winter rain. This spring supplies enough water to drink year round and collects in pools at the bottom of the canyon. Fruit from coffeeberry, toyon, and redberry are all high in water content (68.8, 56.1, and 55.6% respectively), but were not the most frequently consumed fruit. Manzanita, with the lowest water content (10.1%), was most frequently consumed (64.25%) followed by redberry (14.5%), toyon (12.8%), and coffeeberry (10.3%). If water was the main factor in fruit consumption, foxes should choose to eat succulent fruit more frequently. But foxes consumed more dry fruit, leading to the conclusion that, although fruit may supply water, it is primarily consumed for energy. There are other plants in the fox home range that have fruit with high water content, such as white flowered currant (Ribies indecorum), that are not consumed. Over half of the scat collected contained only fruit remains, suggesting that fruit can supply a substantial amount of the fox’s energy requirements. The high use of fruit by gray foxes in chaparral may be an adaptation to reduce competition between the foxes and other carnivores that inhabit the same area. Pequegnat (1951) lists the gray fox as the most abundant carnivore in the chaparral community. A frugivorous diet, in conjunction with tree climbing ability, would allow the gray fox to exploit resources not available to other carnivores. Germination.—Seeds from three of the four plant species tested were viable after gastrointestinal passage, indicating that the fox may be a seed dispersal agent for these species. Seeds from coffeeberry and hollyleaf redberry exhibited more rapid germination after passing through the fox. Earlier germination would allow the seedlings to become established after the first rain of the season. In addition, quicker germination results in less exposure to predation by insects and infection by fungi. Although germination of toyon seeds was not enhanced by the fox’s digestive system, they still might benefit from the increased dispersal provided by foxes. Scat was deposited in the road, usually on low flat rocks. Grinnell and Storer (1924) also note that gray foxes seem to deposit scat on open roads and the tops of rocks. These locations appear to be a poor location for seedling development; however, rain might break up and dissolve scat and wash seeds off the rock and onto the soil (Leiberman et al. 1979). A presumed benefit of animal dispersal is the opportunity for seeds to be de- posited in locations not accessible through other means of dispersal. Without a DIET AND SEED DISPERSAL IN GRAY FOX 12 N mobile disperser, fruit might simply fall off the plant, and seedlings would become established directly under or downslope from the parent plant. An animal disperser provides seeds with the possibility of moving upslope and great distances from the parent plant. A similar effect has been documented in the Machaca (Brycon guatemalensis), a fig eating fish from Costa Rica (Horn 1997). The Machaca feeds on figs that fall into the water and swim upstream depositing the seeds along the river’s bank. The gray fox may provide a means of dispersal for plants in the chaparral in return for the nutrition fruit provides. This interaction may be important in re- generation of the chaparral ecosystem. Orange County, California is currently undergoing extensive development. Foxes may be displaced leaving plants with- out an important dispersal agent. Acknowledgments Funding for this study was obtained from the Department of Biological Science, California State University, Fullerton. Funding was provided by the California State University, Fullerton Departmental Associations Council for funding pre- sentations to the American Society of Mammalogists annual meetings (1996 & 1997). We would like to thank the botanists of The Rancho Santa Ana Botanic Gardens for identifying an unknown seed and providing comparative material. We would also like to thank the following people for helping with plant transects: Jay Wilson, Sarika Thakur, John Yaney, Lance Tayco, and Teo Eng. Literature Cited Applegate, R. D., L. L. Rogers, D. A. Castell, and J. M. Novak. 1979. Germination of cow parsnip from bear feces. J. of Mamm., 60:655. Ball, L. C., and R. T. Golightly. 1992. Energy and nutrient assimilation by gray foxes on diets of mice and himalaya berries. J. of Mamm., 73:840—846. Barnea, A., Y. Yom-Tov, and J. Friedman. 1990. Does ingestion by birds affect seed germination? Functional Ecology 5:394—402. Bennett, L. J.. and P. F English. 1942. Food habits of the grey fox in Pennsylvania. Pennsylvania Game News, 12:10,22. Bustamante, R. Q., and M. Canals. 1995. Dispersal quality in plants—how to measure efficiency and effectiveness of a seed disperser. OIKOS 73:133-—136. Chaddock, T. T. 1939. Report on grey and red fox stomach examinations. Wisc. Cons. Bull. 4:53—54. Delibes, M., L. Hernandez, and F Hiraldo. 1989. Comparative food habits of three carnivores in Western Sierra Madre, Mexico. Z. Saugetierkunde, 54:107—110. Fritzell, E. K., and K. J. Haroldson. 1982. Urocyon cinereoargenteus. Amer. Soc. of Mamm. Spec. Publ. No. 189. 8pgs. Grinnell, J., and T. I. Storer. 1924. Animal life in the Yosemite: an account of the mammals, birds, reptiles, and amphibians in a cross-section of the Sierra Nevada. University of California Press, Berkeley. 720pgs. Hatfield, D. M. 1939. Winter food habits of foxes in Minnesota. J. of Mamm. 20:202—206. Horn, M. 1997. Evidence for dispersal of fig seed by the fruit-eating characid fish Brycon guatema- lensis Regan in a Costa Rican tropical rain forest. Oecologia, 109:259—264. Jaslow, C. R. 1987. Morphology and digestive efficiency of Red Foxes (Vulpes vulpes) and Grey Foxes (Urocyon cinereoargenteus) in relation to diet. Can. J. of Zool. 65:72—78. Krefting, L. W., and E. I. Roe. 1949. The role of some birds and mammals in seed germination. Ecol. Monog. 19:270—286. Korschegen, L. J. 1957. Food habits of coyotes, foxes, house cats, and bobcats in Missouri. Missouri Cons. Comm. PR Ser. 15:1—64. 126 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Lieberman, D., J. B. Hall, M. D. Swaine and M. Lieberman. 1979. Seed dispersal by baboons in the Shai Hills, Ghana. Ecology 60:65—75. Martin, A. C., and W. D. Barkley. 1961. Seed identification manual. University of California Press, Berkeley, California. 221pp. Murray, G. K., S. Russell, C. M. Picone, K. Winnett-Murray, W. Sherwood, and M. L. Kuhlmann. 1994. Fruit laxatives and seed passage rates in frugivores: consequences for plant reproductive success. Ecology, 989-994. Munz, P. A., and D. D. Keck. 1965. A California flora. University of California Press, Berkeley. 1681 pp. Nelson, A. L. 1933. A preliminary report on the winter food of Virginia foxes. J. of Mamm., 14:40—43. Pequegnat, W. E. 1951. The biota of the Santa Ana Mountains. J. of Entomol. and Zool., 42:1—84. Putz, E E. 1993. On diarrhea. Bull. Ecol. Soc. of Ame., 74:177. Rogers, L. L., and R. D. Applegate. 1983. Dispersal of fruit seeds by black bears. J. Mamm., 64:310— SL. Schopmeyer, C. S. 1974. Seeds of woody plants in the United States. Agriculture Handbook No. 450, Forest Service, U.S. Department of Agriculture, Washington DC. Schupp, E. W. 1993. Quantity, Quality, and Effectiveness of seed dispersal by animals. Vegetatio 108: 15-29. Traveset, A., and M. Willson. 1997. Effect of birds and bears on seed germination of fleshy-fruited plants in temperate rainforests of southeast Alaska. OIKOS 80:89—95. Weintraub, J. D., and G. Shockley. 1980. Use of incisors to identify rodent Genera in owl pellets. Bull. So. Calif. Acad. Sci., 79:127—129. Willson, M. 1992. Mammals as seed-dispersal mutualists in North America. OIKOS 67:159-174. Wood, J. E. 1954. Food habits of furbearers of the upland post-oak region in Texas. J. Mamm., 35: 406-415. Wood, J. E., D. E. Davis, and E. V. Komarek. 1958. The distribution of fox populations in relation to vegetation in Southern Georgia. Ecology, 39:160—162. Zhang, S. Y. 1995. Activity and ranging patterns in relation to fruit utilization by brown capuchins in French-Guiana. Internat. J. of Primatol. 16:489—507. Accepted for publication 19 January 1999 Bull. Southern California Acad. Sci. 98(3), 1999, pp. 127-130 © Southern California Academy of Sciences, 1999 Records of the Oarfish Regalecus glesne Ascanius, 1772 in the Eastern Pacific Ocean Felipe Galvan-Magana, L. Andres Abitia-Cardenas and Francisco J. Gutiérrez-Sanchez Centro Interdisciplinario de Ciencias Marinas, Apdo. Postal 592, La Paz, B.C.S. México The monotypic teleost genus Regalecus has a worldwide distribution, mainly in tropical and temperate seas. Although other species have been named in this genus (e.g. Mori 1956; Nishimura 1960; Scott et al. 1980; Lindberg et al. 1980; Fujii 1984; Chavez et al. 1985), it seems likely that Regalecus glesne is the only valid species (Smith and Heemstra 1986; Nelson 1994). The other monotypic regalecid genus, Agrostichthys is represented by Agrostichthys. parkeri in the southeast Atlantic, and off New Zealand and Tasmania. Regalecus glesne has been recorded from the Atlantic Ocean and Mediterranean Sea (Palmer 1973), the In- dian Ocean, and the eastern Pacific (Fitch 1951; Fitch and Lavenberg 1968; Miller and Lea 1972; Hubbs et al. 1979; Eschmeyer et al. 1983). It is believed to inhabit the mesopelagic zone but occasionally is found in the upper level of the epipelagic zone. It is a rare species and almost all records are based on dying or dead, often beached, individuals (Fitch and Lavenberg 1968; Smith and Heemstra 1986). We report unpublished records of 31 Regalecus glesne collected in the Eastern Pacific from 1961 to 1997 (Fig. 1). The fishes were collected by net (one juvenile), La Paz Bay Fig. |. Records of Regalecus glesne in the Eastern Pacific Ocean. 27 128 Table 1. SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Records of Regalecus glesne in the Eastern Pacific Ocean. SIO = Scripps Institution of Oceanography; CICIMAR = Centro Interdisciplinario de Ciencias Marinas; UABCS = Universidad Aut6noma de Baja California Sur. Size Locality Latitude Longitude Date mm Reference 80 mi SW of San Diego 32°45'N 117°13'W 07 Aug 1961 3540 SIO 61-389 Eastern Pacific 06°38'S. 118°59'W 29 Feb 1968 62 SIO 96-31 Eastern Pacific 02°16’S_ 100°34.7'W 26 Nov 1969 590 SIO 69-496 Bahia de La Paz 24°20’N 110°20'W 1979 ? CICIMAR file Bahia de La Paz 24°15’'N 110°20'W = 1982 if CICIMAR file Pichilingue, B.C.S. 24°17'N 110°19'W_ O1 Jul 1984 1970 UABCS(0752) Playa Tecolote, B.C.S. 24°21'N 110°18’W Ol May 1985 3000 CICIMAR file Pichilingue, B.C.S. 24°17'N 110°19'W 06 May 1985 3000 CICIMAR file Playa Tecolote, B.C.S. 24°21’'N 110°18’'W 16 Jun 1985 3930 CICIMAR file S. of San Juan Seamount 32°30'N 121°00’W 42 Jan 1986 4330 SIO 86-1 Bahia de La Paz 24°16’N 110°19'W 1987 ? CICIMAR file Ensenada de La Paz 24°12’N 110°18’'W 28 May 1988 4700 CICIMAR file Playa del Tesoro, B.C.S. 24°16’N 110°19’W 28 May 1988 ? CICIMAR file Punta Colorada, B.C.S. 24°13’N 110°18’'W O1 Jun 1988 4900 UABCS file El Quelele, B.C.S. 24°12’N 110°30'W 02 May 1989 4500 CICIMAR file Bahia de Tenacatita, Jalisco 19°10'N _ 104°50'W_ O01 Apr 1991 4000 CICIMAR file Isla Espiritu Santo, B.C.S. 24°24’N_ 110°20'W_ 16 July 1991 5650 CICIMAR file Punta Prieta, B.C.S. 24°16'N_ 110°19'W_s17 July 1991 4000 CICIMAR file El Mechudo, B.C.S. 24°48’N_ 110°40'W_ 16 Mar 1993. 4780 CICIMAR file Yelapa, Pto. Vallarta, Jalisco 20°30’N_ 105°25’W_ Sept 1993 2000 CICIMAR file Compostela, Nayarit 21°10’N 105°20'W May 1994 2800 CICIMAR file Bahia de La Paz 24°12’N_ 110°30'W Aug 1994 5500 CICIMAR file Los Barriles, B.C.S. 23°40’N_ 109°41’'W 12 July 1995 3720 CICIMAR file Los Barriles, B.C.S. 23°40'N 109°41’W 12 July 1995 5090 CICIMAR file Sea of Cortez 23°40’N_ 109°30’W_ 18 Sept 1995 5000 CICIMAR file Isla Espiritu Santo, B.C.S. 24°24’N_ 110°20’'W 08 Dec 1995 3000 CICIMAR file Puerto Vallarta, Jalisco 20°30’N_ 105°25'Ws. 25 Jan 1996 4050 CICIMAR file Puerto Vallarta, Jalisco 20°30’'N 109°25'W 25 Jan 1996 4080 CICIMAR file Isla Espiritu Santo, B.C.S. 24°24'N 110°20’W_ 16 July 1996 5300 UABCS file Coronado:US Naval Special Warfare 32°40’N 117°14’W_ 19 Sept 1996 7300 SIO 96-82 Ca 17 miles W of Point Loma 32°41’N 117°15’W 26 Oct 1997 5000 SIO 97-226 or beached or weakly swimming from waters close to California to oceanic waters just south of the equator (Table 1). Interestingly the species becomes stranded during the same seasons, the season that these organisms appear in winter through early summer, in the eastern Pacific, in Japanese waters (Nishimura 1962), and on the western coast of Florida (Sal- oman et al. 1973). Nishimura (1962) analyzed the temperature-salinity relationship of the water mass from which oarfish were supposedly taken and suggested that this species inhabits depths between 100—200 m to 400—700 m in the western North Pacific and from 25-50 m to 150—200 m in the Japan Sea. The Gulf of California has deep water close to the tip of the Baja California peninsula, with 3,000 m depths fairly near the coast (Thomson et al. 1979). There are deep chan- nels, 400—700 m, close to Bahia de La Paz and to Cerralvo Island, Espiritu Santo Island, and Los Barriles. Similarly, there are deep waters close to the California and equatorial localities where these fishes were’ collected. Of the 15 oarfish found in Florida between 1920 and 1967 (Taylor and Saloman RECORDS OF REGALECUS GLESNE 129 1968), only one was caught with a net (to 450 m depth) and most were in poor condition. There have been many hypotheses proposed (e.g., parasites, red tide, predation, abrupt temperature changes, storms, and migration for reproduction) for the moribund condition of this species (Goode and Bean 1895; Walters 1959; Hutton 1961; Serventy 1966; Hulley and Rau 1969; Parin 1970), but no substan- tial conclusions. We propose to do more detailed taxonomic and genetic studies in the future to learn more about this rare species. In the southern Gulf of California, the average occurrence is one or two per year, which affords more opportunities to study this species. Acknowledgements We wish to thank the Instituto Politécnico Nacional. Comision de Operacion y Fomento de Actividades Academicas for their support. Also thanks to Dr. Richard Rosenblatt and H.J. Walker Jr. (Scripps Institution of Oceanography) for the data of Regalecus glesne in the SIO-Fish Collection, and the review of the manuscript. Victor Gomez and Jon Elorduy (CICIMAR), Enrique Gonzalez (UABCS) and Ignacio Peha (SEMARNAP) for the information of some specimens. We appre- ciate the review of two anonymous reviewers. Literature Cited Chavez, H., EF Galvan M. and J. R. Torres V. 1985. Primer registro de Regalecus russellii (Shaw) (Pisces:Regalecidae) de aguas mexicanas. Inv. Mar. CICIMAR. 2(2):105—112. Eschmeyer, W. N., E. S. Herald and H. Hammann. 1983. A field guide to Pacific Coast Fishes of North America from the Gulf of Alaska to Baja California. Houghton Mifflin Company, Boston. 336 pp. Fitch, J. E. 1951. Studies and notes on some California marine fishes. Calif. Dept. Fish. and Game. 37(2):111—120. Fitch, J. E. and R. J. Lavenberg. 1968. Deep-water fishes of California. University of California Press, California Natural History guides 25. Berkeley and Los Angeles. 155 pp. Fujii, E. 1984. Family Regalecidae. in Masuda, H. K. Awaoka, C. Araga, T. Uyeno and T. Yoshino (eds.). The fishes of the Japanese archipelago. Tokai University Press, Tokyo, Japan. I: xxii and 437 pp: II: 370 plates. Goode, G. B. and T. H. Bean. 1895. Oceanic ichthyology. Spec. Bull. U.S. Nat. Mus. 533 pp. Hubbs, C. L., W. I. Follet and L. J. Dempster. 1979. List of the fishes of California. Ocais. Papers Cali Acad. Sci. 133: 51 pp. Hulley, P A. and R. E. Rau. 1969. A female Regalecus glesne from Cape Province, South Africa. Copeia 1969(4):835—839. Hutton, R. EF 1961. A plerocercoid (Cestoda: Tetraphyllidea) from the oar-fish, Regalecus glesne (Ascanius), with notes on the biology of the oar-fish. Bull. Mar. Sci. Gulf. Carib. 11(2):309-— ails. Lindberg, G. U., A. S. Heard and T. S. Rass. 1980. Diccionario de nombres de peces marinos comunes de la fauna mundial. Ministry of fisheries of the USSR. Academy of Sciences, Ed. Nauka, Leningrado, USSR. 562 pp. Miller, D. J. and R. N. Lea. 1972. Guide to the coastal marine fishes of California. Calif. Dept. Fish and Game. Fish. Bull. 157:249 pp. Mori, T. 1956. Note on a rare fish Regalecus russellii (Shaw), Regalecidae, from Southern Japan Sea. Hyogo, Prefecture, University of Agriculture, Science Reports. Series, Natural Science. 2(2): 33--36. Nelson, J. S. 1994. Fishes of the world. Third edition John Wiley and Sons, New York. 600 pp. Nishimura, S. 1960. A record of Regalecus russellii (Shaw) from the Sado Straits in the Japan Sea. Ann. Rept. Jap. Sea Reg. Fish Res. Lab. 6:58—68. 130 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Nishimura, S. 1962. Records of the oar-fish in Japanese waters, with notes on some aspects of its distribution. Science Report of the Yokosuka city Museum 7:11-—22. Palmer, G. 1973. Regalecidae. in Hireau, J. C. and Th. Monod (Eds.). Check-list of the fishes of the North-eastern Atlantic and of the mediterranean (CLOFNAM I). Paris. Unesco, xxii and 683 pp. Parin, N. V. 1970. Ichthyofauna of the epipelagic zone. Israel Program for Scientific Translation, Jerusalem, 111 and 206 pp. Saloman, C. H., M. A. Moe and J. L. Taylor. 1973. Observations on a female oarfish (Regalecus glesne). Florida Sci. 36(2—4):187—189. Scott, T. D., C. J. M. Glover and R. V. Southcott. 1980. The marine and freshwater fishes of South Australia. D. J. Woolman, Govi. Printer, South Australia, 392 pp. Serventy, V. 1966. Strange creature of the sea. Pacif. Disc. 19(3):12-15. Smith, M. M. and P. C. Heemstra. 1986. Smith’s Sea Fishes. Springer Verlag. 1047 pp. Taylor, J. L. and C. H. Saloman. 1968. The oarfish Regalecus glesne: a new occurrence and previous records from the Gulf of Mexico. Copeia 1968(2):404—405. Thomson, D., L. Findley and A. Kerstitch. 1979. Reef Fishes of the Sea of Cortez. John Wiley & Sons Inc. New York. 302 p. Walters, V. 1959. The sea serpent that is a fish. Sea frontiers. 5(2):102—104. Accepted for publication 13 March 1999 Bull. Southern California Acad. Sci. 98(3), 1999, pp. 131-136 © Southern California Academy of Sciences, 1999 First Records of ‘Two Tropical Gobies, Awaous tajasica and Ctenogobius sagittula (Pisces: Gobiidae), in the Continental Waters of Baja California, México Gorgonio Ruiz-Campos*, José Luis Castro-Aguirre**, Salvador Gonzaélez-Guzman*, and Sergio Sanchez-Gonziales* *Facultad de Ciencias, Universidad Aut6noma de Baja California, Apdo. Postal 1653, Ensenada, Baja California, 22800, México U.S. Mailing: P.O. Box 189003-064, Coronado, California 92178 ** Departamento de Pesqueritas y Biologia Marina, Centro Interdisciplinario de Ciencias Marinas, Instituto Politécnico Nacional. Apdo. Postal 592, La Paz, Baja California Sur, 23000, México On 23 August and 10 October 1998, two species of gobies (Awaous tajasica and Ctenogobius sagittula), both of Tropical affinity (Follett 1960; Castro-A guirre 1978), were collected in the lower Rio [Arroyo] San Fernando, between Punta San Antonio and Punta San Fernando, Baja California (B.C.), México (29° 43’ 33.7" N, 115° 38’ 49.6” W, Fig. 1). These are the first records of these species in the continental waters of the state of Baja California; a new northern distributional record for A. tajasica, and the first mainland record in the peninsula of Baja California for C. sagittula. Specimens are deposited in the fish collection of the Facultad de Ciencias, Universidad Aut6noma de Baja California (UABOC), at En- senada. The Rio San Fernando (INEGI 1995) is in the northernmost drainage of the Vizcaino faunal district (Nelson 1921). This, and many other sites of this district have not been previously sampled for fishes, as reflected by the absence of con- tinental fish records from south of Rio El Rosario to Rio Santo Dominguito (Fol- lett 1960; Castro-Aguirre 1978; Ruiz-Campos and Contreras-Balderas 1987). The lower Rio San Fernando was sampled with minnow traps, minnow seine and experimental gill net from its mouth to 1 km upstream. Its mouth is blocked by a wide bar of sand and boulders at the high tide level, which impounds a slough approximately 200 m long and 60 m wide. The salinity of the water was measured with an Hydrolab Scout 2 equipment (Hydrolab Co., Austin, Texas) at different points along the slough. The slough is vegetated by dense macrophytes (Chara sp.) and bordered by saltmarsh vegetation (e.g., Salicornia bigelovii, Dis- tichlis spicata, and Juncus acutus). Awaous tajasica (Lichtenstein 1822) Previous range.—This amphiamerican species (Castro-Aguirre et al. 1999) is distributed in the Pacific drainage from the Gulf of California (lower Rio Yaqui at Sonora, México; Hendrickson et al. 1980) to Peri (Miller 1966; Castro-Aguirre 1978). In the peninsula of Baja California, this goby has been previously recorded at: town of La Purisima [at stream of the same name] (Evermann 1908); ‘“‘Boca 131 132 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES US ane MEXICO @ Previous records Mi New record Fig. 1. Previous and new records of Awaous tajasica (Pisces: Gobiidae) in the peninsula of Baja California, México. (1) Arroyo La Purisima at town of La Purisima, (2) Arroyo Las Pocitas at Pozas del Vado, (3) lower Arroyo Grande ca. Todos Santos, (4) Boca de la Sierra at Santiago, and (5) lower Rio San José del Cabo at San José del Cabo. de la Sierra at Santiago”’ [stream of] (De Buen 1942); Rio San José del Cabo [lower part] (Follett 1960; Castro-Aguirre, 1978); Arroyo Las Pocitas [at “‘Pozas del Vado’’] (Espinosa Pérez and Castro-Aguirre 1996); and recently, in the lower part of Arroyo Grande ca. Todos Santos (UABC-771, G. Ruiz-Campos, unpub- lished data) (Fig. 1). Material examined.—Thirty six specimens from three sites in the lower Rio San Fernando, B.C.: UABC-832 (n = 7, 69.0—79.7 mm standard length [SL]), ca. 200 m above mouth (salinity 4.1 ppt), 23 August 1998; UABC-836 (n = 3, 84.0— 98.0 mm SL), ca. 50 m above mouth (salinity 4.3 ppt), 10 October 1998; and UABC-838 (n = 26, 32.3-96.0 mm SL), ca. 1 km upstream (salinity 3.9 ppt), 10 October 1998. Description.—This goby (Fig. 2-A) has the following identifying characters (cf. Jordan and Evermann 1896-1900; Castro-Aguirre 1978): body compressed posteriorly, rather depressed anteriorly; head broader than deep; eyes small, less than interorbital width; mouth large, horizontal, maxillary extending to below anterior part of orbit in adult male, shorter in young; inner edge of the shoulder CONTINENTAL RECORDS OF GOBIES IN BAJA, CALIFORNIA | ae | BNR12 eros Pupry WURDRAReceeeeeerconcusccc Ung oe ca Bic ie oz siz We 20 tie He Oi rr wali rina etucaba lata cake hnivluny Hill bibivarlaaisiicitiisluital sae ry T | IT] y ‘| 4 gue INCHES ole. st se " O1z iz, vie c12 ez , Heng oe toe we rel or se we cee oe te Fig. 2. (A) Awaous tajasica (Lichtenstein, 1822) (UABC-838) and (B) Ctenogobius sagittula (Gunther, 1861) (UABC-837). 134 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES girdle with 2 or 3 rather large papillae; dorsal I-11, anal 11; 59 to 68 scales in lateral series; head length 6.0 times eye diameter; body olivaceous with a series of roundish blotches along the middle of side; dark narrow streaks radiating from eye; and a blackish streak crossing the upper edge of the opercle. Comments.—Two species of Awaous are currently recognized for Tropical America (cf. Castro-Aguirre et al. 1999): Awaous tajasica (Lichtenstein 1822) of amphiamerican distribution, with its junior synonym A. transandeanus (Giinther 1861) [previously assumed to be exclusively from the Pacific drainage] and Awaous banana (Cuvier and Valenciennes 1837), which is confined to the Guy- anas and Brazil. Comparative biochemical or molecular study of populations of ‘A. tajasica”’ from Pacific and Atlantic drainages might clarify relationships and taxonomy. Rio San Fernando is approximately 533 km north of the nearest pre- viously known locality in the peninsula of Baja California (Follett 1960). The finding of this ubiquitous goby (Castro-Aguirre 1978) as far north as Rio San Fernando is likely correlated to the presence of a surface warm water mass related to the “El Nifio”’ oceanographic event. Ctenogobius sagittula (Giinther 1861) Previous range.—From Gulf of California and adjacent waters south to Panama (Jordan and Evermann 1896—1900), with disjunctive coastal records in San Ig- nacio lagoon, B.C.S. (De La Cruz-Agiiero and Cota-Gémez 1998), San Quintin Bay, B.C. (Rosales-Casian 1996), and San Diego Bay, California (Miller and Lea 1972). Previous mainland México records of longtail goby (C. sagittula) are La- guna Caimanero and Escuinapa (Sinaloa), Mar Muerto (Chiapas), and Lagunas Oriental and Occidental at Oaxaca (Castro-Aguirre 1978). It had not been pre- viously reported in continental waters of the peninsula of Baja California (cf. Follett 1960; Castro-Aguirre 1978). Material examined.—Two specimens from the lower Rio San Fernando, B.C-.: UABC-837 (53.2—60.4 mm SL), ca. 200 m above mouth (salinity 3.9 ppt), 10 October 1998. Description.—This taxon (Fig. 2-B) is distinguished by the following character- istics (Jordan and Evermann 1896—1900; Miller and Lea 1972): slender body, quite sharpened from middle of first dorsal to caudal, most compressed posteriorly; cau- dal fin longer than head; body depth 6.1 to 6.3 times in standard length; short head, depressed, and broad; large mouth, nearly horizontal, the maxillary in adults ex- tending beyond middle of eye; teeth in a narrow band in each jaw, those in lower jaw uniform, and the outer series in upper jaw very enlarged and separated by an interspace from the inner band; 65 to 66 scales in a longitudinal series; first and second dorsal fins with 6 soft spines and 13 rays, respectively; anal rays (14); body light yellow-brown with five elongated black blotches on sides. Comments.—This Panamanian gobiid fish is a senior synonym of Gobionellus longicauda Girard 1858 (Robins and Lachner 1966) and Gobius longicauda Jen- kins and Evermarin 1889 (cf. Castro-Aguirre 1978; Castro-Aguirre et al. 1999; and Watson and Horsthemke 1995), which was first reported as far north as San Diego Bay (southern California) in the early 1900’s (Starks and Morris 1906), but without subsequent records (Swift et al. 1993). Its current status in California is considered as extirpated (Swift et al. 1993). Recently, this goby was reported CONTINENTAL RECORDS OF GOBIES IN BAJA, CALIFORNIA 135 in San Ignacio lagoon (De La Cruz-Agiiero and Cota-Gé6mez 1998) and San Quintin Bay (Rosales-Casian 1996). Acknowledgments Our sincere thanks go to German Ruiz-Cota and Juan Diego Flores for their valuable sampling help. Edwin (Phil) Pister and two anonymous reviewers made helpful comments on the manuscript. This contribution was supported by project CONACYT 431100-5-1993PN. Literature Cited Castro-Aguirre, J.L. 1978. Catalogo sistematico de los peces marinos que penetran a las aguas con- tinentales de México con aspectos zoogeograficos y ecol6gicos. Direcci6n General del Instituto Nacional de Pesca, México. Serie Cientifica 19:XI+298 pp. Castro-Aguirre, J.L., H. Espinosa Pérez, and J.J. Schmitter-Soto. 1999. Ictiofauna estuarino-lagunar y vicaria de México, Limusa, S.A., México. (In Press). Cuvier, G, and A. Valenciennes. 1837. Histoire naturelle des poissons. Tome douziéme. Suite du livre quatorzieme. Goboides. Liver quinzieme. Acanthoptérygiens a pectorales pédiculées. Vol. 12: i+xxiv + 1-507 + 1 p., pls. 344-368. De Buen, E 1942. Segunda contribucion al estudio de la ictiologia mexicana. Investigaciones de la Estacion Limnoldgica de Patzcuaro, México, 2:25—55. De La Cruz-Agiiero, J., and V.M. Cota-Gomez. 1998. Ictiofauna de la laguna de San Ignacio, Baja California Sur, México: nuevos registros y ampliaciones de ambito. Ciencias Marinas, 24:353— 358: Espinosa Pérez, H., and J.L. Castro-Aguirre. 1996. A new freshwater clingfish (Pisces: Gobiidae) from Baja California Sur, México. Bull. So. Calif. Acad. Sci., 95:120—126. Evermann, B.W. 1908. Descriptions of a new species of trout (Salmo nelsoni) and a new cyprinodont (Fundulus meeki) with notes on other fishes from Lower California. Proc. Biol. Soc. Wash., 21:19-30. Follett, W.I. 1960. The fresh-water fishes—their origins and affinities. Symposium on the biogeography of Baja California and adjacent seas. Syst. Zool., 9(3—4):212—232. Girard, C.E 1858. Notes upon various genera and new species of fishes, in the museum of the Smith- sonian Institution, and collected with the United States and Mexican boundary survey: Major William Emory, Commissioner. Proc. Acad. Nat. Sci. Phila., 10:167—171. Giinther, A. 1861. Catalogue of the fishes in the British Museum. Catalogue of the acanthopterygian fishes in the collection of the British Museum. Gobiidae . . .[to]. .. Notacanthi. Vol., 3:i-xxv + 1-586 + i-x. Hendrickson, D.A., W.L. Minckley, R.R. Miller, D.J. Siebert, and P. Haddock Minckley. 1980. Fishes of the Rio Yaqui basin, México and United States. J. Arizona-Nevada Acad. Sci., 15:65—106. INEGI [Instituto Nacional de Estadistica, Geografia e Informatica]. 1995. Estudio hidroldgico del Estado de Baja California. INEGI, México, D.F 180 pp. Jenkins, O.P., and B.W. Evermann. 1889. Description of eighteen new species of fishes from the Gulf of California. Proc. U.S.N.M. 11(698):137—158. Jordan, D.S., and B.W. Evermann. 1896—1900. The fishes of north and middle America. Bull. 47, U.S.N.M. (part IID: i-xxiv + 2183a—3136. Lichtenstein, M.H.C. 1822. Die Werke von Marcgrave und Piso Uber die Naturgeschichte Brasiliens, erlautert aus den wieder aufgefundenen Original-Abbildungen. Abh. Akad. Wiss. Berlin, 1820— 21:267-288. Miller, R.R. 1966. Geographical distribution of Central American freshwater fishes. Copeia, 1966:773— 801. Miller, D.J., and R.N. Lea. 1972. Guide to the coastal marine fishes of California. California Depart. Fish and Game, Fish Bull. 157:1—249. Nelson, E.W. 1921. Lower California and its natural resources. Mem. Nat. Acad. Sci., 16:1—194. Robins, C.R., and E.A. Lachner. 1966. The status of Ctenogobius Gill (Pisces: Gobiidae). Copeia, 1966:867-869. 136 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Rosales-Casian, J.A. 1996. Ictiofauna de la Bahia San Quintin, Baja California, México, y su costa adyacente. Ciencias Marinas, 22:443—458. Ruiz-Campos, G, and S. Contreras-Balderas. 1987. Ecological and zoogeographical checklist of the continental fishes of the Baja California peninsula, Mexico. Proc. Desert Fishes Council, 17: 105-117. Starks, E.C., and E.L. Morris. 1906. The marine fishes of southern California. Univ. Calif. Pub. Zool., 3:159-251. Swift, C.C., TR. Haglund, M. Ruiz, and R.N. Fisher. 1993. The status and distribution of the freshwater fishes of southern California. Bull. So. Calif. Acad. Sci., 92:101—167. Watson, R.E., and H. Horsthemke. 1995. Revision of Euctenogobius, a monotypic subgenus of Awaous, with discussion of its natural history (Teleostei: Gobiidae). Rev. Fr. Aquariol., 22:83—92. Accepted for publication 14 March 1999 Bull. Southern California Acad. Sci. 98(3), 1999, pp. 137-138 © Southern California Academy of Sciences, 1999 INDEX TO VOLUME 98 Abitia-Cardenas, Andres, see Felipe Galvan-Magafia Allen, Larry G., see Gregory J. Tranah Armitage, Mark H.: The Euryhaline Gobiid Fish, Gillichthys mirabilis Cooper 1864, Second Intermediate Host of the Trematode, Pygiopsoides spindalis Martin 1951, 75 Castro-Aguirre, José Luis, See Gorgonio Ruiz-Campos Benham, Steven R., see Richard L. Squires Bursey, Charles R., see Stephen R. Goldberg Erickson, Richard A.: Identification and Distribution of Spiny Pocket Mice (Chae- todipus) in Cismontane Southern California, 57 Galvan-Magania, Felipe: Records of the Oarfish Regalecus glesne Ascanius, 1772 in the Eastern Pacific Ocean, 127 Gerlinger, Tom, see Don Maurer Goedert, James L., see Richard L. Squires Goldberg, Stephen R.: Helminths of the Western toad, Bufo boreas (Bufonidae) from Southern California, 39 Gonzalez-Guzman, Salvador, see Gorgonio Ruiz-Campos Grismer, L. Lee: Checklist of Amphibians and Reptiles on Islands in the Gulf of California, Mexico, 45 Gutiérrez-Sanchez, Francisco J., see Felipe Galvan-Magafia Hernandez, Sonia, see Stephen R. Goldberg Heyning, John E., see Joel W. Martin Kelley, Thomas S.: A Hemphillian (Late Miocene) Mammalian Fauna from the Desert Mountains, West Central Nevada, 1 Luganski, Thomas P., see Thomas S. Kelley Martin, Joel W.: First Record of Jsocyamus kogiae Sedlak-Weinstein, 1992 (Crus- tacea, Amphipoda, Cyamidae) from the Eastern Pacific, with Comments on Morphological Characters, a Key to the Genera of the Cyamidae, and a Checklist of Cyamids and Their Hosts, 26 Martinez-Mufioz, Marco A.: Growth and Mortality of the Fantail Sole, Xystreurys liolepis (Jordan and Gilbert 1881) off the Western Coast of Baja California, Mexico, 66 Maurer, Don: Rare Species as Bioindicators in Marine Monitoring, 91 Ortega-Salas, A. A., see Marco A. Martinez-Munoz Patten, Michael A., see Richard A. Erickson 137 138 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Robertson, George, see Don Maurer Ruiz-Campos, Gorgonio: First Records of Two Tropical Gobies, Awaous tajasica and Ctenogobius sagittula (Pisces: Gobiidae), in the Continental Waters of Baja California, Mexico, 131 Sanchez-Gonzales, Sergio, see Gorgonio Ruiz-Campos Squires, Richard L.: First Fossil Record of the Pteropod Limacina from the Pacific Coast of North America, 80 Stadum, Carol J.: Fossil Wood from the Middle Miocene Conejo Volcanics, Santa Monica Mountains, California, 15 Thomas, Barry, see James A. Wilson Tranah, Gregory J.: Morphologic and Genetic Variation Among Six Populations of the Spotted Sand Bass, Paralabrax maculatofasciatus, from Southern Cal- ifornia to the Upper Sea of Cortez, 103 Weigand, Peter W., see Carol J. Stadum Wilson, James A.: Diet and Seed Dispersal Efficiency of the Gray Fox (Urocyon cinereoargenteus) in Chaparral, 119 1 a & INSTRUCTIONS FOR AUTHORS The BULLETIN is published three times each year (April, August, and December) and includes articles in English in any field of science with an emphasis on the southern California area. Manuscripts submitted for publication should contain results of original research, embrace sound principles of scientific investigation, and present data in a clear and concise manner. The current AIBS Style Manual for Biological Journals is recommended as a guide for contributors. Consult also recent issues of the BULLETIN. MANUSCRIPT PREPARATION The author should submit at /east two additional copies with the original, on 8% X 11 opaque, nonerasable paper, double spacing the entire manuscript. Do not break words at right-hand margin anywhere in the manuscript. Footnotes should be avoided. 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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 Rare Species as Bioindicators in Marine Monitoring. Don Maurer, Tom Gerlinger and George Robertson: 0-7 ee Morphologic and Genetic Variation Among Six Populations of the Spotted Sand Bass, Paralabrax maculatofasciatus, from Southern California to the Upper Sea of Cortez. Gregory J. Tranah and Larry G. Allen Diet and Seed Dispersal Efficiency of the Gray Fox (Urocyon cinereoargenteus) in Chaparral. James A. Wilson and Barry Thomas Records of the Oarfish Regalecus glesne Ascanius, 1772 in the Eastern Pacific Ocean. Felipe Galvan-Magana, L. Andres Abitia-Cardenas and Francisco J. Gutiérrez-Sanchez:' 00.0.0 Se First Records of Two Tropical Gobies, Awaous tajasica and Ctenogobius sagittula (Pisces: Gobtidae), in the Continental Waters of Baja California, México. Gorgonio Ruiz-Campos, José Luis Castro- Aguirre, Salvador Gonzalez-Guzman and Sergio Sanchez-Gonzales __. Index to Volume OS. ae COVER: Logo of the Southern California Academy of Sciences. 91 103 i 127