Contributions in Science, 524:31-49 18 November 2016 Paleontology and Stratigraphy of the Miocene Saddleback Valley Limestone, Orange County, Southern California 1 Carol J. Stadum 2 and Kenneth L. Finger 3,4 ABSTRACT. Although rare in California, Neogene limestone outcrops are well exposed within a 10 km 2 4 area of western Saddleback Valley in southern Orange County. These occur as inconsistent, discontiguous sequences of micrite, calcarenite, and limey sands that are informally referred to herein as the Saddleback Valley limestone. Its principal biotic components are cheilostome bryozoans, coralline red algae, and mollusks characteristic of the middle Miocene “Temblor” California Provincial Molluscan Stage (CPMS); hence, previous workers had referred to these beds as the “Temblor” limestone. In the Los Angeles Basin, the shallow-marine “Temblor” macrofauna also occurs in the Topanga Canyon Formation of the Santa Monica Mountains and the Topanga Formation in the Santa Ana Mountains. Nevertheless, geologists mapping the San Juan Capistrano and El Toro quadrangles placed the Saddleback Valley limestone at the base of the local Monterey Formation, presumably on the basis of undescribed field observations. This study combines lithostratigraphic and paleontologic data, including micropaleontology, to determine whether this recently challenged affiliation is justified. Microfossils (ostracodes, foraminifera, and calcareous nannoplankton) in the Saddleback Valley limestone indicate that its conspicuous shallow-water “Temblor” fauna and associated Topanga-like sediments had been transported downslope in the late Relizian (late early Miocene) at —16 Ma. 8 Sr/ 86 Sr analyses confirm the age at 15.9—16.5 My. It is therefore interpreted that the depositional history of the Saddleback Valley limestone began with a rich subtropical invertebrate community inhabiting the margins of a channel that existed between the mainland and a high-relief peninsula or island to the west. Increasing tectonic activity along the plate margin caused large amounts of schistose rock to slide off the island’s eastern slope and into the channel, forming the San Onofre Breccia. Continued subsidence of the area created a deep basin into which turbidity currents transported the unconsolidated sediments, including its “Temblor” biota, which settled on the breccia or where “Monterey” muds had started to accumulate. These displaced sediments filled slope channels that were part of a deep-sea fan complex. In the deep-water Monterey realm, the intermittent deposits of limey sands were buried by hemipelagic muds. Diagenesis transformed isolated calcarenite lenses composed predominately of calcareous algae and invertebrates into nearly pure limestone. As tbe basin emerged in the late Pleistocene, erosion of the overlying strata exposed the Saddleback Valley limestone. Its depositional history and stratigraphy imply that it is the basal “member” of the local Monterey Formation. INTRODUCTION LOCATION, GEOLOGIC SETTING, AND HISTORICAL INTEREST Saddleback Valley is located where the Santa Ana Embayment extended off the southeastern Los Angeles Basin during the Neo¬ gene in southern California (Fig. 1). The valley has been geological¬ ly described as a transgressive marine sequence of Miocene formations folded into a broad, north-trending syncline (Morton et ah, 1974). In the western part of the valley and the adjacent San Joaquin Hills, the stratigraphic succession consists of five geo¬ logic units, which, from oldest to youngest, are the Vaqueros For¬ mation, Topanga Formation, San Onofre Breccia, Monterey Formation, and Capistrano Formation (Fig. 2). Within an area of 10 km 2 , carbonate-rich beds and lenses crop out above the breccia and below the Monterey mudstones. We refer to these limey deposits collectively as the “Saddleback Valley limestone.” The earliest published account of this limestone is that of Bowers (1890:399-400), who described a quarry used to con¬ struct Mission San Juan Capistrano in the 1790s (see Figs. 1 and 3): At this place is a most remarkable bed of fossils. It is about ten miles from the ocean and nearly one and a half miles southwest of El Toro Station. An exposure 1 LJRL: www.nhm.org/scholadypublications 1 San Diego Natural History Museum, Department of Paleontology, 1788 el Prado, San Diego, California 92101, USA. 3 University of California Museum of Paleontology, 1101 Valley Life Sciences Building, Berkeley, California 94720-4780, USA. 4 Corresponding author: Kenneth L. Finger, E-mail: kfinger@berkeley.edu has occurred by excavating into the bed of fossil shells in view of burning for lime. At the exposure the stra¬ tum is about seven feet thick, dipping to the south, and can be traced for nearly a half-mile. It is composed almost wholly of bivalves, Saxidomus gracilis, largely predominating, with occasional Cardium corbis, pec- ten, and casts of univalves. The teeth of sharks are occasionally found. The casts are found in a marly sub¬ stance, which is doubtless the result of the decomposi¬ tion of their shells. They are easily dislodged with a pick, and the bed of a wagon could be filled with them as readily as coal or gravel. What is especially strange concerning them is that the bivalves lie general¬ ly on their side and were fossilized with closed shells. But how did they become heaped up in such vast num¬ bers ? Dr. J.G. Cooper suggests that it is probably the result of an earthquake and tidal wave. At present we can advance no more plausible theory to account for this vast accumulation of fossils in this spot. Still visible in some blocks of the Mission’s buildings are molds of Lyropecten crassicardo (Conrad, 1857), Amussiopecten vanv- lecki (Arnold, 1907), Saxidomus cf. S. vaqnerosensis (Arnold, 1910) (S. gracilis of Bowers, 1890). and Turritella ocoyana Conrad, 1855 (Fig. 3). In addition to these limestone blocks, the limestone was pulverized to make the mortar used in constructing the Mission (Wright, 1950). A century later, tests of Saddleback Valley Limestone showed that it would be a satisfactory source of material for Portland cement, but production never material¬ ized (Logan, 1947). © Natural History Museum of Los Angeles County, 2016 ISSN 0459-8113 ’(Print); 2165-1868 (Online) 32 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone Figure 1 Map of the Saddleback Valley study area delineating city boundaries (thick grey lines) and the three study areas with sample localities. In the 1920s, the short-lived Moulton Marine Shell Fertilizer Company was formed to mine the limestone as soil enrichment, which it marketed to local farmers (Tucker, 1925). A remnant of their activity is a small quarry that remains visible just northwest of Moulton Parkway, as shown on the geologic map of Morton etal. (1974). OCCURRENCES OF OTHER LIMESTONES IN THE MIOCENE OF CALIFORNIA The scarcity of limestone in the thick Tertiary sections of California was noted by Bramlette (1946) in his study of the Monterey For¬ mation, which reported thin beds of impure calcareous and Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone ■ 33 Figure 2 Miocene timescale (derived from Flilgen et al., 2012) with the addition of California biozonations. Shaded horizontal band denotes the age interval of the Saddleback Valley limestone (SVL) and orange sand, and that of younger Monterey mudstone in the present study. Ranges of molluscan stages derived from Prothero (2001) and McCulloh et al. (2002). Intervals of the benthic foraminiferal stages shown as solid black lines are from McDougall (2007); broader correlations with calcareous nannoplankton zones (Crouch and Bukry, 1979:fig. 3) indicated by solid grey extensions for those they range completely through and dashed lines for zones they range into but not beyond. 34 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone Figure 3 Surface of a limestone block at Mission San Juan Capistrano showing mollusk molds and bryozoans: a, cheilostome bryozoan fragments on S. vaquerosensis ; b, Entobia -bored valve of Saxidomus vaquerosensis Arnold, 1910; c, Turritella ocoyana Conrad, 1855. dolomitic rocks, or more commonly as concretions, throughout much of the unit. Bramlette (1946) generalized that limestone occurs at the base of the Monterey Formation in San Luis Obispo, Santa Barbara, Los Angeles, and Orange counties, where it appears as con¬ spicuous carbonate beds. Furthermore, he indicated that these share a fauna similar to that recovered by Arnold and Anderson (1907) in southern Orange County. He also referred to this limestone as a tran¬ sitional middle Miocene horizon in Santa Barbara County. In the lower Monterey Formation near Lompoc, Dibblee (1950) encountered limestone lenses over 30 m thick that are composed mostly of calcareous algae. Nevertheless, when shown samples of the Saddleback Valley limestone, he stated (pers. comm., 2001) that its fossil density, diversity, and preservation did not resemble anything he had observed in the decades he devoted to mapping the geology of central and southern California. In the Santa Monica Mountains, Stanton and Alderson (2013) recently described limey deposits that are interbedded with Con- ejo Volcanics and contain coralline algae, encrusting bryozoans, and the bivalves L. crassicardo and Spondylus scotti Brown and Pilsbry, 1913. GEOLOGICAL STUDIES ON THE LIMESTONE IN SADDLEBACK VALLEY Arnold and Anderson (1907) and Arnold (1909) were the first geologists to suggest an origin for the Miocene limestones that, in some areas, occur between the Vaqueros and the Monterey for¬ mations. They interpreted those carbonates as having formed pri¬ marily in semi-enclosed basins that accumulated alkaline mud. Corby (1922) described and illustrated articulated Pecten crassi¬ cardo from what he referred to as the Temblor Formation near El Toro (now Lake Forest). His locality appears likely to be coin¬ cidental with the “Pecten Reef” locale (SDSNH loc. 4520) in Lake Forest. Woodford (1925) also assigned these limestone deposits to the Temblor Formation based on faunal similarities. He described the unit as occurring between the San Onofre Breccia and the Monterey Formation throughout the Capistrano-El Toro area with a stratigraphic thickness exceeding 30 m and extending northward beyond the pinch-out of the breccia between Wood Canyon and Laguna Woods. He also observed the breccia north of Dana Point, where it intercalates with limey mudstone at the base of the Monterey Formation. Loel and Corey (1932:58) consid¬ ered the limestone fauna as a transition between those of the lower Miocene Vaqueros Formation and middle Miocene Topanga For¬ mation and San Onofre Breccia. They described it as “bryozoan- limestone reefs with P. crassicardo, Ostrea, and Scutella merriamiP Interest in the Saddleback Valley limestone was rekindled in the late 1960s, when commercial developers in Lake Forest and Laguna Hills began blasting the indurated calcarenite, forming fossil-rich spoils and fresh exposures. In 1972-73, construction east of Interstate 5 and south of Lake Forest Drive exposed lime¬ stones rich in fossils. These rocks (Fig. 4) are dominated by the middle Miocene pectinids L. crassicardo, Amusium lompocensis (Arnold, 1906), and A. vanvlecki, which are characteristic of the “Temblor” California Provincial Molluscan Stage (CPMS). Conse¬ quently, the locality became known as the “Pecten Reef” and with¬ in a three-month interval in 1973, paleontologists from the Natural History Museum of Los Angeles County (LACM) collect¬ ed more than 30,000 marine fossils (primarily echinoids, mol- lusks, and shark teeth) from this locality. Southwest of the “Pecten Reef,” Morton et al. (1974) mapped fossiliferous limestones in Laguna Woods, Laguna Hills, Aliso Viejo, and Wood Canyon and estimated the composite stratigraph¬ ic thickness of that discontinuous lithofacies as more than 120 m. Fife (1974) and Morton et al. (1974) placed the limestone at the base of the Monterey Formation; however, there has been no con¬ sensus on whether it belongs to Kleinpell’s (1938) Relizian or Lui- sian California benthic foraminiferal stage (CBFS). For example, Fife (1979) and consulting micropaleontologist R. Boettcher (pers. comm., 2001) considered the unit to be Relizian, whereas Stadum (1982) and Finger (1988) ascribed it to the Luisian. Klein¬ pell’s (1938) differentiation of the Relizian and Luisian, however, is dubious, as differences in taxa, species richness, and apparent depth are particularly subtle and interbasinally inconsistent com¬ pared with the other stages that subdivide the Miocene (Finger, 1992). It is not surprising that Vedder (1971) referred to the fora- minifera present in siltstone interbedded in the San Onofre Breccia in Shell’s Moulton No. 14 well as lower Luisian or Relizian, and that lower Luisian foraminifera occur in strata superjacent to the unit in the Aliso Creek area. Stadum (1982) wrote a thesis on the Saddleback Valley lime¬ stone and its macroinvertebrate fauna and concurred with previ¬ ous workers by referring to it as the basal Monterey Formation. However, Powell and Stadum (2010) recently refuted that correla¬ tion by reassigning the limestone to the Topanga Formation, pri¬ marily because of its “Temblor” molluscan faunal affinity. In this present study, stratigraphic affiliations are reconsidered by further investigating the lithostratigraphy and paleontology of the lime¬ stone. The primary goal, however, is to derive a better understand¬ ing of the depositional history and stratigraphic relationships of this enigmatic unit in the Neogene section of California by inte¬ grating all available data. MATERIALS AND METHODS ABBREVIATIONS Abbreviations used in this study are primarily those that prefix sample locality or specimen numbers: LACMIP, Los Angeles County, Invertebrate Paleontology; LACMVP, Los Angeles County, Vertebrate Paleontology; OCPC, Orange County Paleontological Collection (located in Santa Ana at the John D. Cooper Center for Paleontology and Archaeology, which is herein referred to as the Cooper Center); SDSNH, San Diego Society Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone ■ 35 Figure 4 Well-preserved, mostly articulated Lyropecten crassicardo (Conrad, 1857) in limestone exposed at Fossil Reef Park (SDSNH loc. 4312). Similar occurrences were found at the original “Pecten Reef” site (SDSNH loc. 4520) and near the base of the measured section in the Moulton Parkway road cut (SDSNH loc. 4521). of Natural History (San Diego Natural History Museum [SDNHM] pale¬ ontological collection). METHODS Fieldwork was intermittent during land development that spanned four dec¬ ades. Lab work included removing matrix from the macrofossils and apply¬ ing polyvinyl acetate (Vinac™) to repair and stabilize broken and fragile specimens. Unconsolidated sands were dry-sieved through a series of 20-, 32- and 100-mesh screens to isolate minute fossils, including fish bones and teeth, shell fragments, and echinoid spines. To obtain foraminifers and ostracodes, sediment samples were washed over a 200-mesh screen, then dried and picked under the microscope. Calcareous nannoplankton samples were given to consultant Stanley Kling, who used conventional methods to prepare strewn slides. Thin sections of the indurated limestones were also examined, particularly for microfossils that could be useful in interpreting the depositional history of the local geologic sequence. Latex and silicone peels were made from the molds in indurated limestone to facil¬ itate identifications. FOSSIL LOCALITIES Development has obscured or obliterated many of the exposures, but 40 acres of limestone outcrops remain preserved in the northern Aliso and Wood Canyon Wilderness Park and, in 1982, community interest in the fossiliferous limestone and the efforts of the senior author led to the preservation of a one-acre outcrop in Laguna Hills as “Fossil Reef Park” (Orange County Historical Site No. 28; Fig. 5). Earth-disturbing activities related to the ongoing development of western Saddleback Valley continue to be monitored for paleontological resources. Orange County Resolution No. 87-516, passed in 1987, stipulates that all fos¬ sils recovered from new county excavations and projects belong to Orange County. The fossiliferous outcrops in this study are divided into three areas based on their geographic distribution (Fig. 1) and paleontologic and lithostratigraphic differences (Fig. 6), and referred to herein by number: (1) Wood Canyon, (2) Laguna Hills-Aliso Viejo, and (3) Laguna Woods-Lake Forest. Field observations in each of these areas are pre¬ sented below. All of the following localities are in the Monterey Formation. The nine SDSNH and OCPC localities are in the Saddleback Valley limestone “mem¬ ber,” whereas the two UCMP localities in Area 2 are not. Geographic coor¬ dinates were derived from plotting the localities in Google Earth© utilizing “Street View” mode whenever possible for maximum accuracy. Area 1: Wood Canyon SDSNH LOC. 5752. North canyon wall off Sheep Hills, southern Aliso Creek and Wood Canyon Wilderness Area; San Juan Capistrano quad., 33°33'20.18"N, 117°44T5.33"W, elevation (elev.) 396 ft (121 m). Area 2: Laguna Hills-Aliso Viejo SDSNH LOC. 4312 (OCPC LOC. 0022). One-acre site preserved as Fossil Reef Park, Laguna Hills; San Juan Capistrano quad., 33°35'19.07"N, 117°42'20.24"W, elev. 275 ft (84 m). SDSNH LOC. 4521. Road cut along west side of Moulton Parkway, ~600 m north of Aliso Parkway, Laguna Hills; San Juan Capistrano quad., 33°35'08.74"N, 117°42'33.77"W, elev. 271 ft (83 m). Site now 36 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone Figure 5 Southward oblique aerial view of part of Area 2 showing Fossil Reef Park (SDSNH loc. 4312 [OCPC loc. 0022]) and the Moulton Parkway road cut (SDSNH loc. 4521). greatly reduced. In this report, SDSNH loc. 4521 includes sublocalities SDSNH 4521A and LH-1 to LH-4 of Finger (1988, 1992). UCMP LOC. 12871. Small exposure of steeply dipping dark brown mudstone beds on Aliso Parkway at the east corner of its intersection with Moulton Parkway; Laguna Hills; San Juan Capistrano quad., 33°35'01.07"N, 117°42'23.60"W, elev. 243 ft (74 m). Site no longer exists due to landscaping related to development of adjacent housing community. UCMP LOC. 12873. Orange sand lens (~4 m 3 ) surrounded by green¬ ish-grey mudstone, ~0.4 km northeast of SDSNH 4521, Laguna Hills; San Juan Capistrano quad., 33°35'12.32"N, 117°42'31.64"W, elev. 289 ft (88 m). MAR-254 in Finger (1988, 1992). SDSNH LOC. 6283. Weathered outcrops of Saddleback Valley limestone in North Aliso and Wood Canyon Wilderness Park, Aliso Viejo; San Juan Capistrano quad., 33°35'06.90"N, 117°42'40.42"W, elev. 291 ft (89 m). SDSNH LOC. 6284. Weathered outcrops of Saddleback Valley limestone in North Aliso and Wood Canyon Wilderness Park, Aliso Viejo; San Juan Capistrano quad., 33°35'11.01"N, 117°42'42.65"W, elev. 294 ft (90 m). SDSNH LOC. 6286. Weathered outcrops of Saddleback Valley limes¬ tone in North Aliso and Wood Canyon Wilderness Park, Aliso Viejo. San Juan Capistrano quad., 33°35'02.69"N, 117°42'47.16"W, elev. 295 ft (90 m). Area 3: Laguna Woods and Lake Forest OCPC LOC. 0872. Laguna Woods Self Storage, 24151 Moulton Park¬ way; Laguna Woods, San Juan Capistrano quad., 33°36'37.29"N, 117°43'34.58"W, elev. 389 ft (119 m). Not collected. Site destroyed in 1998. OCPC LOC. 03155. Home Depot, 24332 El Toro Road, Laguna Woods, San Juan Capistrano quad., 33°36'26.52"N, 117°42.67"W, elev. 447 ft (136 m). Site buried in 1997. SDSNH LOC. 4520 (OCPC 0023). Original “Pecten Reef” site. El Toro Quad., 33°37'49"N, 117°42'46"W, elev. ~95 m (311.6'). Site destroyed in 1974. STRATIGRAPHY AND PALEONTOLOGY Fossils characteristic of the outcrops are noted in this section. A total of 106 macroinvertebrate taxa were recovered from nine outcrops of the Saddleback Valley limestone (Table 1): 48 gastropods, 45 bivalves, 4 bryozoans, 4 echinoids, 1 poriferan, 1 polychaete, 1 cirriped, 1 ichno- taxon, and 1 coralline alga. Also present are teeth of cartilaginous and boney fish. Samples from two of the limestone localities in Area 2 were processed for microfossils—SDSNH 4312 for calcareous nannoplankton and SDSNH 4521 for both foraminifera and ostracodes. In addition, microfossil samples were collected from two non-limestone localities in Area 2 for comparative study. Photographs of 38 macroinvertebrate taxa found in this study are shown in Figures 7-52. Area 1: Wood Canyon To the west of the junction of Aliso Creek and Wood Canyon Creek near Laguna Niguel, the Saddleback Valley limestone cropped out as lenses in the north wall of Wood Canyon (Morton et al., 1974). All the Wood Can¬ yon outcrops have been destroyed by development, including the section of five lithofacies at SDSNH loc. 5752 (Fig. 1) that greatly contributed to our understanding of the limestone unit. From top to bottom (Fig. 6), these lithofacies are described below. The sequence begins with Lithofacies A, a 1.2-m-thick lag deposit of sub¬ rounded quartz and blueschist pebbles and cobbles, and abraded mollusk fragments cemented by calcium carbonate. This lithofacies unconformably overlies the >l,000-m-thick San Onofre Breccia, which is subjacent to the limestone in Area 1 and extends northward into western Area 2. Cooper (pers. comm., 1987) described the outcrop in Area 1 as flat-topped with a spoon-shaped base, indicating that it was a channel-fill deposit. From 1.2-5.7 m, Lithofacies B is a 5.4-m-thick layer of bioclast-sup- ported limestone (Fig. 53). Cooper (pers. comm., 1987) described this bed as ranging from coarse limestone (grainstone) to a fine carbonate mud (rudstone) with steinkerns of Saxidomus cf. S. vaquerosensis. The geo¬ petal steinkerns of this lithofacies readily weather, and the fossils, which are elements of a marine shelf fauna, appear to have experienced premortem Contributions in Science, Number 524 AREA 1: Wood Canyon SDSNH loc. 5752 ■14.1 Siliceous mudstone (no macrofossils} 12.1 Palaeophycus oriented horizontally; some pectens near top Coquina of articulated Saxidomus steinkerns 1.2 Conglomerate with lag cobbles & shell fragments “0 AREA 3: Lake Forest SDSNH loc. 4520 25.1 VT-t-ji-! * • ,-L . * » *!■ ■Z^-T - 77 . ’ ■ x- ■T ’ White siliceous mudstone -22.0 Vitricash White siliceous mudstone -20.5 Green mudstone Turbidites in which orange, friable, coarse sandstones alternate with silty mudstones containing fish scales - 12.0 Stadum and Finger: Saddleback Valley Limestone ■ 37 AREA 2: Moulton Parkway Roadcut, SDSNH loc. 4521 Friable sandy limestone with abundant tjz • •—• r m ^ •— • x pectens & other mollusks, Vaqueroseila, ■A' - T -. v Eucidaris spines, cartilagenous & boney fish teeth, & occasional sandy Mn nodules --f-n • A ■.A-'T'.v -0 r Normal fault — | T ■ V ■ ' ' ■ ~ ■ ■' l ' . J 7 * T - aa-taa- w+‘-h- k£dk'-^' ■i?. *W A A = _ -T •_- _u »i « • * _i* * J-.V. ■ I I I I I ea i ~j~ i i ~ iii i . -pv -a Abundant shell fragments, mostly Lyropecten crassicardo -143.5 Normal fault -53.1 (no macrofossils) “50.8 Calcareous shells & fragments 46.0 “45 5 Vaqueroseila, echinoid spines, shell fragments Lyropecten crassicardo (articulated), shell fragments, echinoids, shark teeth -39.0 Lyropecten crassicardo (articulated), L lompocensis, —36.5 echinoid spines, Vaqueroseila, shell fragments (no macrofossils) r32 5 1-32 2 Oy sters ( most| y articulated) Lyropecten lompocensis common; fragments of oysters & barnacles -28.6 ■175.6 (no macrofossils) -vitric ash (no macrofossils) -121.0 Normal fault (no macrofossils) ■ - 92.8 Normal fault Shell hash of fragmented Crassosfrea & Amusium lompocensis ; abundant Vaqueroseila 1-78.2 (Scale is 50% that of lower section) (no macrofossils) 21 q _ 2 ^ Vaqueroseila “button bed” -1-20 8 Fragmented shells & urchin tests Abundant shell fragments x 18 0 Vaqueroseila “button bed" Oyster, bryozoan, pecten, & Vaqueroseila fragments Bryozoans, pectens, shell & barnacle fragments Oyster valves Bryozoan marl 1 0.7 Eucidaris spines, Palaeophycus burrows, bryozoans, Trophon Bryozoan marl Lithology Key mudstone -A-A micrite calcarenite fine to medium calcareous sand conglomerate ’■ VA Figure 6 Examples of stratigraphic columns from each of the three designated areas in Saddleback Valley. Letters on left side of Area 1 column refer to lithofacies described in text. The heterogeneity in and between the sequences supports the hypothesis of localized pulses of transport and deposition. transportation, concentration, and resedimentation in a deeper water envi¬ ronment. A thin section of the limestone reveals foraminifera similar to those found in Area 2 (Fig. 54). Although Cooper’s interpretation of the transport and deposition of the fossil mollusks is admissible, his suggestion that they were transported postmortem is questioned. The Saxidomus vaquerosensis steinkerns are from articulated shells, which indicates that these bivalves were alive when suddenly displaced and rapidly buried by a turbidity current. Soon after death, their internal soft parts decomposed and the surrounding matrix of limey mud and bryozoan fragments infiltrated the shells and filled the voids, forming internal molds that preserve the smooth impressions of distinct muscle scars after the shells dissolved. From 5.7-10.1 m, Lithofacies C is a biosparite recognized in all three areas of Saddleback Valley as a 1.5-m-thick cheilostome bryozoan rud- stone with an irregular, somewhat erosional, basal contact. Cheilostome bryozoans typically form thickets or dense patches at outer-neritic (50- 150 m) depths on sediment-starved hard substrates where weak bottom currents limit lateral transport (Cuffey et ah, 1981). Bryozoan fragments are a major component of the Saddleback Valley limestone, but they are less apparent in the dense biosparite than in the weakly cemented limey sandstone. Broken shells of L. crassicardo and A. vanvlecki are scattered infrequently in the uppermost bryozoan biosparite, whereas whole pectinids and Crassostrea titan (Conrad, 1853) are common throughout the limestones of Areas 2 and 3. Cuffey et al. (1981) referred to the cheilostome bryozoan in the Saddleback Valley limestone as Gemelliporella aff. G. punctata Canu and Bassler, 1919, which was described from outer-neritic depths in the warm waters of the Caribbean. The Saddleback Valley taxon may be its descendant, as the limestone was deposited prior to the closing of the Panama seaway in the Pliocene. Also occurring in this limestone are the encrusting bryozoans Conopeum barbarensis (Gabb and Horn, 1862) and Smittina maccullochae Osborn, 1952, extant taxa previously unknown as fossils. Cuffey et al. (1981:70-71) stated that in “contrast to all the abundant Gemelliporella aff. G. punctata , which is a Caribbean form, these two are more expectable ‘local’ species in the southern California area.” In reference to this limestone, they also noted “few other Cenozoic deposits exhibit as high a bryozoan content as do certain of these lenses.” Living S. maccullochae inhabits shallow substrates at depths down to 35 m off the coasts of southern California and Baja California, Mexico. The presence of Gemelliporella in the Saddleback Valley lime¬ stone appears to be the first fossil occurrence of this genus in the eastern Pacific. A rhodophytic biosparite at 10.1-12.1 m is Lithofacies D. Thin sections of this limestone contain abundant thalli of the red coralline alga 38 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone Table 1 Taxonomic checklist of macrofossil taxa in the Saddleback Valley limestone. Equivalent locality numbers: SDSNH loc. 4312 = OCPC loc. 0022; SDSNH loc. 4520 = OCPC loc. 0023. _ Area 1 Area 2 Area 3 SDSNH SDSNH SDSNH SDSNH SDSNH SDSNH OCPC OCPC SDSNH 5752 4312 4521 6283 6284 6286 0872 03155 4520 Rhodophyta: Corallinales Lithophyllum cf. L. profundum Johnson, 1954 X Bryozoa: Cheilostomatida Conopeurn barbarensis (Gabb and Horn, 1862) X Gemelliporella aff. G. punctata Canu and Bassler, 1919 X Smittina tnaccuUocbae Osborn, 1952 Indeterminate bryozoan X Molhtsca: Gastropoda Antillopbos woodringi Addicott, 1970 X Calyptraea sp. X Cancellaria dalliana Anderson, 1905 X Cancellaria ocoyana Addicott, 1970 X Cancellaria sp. X Chlorostoma sp. X Columbellidae (indeterminate) X Conus sp. X Crepidula sp. X Crucibulum sp. X Cypraeidae Diodora sp. X Fasciolariidae X Felaniella harfordi (Anderson, 1905) X Ficus sp. X Fissurella rixfordi Hertlein, 1928 X Forreria bartoni (Arnold, 1910) X Fusinus sp. X Hexaplex sp. X Kelletia lorata Addicott, 1970 X cf. Leporimetis sp. X cf. Macrarene sp. X Megasurcula keepi (Arnold, 1907) X Melongena californica Anderson and Martin, 1914 X Mitridae X Morula cf. M. grand Addicott, 1970 X Nassariidae (indeterminate) X Naticidae (indeterminate) X cf. Odostomia sequoiana Addicott, 1970 X Opbiodermella cf. O. temblorensis (Anderson and Martin, 1914) X Pleuroploca sp. X Priscofusus geniculus (Conrad, 1849) X Pyruconus cf. P. bayesi (Arnold, 1909) X Pyruconus cf. P. owenianus (Anderson, 1905) X Scaphander cf. S. jugularis (Conrad, 1855) X Sinum cf. S. scopulosum (Conrad, 1849) X Strombus sp. X Terebra cooperi Anderson, 1905 X Teredinidae X Thais edmondi (Arnold, 1907) X Trophon kernensis Anderson, 1905 X Turridae X Turritella ocoyana Conrad, 1855 X Turritella temblorensis Wiedey, 1928 X Turritella spp. X Vermicularia sp. X Mollusca: Bivalvia Amusium lompocensis (Arnold, 1906) X Amussiopecten vanvlecki (Arnold, 1907) X Anodontia sp. X Batillaria sp. X Chlamys sespeensis (Arnold, 1906) X Cbione cf. C. ricbthofeni Hertlein and Jordan, 1927 X X X X X X X X X X X X X X X Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone ■ 39 Table 1 Continued. Area 1 Area 2 Area 3 SDSNH SDSNH SDSNH SDSNH SDSNH SDSNH OCPC OCPC SDSNH 5752 4312 4521 6283 6284 6286 0872 03155 4520 Cbione scbencki Loel and Corey, 1932 X Clementia cf. C. conradiana (Anderson, 1905) X Crassadoma cf. C. gigantea (Gray, 1825) X Crassostrea titan (Conrad, 1853) cf. Crassostrea sp. X X X X X X Crenomytilus expansus (Arnold, 1907) Dosinia sp. X X Felaniella cf. F. barfordi (Anderson, 1905) X cf. Gari sp. X Glycymeris sp. X Lima sp. X Limaria sp. X Litbopbaga sp. X Lucinidae X Lyropecten crassicardo (Conrad, 1857) X X X X X Mactridae X Modiolus ynezensis Arnold, 1907 Mytilus cf. M, coalingensis Arnold, 1910 X X Mytilus cf. M. expansus Arnold, 1907 X Mytilus sp. X X Pacipecten andersoni (Arnold, 1906) X Panopea abrupta (Conrad, 1849) X Panopea tenuis (Wiedey, 1928) X Panopea sp. X X Pycnodonte cf. P. bowelli (Wiedey, 1928) X X Pycnodonte cf. P. wiedeyi (Hertlein, 1928) X X X Penitella sp. X Prototbaca sp. Saxidomus vaquerosensis Arnold, 1910 X X Saxidomus sp. Solecurtidae? Spondylus perrini Wiedey, 1928 X X X X Spondylus scotti Brown and Pilsbry 1913 Spondylus sp. X X Trachycardium cf. T. vaquerosensis (Arnold 1908) X Tracbycardium sp. X X X Tresus sp. X Solencurtidae (indeterminate) X Veneridae X X Annelida: Polychaeta Spirorbis sp. Arthropoda: Crustacea: Cirripedia X Megabalanus tintinnabulum (Linnaeus, 1758) Echinodermata: Echinoidea X X Eucidaris cf. E. thouarsii (Valenciennes, 1846) X X X X Strongylocentrotus sp. Vaquerosella cf. V. andersoni (Twitched, 1915) X X VaqneroseUa merriami Anderson, 1905 Ichnotaxa X X X X X X X Entobia isp (burrows) X Palaeopbycus isp. Chordata: Vertebrata X X X X X Chondrichthyes X X X X Osteichthyes X X Lithophyllum cf. L. profundum Johnson, 1954 (Fig. 55), as well as benthic foraminifers, ostracodes, echinoid spines, and mollusk shell fragments. Another feature of Lithofacies D are clusters of linear bioturbations (Fig. 56) that Cuffey et al. (1981) described as burrows of polychaete colonies composed of cylindrical, micrite-filled tubes arranged parallel to bedding and each other. Loel and Corey (1932:275) referred to these as Serpula coreyi Wiedey, 1928, recognizing that its original designation as S. careyi was a typographical error. Loel and Corey also reported large colonies of “worms” in the Vaqueros and Temblor formations in the San Joaquin Hills in Orange County, the Santa Monica Mountains in Los Angeles County, the Ventura River “Reef” beds in Ventura County, and the west¬ ern Santa Ynez Mountains in Santa Barbara County. Lithofacies D bur¬ rows are herein referred to Palaeophycus isp. These parallel bioturbations range from 2 to 4 nnn in diameter and up to 20 cm in length and are straight or, more rarely, slightly curved. There is no evidence of backfilling or setae impressions in any of the burrow walls; however, faint concentric rings, possibly from somite segments, with a thin linear ridge, extend the length of the burrow. Sullwold (1940) also noted thin rings on the internal walls of his specimens from Wood Canyon (LACMIP loc. 22004). Having examined specimens from Area 1, A. Myra Keen (pers. 40 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone Figures 7-34 Saddleback Valley limestone gastropods (Figs. 7-29) and other invertebrates (Figs. 30-34). SDSNH numbers are locality/specimen; H=height, W=width, L=length. 7, 8. Fissurella rixfordi Hertlein, 1928: 7, internal mold, SDSNH 4312/136497, L=35 mm, W=24 mm; 8, external mold, SDSNH 4312/140187, L=30 mm, W=20 mm; 9. Chlorostoma sp., external cast, SDSNH 4312/93707, H=10 mm, W=10 mm; 10. Turritella ocoyana Conrad, 1855, internal mold, SDSNH 4312/140188, H=75 mm, bottom whorl W=20 mm; 11. Turritella sp., latex peel of external cast, SDSNH 4312/85244, H=33 mm, W=10 mm; 12. Forreria bartoni Arnold, 1910, original shell (missing upper part of spire), SDSNH 4521/83243, H=24 mm, W=33 mm; 13. Calyptrea sp., internal mold, SDSNH 4312/85244, H=25 mm, W=35 mm; 14. Crucibulum sp., apical view of internal mold showing septum partially attached to shell; 15. naticid, internal mold, SDSNH 4312/140186, H=35 mm, ultimate whorl W=32 mm; 16. Sinum cf. S. scopulosum (Conrad, 1849), internal mold, SDSNH 4312/93721, H=25 mm, ultimate whorl W=28 mm; 17. Trophon kernensis Anderson, 1905, latex peel of Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone ■ 41 comm., 1979) commented that the “Reef” looks like a stratum of lime mud in which these annelids ivere burrowing, and as the deposit thickened, they moved upward to form long tubes. The upper part of the section, from 12.1-14.1 m, is Lithofacies E, a white Monterey mudstone unconformably above the biosparite of Lithofa¬ cies D. It was exposed on a grassy hillside that extended downward for more than 6 m. Area 2: Laguna Hills-Aliso Viejo Area 2 is 5.2 km north of Wood Canyon and extends west from Via Lomas in Laguna Hills (SDSNH loc. 4312) and into Aliso Viejo (SDSNH Iocs. 4521, 6283, 6284, 6286) (Figs. 1, 57). In this area, the Saddleback Valley limestone consists primarily of highly indurated fossiliferous limestone and friable calcareous sandstone. Mollusk molds (Fig. 58) are visible in the limestone at Fossil Reef Park (SDSNH loc. 4312; Fig. 5), which is part of a small fault block displaced 300 m southward from an east-west ridge (Morton et al., 1974). The ridge structure is characterized by minor east-west normal faults along both its northern and southern flanks. The nearby Moulton Parkway road cut (SDSNH loc. 4521; Fig. 5) exposed a 201-m-long cross section of the ridge. South of the first fault, 88.2 stratigraphic meters of the Saddleback Valley limestone arc exposed. It comprises 15 beds differentiated by lithology and fossils (Fig. 6). Most of the beds are calcareous sands to friable calcare- nites, but also present are two marls composed almost entirely of abraded cheilostome bryozoan fragments, two “button beds” (sensu Anderson, 1905) densely packed with the small sand dollar Vaquerosella merriami Anderson, 1905, and two beds of highly indurated limestone composed of 93-95% calcium carbonate (Morton et ah, 1974). From 0 to 21.9 m, the fauna is characterized by intermittent appearances of Eucidaris cf. E. thouarsii (Valenciennes, 1846), Palaeophycns isp. burrows preserved as biosparite intraclasts, bryozoans, oysters, pectens, gastropods, echinoids, and barnacles. The 28.6-45.5-m interval is characterized by articulated pectens and oysters and include large well-preserved L. crassicardo in pre¬ ferred orientation, similar to their occurrence in fine calcarenite near Fossil Reef Park in Area 2 (Fig. 4). The road cut section is then dominated by unconsolidated calcareous sand with shell fragments. Sands between the first and second fault are finer, and between the second and fourth fault they are devoid of macrofossils. The latter fault separates the calcareous sands of the Saddleback Valley limestone from the mudstone that is most characteristic of the lower Monterey Formation. A thin layer of vitric vol¬ canic ash is present in the northernmost mudstone block, but attempts in this study to isotopically date it were unsuccessful. The “Temblor” molluscan fauna in Area 2 is characterized by the bivalves S. vaquerosensis , L. crassicardo, Crenomytilus ex pans ns (Arnold, 1907), Panopea abrupta (Conrad, 1849), Modiolus ynezensis Arnold, 1907, and C. titan, and the gastropods T. ocoyana, Turritella temblorensis Wiedey, 1928, and Pyruconus cf. P. bayesi (Arnold, 1909). Throughout Area 2, molds account for the highest numbers of specimens and species of any molluscan assemblage collected in this study. A hillside outcrop immediately west of Moulton Parkway displays what appears to be a scries of channelized, partial turbidite sequences in which dense limestone alternates with friable sandy limestone (Fig. 59). The sandy limestone contains V. merriami , cchinoid spines, Gemelliporella aff. G. punctata, and Megabalanus tintinnabulum (Linnaeus, 1758). In contrast, 90% of the dense limestone consists of unsorted invertebrates, predominately the bivalves C. titan , L. crassicardo, Pycnodonte cf. P. howelli (Wiedey, 1928), 5. vaquerosensis, and Tresus sp. This sequence was apparent in western Area 2 localities SDSNH 4521, 6283, 6284, and 6286. On the west side of the northern Aliso and Wood Canyon Wilderness Park, south of the intersection of Aliso Viejo Parkway, a paved path leads up from Cedarbrook past weathered dense limestone to an undeveloped outcrop where mudstone unconformably caps the limestone. Morton et al. (1974) mapped similar contacts elsewhere in the area, but this is the only one that remains visible. Three exploratory wells drilled in the mid-20th century encountered limestone in the subsurface of western Saddleback Valley. In Shell Oil’s Moulton No. 88-4, adjacent to SDSNH loc. 6283 in Area 2, a thick coqui¬ na of megafossil fragments that includes some blueschist clasts is at 0-300 ft, and a much thinner calcareous bed containing shell fragments, small bivalves, and cchinoid spines is at 488-491 ft. About a quarter-mile north-northeast of the Area 2 fossil localities. South Fullerton Oil’s No. 1 well has a hard shell bed at 840-895 ft. In the vicinity of Area 1, Shell’s Moulton No. 14 well drilled through Monterey mudstone and into a fos¬ siliferous limestone 239-253 ft above the San Onofre Breccia. Area 3: Laguna Woods and Lake Forest Limestone localities in Area 3 include the original “Pecten Reef” in Lake Forest (SDSNH loc. 4520/OCPC loc. 0023) and the Home Depot (OCPC loc. 03155) and Laguna Woods Self-Storage (OCPC loc. 0872) sites. No basal contacts of the limestone were exposed in this area, but infrequent blueschist and quartz cobbles in the sandy limestone reveal that the San Onofre Breccia extended into OCPC loc. 03155. Narrow lenses of friable calcareous sandstone in Lake Forest (SDSNH loc. 4520), containing echinoids (“button” sand dollars, eucidarid spines), cartilaginous and boney fish teeth, and occasional sandy manganese nodules up to 6 cm in width, exuded a strong petroliferous odor when exposed. Bonatti and Nayudu (1965:26) contend that the rate of manga¬ nese oxide deposition can vary, and although manganese nodules generally represent low rates of growth in the deep sea, nodules have also been found in several localities where sedimentation rates are relatively high, such as the Gulf of California, and at shallow depths (Calvert and Price, 1977). The friable calcareous sandstones are overlain by a sequence of three turbidites in which 60-72-cm-thick, orange, friable, coarse sandstones alternate with 200-210-cm-thick greenish-gray silty mudstones containing scattered fish scales. The sandstones (SDSNH loc. 4964; LACMVP loc. 3414) yielded abundant fish teeth, as well as a croco¬ dile tooth. In 1973, development of a housing tract that includes Mountain View Park covered the limestone and sandstone, although a few boul¬ ders of the indurated limestone remain as decorative features of the community landscape, and a small cut-slope east of the park exposes the mudstones. West of Lake Forest, in Laguna Woods, grading for the Leisure World community in the 1960s exposed an orange, friable, coarse sandstone that yielded fish, sea turtle, and bird fossils at three localities north of Aliso Creek. A rich marine vertebrate fauna recovered from a later exposure (LACMVP loc. 1945) includes 19 bird and 20 shark species dated at 12- 14 Ma (Howard, 1968; Howard-Wylde, 1980), which is younger than the limestone. Howard-Wylde (1980) characterized this sandstone as hav¬ ing the densest concentration of shark teeth in western North America. Grading for the Laguna Woods Home Depot (OCPC loc. 03155; 3.3 km west of SDSNH loc. 4520) exposed a limestone with poorly preserved mollusks, notably L. crassicardo and C. titan and fragments of S. vaquer¬ osensis, Pyruconus cf. P. hayesi, Forreria bartoni (Arnold, 1910), cheilos¬ tome bryozoans, and Palaeophycns isp. Also present are unbroken V. merriami and random small shark teeth. Along the north side of the Home Depot site (OCPC loc. 03155), there was a small outcrop of gritty, poorly consolidated Topanga siltstone that Morton et al. (1974) mapped as the Vaqucros Formation. The unit contains Acila sp., Scapharca sp., A. lompocensis, and well-preserved clusters of translucent brown brachiopods identified as a Glottidia sp. This association was also seen in a dark siltstone laterally adjacent to external cast; 27, internal mold; 28. Calyptraea sp., internal mold, SDSNH 4520/98688, L=21 mm, W= 15 mm; 29. vermetid, internal mold, SDSNH 4512/142016, image H=30 mm; 30. Gemelliporella aff. G. punctata Canu and Bassler, 1919, fragments, SDSNH4521/98668, image H=63 nun; 31, 32. Eucidaris cf. E. thouarsii (Valenciennes, 1846): 31, original spine with regularly spaced knobs, SDSNH 4521/140192, L=30 mm; 32, original test fragment, SDSNH 4521/98680, L=19 mm; 33. Vaquerosella merriami (Anderson, 1905), external dorsal view, SDSNH 4521/83249, H = 18 mm, W = 18 mm; note anal aperture (notch) at ventral margin; 34. Megabalanus tintinnabulum (Linnaeus, 1758), original shells, side view of attached pair, SDSNH 4312/140191, H=25 mm, W=20 mm. 42 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone Figures 35-52 Saddleback Valley limestone bivalves. Figured specimens are external casts unless indicated otherwise. LV=left valve, RV=right valve; SDSNH numbers are locality/specimen; H=height (dorsal to ventral), L=length (anterior to posterior). 35. Crenomytilns expansus (Arnold, 1907), LV, SDSNFI4312/83246, H=130 mm, L=75 mm; 36,37. Crassostrea titan (Conrad, 1853), RV, SDSNH 4312/83238, H=97 mm, L=52 mm: 36, exterior; 37, interior; 38. Pycnodonte cf. P. bowelli (Wiedey, 1928), SDSNH 4312/83234, H=70 mm, L=70 mm; 39. Amussiopecten vanvlecki (Arnold, 1907), LV with faintly raised ribs, SDSNH 4312/83232, H=70 mm, L=75 mm; 40. Amusium lompocensis (Arnold, 1906), crushed shell with smooth surface, SDSNH 4512/98676, H=104 mm, W=68 mm; 41, 42. Lyropecten crassicardo (Conrad, 1857), original shells, external views: 41, LV, SDSNH 4521/98672, H=97 mm, L= 107 mm; 42, juvenile, LV, SDSNH 4521/98671, H=30 mm, L=30 mm; 43. Pacipecten andersoni (Arnold, 1906), original shell, RV, external view, SDSNH 4312/140181, L=30 mm, W=30 mm; 44. Crassadoma cf. C. gigantea (Gray, 1825), external cast, LV, SDSNH 4312/140354, H=42mm, W=32 mm; 45. Lima sp., external cast in bioturbated matrix, RV, SDSNH 4312/140182, L=47 mm, W=35 mm; 46. Limaria sp., external cast, RV, SDSNH 4312/121368, L=30 mm, W=21 mm; 47. Spondylus scotti (Brown and Pilsbry, 1913), external cast, SDSNH 4520/83235, L=124 mm, W=100 mm; 48. Trachycardium sp., LV, external cast, SDSNH 4312/93837, L=86 mm, W=80 mm; 49. Chione cf. C. ricbtbofeni Hertlein and Jordan, 1927, latex peel of external cast, LV, SDSNH 4312/83248, L=20 mm, W=25 mm; 50. Saxidomus vaquerosensis Arnold, 1910, RV view of steinkern, SDSNH 4312/180184, L=70 mm, W=100 mm. Note anterior and posterior muscle scars (light patches); 51. Panopea tenuis (Wiedey, 1928), external cast, LV, SDSNH 4312/ 140185, L=45 mm, W=85 mm; 52. Panopea abrupta (Conrad, 1849), external cast, LV, SDSNH 4312/83237, L=40 mm, W=58 mm. Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone ■ 43 Figure 53 Thick limestone of Lithofacies B dominated by steinkerns of Saxidomus vaquerosensis in Wood Canyon (SDSNFI loc. 5752). W.J. Edgington stands on a lag deposit (Lithofacies A) consisting of abraded and broken mollusk shells and blueschist cobbles derived from the San Onofre Breccia. Image taken in 1976 before site was destroyed by development. indurated limestone, 15 km to the south near the junction of Wood Canyon and Aliso Canyon in Area 1 (Sullwold Collection, LACMIP loc. 21981). Thin sections of this sandy limestone display articulated ostracodes. In the vicinity of the Home Depot site is the Laguna Woods Self Storage facility (OCPC loc. 0872), located at the northwest junction of El Toro Road and Moulton Parkway. Grading on this site in 1997 exposed an indurated sandy limestone containing abraded fragments of L. crassi- cardo and C. titan. East of SDSNH loc. 4520, grading in 1975 had also revealed a small limestone lens near Muirlands Boulevard and Lake Forest Drive. No fossils or data were collected from either of these two localities. DISCUSSION DEPOSITIONAL PALEOENVIRONMENT The fossils of Saddleback Valley limestone (Table 1) indicate that around 16 Ma, middle Miocene coastal waters were warmer than those off southern California today. Fife (1979) suggested that the Figure 54 Microphotograph of thin-sectioned biosparite from Lithofa¬ cies B in Wood Canyon (Area 1, SDSNH loc. 5752). Faint microcrystalline calcite surrounds benthic foraminifera that are likely among those taxa isolated from the matrix of SDSNH loc. 4521 in Area 2. Plausible identifications based in Finger (1990) are Bolivina sp., (upper left), Anomalinoides salinasensis (Kleinpell, 1938) (below center), and Valvu- lineria miocenica Cushman, 1926 (upper right). limestone was an “allochthonous fossiliferous deposit” in which marine benthic organisms were transported by longshore currents, storm waves, and gravity to lower areas on the shallow substrate. However, the microfossil component described in the present study reveals that the shallow-water benthos were redeposited at much greater depths, most likely due to subsidence and turbidity currents. Those phenomena are responsible for much of the deep-water deposition off the coast of California during the Mio¬ cene, particularly that of the Monterey Formation and its equiva¬ lent units in the Los Angeles Basin. Woodford (1925) interpreted the depositional environment as a gulf or strait bounded on the west by a region of high relief, with an arid climate that permitted the thick, poorly sorted San Onofre Breccia to accumulate without muddying the waters. Figure 55 Thallus of the red coralline algae Lithophyllum cf. L. profundum Johnson, 1954. Photomicrograph of thin-sectioned biosparite of Lithofacies D, SDSNH loc. 5752. 44 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone Figure 56 Palaeophycus isp. (SDSNH 93703) in limey mudstone of Lithofacies D, SDSNH loc. 4512. Loel and Corey (1932) proposed that the widespread uniformity of marine conditions that characterize the “Temblor” CPMS resulted from an abrupt subsidence of the California coast and that the resulting transgression that modified the coastline was accompanied by a rapid increase in the number and diversity of invertebrates as tropical species expanded their ranges northward. Similar to the rhodophytic rudite described from Wood Canyon, Johnson and Kaska (1965) found abundant red coralline algae in the Miocene El Peten Limestone of Guatemala, which they described as a biosparite with a rich megainvertebrate fauna of mollusks, echinoids, and bryozoans. Modern colonies of coralline red algae inhabit the photic zone of calm waters, such as those in the Gulf of California, around Cedros Island west of central Baja California, and on the leeward side of Santa Catalina Island in the San Pedro Channel off Los Angeles. Cuffey et al. (1981:65) suggested that the Saddleback Valley limestone “developed on the shallow part of a submarine slope, from a gravel flanked offshore island, down into a mud-floored deep-marine basin to the east.” They noted that foraminifers and nannofossils in the limestones (from all three areas referred to in the present study) indicate deposition at outer-neritic to upper- bathyal depths, but the associated ostracodes and mollusks are characteristic of intertidal to inner-neritic depths. These mixed- depth associations confirm that the limestone was formed mostly from sediments that had been displaced downslope. Linger (1988) documented the occurrence of shallow-marine ostracodes in Laguna Hills and interpreted them also as having been displaced downslope based on their association with deep¬ water foraminifera in accordance with the upper depth limits (UDLs) assigned by Ingle (1980). Linger (1992) later provided a list of those foraminiferal assemblages, three of which were from the original Moulton Parkway road cut (SDSNH loc. 4521). The composite foraminiferal assemblage in the road cut consists of 35 species. Each of the three assemblages are similarly mixed, as indicated by the association of multiple species representing four paleobathymetric biofacies ranging from inner-neritic to upper- middle bathyal depths (Table 2). The deepest dwelling species indicate that final deposition of the Saddleback Valley limestone sediments was at upper middle-bathyal depths. In association with the foraminifera are nine species of ostra¬ codes that Linger (1988) assigned to seven genera characteristic of a shallow-marine biofacies: Aurila (3 species), Cytberella, Her- manites , Loxoconcba, Loxocorniculum, “Paijenborcbella ,” and Paracytberidea. About 0-.4 km northeast of, and stratigraphically above, the road cut section, an orange sand lens (~4 m 2 ) within a greenish-grey silty mudstone (UCMP loc. 12873) yielded a similar microfauna. Ostracodes were also observed in thin sections from Area 1, but none have been recorded from Area 3. Linger (1988, 1992) also sampled the mudstone that was exposed across the nearby intersection of Alicia and Moulton parkways, and they yielded Luisian foraminiferal assemblages more typical of the deeper water Monterey Lormation, with the Saddleback Mtn. Figure 57 Saddleback Valley looking N79E from Aliso and Wood Canyons Wilderness Park (Area 2). The Santa Ana Mountains are visible on the horizon. Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone ■ 45 Figure 58 Fossiliferous biosparite (OCPC 50310) salvaged in 1979 from a spoils pile at SDSNFI loc. 4312 (OCPC loc. 0022) in Laguna Hills (Area 2). Note the fossils are molds not in preferred orientation or size- sorted. Visible in this photograph: a, gastropods; b, Saxidomus vaquer- osensis (steinkern, hinge-up); c, mold of S. vaquerosensis valve with calcite-filled Entobia isp. burrows. inclusion of species such as Rectuvigerina branneri (Bagg, 1905) and Pullenia miocenica Kleinpell, 1938, and a much lower compo¬ nent of shallow-water species. No ostracodes were recovered from these mudstones. Although they did not analyze microfossils, Stanton and Alder- son (2013) concluded that the limestone interbedded with the Conejo Volcanics in the Santa Monica Mountains is similar to the limestone in Saddleback Valley, in that both formed from shal¬ low-water sediments displaced into deep water during the early to middle Miocene. Hall (2002) considered the bivalve L. crassicardo and the gas¬ tropod Tropbosycon kerniana (Cooper, 1894), both of which are abundant in some of the Saddleback Valley limestone out¬ crops, as representatives of his middle Miocene “outer tropical range.” He placed the region approximately 750 km to the south, in subtropical waters where Cedros Island is today, during the global Middle Miocene Climate Optimum (MMCO) at 17-15 My (Flower and Kennett, 1994; Zachos et al., 2001). This Figure 59 Alternating beds of indurated limestone and friable sandy limestone, North Aliso and Wood Canyon Wilderness Park (Area 2, SDSNH loc. 6283). The more resistant limestone that forms the ledges shown here correlates with the indurated limestone exposed along Moulton Parkway (SDSNH loc. 4521), which is on the east side of this hill (to the left of photo). Table 2 Upper depth limits of benthic foraminifera recovered from the Saddleback Valley limestone in the Moulton Parkway road cut (SDSNH loc. 4521). Inner neritic (50-150 m) Buccella oregonensis (Cushman, Stewart and Stewart, 1948) Buliminella elegantissima (d’Orbigny, 1839) Elphidium grand Kleinpell, 1938 Gaudryina pliocenica Cushman, Stewart and Stewart, 1949 Gaudryina subglabrata Cushman and McCulloch, 1939 Nonionella miocenica Cushman, 1926 Pseudononion cosdferum (Cushman, 1926) Outer neritic (50-150 m) Bolivina advena ornata Cushman, 1925 Buliminella subfusiformis Cushman, 1925 Islandiella modeloensis (Rankin, 1934) Marginulinopsis beali (Cushman, 1925) Valvulineria californica Cushman, 1926 Valvulineria miocenica Cushman, 1926 Upper bathyal (150-500 m) Baggina calif ornica Cushman, 1926 Bolivina brevior Cushman, 1925 Bolivina tumida Cushman, 1925 Cancris baggi Cushman and Kleinpell, 1934 Pseudoparrella subperuviana (Cushman, 1926) Kleinpella californiensis (Cushman, 1925) EJvigerinella calif ornica ornata Cushman, 1926 Upper-middle bathyal (500-1,500 m) Bolivina calif ornica Cushman, 1925 Bolivina imbricata Cushman, 1925 MMCO was the warmest interval in the Neogene, as evidenced by the many molluscan ranges that extended their farthest north since the middle Eocene (Oleinik et al., 2008). BIOSTRATIGRAPHIC AGE The shallow-marine invertebrate fauna of the Saddleback Valley limestone represents the “Temblor” CPMS, which Smith (1991) placed at approximately 19.5-12 My, an interval spanning the middle Saucesian to early Mohnian benthic foraminiferal stages of Kleinpell (1938) (see Fig. 2). McCulloh et al. (2002) shortened this CPMS in the Los Angeles Basin to approximately 16.65-14.2 (late Saucesian to late Luisian). Among the mollusks, the bivalve Pacipecten andersoni and the gastropods Pyruconus cf. P. bayesi, Megasurcula keepi (Arnold, 1906), and Priscofusus geniculus (Conrad, 1849) are restricted to the “Temblor” CPMS. Although most of the species in the Saddleback Valley limestone are known to occur in the “Temblor” CPMS, P. geniculus and the bivalve Pycnodonte cf. P. wiedeyi (Hertlein, 1928) had not been previous¬ ly recognized above the Vaqueros CPMS. The benthic foraminiferal fauna recovered from the limestone at Moulton Parkway (Area 2) characterizes the Relizian-Luisian interval, while the absence of species that first appear in the Luisian Stage favors the older part of that range. Also, in Area 2 (SDSNH 4312), Kling (pers. comm., 2002) identified coccoliths of Calcidis- cus leptoporus (Murray and Blackman, 1898), Coccolithus pelagi- cus (Wallich, 1877) Kamptner 1954, Dictyococcites minutus (Haq, 1971), Helicosphaera carteri (Wallich 1877), and Helico- spbaera scissura Miller, 1981. The concurrent calcareous nanno- plankton zonal range is delimited by the range of H. scissura at calcareous nannofossil (CN) zones CN1-CN3 (late Saucesian- late Relizian). The concurrent biostratigraphic data therefore place the limestone in the Relizian and within CN3. The Monterey For¬ mation mudstones at the intersection of Alicia and Moulton park¬ ways (just south of the Moulton Parkway road cut) yielded a 46 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone Table 3 Strontium isotope data and ages for the Saddleback Valley limestone (three SDSNH localities), the orange sand lens in mudstone (UCMP loc. 12873), and dark brown mudstone (UCMP loc. 12873). Ages interpreted from unpublished reference data provided by J. McArthur (LOWESS 5) and D. DePaolo. 87 Sr/ 86 Sr normalized +2 SE Age Sample Minimum Mean Maximum LOWESS 5 DePaolo UCMP 12871 .708830 .708837 .708844 12.00+2.5 13.5+2.0 SDSNH 4521A .708764 .708774 .708784 15.00+0.7 15.9+0.1 UCMP 12873 .708756 .708765 .708774 15.40 -0.7/+0.5 16.0+0.1 SDSNH 4521 .708722 .708730 .708738 15.92+0.6 16.3+0.1 SDSNH 4312 .708717 .708727 .708737 15.96+0.6 16.3+0.1 distinctly Luisian foraminiferal fauna, which suggests that the older limestone could be within the late Relizian Stage, but that interval overlaps with the Luisian Stage (Fig. 2). Although McCulloh et al. (2002) places the Relizian/Luisian boundary at 15.6 Ma, just above the 15.97 Ma boundary between the early and middle Miocene, Crouch and Bukry (1979) had previously indicated that the Relizian ranges into CN5 (13.6-11.7 My). For the most part, the relative age, lithology, and paleontology of the Saddleback Valley limestone are consistent with the “Tem¬ blor” CPMS in the Topanga Formation. Coccoliths are elements of oceanic plankton that are more likely to be deposited in a deep marine embayment than along the shallow margin of a restricted fluviomarine inlet or channel. Hence, it is unlikely that H. scis- sura was deposited in the Topanga sediment before its down- slope displacement, implying that deposition of the Monterey Formation in the study area commenced in the Relizian. It is unfortunate that the Wood Canyon exposure (Area 1) was among the first destroyed by development; had microfossil sam¬ ples been taken at or just below the level of its limestone lenses, they could have added credence to the late Relizian age of the limestone. The California Miocene benthic foraminiferal zonation of Klein- pell (1938) began morphing in the 1970s with the development of global planktic microfossil zonations. Local studies on planktic microfossils (i.e., foraminifera, diatoms, radiolarians, calcareous Seawater Sr Data Normalized Age (Ma) Figure 60 Determination of Miocene isochronologic ages for five samples of the Monterey Formation in Saddleback Valley using DePaolo’s latest (unpublished) version of the Richter and DePaolo (1988) 87 Sr/ 86 Sr curve for the Miocene. The three SDSNH samples are from limestones, UCMP 12873 is an orange sand lens within mudstone, and UCMP 12871 is mudstone. nannofossils) associated with “stage-diagnostic” benthic foraminif¬ eral assemblages evidenced the time-transgressive nature of the Oppelian-based benthic stages (see introduction in McDougall, 2007, for elaboration). The implied geographical inconsistency is simply because benthic organisms are controlled by local bottom facies. Problems with the CBFS are most evident when attempting interbasinal correlation (Finger, 1992), and a similar situation restricts the utility of the CPMS. The biostratigraphic placement of the Saddleback Valley in the late early Miocene therefore needed to be assessed by a geochronologic method that could produce high-resolution dates, as reported in the following section. CHRONOSTRATIGRAPHIC AGE Five well-preserved samples of biogenic calcite from Area 2 were sent to the Pacific Centre for Isotopic and Geochemical Research (PCIGR) at the University of British Columbia for strontium iso¬ tope ( 87 Sr/ 86 Sr) analysis. These were bivalve (L. crassicardo) shells from SDSNH 4312 (Fossil Reef Park), gastropod (F. bartoni ) spines from SDSNH 4521, and cheilostome bryozoans ( Gemelli - porella aff. G. punctata) from SDSNH 4521A (both in the Moul¬ ton Parkway road cut), foraminifera ( Valvulineria miocenica Cushman, 1926) from UCMP loc. 12873 (orange sand lens in Monterey mudstone), and foraminifera ( R. branneri) from UCMP loc.12871 (dark brown mudstone of Monterey Formation that was exposed at the eastern corner of Moulton and Alicia parkways). Table 3 presents the analytical data ( 87 Sr/ 86 Sr ratios) and two sets of age interpretations according to McArthur’s LOWESS 5 curve (unpublished version “LOWESS 5 Fit 26 03 13” of the LOWESS 3 curve in McArthur et al., 2001) and DePaolo’s revi¬ sion (unpublished) of the Miocene 87 Sr/ 86 Sr curve of Richter and DePaolo (1988) that DePaolo and Finger (1991) used to evaluate the biostratigraphic correlations in the Monterey Formation of central California. That 87 Sr/ 86 Sr curve is offset from the more commonly used LOWESS curve that indicates younger ages for the same values. DePaolo’s curve is favored here as the more accu¬ rate of the two, because the pertinent part of it was derived solely from the datasets of two Deep Sea Drilling Project (DSDP) cores (575B and 590B), and diagenetic effects, which were evident in the poor match of the original datasets, were corrected with the methods described by Richter and DePaolo (1988). This correc¬ tion can only be done on a continuous section that has both pore- water and carbonate fossil data; hence, most 87 Sr/ 86 Sr data incorporated into the LOWESS curve is uncorrected. Data from the 13-18 My interval, within which the curve is kinked at about 15 Ma, are particularly susceptible to shifting by diagenesis. The inaccuracy of the LOWESS curve was most evident in the impos¬ sible age of 12.0 Ma it indicated for Rectuvigerina tests from the Monterey mudstone (UCMP loc. 12871), as that genus had disap¬ peared from California at the mid-Miocene cooling event that trig¬ gered the biotic turnover demarcating the Luisian/Mohnian boundary at 13.6 Ma. Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone ■ 47 Correlation with DePaolo’s curve (Table 3; Fig. 60) age-dates the “Temblor” fauna in the Saddleback Valley limestone within the interval 16.3-15.9 ± 1.0 My (see Fig. 2). Determination of this relatively narrow age interval is possible because the 8 Sr/ 86 Sr values correlate with the steepest part of the Cenozoic curve. In contrast, the higher ratio of the mudstone from UCMP loc. 12871 is within the flattest section of the curve and yields an imprecise age of 15.4-11.4 My, but its Luisian foraminiferal assemblage indicates that it is unlikely to be younger than 13.6 Ma, refining the age of the mudstone to 14.5 ± 0.9 Ma. The part of the DePaolo curve relative to this study remains intact despite the latest revision (Hilgen et al., 2012) of the geologic time scale and its CN zonation in particular. TAPHONOMY Thin sections of the indurated limestone exhibit a microcrystalline calcite matrix with foraminifers, ostracodes, coccoliths, siliceous sponge spicules, echinoid spines, coralline red algae thalli, and fragments of mollusks and bryozoans. Recrystallization has degraded the preservation quality of the calcareous microfossils with the exception of the coralline red algae thalli. In all three areas, indurated biosparite has mollusk molds displaying detailed internal and exterior shell structures. Distinctive chains of round chambers ascribed to Entobia isp, occur in molds of Saxidomus vaquerosensis. As the biosparite grades into a sandy limestone, the form of preservation changes from molds to well-preserved, articulated shells that are primarily pectinids but also include C. titan, Pycnodonte cf. P. howefli, and Pycnodonte cf. P. wiedeyi. The limey sandstones of Area 2 contain scattered shell fragments of S. scotti, Tracbycardium sp., and F. bartoni ; spines of the latter species retain some original pigmentation visible as a faint pink. In sandier limestone matrices, abundant well-preserved echinoids (Eucidaris sp,, V. merriami) occur with shark teeth and red-tinted barnacles (M. tintinnabulnm). Also common in the limey sand¬ stone are bryozoan branches, especially in Lithofacies C of Area 1 (SDSNH loc. 5752) and in the upper friable limey sandstones beds of Area 2 (SDSNH loc. 4521). The poor preservation of fos¬ sils in limestone at the western Area 3 site (OCPC loc. 03155) is attributed to groundwater infiltration along a fault that extends across the development; for example, articulated pectinids, well preserved in other limestone localities, are crumbly and stained orange. COMPARISON WITH “TEMBLOR” FAUNA IN THE LOS ANGELES BASIN The “Temblor” fauna in the Saddleback Valley limestone differs from those of the Topanga Formation in the northern foothills of the Santa Ana Mountains (Schoellhamer et al., 1981) and the Cold Creek Member of the Topanga Canyon Formation in the Santa Monica Mountains (Alderson, pers. comm., 2014) by hav¬ ing abundant coralline algae, cheilostome bryozoans, common small gastropods, and isolated concentrations of pectinids. Addi¬ tionally, V. merriami , which is abundant in all three Saddleback Valley limestone areas, has not been reported from the Santa Mon¬ ica Mountains, but it occurs in the northern Santa Ana Mountains (SDSNH loc. 4308A) in association with T. ocoyana , T. temblor- ensis , Dosinia sp., and Antillopbos posunculensis (Anderson and Martin, 1914). Although no bioturbated mudstones have been reported in the Topanga and Topanga Canyon formations, clusters of Palaeophy- cus isp. were observed in Topanga Formation siltstone along the eastern flank of SDSNH loc. 4312 in Area 2. Similar parallel bur¬ rows with internal ringlike impressions also occur in gray biosparite in Wood Canyon (LACMIP loc. 22004) and in the San Joaquin Hills just south of Saddleback Valley in the Bommer Member of the Topanga Formation (SDSNH loc. 6836). These burrows are also in the Topanga Formation near Ortega Highway in southern Rancho Mission Viejo (OCPC loc. 3266) associated with cheilostome bryozoans and the majority of mollusk species found in the Saddleback Valley limestone. Foraminifera do not appear to be abundant in the Topanga For¬ mation in the San Joaquin Hills, as first noted by Vedder et al. (1957) when they split it into, from oldest to youngest, the Bom¬ mer, Los Trancos, and Paularino members. Their only mention of foraminifera referred to them as sparse in the Los Trancos Member. Smith’s (1960) foraminiferal study focused on the Santa Ana Mountains but included the San Joaquin Hills. She reported four assemblages in the Topanga Formation of the Santa Ana Mountains that yielded a total of 11 species and noted that fora¬ minifera are abundant but highly dominated by Bolivina advena Cushman, 1925, var.; Nonion costiferum Cushman, 1926; N. aff. N. costiferum; and Valvulineria depressa Cushman, 1926. Smith (1960) documented Luisian foraminiferal assemblages from the Monterey Formation overlying the San Onofre Breccia in the San Joaquin Hills. Although she noted that the Monterey Formation exposed west of Oso Creek consists of diatomaceous shale and siltstone containing abundant foraminifera, there was no mention of limestone, Relizian foraminifera, or the foraminif¬ era from Shell’s Moulton No. 14 well, which was located almost 1 mi (1.6 km) south of Area 2 and revealed the Luisian/Mohnian boundary at a depth of 410 ft. A paucity of foraminifera in the Topanga Formation in the Santa Ana Mountains was noted by Yerkes and Campbell (1979) and Schoellhamer et al. (1981). Their reports indicated that the foraminifera are represented by relatively few species, all of which indicate shelf-depth environments. This is in stark con¬ trast with the four assemblages recorded by Finger (1992) from the limestone section in the Moulton Parkway road cut. Ostra¬ codes probably occur in the Topanga Formation in Orange Coun¬ ty, but none have been reported. CONCLUSIONS The Saddleback Valley limestone occurs only in western Saddle¬ back Valley. It does not represent a reef, biostrome, or bioherm as had been reported previously. Rather, it comprises related but inconsistent and discontinuous sequences of calcareous beds that locally vary in lithology, lithification, paleontology, and stratigra¬ phy. Its lithologies range from unconsolidated limey sands to calcarenites and micrites. Conspicuous “Temblor” CPMS macro¬ fauna and sandy layers in the Saddleback Valley limestone resemble those of the Topanga Formation and indicate a warm, shallow-marine environment. Accordingly, microfossils correlate the limestone with the late Relizian Stage and MMCO between 17 and 15 My, and Sr s7 /Sr 86 analysis refines its age within the interval of 16.5-15.9 My, with a maximum mean age of 15.85 Ma, which is at the base of the middle Miocene. Mixed-depth ben¬ thic foraminiferal assemblages, however, indicate that these bio- clastic sediments had been dislodged from their provenance and transported to bathyal depths. The limey deposits thus represent the basal Monterey Formation, which is supported by field observations of limestone lenses within Monterey Formation mudstones and channel structures. A summarization of the geo¬ logic history of the limestone is as follows: At the end of the early Miocene, the Saddleback Valley area was located about 750 km south of its present location, at shallow depths in a marine channel between the mainland and a high-relief island or peninsula. Its rich subtropical “Temblor” biota included 48 ■ Contributions in Science, Number 524 Stadum and Finger: Saddleback Valley Limestone dense bryozoan patches that were baffles against which inverte¬ brates, in particular, accumulated in shoals. Coastal tectonics related to the San Andreas fault system caused landslides off the schistose highlands, which migrated across the subsiding channel and formed the San Onofre Breccia. Subsequent high-energy events (i.e., earthquakes and storms) triggered gravity flows (e.g., turbidity currents) that displaced the shallow-water Topanga-like sediments with live and dead elements of the “Temblor” biota down the slope of the newly created and other¬ wise sediment-starved basin, part of which had already started accumulating the characteristic hemipelagic muds of the Mon¬ terey Formation. Diagenesis transformed some of the richest carbonate deposits into an indurated and nearly pure limestone, while others formed unconsolidated calcaremtes with macro¬ fossils that are often highly concentrated and well preserved. In conclusion, recognition of channel-fill structures and mixed- depth microfossil assemblages support the long-held but recently challenged notion that the Saddleback Valley limestone is the bas¬ al subunit or informal member of the local Monterey Formation. Despite its affinities with the “Temblor” CPMS fauna and Topanga Formation, the limestone represents the initial subsi¬ dence of the basin into the deep-water realm associated with the Monterey Formation. ACKNOWLEDGMENTS This study is dedicated to our late colleagues Drs. John D. Cooper (Cali¬ fornia State University, Fullerton) and Takeo Susuki (University of Cali¬ fornia, Los Angeles), who assisted in both the field study and the effort that succeeded in preserving two limestone exposures as parklands. Charles L. Powell II (U.S. Geological Survey, Menlo Park) generously assisted in molluscan taxonomy, informed us of his observations in the study area, and provided provocative comments that enhanced the focus of this study. We thank independent consultants Richard S. Boettcher and Stanley A. Kling for providing supplementary micropaleontologic data, John McArthur (University College London) for his unpublished Sr data table, and Donald J. DePaolo (University of California, Berkeley) for assisting in the Sr age determinations. The University of California Museum of Paleon¬ tology (UCMP) granted funding for the Sr analyses, which were arranged through Dominique Weis and performed by Bruno Kieffer (University of British Columbia). We are also grateful to our many other colleagues who discussed various aspects of the Saddleback Valley limestone with us: Thomas A. Demere, Patricia Don Vito, and N. Scott Rugh (San Diego Natural History Museum), Paul K. Morton (TerraMins Inc,), Jere H. Lipps (John D. Cooper Archaeological and Paleontological Center), Richard L. Squires (California State University Northridge), the late Thomas W. Dibblee Jr. (Dibblee Institute), the late A. 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Notes on the Vaqueros and Temblor formations of the California Miocene with descriptions of new species. San Diego Soci¬ ety of Natural History Transactions 5(10):95— 1S2. Woodford, A.O, 1925. The San Onofre Breccia. University of California Publications, Bulletin of the Department of Geological Sciences 15:159-280. Wright, R.B. 1950. California's missions. Los Angeles: California Mission Trails Association, 96 pp. Yerkes, R.F., and R.H. Campbell. 1979. Stratigraphic nomenclature of the central Santa Monica Mountains, Los Angeles County, California. United States Government Printing Office, Washington, DC: U.S. Geological Survey Bulletin 1457-E, 39 pp. Zachos, J., M. Pagani, L. Sloan, £. Thomas, and K. Billups. 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292:686-693. Received 1 September 2015; accepted 14 March 2016. Contributions in Science, 524:50 2016 ERRATA Paleontology and Stratigraphy of the Miocene Saddleback Valley Limestone, Orange County, Southern California 1 Carol J. Stadum 2 and Kenneth L. Finger 3,4 PAGES 40-41 CORRECTED FIGURE CAPTION Figures 7-34 Saddleback Valley limestone gastropods (Figs. 7-29) and other invertebrates (Figs. 30-34). SDSNH numbers are locality/ specimen; H=height, W=width, E=length. 7, 8. Fissurella rixfordi Hertlein, 1928: 7, internal mold, SDSNFI 4312/136497, D=35 mm, W=24 mm; 8, external mold, SDSNH 4312/140187, D=30 mm, W=20 mm; 9. Cblorostoma sp., external cast, SDSNH 4312/93707, H=10 mm, W=10 mm; 10. Turritella ocoyana Conrad, 1855, internal mold, SDSNH 4312/140188, H=75 mm, bottom whorl W=20 mm; 11. Turritella sp., latex peel of external cast, SDSNH 4312/83244, H=33 mm, W=10 mm; 12. Forreria bartoni Arnold, 1910, original shell (missing upper part of spire), SDSNH 4521/83243, H=24 mm, W=33 mm; 13. Calyptraea sp., internal mold, SDSNH 4312/136498, H=21 mm, W=53 mm; 14. Crucibulum sp., apical view of internal mold showing septum partially attached to shell, SDSNH 4312/93719; 15. naticid, internal mold, SDSNH 4312/140186, H=35 mm, ultimate whorl W=32 mm; 16. Sinum cf. S. scopulosum (Conrad, 1849), internal mold, SDSNH 4312/93721, H=25 mm, W=28 mm (ultimate whorl); 17. Troph on kernensis Anderson, 1905, latex peel of external cast, SDSNH 4312/93835, H=24 mm, W=l4 mm; 18, 19. Trophosycon sp., internal mold: 18, apical view, SDSNH 4520/98691, W=62 mm; 19, side view, SDSNH 4312/83239, H=68 mm, penultimate whorl W=66 mm; 20, 21. fasciolariid, internal mold, SDSNH 4312/93739: 20, apical view, W=80 mm; 21, side view H=97 mm; 22. Antillophos woodringi Addicott, 1970, latex peel of external cast, SDSNH 4312/ 83242, H=15 mm, W=9 mm; 23. Pyruconus cf. P. owenianus (Anderson, 1905), SDSN 4312/83241, H=36 mm, W=ll mm; 24. Pyruconus cf. P. hayesi (Arnold, 1909), internal mold, SDSNH 4312/ 140190, H=42 mm, W=30 mm; 25. Priscofusus geniculus (Conrad, 1849), peel of exterior cast, SDSNH 4312/83245, H=44 mm, W=23 mm; 26, 27. Megasurcula keepi (Arnold, 1907), internal mold, SDSNH 4312/83240, H=60 mm, W=30 mm: 26, latex peel of external cast; 27, internal mold; 28. cypraeid, internal mold, SDSNH 4520/98688, D=21 mm, W=15 mm; 29. vermetid, internal mold, SDSNH 4512/142016, image H=30 mm; 30. Gemelliporella afif. G. punctata Canu and Bassler, 1919, fragments, SDSNH4521/98668, image H=63 mm; 31, 32. Eucidaris cf. E. thouarsii (Valenciennes, 1846): 31, original spine with regularly spaced knobs, SDSNH 4521/120893, E=30 mm; 32, original test fragment, SDSNH 4521/140192, D=19 mm; 33. Vaquerosella merriami (Anderson, 1905), external dorsal view, SDSNH 4521/83249, H=18 mm, W=18 mm, note anal aperture (notch) at ventral margin; 34. Megabalanus tintinnabulum (Linnaeus, 1758), original shells, side view of attached pair, SDSNH 4312/140191, H=25 mm, W=20 mm. PAGE 42 CORRECTIONS IN CAPTION FOR FIGURES 35-52 40. SDSNH 4521/98676 should read SDSNH 4312/98676. 42. SDSNH 4521/98671 should read SDSNH 4521/140355. 49. SDSNH 4312/83248 should read SDSNH 4312/93791. 50. SDSNH 4312/180184 should read SDSNH 4312/140184. PAGE 44 CORRECTED FIGURE CAPTION Figure 56 Palaeophycus isp. (SDSNH 137452) in limey mudstone of Lithofacies D, SDSNH loc. 4312. 1 URL: www.nhm.org/scholarlypublications San Diego Natural History Museum, Department of Paleontology, 1788 El Prado, San Diego, California 92101, USA. 3 University of California Museum of Paleontology, 1101 Valley Life Sciences Building, Berkeley, California 94720, USA. 4 Corresponding author: Kenneth L. Finger, e-mail: kfinger@berkeley.edu © Natural History Museum of Los Angeles County, 2017 ISSN 0459-8113 (Print); 2165-1868 (Online) Contributions in Science, 524:51-109 9 December 2016 Revision of Aerophilus Szepligeti (Hymenoptera, Braconidae, Agathidinae) from Eastern North America, with a Key to Nearctic Species North of Mexico 1 Michael J. Sharkey , 2 ’ 4 Eric G. Chapman , 2 and Giulia Iza De Campos 3 ABSTRACT. The Nearctic species of Aerophilus Szepligeti, 1902, are revised with an emphasis on the fauna of the eastern USA. The generic name Lytopylus Foster, 1862, is shown to have been misapplied to the group revised here and it is replaced by Aerophilus. The following genera are synonymized with Aerophilus: Neomicrodus Szepligeti, 1908, syn. n.; Aerophilopsis Viereck, 1913, syn. n.; Aerophilina Enderlein, 1920, syn. n.; loxia Enderlein, 1920, syn. n.; Hormagathis Brues, 1926, syn. n.; Obesomicrodus Papp, 1971, syn. n.; Facilagathis van Achterberg and Chen, 2004, syn. n. The type species of Lytopylus (L. azygos Viereck, 1905) fits the generic concept of Austroearinus Sharkey, 2006; the latter is therefore synonymized with Lytopylus, and all species included in Austroearinus are transferred to Lytopylus as new combinations. Agathellina Enderlein, 1920, and Ditropia Enderlein, 1920, are synonymized with Lytopylus, syn. n. A list of all new combinations is included. Thirty-five species of Aerophilus are treated, with 16 described as new (i.e., A. arthurevansi, A. chapmani, A. davidsmithi, A. hopkinsensis, A. jdherndoni, A. klastos, A. kowlesae, A. malus, A. minys, A. pookae, A. rayfisheri, A. reginae, A. robertcourtneyi, A. stoelbae, A. terrymoyeri, A. tommurrayi). The senior author (M.J.S.), is the sole authority of these species. All 19 previously described species have new combinations (i.e., A. abdominalis, A. aciculatus, A. acrobasidis, A. bakeri, A. binominatus, A. buttricki, A. calcaratus, A. crassicornis, A. difficilis, A. erythrogaster, A. nigripes, A. ninanae, A. nucicola, A. perforator, A. reticulatus, A. rugareolatus, A. tenuiceps, A. usitatus, A. wyomingensis ). Several new synonyms are proposed (i.e., Agathis atripes Cresson is synonymized under Agathis nigripes Cresson, Bassus pini Muesebeck is synonymized under Aerophilopsis erythrogaster Viereck, and Agathis wyomingensis Viereck is removed from synonymy with Agathis nigripes Cresson). An illustrated key, image plates, and distribution maps are included for each species. The revision is primarily based on newly collected material from Kentucky, Virginia, and West Virginia, for which molecular data were available. A phylogenetic analysis of Aerophilus, based on 28S and cytochrome c oxidase subunit I (COI), with representatives from all major biomes, is included. Another tree, based solely on COI data is included to show species divergences, which was used in conjunction with morphological data to delimit species. INTRODUCTION Agathidinae are a moderately diverse subfamily of Braconidae with about 1,200 described species (Yu et ah, 2015), and many times that number are yet to be named. Larvae are parasitoids of lepidopteran caterpillars of a multitude of families. Most agathidine genera, including Aerophilus , attack an early instar caterpillar and are quiescent until the host has reached the final instar and is ready to spin a cocoon. At this point in time the parasitoid larva becomes active and quickly consumes the host, i.e., they are koinobiont endoparasitoids. Aerophilus is world¬ wide in distribution, with the exception of Antarctica. Aerophilus is an unusual genus of Agathidinae in that host Lepidoptera are in a wide range of higher taxa. Most genera of Agathidinae seem to be restricted to one or a few closely related families of Lepidoptera. The collective set of Aerophilus species attack a wide range of host families. Even within the small sample of 10 reared species of Aerophilus represented in the revision of Costa Rican species (Sharkey et ah, 2011), five host families are attacked (i.e., Crambidae, Elachistidae, Pyralidae, Thyrididae, and Tortricidae). However, all of these are leaf¬ rolling and leaf-tying small caterpillars. Members of Aerophilus are conspicuous in not using species of caterpillars that feed exposed on leaf surfaces (e.g., butterflies and macro-moths). Individual species of Aerophilus tend to be host specific. Of the 1 URL: www.nhm.org/scholarlypublications 2 Department of Entomology, University of Kentucky, Lexington, Kentucky 40506, USA. 3 Universidade Federal de Sao Carlos-UFSCar, Departamento de Ecologia e Biologia Evolutiva, Sao Carlos, SP, Brasil. 4 Corresponding author: Michael J. Sharkey, E-mail: msharkey@uky.edu 10 species treated from Costa Rica, all but one, A. jessiehillae, are host specific (i.e., each attacks only one host species). And even the hosts of A. jessiehillae are very closely related to each other. The results of the phylogenetic analyses (fig. 1 in Sharkey et ah, 2011) show that host family is constrained by phylogenetic history, with sister species of Aerophilus sharing the same host family in every case (three). Several species of Aerophilus have been employed in biological control attempts. Aerophilus rufipes (as Bassus diversus) was introduced into the eastern USA from Japan to combat the oriental fruit moth, Grapholitha molesta (Busck), Tortricidae, over a period of a number of years in the 1930s (Allen and Yetter, 1949). Despite initial signs of success it does not appear to have become established. Aerophilus rufipes was also imported to California in the 1990s to control the codling moth, Cydia pomonella L., Tortricidae. However it too failed to become established (Mills, 2005). Aerophilus acrobasi¬ dis has been used in the biocontrol of Acrobasis nuxvorella (the pecan nut casebearer) and Cydia caryana (the hickory shuck- worm) (Ellington et ah, 1995; Romero et ah, 2001) with unknown results. Although this revision is primarily based on newly collected material from Kentucky, Virginia, and West Virginia, all Nearctic species are included in the key. There are two principal reasons for the emphasis on eastern species. The first is that there are very few if any species with distributions that span the USA or Canada from east to west, therefore including both faunas would pointlessly complicate the key. The second reason for the emphasis on eastern species is that many species are difficult to separate on morphological grounds and molecular data from fresh specimens was necessary. Intensive Malaise trap collecting has been conducted over the past decade in Kentucky by members of the Sharkey lab at the University of Kentucky and © Natural Elistory Museum of Los Angeles County, 2016 ISSN 0459-8113 (Print); 2165-1868 (Online) 52 ■ Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus Figure 1 Lytopylus azygos, holotype: A. lateral habitus, B. tarsal claw, C. forewing, D. dorsal mesosoma, E. lateral mesosoma, F. propodeum showing sculpture and narrow sclerite between metasomal and hind coxal foramina. by Dr. David Smith and colleagues in Virginia and West Virginia. Therefore, we have a fair representation of some eastern species, especially those at mid-latitudes, for which molecular data were obtained. The previously described western species are included in the key, but they are clearly noted as being western, so they can be quickly passed over in most cases. There are many more undescribed western species, some of which are included in the cladograms (Figs. 1 and 2), and a separate revision on these is in preparation (Sharkey and Chapman, in prep.). The species with the widest east-west distribution confirmed here is A. rayfisberi, which occurs in North Dakota and Kentucky. It is likely that a good number of species are midwestern with this magnitude of range. Other species such as A. nigripes have a wider published distribution; however, in these cases the species identifications are suspect. Six of the new eastern species are represented by singletons, and several more species are known from fewer than four specimens. These data suggest, despite adding 16 species to the eastern fauna, that there are many more eastern species yet to be discovered. New World members of Aerophilus may be distinguished from other agathidines by the almost universal presence of longitudinal striae on the third metasomal median tergite, at least in some transverse depressions, and by the structure of the propodeal foramen (i.e., a relatively wide space between the metasomal and hind coxal foramina and a strong transverse carina connecting the dorsal margins of the coxal foramina). A key to the Nearctic genera of Agathidinae is provided in Sharkey and Chapman (2015); unfortunately, in this key, the name Lytopylus must be replaced by Aerophilus and Austroearinus must be replaced by Lytopylus to conform to the corrections made here. The sister group to Aerophilus is Braunsia Kriechbaumer, which is re¬ stricted to the Old World and is mostly tropical or subtropical (Sharkey et al., 2006; Sharkey and Chapman, 2015). Muesebeck (1927) revised the species of Aerophilus (as members of Bassus) in his revision of the species of Agathidinae north of Mexico. He treated 16 nominal species in this revision and described several more in subsequent publications (Muese¬ beck, 1932, 1940). No Nearctic species has been described since 1940. Because Muesebeck’s (1927) key includes what are now considered other genera (i.e., Alahagrus, Agathis s.s., Aphela- gathis, Lytopylus, Pneumagathis, Therophilus, and Neothlipsis), it is unduly complicated; it is also missing the species described since 1927. Simbolotti and van Achterberg (1992) included the six species of Aerophilus from the western Palearctic under the umbrella of Bassus. Sharkey (1996) included two species from Japan, and Sharkey et al. (2009) recorded two species occurring in Thailand. Van Achterberg and Long (2010) recorded two species for Vietnam, one under Lytopylus and one under Facilagathis. Farahani et al. (2014) described a new species from Iran and included a key to the West Palearctic species of Aerophilus (as Lytopylus ). Sharkey et al. (2011) revised the 10 species reared in Costa Rica, as Lytopylus. One species has been recorded from Australia (Stevens et al., 2010). Aerophilus is diverse in Africa (where it has never been revised) and throughout the New World. The genus is not species-rich in Eurasia or in the Australian and Oriental regions, and it is in unknown in Pacifica. For a world distribution of the Aerophilus specimens we have included in our Symbiota database, see our map online (http://bit.ly/lM9DhqM). Figure 2 Lytopylus azygos: A. dorsal metasoma, B. anterior head. Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus ■ 53 METHODS Morphological terms are from Sharkey and Wharton (1997) and are matched to the Hymenoptera Anatomy Ontology (HAO; http://glossary.hymao.org; Yoder et al., 2010). Anatomical concepts in HAO are provided to enable readers to confirm their understanding of the anatomical structures being referenced. To find out more about a given structure, including images, references, and other metadata, simply search for the anatomical structure at hand and select the best match from the list that appears (e.g., typing “frons” returns a list of 11 possible matches). In electronic versions of this paper, terms are hyperlinked to the ontology the first time they appear, either in the text, key, or subsequent species descriptions. Museum acronyms found in the “Specimens Examined” sections of this paper are taken from “Abbreviations for Insect and Spider Collections of the World” (Evenhuis, 2014). Host records for each species were taken from Taxapad (Yu et al., 2015). All species are treated with a diagnosis and distributional data. For previously published species, the states and provinces from which they are recorded are listed; however, due to the degree of misidentifica- tions that are present in collections, only those specimens determined by M.J.S. are included in the linked distribution maps. These records are stored in the Symbiota database (Gries et al., 2014) under the Hymenoptera Institute Collection (HIC). The maps were generated by conducting a map search for each species on the SCAN portal (Symbiota Collections of Arthropods Network; http://symbiota4.acis.ufl.edu/scan/portal/), querying only specimens in the HIC. This generates a map URL that contains a search query for the taxon at hand in the HIC database. Mouse over any data point on the maps to access the details of each record (including images if present). These are not static maps; therefore, as georeferenced specimens are added to the HIC Symbiota database, they will automatically be added to the map in real time. Also, because these data are public, anyone can generate a map search for any taxon in our Symbiota database: From the Symbiota link above, under the Search menu, select Map Search. This will open a new tab with a map that has an Open icon at the top left, which opens a dialog box with many options for searching any or all of the databases in the Symbiota network. If one wishes only to search within the HIC database, select only HIC under the Collections tab. All species are illustrated with color photos using a JVC digital camera mounted on a Leica MZ16 microscope and Auto- Montage® stacking software. Species descriptions are of the holotype, and variation is given in parentheses. DNA EXTRACTION, PCR, AND SEQUENCING DNA was extracted from individual legs with the Qiagen DNeasy Blood and Tissue Kit using the animal tissue protocol (Qiagen Inc., Chatsworth, California, USA). The mitochondrial cyto¬ chrome c oxidase subunit I (COI) gene was amplified with the COI primer pairs LepFl and LepRl (—655 bp between the primers; Hebert et al., 2004), and when this fragment did not amplify, we employed LepFl and Creml55R (Tucker et al., 2015) to amplify a smaller fragment ( — 135 bp). We also sequenced the D2-D3 regions of 28S using the primer pairs 28SD2F (Belshaw and Quicke, 1997) and D3R (Harry et al., 1996). PCR was conducted using Takara reagents for COI, with each reaction consisting of IX buffer, 0.3 mM nucleotides, 0.4 pM of each primer, 0.625 U Takara Ex Taq, ddH 2 0, and 1-3 pL template DNA in a total reaction volume of 25 pL. The thermal cycling protocol had an initial denaturation period at 95 °C for 2.5 min, followed by 40 cycling steps which denatured at 95 °C for 30 s, annealed at 44 °C for 30 s and extended at 68 °C for 45 s, with a final extension step of 72 °C for 7 min. For 28S, PCR reactions consisted of Qiagen IX buffer, 4 mM MgS0 4 , 0.3 mM dNTP, 0.4 pM of each primer, 0.625 U Qiagen Taq, ddH 2 0, and 1-3 pL template DNA with a total reaction volume of 25 pL. Thermal cycling was as above except annealing at 53 °C, extending for 70 s, and a total of 35 cycles. PCR products were outsourced for Sanger sequencing either by the Advanced Genetic Technologies Center (University of Kentucky, Lexington, Kentucky, USA) or Beckman Coulter Genomics (Danvers, Massachusetts, USA) using labeled dideox- ynucleotides with ABI 3730 BigDye Terminator mix v. 3.0 or with ABI PRISM 3730x1, BigDye Terminator mix v. 3. 1 (Applied Biosystems, Foster City, California, USA). DNA ASSEMBLY AND PHYLOGENETIC ANALYSIS Bidirectional sequences were aligned and edited using Geneious Pro (v. 6.1.5; Drummond et al., 2009) and multiple alignments were assembled using the default settings on the MAFFT server (http://www.ebi.ac.uk/Tools/msa/mafft/; v. 7; Katoh et al., 2006). We conducted maximum likelihood (ML) analyses on two data sets: COI (655 nt, 110 operational taxonomic units [OTU]) and a concatenated (using MacClade v. 4.08; Maddison and Maddison, 2005) COI+28S (1,161 nt, 127 OTU) data set using Garli (v. 2.01; Zwickl, 2006). The data were partitioned by gene region and codon position for 28S+COI (total of four partitions) and by codon position for COI (three partitions). We applied the most complex model available (GTR+I+G; Rodriguez et al., 1990) to each partition as per recommendations of Huelsenbeck and Rannala (2004) for likelihood-based analyses. Garli applies separate parameter estimates to each partition. For both data sets, a 200-replicate ML analysis was conducted using the default settings. Nodal support was assessed by conducting a 500- replicate ML bootstrap analysis (Felsenstein, 1985) on each data set using the default settings, with three independent search replicates per bootstrap replicate. The COI analyses (Supp. Figs. 1 and 2; summarized in Fig. 3) was used to help make decisions regarding species delimitation, as 28S is not variable enough to reliably separate closely related lineages, whereas the analyses of the COI+28S data set (Supp. Figs. 3 and 4; summarized in Fig. 4) was conducted to find the best estimate of phylogeny. A Bayesian inference (BI) phylogenetic analysis was also conducted on the COI+28S data set with MrBayes (v. 3.1.2; Huelsenbeck and Ronquist, 2001; Ronquist and Huelsenbeck, 2003). As in the Garli analyses, the data were partitioned by gene region and codon position. To allow each partition to have its own set of parameter estimates, revmat, tratio, statefreq, shape, and pinvar were all unlinked during the analyses. To obtain the most accurate branch length estimates possible, the option prset ratepr = variable (which assigns a separate branch length parameter for each partition) was employed as per the recommendations of Marshall et al. (2006). Two independent, simultaneous BI searches were run for 100 million generations, saving a tree every 1,000 generations, with four search chains each. The average standard deviation of split frequencies fell below 0.02 just before 80 million generations. The 20,000 post¬ burn-in trees from each run (40,000 total), determined by examination of the log probability of observing the data by generation plot with Tracer (v. 1.5; Rambaut and Drummond, 2009), were used to calculate the majority rule consensus tree using PAUP* (v. 4.0(310; Swofford, 2003). The tree of highest posterior probability from the Bayesian analysis is shown in Supplemental Figure 5, and the majority rule consensus tree is shown in Supplemental Figure 6. The data sets analyzed herein are available from the authors upon request. 54 ■ Contributions in Science, Number 524 Sharkey et ah: Revision of Aerophilus RESULTS N OMEN CL AT ORIAL CONSIDERATIONS The senior author (M.J.S.) recently had the opportunity to view the type of Lytopylus Forster, 1862, Lytopylus azygos Viereck, 1905. Forster (1862) did not include any species under Lytopylus, therefore the type species by monotypy became the first included species, in this case L. azygos. Unfortunately this species does not conform to the generic concept that has been applied to it in recent years (Sharkey et ah, 2009, 2011; van Achterberg, 2011). Rather, it fits the concept of Austroearinus (Figs. 1 and 2) proposed by Sharkey et al. (2006). This necessitates the synonymy of Austroearinus under Lytopylus n. syn. Furthermore, two other nominal genera fit my (M.J.S.) concept of Austroearinus that must be synonymized under Lytopylus , namely, Ditropia Enderlein, 1920, n. syn. and Agathellina Enderlein, 1920, n. syn. Figure 1 is a plate of the type species of Lytopylus. The type is missing both the head and metasoma; therefore, Figure 2 is included to show these body parts in what I (M.J.S.) believe to be a conspecific specimen. Characters consistent with the concept of Austroearinus are: sessile second submarginal cell, that is, lacking a petiole (Fig. 1C); notauli not impressed (Fig. ID); propodeum with sculpture confined to the midline posteriorly (Fig. IF); terga of metasoma smooth except for pair of longitudinal carinae on first median tergite (Fig. 2A); interantennal space with a weak medial depression (Fig. 2B); sclerite between metasomal and coxal foramina narrow. The oldest name that applies to the old (misinformed) concept of Lytopylus is Aerophilus Szepligeti, 1902. All species recently described under Lytopylus are transferred to Aerophilus. The new combinations for both Aerophilus and Lytopylus are presented below. Aerophilus Szepligeti, 1902 TYPE SPECIES. Aerophilus brullei Szepligeti, 1902 (by monotypy). Aerophilina Enderlein, 1920, syn. n. Type species: Aerophilina bicristata Enderlein, 1920. Aerophilopsis Viereck, 1913, syn. n. Type species: Bassus erythrogaster Viereck, 1913. Facilagathis van Achterberg and Chen, 2004, syn. n. Type species: Facilagathis spinulata van Achterberg and Chen, 2004. Hormagathis Brues, 1926, syn. n. Type species: Hormagathis mellea Brues, 1926. Ioxia Enderlein, 1920, syn. n. Type species: Ioxia faceta Enderlein, 1920. Neomicrodus Szepligeti, 1908, syn. n. Type species: Neomicro- dus boliviensis Szepligeti, 1908. Obesomicrodus Papp, 1971, syn. n. Type species: Obesomicro- dus niger Papp, 1971. NEW COMBINATIONS (alphabetized by epithet) Microdus astioles Nixon, 1950, to Aerophilus astioles Bassus barbieri Simbolotti and van Achterberg, 1992, to Aerophilus barbieri Metriosoma bicarinatum Enderlein, 1920, to Aerophilus bicari- natum Aerophilina bicristata Enderlein, 1920, to Aerophilus bicristatus Neomicrodus boliviensis Szepligeti, 1908, to Aerophilus boli¬ viensis Lytopylus bradzlotnicki Sharkey, 2011, to Aerophilus bradzlot- nicki Metriosoma brasiliense Enderlein, 1920, to Aerophilus brasi- liense Lytopylus brevitarsus van Achterberg, 2011, to Aerophilus brevitarsus Lytopylus colleenhitchcockae Sharkey, 2011, to Aerophilus colleenhitchcockae Agathis ebula Nixon, 1950, to Aerophilus ebulus Agathis burmensis Bhat and Gupta, 1977, to Aerophilus burmensis (junior synonym of Agathis ebula) Ioxia faceta Enderlein, 1920, to Aerophilus facetus Microdus femoratus Cameron, 1887, to Aerophilus femoratus Metriosoma flavicalcar Enderlein, 1920, to Aerophilus flavical- car Microdus fortipes Reinhard, 1867, to Aerophilus fortipes Lytopylus gregburtoni Sharkey, 2011, to Aerophilus gregburtoni Microdus infumatus Granger, 1949, to Aerophilus infumatus Lytopylus jessicadimauroae Sharkey, 2011, to Aerophilus jessi- cadimauroae Lytopylus jessiehillae Sharkey, 2011, to Aerophilus jessiehillae Microdus leucotretae Nixon, 1941, to Aerophilus leucotretae Bassus macadamiae Briceno and Sharkey, 2000, to Aerophilus macadamiae Microdus melanocephalus Cameron, 1887, to Aerophilus mela- nocephalus Hormagathis mellea Brues, 1926, to Aerophilus melleus Lytopylus mingfangi Sharkey, 2011, to Aerophilus mingfangi Obesomicrodus niger Papp, 1971, to Aerophilus niger Cremnops nigrobalteatus Cameron, 1911, to Aerophilus nigro- balteatus Bassus pastranai Blanchard, 1952, to Aerophilus pastranai Lytopylus persicus Farahani and Talebi, 2014, to Aerophilus persicus Agathis philippinensis Bhat and Gupta, 1977, to Aerophilus philipp inensis Microdus pilosus Tobias, 1976, to Aerophilus pilosus (this species fits well with the concept of Aerophilus, with the exception of a shorter distance between the hind coxal cavities and the metasomal foramen.) Microdus rugulosus Nees, 1834, to Aerophilus rugulosus Lytopylus rebeccashapleyae Sharkey, 2011, to Aerophilus rebeccashapleyae Lytopylus robpringlei Sharkey, 2011, to Aerophilus robpringlei Microdus romani Shestakov, 1940, to Aerophilus romani Bassus ater Chou and Sharkey, 1989, to Aerophilus ater (junior synonym of Microdus romani) Microdus rufipes Nees, 1812, to Aerophilus rufipes Microdus amurensis Shestakov, 1940, to Aerophilus amurensis (junior synonym of Microdus rufipes) Bassus diversus Muesebeck, 1933, to Aerophilus diversus (junior synonym of Microdus rufipes) Braunsia germanica Enderlein, 1904, to Aerophilus germani- cus (junior synonym of Microdus rufipes) Lytopylus sandraberriosae Sharkey, 2011, to Aerophilus sandra- berriosae Microdus sculptilis Tobias, 1986, to Aerophilus scuptilis Facilagathis spinulata van Achterberg and Chen, 2004, to Aerophilus spinulatus Bassus tayrona Campos, 2007, to Aerophilus tayrona Lytopylus vaughntani Sharkey, 2011, to Aerophilus vaughntani REINSTATED ORIGINAL COMBINATIONS Aerophilus brullei Szepligeti, 1902 (from Bassus) Aerophilus lamelliger Granger, 1949 (from Bassus) Contributions in Science, Number 524 Sharkey et ah: Revision of Aerophilus ■ 55 58 54 5£ — i Braunsia spp. (17) Aerophilus sp. USA: AZ -Aerophilus chapmani USA: KY -Aerophilus abdominalis USA: KY 98 Aerophilus klastos USA: KY <] Aerophilus rufipes France (5) 100 1 Aerophilus reginae USA: KY (2) 89 Aerophilus rugulosus Hungary -Aerophilus sp. France (2) Aerophilus sp. Congo 99 66 < Aerophilus sp. Congo (4) 62 Aerophilus difficilis USA: FL 87 < Aerophilus malus USA: WV (2) 54 89 u 1 Aerophilus arthurevansi USA: VA (3) Aerophilus davidsmithi USA: WV 74 ♦" <3 Aerophilus sp. Mexico: Yucatan (4) □ Aerophilus calcaratus USA: FL, KY, WV (9) 59 Aerophilus robertcourtneyi USA: KY -Aerophilus stoelbae USA: KY — Aerophilus terrymoyeri USA: IL 93 93 < Aerophilus sp. Mexico: Yucatan (2) < Aerophilus sp. USA: AZ (3) 87 < Aerophilus minys USA: KY, VA, WV (12) Aerophilus spp. USA: AZ, CA (4) 51 Aerophilus sp. USA: CA Aerophilus sp. USA: CA Aerophilus sp. USA: CA (2) 61 Aerophilus sp. USA: CA -Aerophilus rayfisheri USA: KY, ND (2) Aerophilus sp. USA: CA —^ Aerophilus sp. USA: AZ (2) W MI 4; stream rot loi;s 25‘. i-5.vn.2007. 14^60 Hymenoptcra Institute Figure 32 Aerophilus reginae , holotype: A. lateral habitus, B. anterior head, C. wings, D. dorsal habitus. Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus ■ 93 Figure 33 Aerophilus reticulatus, holotype: A. lateral habitus, B. wings, C. anterior head, D. dorsal head and thorax, E. propodeum, F. tergite 1, G. posterior tergites, tergites 2-7. HI 252- 1 USA: KY Fayell9Co Lexington. Scuth Farm 54-32 18’W 37’S3 58'V Malaise Pitz & Seltmann 30 Aug - 4 Sep 2002 ETYMOLOGY. Named in honor of Ray Fisher, former graduate student of M.J.S. and collector of the type specimen. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female: Kentucky, Fayette Co., Lexington, Brier E Rd., 38.00867°N, 84.38778°W, 3.viii.2008, Ray Fisher, (HIC H1212). Paratype female: North Dakota, Randsom Co., 7 mi SE Sheldon, 23.vii.1995, J.R. Powers (CISC, specimen H4922). For a map of the examined material see http://bit.ly/lkWe7Fj. Aerophilus reginae Sharkey n. sp. Figure 32 DIAGNOSIS. Face not elongate; wings hyaline; face yellow, occiput mostly melanic; first metasomal median tergite pre¬ dominantly melanic; syntergite 2+3 partly to entirely yellow or tan colored with some more posterior terga partly or entirely melanic. DESCRIPTION. Length 4.4 mm. Ovipositor length 5.5 mm. Flagellomere number 29 (28-29). Gena not elongate; ratio of length of malar space to eye height, viewed laterally, 0.3. Notauli barely perceptible. Propodeum with distinct cells, median cell narrow, some rugae in median cell and elsewhere. Forewing hyaline. First metasomal median tergite clearly longer than posterior width. First median tergite weakly rugosostriate over much of surface, smooth posterolaterally; pair of longitudinal carinae weak and extending to midlength of tergite. Median syntergite 2+3 mostly smooth with longitudinal striae in the three transverse depressions. HOSTS. Gelechiidae: Coleotechnites apicitripunctella, Coleo- technites gibsonella. ETYMOLOGY. Named in honor of the junior author’s (G.I.d.C.) mother, Regina. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female: Kentucky, Harlan Co., Blanton Forest N.P., 36.86370°N, Figure 34 Aerophilus robertcourtneyi, holotype: A. lateral habitus, B. anterior head, C. wings, D. dorsal habitus. 94 ■ Contributions in Science, Number 524 Sharkey et ah: Revision of Aerophilus Figure 35 Aerophilus rugareolatus, holotype: A. lateral habitus, B. wings, C. lateral head and dorsal mesosoma, D. anterior head, E. propodeum. 83.36998°W, 25.iv-5.vii.2007,1,476 ft. [450 m.], (HIC H1280). Paratypes: 58 specimens, 36 females, 22 males. Canada: Ontario, Quebec, New Brunswick. USA: New York, Kentucky, North Carolina, Florida. Late April to late August. All paratypes except one from Kentucky are deposited in the CNC. For a map of the examined material see http://bit.ly/lNDuvnJ. Aerophilus reticulatus (Muesebeck, 1932) n. comb. Figure 33 Bassus reticulatus Muesebeck, 1932:332. Other combinations: Agathis. DIAGNOSIS. Forefemur pale in apical 2/3, melanic in basal 1/3; raised areas of metasomal median syntergite 2+3 longitudi¬ nally striogranulate or granulate; propodeum areolate rugose with some granulae. DESCRIPTION. Length 4.7 mm. Ovipositor length 5.1 mm. Flagellomere number 20, contrary to the Muesebeck’s (1932) description, which states 21 segments, indicating 19 flagello- meres (two female paratypes with 18 flagellomeres). Gena elongate; ratio of length of malar space to eye height, viewed laterally, 1.1. Notauli absent and lacking pits. Propodeum entirely rugosoreticulate with a deep irregular median longitu¬ dinal cell/depression. Forewing infuscate. First metasomal median tergite about as long as, or slightly longer than, posterior width. First median tergite almost entirely rugoso- granulate with a hint of irregular longitudinal striation; pair of carinae short and blunt. Median syntergite 2+3 entirely sculptured except for smooth posterior margin; sculptured mostly with granulate microstriae, striae deeper and more pronounced in transverse depressions. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female, Illinois (southern), C. Robertson (USNM type 44083). Published state records: Illinois, Kansas, Missouri. For a map see http://bit.ly/lM01boK. Aerophilus robertcourtneyi Sharkey n. sp. Figure 34 DIAGNOSIS. Face slightly elongate; head color entirely pale, orange to yellow; hind femur entirely pale; similar to A. abdominalis and A. klastos but differs from those species in the smooth, barely perceptible notauli. DESCRIPTION. Length 4.4 mm. Ovipositor length 4.0 mm. Flagellomere number 26 (26-27). Gena slightly elongate; ratio of length of malar space to eye height, viewed laterally, 0.6. Notauli barely perceptible and lacking pits. Propodeum with distinct cells, cells are filled with smooth rugae. Forewing infuscate. First metasomal median tergite about as long as, or slightly longer than, posterior width. First median tergite entirely striate; pair of carinae weak and almost reaching midlength of tergite. Median syntergite 2+3 longitudinally striate except extreme apex smooth. ETYMOLOGY. Named in honor of Mr. Robert Courtney, owner of the horse farm where the paratype was collected. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female: Kentucky, Fayette Co., South Farm, 37.9755°N, 84.53633°W, 30.viii-4.ix.2002, Pitz and Seltmann, (HIC, spec¬ imen H1252). Paratype female, Kentucky, Fayette Co., Stone- bridge Horse Farm, 38.00467°N, 84.36817°W, 21.ix-8.x.2012 (HIC, specimen HI 0026). For a map of the examined material see http://bit.ly/ljUQsFa. Aerophilus rugareolatus (Viereck, 1917) n. comb. Figure 35 Bassus ( Lytopylus) rugareolatus Viereck, 1917:228. Other combinations: Agathis. DIAGNOSIS. Gena intermediate in length; hind femur entirely pale; head color mostly or entirely black, with some pale color on eye orbits or on gena; propodeum areolate rugose; metasoma entirely pale (according to Viereck, 1917). Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus ■ 95 Figure 36 Aerophilus stoelbae, holotype: A. lateral habitus, B. anterior head, C. wings, D. dorsal habitus. DESCRIPTION. Length 4.8 mm. Ovipositor length unknown, the sole known specimen is the holotype male. Flagellomere number unknown. Gena slightly elongate; ratio of length of malar space to eye height, viewed laterally, 0.6. Notauli very weakly impressed and lacking pits. Propodeum entirely rugoso- reticulate with an irregular median cell. Forewing infuscate. Metasoma missing from holotype. MATERIAL EXAMINED AND DISTRIBUTION. Holotype male, Connecticut, New Haven, 12.09.1904, B.H. Walden (USNM type 66253). Published state records: Connecticut. For a map of the examined material see http://bit.ly/limWNHB. Aerophilus stoelbae Sharkey n. sp. Figure 36 DIAGNOSIS. Small specimens 3.2-3.4 mm.; antenna with 22- 24 flagellomeres; notauli barely impressed; metasomal median tergites 1-3 often mostly striate. 96 ■ Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus Figure 37 Aerophilus tenuiceps, holotype: A. lateral habitus, B. wings, C. dorsal habitus, D. anterior head, E. dorsolateral head, F. lateral head and mesosoma, G. dorsal metasoma. DESCRIPTION. Length 3.2 mm (3.2-3.4). Ovipositor length 3.0 mm (3.0-3.2). Flagellomere number 22 (22-24). Gena not elongate; ratio of length of malar space to eye height, viewed laterally, 0.5 (0.4-0.5). Notauli barely perceptible and lacking pits. Propodeum with carinae forming moderately regular cells, median cell with weak carinae. Forewing infuscate. First metasomal median tergite about as long as, or slightly longer than, posterior width. First median tergite with weak shallow striae over half of surface (varying to entirely striate), smoother antero- and posteromedially; pair of longitudinal carinae extending to midlength of tergite (or slightly past). Median syntergite 2+3 mostly smooth with longitudinal striae in the three transverse depressions and with weak smooth striae in anterior lobe of median tergite 2; sometimes striate anterome- dially, anteriad first transverse depression and on first raised area (varying to entirely striate except posterior lobe of median tergite 3). ETYMOLOGY. Named in honor of Stephanie Stoelb, former technician extraordinaire in the Department of Entomology, University of Kentucky. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female: Kentucky, Hopkins Co., Thomas Farm, 37.33928°N, 87.68802°W, 22.viii-13.ix.2010, MT 4: swamp 2 (HIC H7655). Paratypes: 4 females, Kentucky, Breathitt Co., Robertson forest, 17- 18.ix.2010, 37.7883°N, 83.23833°W, (HIC 11799, 6502, 6695). For a map of the examined material see http://bit.ly/lP89enA. Aerophilus tenuiceps (Muesebeck, 1927) n. comb. Figure 37 Bassus tenuiceps Muesebeck, 1927:47. Other combinations: Agathis. DIAGNOSIS. Head, metasoma, and body of mesosoma entirely pale; notauli weakly impressed, barely perceptible; second median tergite smooth in raised areas. DESCRIPTION. Length 5.7 mm. Ovipositor length 4.9 mm. Flagellomere number 32. Gena not elongate; ratio of length of malar space to eye height, viewed laterally, 0.4. Notauli barely perceptible or weakly impressed. Propodeum with irregular cells, smooth weak rugae in cells. Forewing hyaline. First metasomal median tergite about as long as, or slightly longer than, posterior width. First median tergite weakly striate over much of surface, smooth posterolaterally; pair of carinae weak and extending to midlength of tergite. Median syntergite 2+3 mostly smooth with longitudinal striae in the three transverse depressions. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female: New Mexico, Wild Horse Canyon, Animas Mts. 5,000 ft. [1524 m] (USNM type 28687). Published state records: Arizona, New Mexico. For a map of the examined material see http://bit.ly/lXCkImp. Contributions in Science, Number 524 Sharkey et ah: Revision of Aerophilus ■ 97 USA: Illinois: Lee Co. Richardson Wildlife Foundation 41 c 42'26.91 "N. 89°11'12.79"W 19.vni-2.ix.2010. 252m ^7685 coll. Terry Moyer Figure 38 Aerophilus terrymoyeri, holotype: A. lateral habitus, B. anterior head, C. wings, D. dorsal habitus. Aerophilus terrymoyeri Sharkey n. sp. Figure 38 DIAGNOSIS. Second median tergite completely striate in raised areas; gena slightly elongate. DESCRIPTION. Length 4.6 mm. Ovipositor length 5.5 mm. Flagellomere number 30 (28-30). Gena slightly elongate; ratio of length of malar space to eye height, viewed laterally, 0.6. Notauli weakly impressed and lacking pits. Propodeum mostly smooth with well-defined cells, median cell with some rugae. Forewing infuscate. First metasomal median tergite about as long as, or slightly longer than, posterior width. First median tergite entirely striate except anteromedially between longitudinal carinae; pair of longitudinal carinae extending to midlength of tergite. Median syntergite 2+3 longitudinally striate except posterior half of tergite 3 smooth. ETYMOLOGY. Named in honor of Terry Moyer, manager of The Richardson Wildlife Foundation Reserve. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female: Illinois, Lee Co., Richardson Wildlife Foundation, 41.70748°N, 89.18689°W, 19.viii-2.ix.2010, 252 m, coll. Terry Moyer (HIC, specimen H7685). Paratype female, same data as holotype except 2-15.ix.2010 (HIC, specimen H7550). For a map of the examined material see http://bit.ly/lMl2oWL. Aerophilus tommurrayi Sharkey n. sp. Figure 39 DIAGNOSIS. Face moderately elongate; head, body of mesosoma, and all coxae melanic; first metasomal median tergite entirely striate. DESCRIPTION. Length 4.4 mm. Ovipositor length unknown, the sole known specimen is the holotype male. Flagellomere number 28. Gena elongate; ratio of length of malar space to eye height, viewed laterally, 0.7. Notauli well impressed with pits in anterior 1/3. Propodeum with irregular cells broken by irregular carinae and rugae. Forewing infuscate. First metasomal median tergite about as long as, or slightly longer than, posterior width. 98 ■ Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus Figure 39 Aerophilus tommurrayi, holotype: A. lateral habitus, B. wings, C. anterior head, D. dorsal habitus. First median tergite entirely striate with pair of weak carinae not extending past midlength of tergite. Median syntergite 2+3 with all transverse depressions and anterior-most elevated area striate. ETYMOLOGY. Named in honor of the collector of the type specimen. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female: Massachusetts, Groton, 42.60°N, 71.5667°W, 5.ix.2010, T. Murray (HIC, specimen H8496). For a map see http://bit.ly/limXnFs. Aerophilus usitatus (Gahan, 1919) n. comb. Figure 40 Bassus tenuiceps Gahan, 1919:119. Other combinations: Agathis. DIAGNOSIS. Propodeum entirely pale; transverse grooves of syntergite 2+3 all smooth; notauli with crenulae/pits in anterior 1/2 or more; first metasomal median tergite smooth except for pair of carinae; forefemur almost entirely melanic with a pale patch apically. DESCRIPTION. Length 4.7 mm. Ovipositor length 3.5 mm. Flagellomere number undetermined (broken after fla- gellomere 14). Gena not elongate; ratio of length of malar space to eye height, viewed laterally, 0.2. Notauli well impressed with pits in anterior 1/2. Propodeum smooth with well-defined cells. Forewing infuscate. First metasomal median tergite about as long as, or slightly longer than, posterior width. First median tergite smooth; pair of carinae extending past Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus ■ 99 Figure 40 Aerophilus usitatus, holotype: A. lateral habitus, B. anterior head, C. lateral head and thorax, D. wings, E. tergites 1-3, F. dorsal head and mesosoma, G. dorsal habitus. midlength of tergite. Median syntergite 2+3 smooth, transverse depressions lacking striae. HOSTS. Pyralidae: Acrobasis vaccinii. MATERIAL EXAMINED AND DISTRIBUTION. Holotype female: Massachusetts, East Wareham, 15.vii.1916, parasite of cranberry fruit-worm [sic] (USNM type 21609). Published state records: Massachusetts, Michigan. For a map of the examined material see http://bit.ly/lNbdfrn. Aerophilus tvyomingensis (Viereck, 1905) n. comb. Figure 41 Agathis tvyomingensis Viereck, 1905:284. Other combina¬ tions: Bassus. SYNONYMS. Synonymized under Agathis nigripes Cresson, 1865:297 by Muesebeck (1927) but reinstated here. Figure 41 Aerophilus wyomingensis, holotype: A. lateral habitus and wings, B. dorsal habitus, C. lateral head and mesosoma, D. anterior head, E. apex of hind tibia. 100 ■ Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus DIAGNOSIS. Western species. Similar to A. nigripes both of which have elongate faces. Unlike A. nigripes , A. wyomingensis lacks notauli and has shorter antennae with an average of 24 flagellomeres, rather than the 32-34 flagellomeres typical of A. nigripes. A number of undescribed western species further complicate the identity of both A. nigripes and A. wyomingensis. DESCRIPTION. Length 6.5 mm. Ovipositor length 6.0 mm. Flagellomere number (24-25) broken on holotype. Gena elongate; ratio of length of malar space to eye height, viewed laterally, 0.8. Notauli barely indicated (to completely absent) and lacking pits. Propodeum mostly smooth with a narrow spindle- shaped median longitudinal cell. Forewing infuscate. First metasomal median tergite about as long as, or slightly longer than, posterior width. First median tergite smooth with carinae extending to midlength of tergite. Median syntergite 2+3 smooth, lacking microsculpture in transverse depressions. HOSTS. Due to some historic confusion over the identity of this species the two hosts ( Homoeosoma electellum and Phaneta bucephaloides) attributed to A. nigripes may rather be hosts of A. wyomingensis. MATERIAL EXAMINED AND DISTRIBUTION. Holo¬ type female: Wyoming, Lusk,vii.l895 (SEMC). Published state records: Wyoming. For a map of the examined material see http://bit.ly/ljURNMO. ACKNOWLEDGMENTS We thank the collectors of all of the specimens employed in this revision, especially David Smith, who has been sharing Malaise trap residues for many years. This project was made possible in part by the support of Southwest Collections of Arthropods Network (SCAN) NSF EF 1207371. Funding for this research was provided by Flatch projects KY008041 and KY008065 (to M.J.S.). The information reported in this paper (15-08-127) is part of a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director. We also thank Dominique Zimmermann for the translation of some German text; Dicky Yu for help with literature; Brian Brown, Paul Hanson, and an anonymous reviewer for comments on a draft; Doug Yanega for lessons on the Code of Zoological nomenclature; and Kees van Achterberg for advice. LITERATURE CITED Allen, H.W., and W.P. Yetter. 1949. Bassus diversus, an oriental fruit moth parasite established in the United States. Journal of Economic Entomology 42(3):540. Ashmead, W.H. 1889. Descriptions of new Braconidae in the collection of the U.S. National Museum. 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H2373 Congo: Pool KU059046 — Braunsia sp. H2375 Congo: Pool KU059050 — Braunsia sp. H2376 Congo: Pool KU059044 — Braunsia sp. H2377 Congo: Pool KU059047 — Braunsia sp. H2378 Congo: Pool KU059051 — Braunsia sp. H2379 Congo: Pool KU059048 — Braunsia sp. H2380 Congo: Pool KU059049 — Braunsia sp. H2381 Congo: Pool KU059052 — Braunsia sp. H1895 Equatorial Guinea — KU058995 Braunsia sp. H1889 Kenya: Eastern — KU058991 Braunsia sp. H1884 Kenya: Nyanza ATRMK333-11 KU058996 Braunsia sp. H1890 Kenya: Nyanza — KU058994 Braunsia sp. H1892 Kenya: Nyanza ATRMK338-11 KU058992 Braunsia sp. H2384 Madagascar: Antsiranana ATRMK543-11 — Braunsia sp. H1297 Madagascar: Toliara ATRMK447-11 — Braunsia sp. H1893 Uganda ATRMK435-11 KU058999 Aerophilus abdominalis H1253 USA: GA — KU059000 Aerophilus abdominalis H1313 USA: KY ATRMK294-11 KP943685 Aerophilus arthurevansi H5615 USA: VA Paratype KP943622 — Aerophilus arthurevansi H8554 USA: VA Holotype KP943628 — Aerophilus arthurevansi H8556 USA: VA Paratype KP943629 — Aerophilus calcaratus H209 USA: KY KU059053 KU059001 Aerophilus calcaratus H1220 USA: WV ATRMK559-11 — Aerophilus calcaratus H1246 USA: WV ATRMK566-11 — Aerophilus calcaratus H1247 USA: FL ATRMK567-11 — Aerophilus calcaratus H1249 USA: WV ATRMK280-11 KU059002 Aerophilus calcaratus H1250 USA: TN ATRMK569-11 — Aerophilus calcaratus H1457 USA: WV ATRMK307-11 KP943692 Aerophilus calcaratus H1461 USA: KY — KU059003 Aerophilus calcaratus H1478 USA: WV ATRMK308-11 KU059004 Aerophilus calcaratus H1487 USA: WV ATRMK310-11 KU059005 Aerophilus chapmani H1350 USA: KY Holotype ATRMK302-11 KP943689 Aerophilus davidsmithi H1267 USA: WV Holotype ATRMK289-11 KP943679 Aerophilus difficilis H11965 USA: FL KP943633 — Aerophilus erythrogaster H1216 USA: KY KP943615 KU059006 Aerophilus erythrogaster H8274 USA: KY KP943626 — Aerophilus hopkinsensis H7424 USA: KY Paratype KP943623 KU059007 Aerophilus hopkinsensis H7438 USA: KY Holotype KP943624 KU059008 Aerophilus jdherndoni H1217 USA: KY Holotype KU059054 KU059009 Aerophilus jdherndoni H1218 USA: KY Paratype ATRMK557-11 KP943676 Aerophilus jdherndoni H1256 USA: KY Paratype ATRMK285-11 KU059010 Aerophilus jdherndoni H1265 USA: KY Paratype KU059055 KU059011 Aerophilus jdherndoni H1268 USA: KY Paratype — KU059012 Aerophilus jdherndoni H1307 USA: KY Paratype ATRMK293-11 KU059013 Aerophilus jdherndoni H1311 USA: KY Paratype — KU059014 Aerophilus jdherndoni H1380 USA: KY Paratype — KU059015 Aerophilus jdherndoni H1400 USA: KY Paratype ATRMK306-11 KU059016 Aerophilus jdherndoni H1486 USA: KY Paratype — KU059017 Aerophilus jdherndoni H6033 USA: WI Paratype KU059056 KU059018 Aerophilus jdherndoni H7682 USA: KY Paratype — KU059019 Aerophilus jdherndoni H8848 USA: KY Paratype KU059057 — Aerophilus jdherndoni H10030 USA: KY Paratype KU059058 — Aerophilus klastos H205 USA: KY Holotype KP943612 KP943664 Aerophilus kowlesae H7683 USA: KY Holotype KP943625 KU059020 Aerophilus malus H1248 USA: WV Paratype ATRMK568-11 — Aerophilus malus H1484 USA: WV Holotype ATRMK309-11 KP943693 Aerophilus minys H206 USA: KY Paratype KU059059 KU059021 Aerophilus minys H966 USA: KY Paratype ATRMK267-11 KU059022 Aerophilus minys H1215 USA: KY Paratype ATRMK279-11 KU059023 Aerophilus minys H1219 USA: WV Paratype ATRMK558-11 KU059024 Aerophilus minys HI 222 USA: KY Paratype — KU059025 Aerophilus minys H1251 USA: KY Paratype ATRMK281-11 KU059026 Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus ■ 103 Appendix Table [Continued] Taxon name No. Country: region Type status COI 28S Aerophilus minys H1254 USA: KY Holotype ATRMK283-11 KU059027 Aerophilus minys H1255 USA: WV Paratype ATRMK284-11 KP943678 Aerophilus minys H1257 USA: KY Paratype — KU059028 Aerophilus minys H1261 USA: KY Paratype ATRMK286-11 KU059029 Aerophilus minys H1262 USA: KY Paratype — KU059030 Aerophilus minys H1263 USA: WV Paratype ATRMK287-11 KU059031 Aerophilus minys H1266 USA: WV Paratype ATRMK288-11 KU059032 Aerophilus minys H6029 USA: KY Paratype — KU059033 Aerophilus minys H7425 USA: KY Paratype ATRMK380-11 KU059034 Aerophilus minys H11343 USA: VA Paratype KU059060 — Aerophilus perforator H12241 USA: KY KP943634 — Aerophilus perforator HI 2242 USA: KY KP943635 — Aerophilus rayfisheri H1212 USA: KY Holotype ATRMK278-11 KP943675 Aerophilus rayfisheri H4922 USA: ND Paratype KU059061 — Aerophilus reginae H1213 USA: KY KU059062 — Aerophilus reginae H1280 USA: KY Holotype KP943616 KP943681 Aerophilus robertcourtneyi H1252 USA: KY Holotype ATRMK282-11 KP943677 Aerophilus rufipes Evl0-8944 France KP402064 — Aerophilus rufipes Evl 1-1-291 France KP402062 — Aerophilus rufipes Evl 1-3-4098 France KP402063 — Aerophilus rufipes Val05-328 France KP402060 — Aerophilus rufipes Val05-330 France KP402061 — Aerophilus stoelbae H7655 USA: KY Holotype ATRMK418-11 — Aerophilus terrymoyeri H7685 USA: IF Holotype ATRMK433-11 KU059035 Aerophilus tommurrayi H8496 USA: MA Holotype KP943627 — Aerophilus sp. H877 Mexico: Yucatan ATRMK249-11 KU059036 Aerophilus sp. H884 Mexico: Yucatan ATRMK252-11 KU059037 Aerophilus sp. H2200 Congo: Pool KU059063 KU059038 Aerophilus sp. H2201 Congo: Pool KU059067 KU059039 Aerophilus sp. H2202 Congo: Pool KU059064 — Aerophilus sp. H2206 Congo: Pool KU059065 — Aerophilus sp. H2208 Congo: Pool KU059066 — Aerophilus sp. H1214 France: Fanguedoc ATRMK496-11 — Aerophilus sp. H1221 France: Fanguedoc ATRMK560-11 — Aerophilus sp. H7611 Mexico: Yucatan ATRMK398-11 KU059040 Aerophilus sp. H7612 Mexico: Yucatan ATRMK399-11 KP943707 Aerophilus sp. H7613 Mexico: Yucatan ATRMK400-11 KP943708 Aerophilus sp. H7614 Mexico: Yucatan ATRMK401-11 KU059041 Aerophilus sp. H7615 Mexico: Yucatan — KU059042 Aerophilus sp. H4919 USA: AZ KU059070 — Aerophilus sp. H4924 USA: AZ KU059074 — Aerophilus sp. H4925 USA: AZ KU059075 — Aerophilus sp. H14484 USA: AZ KP943639 — Aerophilus sp. H14844 USA: AZ KP943640 — Aerophilus sp. H14888 USA: AZ KP943642 — Aerophilus sp. H14891 USA: AZ KP943641 — Aerophilus sp. H15097 USA: AZ KP943645 — Aerophilus sp. H15104 USA: AZ KU059081 — Aerophilus sp. H4911 USA: CA KU059068 — Aerophilus sp. H4912 USA: CA KU059069 — Aerophilus sp. H4920 USA: CA KU059071 — Aerophilus sp. H4921 USA: CA KU059072 — Aerophilus sp. H4923 USA: CA KU059073 — Aerophilus sp. H4926 USA: CA KU059076 — Aerophilus sp. H4930 USA: CA KU059077 — Aerophilus sp. H4931 USA: CA KU059078 — Aerophilus sp. H4938 USA: CA KU059079 — Aerophilus sp. H4943 USA: CA KU059080 — 104 Contributions in Science, Number 524 Sharkey et ah: Revision of Aerophilus — H2374 Braunsia sp Central African Republic H2376 Braunsia sp Congo H1891 Braunsia sp Congo HI892 Braunsia sp Kenya H2380 Braunsia sp Congo -H2371 Braunsia sp Congo - H2373 Braunsia sp Congo H2377 Braunsia sp Congo H2379 Braunsia sp Congo 82j- HI 297 Braunsia sp Madagascar H2384 Braunsia sp Madagascar - HI884 Braunsia sp Kenya 98r HI876 Braunsia sp Congo *- H2375 Braunsia sp Congo - HI893 Braunsia sp Uganda H2378 Braunsia sp Congo Cq 58 54 55 r H2381 Braunsia sp Congo HI4844 Aerophilus sp USA: AZ -HI350 Aerophilus chapmani Holotype USA: KY H1313 Aerophilus abdomina USA: KY H205 Aerophilus klastos Holotype USA: KY 98l 59 I Evl0-8944 Aerophilus rufipes France i Val05-328 Aerophilus rufipes France U Evl 1-3-4098 Aerophilus rufipes France Evl 1-1-291 Aerophilus rufipes France - Val05-330 Aerophilus rufipes France 100 ' ' " ' “ . HI213 Aerophilus reginae USA: KY 1 HI 280 Aerophilus reginae Holotype USA: KY 891 -H185 Aerophilus rugulosus Hungary —|_85. H1214 Aerophilus sp France L H1221 Aerophilus sp France 99 H2201 Aerophilus sp Congo 66 62 H2200 Aerophilus sp Congo - H2202 Aerophilus sp Congo H2206 Aerophilus sp Congo H2208 Aerophilus sp Congo H11965 Aerophilus difficilis USA: FL 54 87i HI248 Aerophilus malus Paratype USA: WV L Apmnhilnc: malim Hnlnfvnp I 1AA 89 HI484 Aerophilus malus Holotype USA: WV H5615 Aerophilus arthurevansi Paratype USA: VA H8554 Aerophilus arthurevansi Holotype USA: VA H8556 Aerophilus arthurevansi Paratype USA: VA — H1267 Aerophilus davidsmithi Holotype USA: WV 741 — H7611 Aerophilus sp Mexico: Yucatan 4 1 H877 Aerophilus sp Mexico: Yucatan H H7613 Aerophilus sp Mexico: Yucatan 1 H7614 Aerophilus sp Mexico: Yucatan 591 ■ H7614 Aerophilus sp Mexico: H1250 Aerophilus calcaratus USA: TN j- H1220 Aerophilus calcaratus USA: WV l_r HI247 Aerophilus calcaratus USA: FL “— H1457 Aerophilus calcaratus USA: WV r H209 Aerophilus calcaratus USA: KY H1487 Aerophilus calcaratus USA: WV I H1246 Aerophilus calcaratus USA: WV 4 HI249 Aerophilus calcaratus USA: WV 52 H1478 Aerophilus calcaratus USA: WV HI252 Aerophilus robertcourtneyi Holotype USA: KY - H7655 Aerophilus stolbae Holotype USA: KY H7o85 Aerophilus terrymoyeri Holotype USA: IL 93r 93 53 H884 Aerophilus sp Mexico: Yucatan L H7612 Aerophilus sp Mexico: Yucatan R7 i H14891 Aerophilus sp USA: AZ -M H15097 Aerophilus sp USA: AZ *- H15104 Aerophilus sp USA:AZ HI 251 Aerophilus minys Paratype USA: KY HI255 Aerophilus minys Paratype USA: WV H7425 Aerophilus minys Paratype USA: KY H11343 Aerophilus minys Paratype USA: VA H966 Aerophilus minys Paratype USA: KY H1219 Aerophilus minys Paratype USA: WV HI266 Aerophilus minys Paratype USA: WV H206 Aerophilus minys Paratype USA: KY H1215 Aerophilus minys Paratype USA: KY HI254 Aerophilus minys Holotype USA: KY HI261 Aerophilus minys Paratype USA: KY HI263 Aerophilus minys Paratype USA: WV - H4921 Aerophilus sp USA: CA H4912 Aerophilus sp USA: CA 51 H4919 Aerophilus sp USA: AZ H4943 Aerophilus sp USA: CA H4931 Aerophilus sp USA; CA H4926 Aerophilus sp USA: CA H4911 Aerophilus sp USA: CA H4920 Aerophilus sp USA: CA I H4938 Aerophilus sp USA: CA I 83. HI212 Aerophilus rayfisheri Holotype USA: KY “ H4922 Aerophilus rayfisheri Paratype USA: ND — H4923 Aerophilus sp USA: CA _57. H4924 Aerophilus sp USA: AZ 1 H4925 Aerophilus sp USA: AZ 61, _r- HI216 Aerophilus erythrogaster USA: KY 1 — H8274 Aerophilus erythrogaster USA: KY H7683 Aerophilus kowlesae Holotype USA: KY jn/oo HUE, — 0.005 substitutions/site H8496 Aerophilus tommurrayi Holotype USA; MA H7424 Aerophilus hopkinsensis Paratype USA: KY H7438 Aerophilus hopkinsensis Holotype USA: KY — H14484 Aerophilus sp USA: AZ H4930 Aerophilus sp USA: CA H14888 Aerophilus sp USA: AZ H1217 Aerophilus jdherndoni Holotype USA: KY H1218 Aerophilus jdherndoni Paratype USA; KY - H1265 Aerophilus jdherndoni Paratype USA: KY H1400 Aerophilus jdherndoni Paratype USA: KY H6033 Aerophilus jdherndoni Paratype USA: Wl HI 0030 Aerophilus jdherndoni Paratype USA: KY i— HI 307 Aerophilus jdherndoni Paratype USA: Ky ■ H8848 Aerophilus jdherndoni Paratype USA: KY 4 | HI 256 Aerophilus jdherndoni Paratype USA: KY n___65, H12241 Aerophilus perforator USA: KY ' -HI2242 Aeropnilus perforator USA: KY Supplemental Figure 1 Tree of highest log-likelihood from a 200-replicate ML analysis of the combined COI data set (summarized in ML bootstrap values are plotted above the branches (see Supplemental Figure 2). Figure 3). Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus ■ 105 Majority rule 66 51 86 83 89 99 58 54 98 51 93 62 54 65 r 82 r 98 r 50 61 56 65 100 59 93 85, 87 66 55, 59 64 83, 57 r 50 53 87 89 52 74 HI884 Braunsia sp Kenya HI893 Braunsia sp Uganda H2378 Braunsia sp Congo H2381 Braunsia sp Congo H2374 Braunsia sp Central African Republic H2376 Braunsia sp Congo HI 297 Braunsia sp Madagascar H2384 Braunsia sp Madagascar HI 876 Braunsia sp Congo H2375 Braunsia sp Congo H1891 Braunsia sp Congo HI892 Braunsia sp Kenya H2380 Braunsia sp Congo H2371 Braunsia sp Congo H2373 Braunsia sp Congo H2377 Braunsia sp Congo H2379 Braunsia sp Congo H7683 Aerophilus kowlesae Holotype USA: KY H1252 Aerophilus robertcourtneyi Holotype USA: KY H8496 Aerophilus tommurrayi Holotype USA: MA H4912 Aerophilus sp USA: CA H4919 Aerophilus sp USA: AZ H4921 Aerophilus sp USA: CA H4930 Aerophilus sp USA: CA H4943 Aerophilus sp USA: CA H14888 Aerophilus sp USA: AZ H14484 Aerophilus sp USA: AZ H1216 Aerophilus erythrogaster USA: KY H8274 Aerophilus erythrogaster USA: KY H7424 Aerophilus hopkinsensis Paratype USA: KY H7438 Aerophilus hopkinsensis Holotype USA: KY H12241 Aerophilus perforator USA: KY H12242 Aerophilus perforator USA: KY H1213 Aerophilus reginae USA: KY H1280 Aerophilus reginae Holotype USA: KY H7655 Aerophilus stolbae Holotype USA: KY H7685 Aerophilus terrymoyeh Holotype USA: IL H884 Aerophilus sp Mexico H7612 Aerophilus sp Mexico H185 Aerophilus rugulosus Hungary H1214 Aerophilus sp France H1221 Aerophilus sp France H14891 Aerophilus sp USA: AZ H15097 Aerophilus sp USA: AZ H15104 Aerophilus sp USA: AZ H2201 Aerophilus sp Congo H2200 Aerophilus sp Congo H2202 Aerophilus sp Congo H2206 Aerophilus sp Congo H2208 Aerophilus sp Congo HI4844 Aerophilus sp USA: AZ H1350 Aerophilus chapmani Holotype USA: KY H1313 Aerophilus abcfominalis USA: KY H205 Aerophilus klastos Holotype USA: KY Val05-330Aerophilus rufipes EvlO-8944Aerophilus rufipes Val05-328 Aerophilus rufipes Evll-3-4098 Aerophilus rufipes Evll-1-291 Aerophilus rufipes H1217 Aerophilus jdherndoni Holotype USA: KY H1218 Aerophilus jdherndoni Paratype USA: KY H1256 Aerophilus jdherndoni Paratype USA: KY H1265 Aerophilus jdherndoni Paratype USA H1307 Aerophilus jdherndoni Paratype USA: KY H1400 Aerophilus jdherndoni Paratype USA: KY H6033 Aerophilus jdherndoni Paratype USA: Wl H8848 Aerophilus jdherndoni Paratype USA: KY H10030 Aerophilus jdherndoni Paratype USA: KY H4923 Aerophilus sp USA: CA H4931 Aerophilus sp USA: CA H4938 Aerophilus sp USA: CA H1212 Aerophilus rayfisheri Holotype USA: KY H4922 Aerophilus rayfisheri Paratype USA: ND H4924 Aerophilus sp USA: AZ H4925 Aerophilus sp USA: AZ H4911 Aerophilus sp USA: CA H4920 Aerophilus sp USA: CA H4926 Aerophilus sp USA: CA H966 Aerophilus minys Paratype USA: KY H1219 Aerophilus minys Paratype USA: WV H1251 Aerophilus minys Paratype USA: KY H1255 Aerophilus minys Paratype USA: WV H1266 Aerophilus minys Paratype USA: WV H7425 Aerophilus minys Paratype USA: KY H11343 Aerophilus minys Paratype USA: VA H206 Aerophilus minys Paratype USA: KY H1215 Aerophilus minys Paratype USA: KY HI254 Aerophilus minys Holotype USA: KY H1261 Aerophilus minys Paratype USA: KY H1263 Aerophilus minys Paratype USA: WV H11965 Aerophilus difficilis USA: FL H209 Aerophilus calcaratus USA: KY H1220 Aerophilus calcaratus USA: WV H1247 Aerophilus calcaratus USA: FL H1250 Aerophilus calcaratus USA: TN HI457 Aerophilus calcaratus USA: WV H1487 Aerophilus calcaratus USA: WV H1267 Aerophilus davidsmithi Holotype USA: WV H1248 Aerophilus malus Paratype USA: WV H1484 Aerophilus malus Holotype USA: WV H5615 Aerophilus arthurevansi Paratype USA: VA H8554 Aerophilus arthurevansi Holotype USA: VA H8556 Aerophilus arthurevansi Paratype USA: VA H1246 Aerophilus calcaratus USA: WV H1249 Aerophilus calcaratus USA: WV HI478 Aerophilus calcaratus USA: WV H877 Aerophilus sp Mexico H7611 Aerophilus sp Mexico H7613 Aerophilus sp Mexico H7614 Aerophilus sp Mexico Supplemental Figure 2 ML bootstrap analysis (500 replicates) of the COI data set. 106 ■ Contributions in Science, Number 524 Sharkey et ah: Revision of Aerophilus H2374 Braunsia sp Central African Republic -H2376 Braunsia sp Congo HI 889 Braunsia sp Kenya H1891 Braunsia sp Congo j mo m: 83 j HI892 Braunsia sp Kenya 1 H1115 Braunsia bilunata S Sao Tome & Principe 91 54 f; I H H2380 Braunsia sp Congo H2371 Braunsia sp Congo H2373 Braunsia sp Congo H2377 Braunsia sp Congo H2379 Braunsia sp Congo 821 HI297 Braunsia sp Madagascar H2384 Braunsia sp Madagascar HI890 Braunsia sp Kenya HI895 Braunsia sp Equatorial Guinea H1884 Braunsia sp Kenya 66 52 H2370 Braunsia sp Congo 97 r- Hi876 Braunsia sp Congo ■— H2375 Braunsia sp Congo _jsp(_ HI 893 Braunsia sp Uganda 971 H185 Aerophilus ruaulosus Hungary _I H1214 Aerophitus Sp H2381 Braunsia sp Congo H2378 Braunsia sp Congo « n i z. i *+ MtJi upniius sp Franee 82*— H1221 Aerophilus sp France 100 63 H2201 Aerophilus sp Congo H22 Aerophilus sp Congo - H2202 Aerophilus sp Congo H2206 Aerophilus sp Congo H2208 Aerophilus sp Congo 88ll| I U'. . HI4844 Aerophilus sp USA: AZ - H1350 Aerophilus chapmani Holotype USA: KY i-H205 Aerophilus 100 100 Aerophilus klastos Holotype USA: KY 98 1 H1253 Aerophilus abdominalis USA: GA 1 HI313 Aerophilus abdominalis USA: KY Evl 0-8944 Aerophilus rufipes France Val05-328 Aerophilus rufipes France Evl 1-3-4098 Aerophilus rufipes France Evl 1-1-291 Aerophilus rufipes France Val05-330 Aerophilus rufipes France I tv ki 56 i H1213 Aerophilus reginae USA: KY ' HI280 Aerophilus reginae Holotype USA: KY -H11965 Aerophilus difficilis USA: FL ^Hl248 Aerophilus malus Paratype USA: WV 57 78 u HI484 Aerophilus malus Holotype USA: WV act i H5615 Aerophilus arthurevansi Paratype USA: VA -4l£j H8554 Aerophilus arthurevansi Holotype USA: VA 1 H8556 Aerophilus arthurevansi Paratype USA: VA H1487 Aerophilus calcaratus USA: WV I H1246 Aerophilus calcaratus USA: WV j- H1249 Aerophilus calcaratus USA: WV 1 H1478 Aerophilus calcaratus USA: WV H209 Aerophilus calcaratus USA: KY -HI 250 Aerophilus calcaratus USA: TNI — H1247 Aerophilus calcaratus USA: FL H1461 Aerophilus calcaratus USA: KY HI220 Aerophilus calcaratus USA: WV - H8496 Aerophilus tommurrayi Holotype USA: MA H7424 Aerophilus hopkinsensis Paratype USA: KY H7438 Aerophilus hopkinsensis Holotype USA: KY H7683 Aerophilus kowlesae Holotype USA: KY I— HI216 Aerophilus erythrogaster USA: KY 60 1 -H8274 Aerophilus erythrogaster USA: KY - H14484 Aerophilus spuSA: AZ _i- H4930 Aerophilus sp USA: CA ■-H14888 Aerophilus sp USA: AZ HI217 Aerophilus jdherndoni Holotype USA: KY H1218 Aerophilus jdherndoni Paratype USA: KY - HI 265 Aerophilus jdherndoni Paratype USA: KY - H14 Aerophilus jdherndoni Paratype USA: KY H6033 Aerophilus jdherndoni Paratype USA: Wl HI 30 Aerophilus jdherndoni Paratype USA: KY r— H1307 Aerophilus jdherndoni Paratype USA: KY I H8848 Aerophilus jdherndoni Paratype USA: KY T HI256 Aerophilus jdherndoni Paratype USA: KY U H1268 Aerophilus jdherndoni Paratype USA: KY H1311 Aerophilus jdherndoni Paratype USA: KY _ H1380 Aerophilus jdherndoni Paratype USA: KY H1486 Aerophilus jdherndoni Paratype USA: KY H7682 Aerophilus jdherndoni Paratype USA: KY H1457 Aerophilus calcaratus USA: WV H1267 Aerophilus davidsmithi Holotype USA: WV H7611 Aerophilus sp Mexico: Yucatan H877 Aerophilus sp Mexico: Yucatan H7613 Aerophilus sp Mexico: Yucatan H7614 Aerophilus sp Mexico: Yucatan H7615 Aerophilus sp Mexico: Yucatan ft 76, HI2241 Aerophilus perforator USA: KY -HI2242 r r ' ' -H4921 Aerophilus sp USA: CA I H4912 Aerophilus sp USA: CA -I——-H4919 Aerophilus sp USA: AZ I Aerophilus perforator USA: KY 58 1 H4943 Aerophilus sp USA: CA H4931 Aerophilus sp USA: CA 56 1 92 1 — 0.5 substitutions/site H4926 Aerophilus sp USA: CA . H49§8 Aerophilus sp USA: CA _82. HI212 Aerophilus rayfisheri Holotype USA: KY ^ H4922 Aerophilus rayfisheri Paratype USA: ND I H4920 Aerophilus sp USA: CA -1—- H4911 Aerophilus sp USA: CA I-H4923 Aerophilus sp USA: CA 1 - 1 60, H4924 Aerophilus sp USA' AZ 1 H4925 Aerophilus sp USA: AZ - H1252 Aerophilus robertcourtneyi Holotype USA: KY -H7655 Aerophilus stolbae Holotype USA: KY - H7685 Aerophilus terrymoyeri I lolotype USA: IL i- H884 Aerophilus sp Mexico: Yucatan *— H7612 Aerophilus sp Mexico: Yucatan mi H14891 Aerophilus sp USA: AZ -22J HI5097 Aerophilus sp USA: AZ 1 - H15104 Aerophilus sp USA: AZ H1222 Aerophilus minys Paratype USA. KY H1251 Aerophilus minys Paratype USA: KY - H1255 Aerophilus minys Paratype USA: WV H1257 Aerophilus minys Paratype USA: KY H6029 Aerophilus minys Paratype USA: KY H7425 Aerophilus minys Paratype USA: KY H11343 Aerophilus minys Paratype USA: VA i- H966 Aerophilus minys Paratype USA: KY HI219 Aerophilus minys Paratype USA: WV H1266 Aerophilus minys Paratype USA: WV HI262 Aerophilus minys Paratype USA: KY - H206 Aerophilus minys Paratype USA: KY H1215 Aerophilus minys Paratype USA: KY H1254 Aerophilus minys Holotype USA: KY H1261 Aerophilus minys Paratype USA: KY H1263 Aerophilus minys Paratype USA: WV 91 Supplemental Figure 3 Tree of highest log-likelihood from a 200-replicate ML analysis of the combined COI+28S data set (summarized in Figure 4). ML bootstrap values are plotted at the nodes when supported by values >50 (see Supplemental Figure 4). Contributions in Science, Number 524 Sharkey et al.: Revision of Aerophilus ■ 107 Majority rule 87 65 83 82 66 91 54 52 97. 50.- 60 76 100 56 92 97 82 86 58 77 58 82 60 98 100 100 78 -- 90 HI889 Braunsia sp Kenya HI891 Braunsia sp Congo HI890 Braunsia sp Kenya H1895 Braunsia sp Equatorial Guinea H2374 Braunsia sp Central African Republic H2376 Braunsia sp Congo HI892 Braunsia sp Kenya H1115 Braunsia bilunata SaoTome & Principe HI297 Braunsia sp Madagascar H2384 Braunsia sp Madagascar HI884 Braunsia sp Kenya H2370 Braunsia sp Congo H2380 Braunsia sp Congo H2371 Braunsia sp Congo H2373 Braunsia sp Congo H2377 Braunsia sp Congo H2379 Braunsia sp Congo H2378 Braunsia sp Congo HI876 Braunsia sp Congo H2375 Braunsia sp Congo HI893 Braunsia sp Uganda H2381 Braunsia sp Congo HI252 Aerophilus robertcourtneyi Holotype USA: KY H4912 Aerophilus sp USA: CA H4919 Aerophilus sp USA: AZ H4921 Aerophilus Sp USA: CA H4930 Aerophilus sp USA: CA H4943 Aerophilus sp USA: CA HI4888 Aerophilus sp USA: A 7 H14484 Aerophilus sp USA: AZ H1216 Aerophilus erythrogaster USA: KY H8274 Aerophilus erythrogaster USA: KY HI2241 Aerophilus perforator USA: KY HI2242 Aerophilus perforator USA: KY H1213 Aerophilus reginae USA: KY HI280 Aerophilus reginae Holotype USA: KY H7655 Aerophilus stcflbae Holotype USA: KY H7685 Aerophilus terrymoyeri Holotype USA: IL H884 Aerophilus sp Mexico: Yucatan H7612 Aerophilus sp Mexico: Yucatan HI85 Aerophilus rugulosus Hungary HI214 Aerophilus sp France H1221 Aorophilus sp France H14891 Aerophilus sp USA: AZ HI5097 Aerophilus sp USA: AZ H15104 Aerophilus sp USA: AZ H8496 Aerophilus tommurrayi Holotype USA: MA H7424 Aerophilus hopkinsensis Paratype USA: KY H7438 Aerophilus hopkinsensis Holotype USA: KY H7683 Aerophilus kowlesae Holotype USA: KY H4911 Aerophilus sp USA: CA H4920 Aerophilus sp USA: CA H4923 Aerophilus sp USA: CA H4926 Aerophilus sp USA: CA H4931 Aerophilus sp USA: CA H4938 Aerophilus sp USA: CA H1212 Aerophilus rayfisheri Holotype USA: KY H4922 Aerophilus rayfisheri Paratype USA: ND H4924 Aerophilus sp USA: AZ H4925 Aerophilus sp USA: AZ HI217 Aerophilus jdherndoni Holotype USA: KY H1218 Aerophilus jdherndoni Paratype USA: KY HI 256 Aerophilus jdherndoni Paratype USA: KY HI265 Aerophilus jdherndoni Paratype USA: H1268 Aerophilus jdherndoni Paratype USA: KY H1307 Aerophilus jdherndoni Paratype USA: KY H1311 Aerophilus jdherndoni Paratype USA: KY H1380 Aerophilus jdherndoni Paratype USA: KY H1400 Aerophilus jdherndoni Paratype USA: KY HI486 Aerophilus jdherndoni Paratype USA: KY H6033 Aerophilus jdherndoni Paratype USA: WI H7682 Aerophilus jdherndoni Paratype USA: KY H8848 Aerophilus jdherndoni Paratype USA: KY H10030 Aerophilus jdherndoni Paratype USA: KY HI350 Aerophilus chapmani Holotype USA: KY H205 Aerophilus klastos Holotype USA: KY HI4844 Aerophilus sp USA: AZ HI253 Aerophilus abaominalis USA: GA H1313 Aerophilus abdominalis USA: KY Ev10-8944Aerophilus rufipes France Val05-328Aerophilus rufipes France Evl 1-3-4096Aerophilus rufipes France Ev11-1-291Aerophilus rufipes France Val05-330 Aerophilus rufipes France H2201 Aerophilus sp Congo H2200 Aerophilus sp Congo H2202 Aerophilus sp Congo H2206 Aerophilus sp Congo H2208 Aerophilus sp Congo H206 Aerophilus minys Paratype USA: KY H966 Aerophilus minys Paratype USA: KY H1215 Aerophilus minys Paratype USA: KY H1219 Aerophilus minys Paratype USA: WV HI222 Aerophilus minys Paratype USA: KY HI251 Aerophilus minys Paratype USA: KY H1254 Aerophilus minys Holotype USA: KY HI255 Aerophilus minys Paratype USA: V\A/ H1257 Aerophilus minys Paratype USA: KY HI261 Aerophilus minys Paratype USA: KY H1262 Aerophilus minys Paratype USA: KY H1263 Aerophilus minys Paratype USA: WV H1266 Aerophilus minys Paratype USA: WV H6029 Aerophilus minys Paratype USA: KY H7425 Aerophilus minys Paratype USA: KY H11343 Aerophilus minys Paratype USA: VA H11965 Aerophilus difficilis USA: FL H209 Aerophilus calcaratus USA: KY HI220 Aerophilus calcaratus USA: WV HI246 Aerophilus calcaratus USA: WV HI247 Aerophilus calcaratus USA: FL HI249 Aerophilus calcaratus USA: WV HI250 Aerophilus calcaratus USA: TN H1457 Aerophilus calcaratus USA: WV H1461 Aerophilus calcaratus USA: KY HI478 Aerophilus calcaratus USA: WV HI487 Aerophilus calcaratus USA: WV H1267 Aerophilus davidsmithi Holotype USA: WV H877 Aerophilus sp Mexico: Yucatan H7611 Aerophilus sp Mexico: Yucatan H7613 Aerophilus sp Mexico: Yucatan H7614 Aerophilus sp Mexico: Yucatan H7615 Aerophilus sp Mexico: Yucatan HI248 Aerophilus malus Paratype USA: WV HI484 Aerophilus malus Holotype USA: WV H5615 Aerophilus arthurevansi Paratype USA: VA H8554 Aerophilus arthurevansi Holotype USA: VA H8556 Aerophilus arthurevansi Paratype USA: VA Supplemental Figure 4 ML bootstrap analysis (500 replicates) of the combined COI+28S data set. 108 Contributions in Science, Number 524 Sharkey et ah: Revision of Aerophilus H2374 Braunsia sp Central African Republic • H2376 Braunsia sp Congo HI889 Braunsia sp Kenya -H1891 Braunsia sp Congo HI892 Braunsia sp Kenya H1115 Braunsia bilunata Sao Tome & Principe H2380 Braunsia sp Congo 93 H2379 Braunsia sp Congo -H2371 Braunsia sp Congo H2373 Braunsia sp Congo H2377 Braunsia sp Congo h 99 r H1895 Braunsia sp Equatorial Guinea HI890 Braunsia sp Kenya HI884 Braunsia sp Kenya H1297 Braunsia sp Madagascar H2384 Braunsia sp Madagascar 92 H2370 Braunsia sp Congo L HI876 Braunsia sp Congo H2375 Braunsia sp Congo \ 90 r 67 r HI893 Braunsia sp Uganda 100 r 58 HI85 Aerophjlus ru^ujqsus Hungary H2381 Braunsia sp Congo H2378 Braunsia sp Congo 100 . 214 Aerophilus sp France H1221 Aerophilus sp France 57 56 H4912 Aerophilus sp USA: CA -H491S “ r H4921 Aerophilus sp USA: CA 19 Aerophilus sp USA: AZ 100 95r 84 97j - H2201 Aerophilus sp Congo „ | H2200 Aerophilus sp Congo _ 100 H2208 Aerophilus sp Congo — I— H2202 Aerophilus so Congo 1 H2206 Aerophilus sp Congo HI350 Aerophilus chapmani Holotype USA: KY -H205 Aerophilus klastos Holotype USA: KY j HI253 Aerophilus abdominalis USA: GA 1 H1313 Aerophilus abdominalis USA: KY 100 . HI4844 Aerophilus USA: AZ 100 Evl 1-1-291 Aerophilus rufipes France Evl 1-3-4098 Aerophilus rufipes France — Val05-330 Aerophilus rufipes France — EvlO-8944 Aerophilus rufipes France Val05-328 Aerophilus rufipes France 100 89 63 H1213 Aerophilus reginae USA: KY HI280 Aerophilus reginae Holotype USA: KY H11965 Aerophilus difficilis USA: FL I H8556 Aerophilus arthurevansi Paratype USA: VA H5615 Aerophilus arthurevansi Paratype USA: VA H8554 Aerophilus arthurevansi Holotype USA: VA 99 1 H1248 Aerophilus malus Paratype USA: WV L H1484 Aerophilus malus Holotype USA: WV r H1487 Aerophilus calcaratus USA: WV - H209 Aerophilus calcaratus USA: KY 86 _ - HI246 Aerophilus calcaratus USA: WV HI249 Aerophilus calcaratus USA: WV H1478 Aeropnilus calcaratus USA: WV 7/ .-H1250 Aerophilus calcaratus USA: TN H1220 Aerophilus calcaratus USA: WV H1247 Aerophilus calcaratus USA: FL — H1457 Aerophilus calcaratus USA: WV ■ H1461 Aerophilus calcaratus USA: KY yg 1 HI267 Aerophilus davidsmithi Holotype USA: WV 99 1 - ha 9oL§2j 62 66 56 61 H8496 Aerophilus tommurrayi Holotype USA: MA H7683 Aerophilus kowlesae Holotype USA: KY 87. H7424 Aerophilus hopkinsensis Paratype USA: KY ■ H7438 Aerophilus hopkinsensis Holotype USA: KY H7682 Aerophilus jdherndoni Paratype USA: KY H14484 Aerophilus sp USA: AZ 90 H7615 Aerophilus sp Mexico: Yucatan — 1 H7614 Aerophilus sp Mexico: Yucatan — H7611 Aerophilus sp Mexico: Yucatan ■ H877 Aerophilus sp Mexico: Yucatan H7613 Aerophilus sp Mexico: Yucatan 50i H4943 Aerophilus sp USA: CA I— H1216 Aerophilus erythrogaster USA: KY I-H8274 Aerophilus erythrogaster USA: KY ■ HI4888 Aerophilus sp USA: AZ 56 -H4930 Aerophilus sp USA: CA HI 268 Aerophilus jdherndoni Paratype USA: KY HI 311 Aerophilus jdherndoni Paratype USA: KY r H1400 Aerophilus jdherndoni Paratype USA: KY HI0030 Aerophilus jdherndoni Paratype USA: KY H8848 Aerophilus jdherndoni Paratype USA: KY 96 59 57 j-■ noo*to Merupmius juriemuurn raraiype uon. i\t p— H1256 Aerophilus jdherndoni Paratype USA: KY I— HI307 Aerophilus jdherndoni Paratype USA: KY H1217 Aerophilus jdherndoni Holotype USA: KY HI 380 Aerophilus jdherndoni Paratype USA: KY H6033 Aerophilus jdherndoni Paratype USA: Wl H1218 Aerophilus jdherndoni Paratype USA: KY 57L1 H1265 Aerophilus jdherndoni Paratype USA “ H1486 Aerophilusjdherndoni Paratype USA: KY H4931 Aerophilus sp USA: CA 100. H1212 Aerophilus rayfisheri Holotype USA: KY H4922 Aerophilus rayfisheri Paratype USA: ND 78 H4923 Aerophilus sp USA: CA H4925 Aeropnilus sp USA: AZ 78 1 -H4911 Aerophilus sp USA: CA H4920 Aerophilus sp USA: CA H4924 Aerophilus sp USA: AZ H4926 Aerophilus sp USA: CA -H4938 Aerophilus sp USA: CA 57 59 58 HI252 Aerophilus robertcourtneyi Holotype USA, KY L H884 Aerophifus sp Mexico 1 - ' ' 100 1- H7612 Aerophilus sp Mexico 97 . H12241 Aerophilus perforator USA: KY 93 r 99 1 H14891 Aerophilus sp USA: AZ - HI5097 Aerophilus sp USA: AZ — H15104 Aerophilus sp USA: H7655 Aerophilus stolbae Holotype USA: KY H7685 Aerophilus terrymoyeri Holotype USA: IL HI2242 Aerophilus perforator USA: KY 791 95 57 0.005 substitutions/site HI262 Aerophilus minys Paratype USA: KY ' H1222 Aerophilus minys Paratype USA: KY H7425 Aerophilus minys Paratype USA: KY H1251 Aerophilus minys Paratype USA: KY - H1255 Aerophilus minys Paratype USA: WV H966 Aerophilus minys Paratype USA: KY H1266 Aerophilus minys Paratype USA: VA/ H1219 Aerophilus minys Paratype USA: WV H6029 Aerophilus minys Paratype USA: KY HI257 Aerophilus minys Paratype USA: KY H1215 Aerophilus minys Paratype USA: KY 57 - H206 Aerophilus minys Paratype USA: KY HI261 Aerophilus minys Paratype USA: KY H11343 Aerophilus minys Paratype USA: VA H1254 Aerophilus minys Holotype USA: KY HI263 Aerophilus minys Paratype USA: WV Supplemental Figure 5 Tree of highest posterior probability from a Bayesian analysis of the combined COI+28S data set. Posterior probabilities plotted at the nodes (see Supplemental Figure 6). are