THE WASMANN JOURNAL OF BIOLOGY 35(1), 1977, pp. 87-114 THE OF STANFORD UNIVERSITY'S JASPER RIDGE AND NEIGHBORING PALO ALTO, CALIFORNIA

David B. Weissman and David C. F. Rentz

Abstract.-This paper reports on the and ecology of the Orthoptera of Stanford University's Jasper H.idge Biological Preserve and nearby Palo Alto. The habitat preference, abundance, and bionomics of 44 species are discussed. A key to the species and photographs of study sites illustrating different habitats are provided.

This paper reports upon the Orthopteran fauna of Jasper Ridge, San Mateo County (Fig. 1), and neighboring Palo Alto, Santa Clara County, California (including Stanford University). Systematic and bionomic ob­ servations over more than a five year period for 44 species are presented . The Orthopteran fauna of the above areas is of interest for several rea­ sons: 1) the type localities of several species arc included within this study area or are located in nearby regions; 2) the San Francisco Bay Area, in general, has extensive entomological potential because of the presence of several major academic institutions specializing in entomology; and 3) Jasper Ridge, Stanford University's Biological Preserve, is unique among academic shtdy areas with reference to diversity of habitats and proximity (only 10 minutes by auto from the University campus). This research facility has the additional benefit of a long-term documented history and intensive research utilization. Historically, the Preserve was grazed until 1960; docu­ mentation of floral changes since the removal of cattle now exists and, with the completion of this study, Orthopteran faunal changes through time can likewise now be monitored. This report concentrates on Jasper Ridge because of its species diversity and importance as a Preserve. Attention also has been given to areas of Palo Alto and the Stanford University Campus because they contain species not located on Jasper Ridge at present, but the species represent fuhtre pos­ sible Jasper ridge colonizers. All specimens collected will be deposited at the California Academy of Sciences.

Study Areas Jasper Hidge is a low-lying ridge situated in the foothills of the Santa Cruz Mountains, a branch of the Coast Range. It lies some 40 km south­ cast of San Francisco, and 11.2 km due west of Palo Alto. Jasper Ridge is 4 km long and from 1-2 km wide. Stanford University's Jasper Ridge Biological Preserve encompasses only the northern half of the ridge and ex- 00 00

~ iii

Ul~ 7'.... > L: 7. 0c !Jj > l' 0 "fj tl:l 0 t'"' 0 (l Fig. 1. Aerial view of Jasper Ridge. l\'umhers refer to sites described in text. --<: VOLUME 35, NUMBER 1 89 tends 2 km in length. The Preserve consists of 389 hectares with the central axis running a northwesterly-southeasterly direction. Maximum elevation is 188 111. Seven of the ten plant communities present in the entire Coast Range of California are found on Jasper Ridge: these include chaparral (Fig. 15, site 12); mixed evergreen forest; oak woodlands (Fig. 3, site 4); fresh­ water marsh; riparian (Fig. 14, site 14); grasslands (both serpentine, Fig. 9, site 8, and sandstone, Fig. 8, site 7); and redwood forest (Fig. 16, site 14). Over the past 45 years, Searsville L ake (elevation 105 m ) on the west side of the Ridge (Fig. 1, site 13) has averaged 71.1 em (SD ± 20.3 em) precipitation per year, while Palo Alto, lower (elevation 8 m) and more in­ land, averaged approximately half that amount. Collections in Palo Alto were limited to grazed fields and watered gar­ dens around human dwellings. Much of the Stanford University Campus consists of seminatural oak woodland and fields of introduced European annual grasses.

Materials and Methods Sampling periods.-Collecting was conducted throughout the year. Collection sites.- These are described below and given specific reference numbers for use in the text.

Jasper Ridge (Fig. 1) Site l. Sandstone grassland (Fig. 2). Site 2. Serpentine grassland (Fig. 4), corresponds to part of Area H of Ehrlich (1965). Site 3. Serpentine grassland, part of Area I-I of Ehrlich (1965), (Fig. 6). Site 4. Oak woodland (Fig. 3). Site 5. \iVeathcr station, sandstone grassland (Fig. 7). Site 6. Serpentine grassland, Area G of Ehrlich (1965). Site 7. Sandstone grassland along trail to SE of large oak (Quercus lohata Nee) (Fig. 8). Site 8. Serpentine grassland along trail to NW of Q. lobata extending to Site 10 (Fig. 9). Site 9. Interface of chaparral and serpentine grassland (left side, b ack- ground of Fig. 10). Site 10. Oak woodland. Site 11. Oak woodland. Site 12. Chaparral area above Scarsville Lake (Fig. 15). Site 13. Scarsville Lake- Freshwater marsh. Site 14. San Francisquito Creek- ripalian, redwoods (Figs. 14, 16). 90 THE WAS~'IAl\'1\ JOUHNA L OF BIOLOGY

Fig. 2. Site 1. Cracks in ground in sandstone grassland. Neonemobius eurynotus occurs in small numbers in the cracks. The adjacent tarweed-grasslancl harbors dense populations of Conoceplwlus occidentalis. Fig. 3. Site 4. Oak woodland area. Pristoceuthophilus marmoratus is found here.

Stanford Campus Site 15. Corner of Palm Dr. and Campus Dr.- Oak woodland-grassland (Fig. 11). Site 16. Lake Lagunita-Freshwater marsh, rank vegetation (Fig. 12). Site 17. Short grassland area behind Anatomy Building (Fig. 17).

Palo Alto Site 18. General residential area-backyards and ploughed fields. Site 19. Grazed field above Foothill Expressway at Stanford Avenue (Fig. 13).

7 Fig. 4. Site 2. Serpentine grassland (Area H ) near entrance to Jasper Ridge at Escobar Road. Fig. 5. Between Sites 3 and 6. Serpentine grassland (Area H). Area to left harbors small numbers of occidentalis. Grassy area to right contains densest populations of Dissosteira ·pictiperutis on Jasper Ridge. 96 THE WASMANi\ JOURNAL OF BIOLOGY VOLUME 35, !\UMBER 1 91 92 THE WASMANN JOURNAL OF BIOLOGY VOLUME 35, i\Ut-IBE R l 93

Abundance.-After the locality site number of some species, we give another number (in parenthesis) which gives an estimate of that species' abundance at that site. Such inf01ma tion will permit comparisons with future studies and facilitate monitoring of faunal composition changes. A useful indicator of abundance is the number of individuals which an expert can see or collect within a two-hour peliod. On this basis we em­ ploy the following numerical indicators: 1-less than 5 individuals; 2-- 5-10 individuals; 3-11-25 individuals; 4- 26-50 individuals; 5-over 50 in­ dividuals easily seen or caught. Additional specific information, such as cytology, oscillogram analysis, re­ productive biology, and stridulatory apparatus, will be included in a paper by Rentz and Weissman (1978).

Key to the Orthopteroid of the Jasper Ridge­ Palo Alto Vicinity l. Hind femur enlarged, suitable for jumping; tarsi with 4 or fewer articles 2 - Hind femur not enlarged for jumping; tarsi with 5 articles (excep­ tion: Timema californicum with 3-jointed tarsi, easily identified by soft body and males with intra-cereal dextral process) (mantids, roaches) 42 2. Pronotum extending posteriorly, entirely covering abdomen, apex acute (pygmy grasshoppers, Tel:ligidae) mexicanus (Saussure) Pronotum at most covering only first few abdominal segments, apex obtuse 3 3. Antenna short, less than half as long as body; ovipositor reduced, barely extending from apex of abdomen; auditory structme present on sides of first abdominal segment (grasshoppers, ) 4 - Antenna long, much longer than length of body; ovipositor long, needle- or sickle-shaped; auditory stmcture, if present, located on front tibia 17 4. Presternum with a cortical process between bases of front legs 5 - Presternum without a process 8 5. Adults fully winged, capable of flight Melanoplus deoastator Scudder

Fig. 6. Site 3. Serpentine grassland (Area H) with a southern exposure. Contains densest population of Trimerotropis occidentalis on Jasper Ridge. Fig. 7. Site 5. Sandstone area near weather station. Grassland harbors the densest population of Chimaroceplwla p. ·pacifica on Jasper Ridge. 94 THE WASMANN JOURNAL OF BIOLOGY VOLUME 35, NUMBER 1 95

- Adults short-winged, incapable of flight 6 6. Basic body color green, brown forms rare; dorsum of body with a narrow white median longitudinal stripe; dorsal surface of hind femur reddish brown H esperotettix pacific us pocificus Scudder Combination of characters not as above 7 7. Transverse pronotal sulci deep, traversing median carina Oedaleonotus brocki (Stal) Transverse pronotal sulci shallow, interrupted by median carina Melanoplus fricki Strohecker 8. Tegmina short, never attaining apex of abdomen; hind wings rudi­ mentary; male stridulatory mechanism a series of pegs on inner surface of hind femur Eupnigodes rnegocephala (McNeill) Fully winged, hind wings brightly colored; males of most species crepitate in flight (Oedipodinae) 9 9. Hind wings brilliant red-pink Dissosteiro pictipennis Brunner Hind wings variously colored, not red-pink 10 10. Hind wings opaque, not translucent, orange or yellow orange; adults appeming in spring Arphia conspersa m mona Rehn - Com hi nation of characters not as above 11 11. Hind wings colorless or smoky, not distinctly yellow, green or bluish 12 - Hind wings distinctly colmed, yellow, greenish or blue green 13 12. F emales flightless highly variable in color, ranging from uni­ form dark brown to reddish brown, green or various combinations thereof; males uniform dark brown; median carina of pronotum high; tegmina nmmally concolorous Chimarocephala pacifica pacifica (Thomas) - Both sexes equally capable of flight; color rather uniform straw brown with a series of large dark brown blotches on tegmina in both sexes; median ca1ina of pronotum low Camnula pellucida Scudder 13. Hind wing bright yellow; hind tibia yellow Trimerotropis pallidipennis pollidipennis (Burmeister) - Hind wing not b1ight yellow; hind tibia bluish or greenish 14 14. F01m very slender; face with a narrow black band beneath and

Fig. 8. Site 7. "Backbone" area of sandstone grassland, principal habitat of Camnula pellucida. Fig. 9. Site 8. Serpentine grassland along "backbone" (note difference in amount of vegetation along trail in comparison with Site 7). Camnula pellucida fly here earlier than at contiguous Site 7. VOLUME 35, NUMBER 1 97

between eyes; wing disc yellow green Tri·merotropis koebelei (Bruner) - Combination of characters not as above 15 15. Basic body color dark, sometimes black; tegmina! bands distinct; wing disc color greenish yellow, apex smoky; median calina of pronotum short, elevated on prozona (Thomas) - Basic body color grey; tegmina! bands indistinct or absent; wing disc color faint greenish yellow with bluish or greyish overcast; median carina of pronotum low, poorly defined 16 16. Form slender; tegmen with banding faint or absent; wing disc faint yellow green, highly translucent, black band present but ill-defined, especially in median portion, band often broadly expanded to apex of disc Trimerotropis thalassica Bruner - Fom1 robust, stout; tegmen often with distinct bands, usually with brown speckles; wing disc very faint yellow green with distinct bluish or greyish overcast, translucent, black band narrow, distinct Trimerotropis occidentalis (Bmner) 17. Tarsi, at least those of the middle legs, 4-segmented; ovipositor compressed, sword-shaped 18 - All tarsi 3-segmented; ovipositor tubular, needle-shaped 26 18. Tegmina always present in adults; males with stridulatory sh·uc­ ture on tegmen; front tibia with audit01y structure (katydids, Tet­ tigoniidae) 19 -Tegmina absent, stridulatory stmcture, if present, located on in­ ternal smface of hind femur and opposing portion of thorax and abdomen 36 19. Auditory structure of front tibia circular, tambomine-sbaped (bush katydids, Phaneroptcrinae) 20 - Auditory structme of front tibia consisting of two slits separated by a bridgelike structure 22 20. Small, robust; tegmina of male with brownish areas dorsally in region of stridulatory vein Platylyra califomica Scudder

Fig. 10. View toward Searsville Lake. Oak woodland, c.-e nter background, contains densest population of Arphia c. ramona. Chaparral-grassland interface harbors both T rimerotropis occidentalis and T. thalassica. A cytologically confitmcd hybrid male was found in this area in 1974. Fig. 11. Site 15. Stanford campus, comer Palm Drive and Campus Drive. Oak woodland. Harbors Gryllus III, Neonemobit1s emynotus, Ceuthophilt1s califomimaus, Pristoceuthoplzilus sp . 98 THE WASMANN jOURNAL OF BIOLOGY

Fig. 12. Sjte 16. Lake Lagunita, dry in summer. Grullus VII is abundant during spring and early summer in rank vegetation on sides of lake. In summer, Grullus VI can be found in cracks on dry lake bottom. VOLUME 35, NUMBER 1 99

- Larger, elongate; tegmen of male immaculate dorsally, or with 4 black spots 21 21. Male with 4 small black spots dorsally on tegmina; tenth abdominal tergite unmodified; female with ovipositor uniform green Phcmeroptera ncma nana (Fieber) - Male without any dorsal black spots on tegmina; tenth tergite with elongate median projection; female with ovipositor reddish brown Scudderia furcata (Brunner) 22. Small, rather cylindrical; body surface highly polished; ovipositor nearly as long as hind femur (Conocephalinae) Conocephalus occiclentalis (Morse) - Combination of characters not as above (shield backs, Decticinae) 23 23. Adults fully-winged; tegmina with several brown spots d01·sally Platycleis tessellata (Charpentier) - Adults micropterous; color not as above 24 24. Body of adults highly polished, lustrous; ovipositor short scythe- shaped Decticita brevicaucla (Caudell) - Body of adults dull; ovipositor elongate, s\>vord-shaped 25 25. Pronotum very large, in male completely concealing tegmina; apex of ovipositor toothed Neduba cliabolica (Scudder) - Length of exposed portion of tegmina at least % that of the pro- notum; apex of ovipositor smooth Clinopleura minuta Caudell 26. Size minute, adult under 5 mm in total length; body covered with scales (ant crickets, Mym1ecophilidae) Myrmecophilus oregonensis Bruner - Adult size much larger; body not healing scales 27 27. Adult male body form deltoid, female tubular; general body coloration pale lime green or light brown, never black (tree c1ickets, Oecantbidae, Oecanthus) 28 - Adult male and female body form cylinillical or dorsoventrally flattened, never deltoid; general coloration black or dark brown, never green (Gryllidae) 32 28. Inner margin of first antenna! article with a pale yellow swelling; calling song a broken h·ill or chirp 29 - Inner margin of first antenna! article without any trace of a swelling; calling song a long continuous trill 30 29. First antenna! article with a small ovate black mark over swelling

Fig. 13. Site 19. Grazed field above Foothill Expressway at Stanford Ave. Dense populations of Neonemobius eurynotus, along with Gryllus III, Gryllus VI, Gryllus Vlli, and Stenopelmatus fuscus occur here. 100 T HE WAS MANN JOURNAL OF BIOLOGY

Fig. 14. Site 14. Area of San Francisquito Creek where Paratettix mexicanus was found. Fig. 15. Site 12. Chaparral area above Searsv ille Lake. Trimerotropis occidentalis, T. tlwlassicct, and T . koebelei occur in this area. VOLUME 35, NUMBER 1 101

Fig. 16. Site 14. Redwood grove along San Francisquito Creek. P.ristoceuthophilus sp. is abundant here.

or no mark at all; calling song about 90 chirps per minute at 21 °C Oecanthus rileyi Baker First antenna} article with a large ovate black mark; calling song at 2PC at least 150 chirps per minute Oecanthus fultoni T . Walker 30. Third article of antenna darker than second, frequently a dark line on inner edge of first two articles; entire basal portion of an­ tenna generally dark reddish brown , frequently black; calling song produced chiefly at night Oecanthus californicus Saussure - Third article of antenna not darker than second; entire antenna greenish ; calling song heard both day and night 31 31. Black marks on second antennal article contiguous or separated by no more than % of the width of the inside mark Oecanthus argentinus Saussure - Black marks of second antenna! article not contiguous, separated by 102 THE WASMANN JOURNAL OF BIOLOGY

Fig. 17. Site 17. Stanford Campus, disturbed area between Anatomy Building and Medical Center. G1'yllus III is found in cracks in ground in right foreground.

more than ¥s the width of the inside marks Oecanthus quadripunctatus Beutenmuller 32. Spines of bind tibia long, moveable, pilose; terminal article of maxillaty palpus at least twice the length of penultimate article; length of body less than 12 nun Neonemobius eurynotus (Rehn & Hebard) - Spines of hind tibia immoveable, not pilose; terminal article of maxillary palpus not more than ¥s longer than penultimate article; length of body more than 14 mm (Gryllus) 33 33. Calling song a bill, macropterous Gryllus VI - Calling song a chirp; usually micropterous 34 34. Medium large (body >19 mm long); calling song medium, about 160 chirps per minute, metallic-sounding; 6-10 pulses per chirp; shidulatory file less than 40 teeth per mm; habitat cracks in ground Gryllus III - Small (body < 19 mm long); calling song a slow chirp, 100-125 chirps per minute; 2-6 pulses per chirp; file with more than 40 teeth per mm 35 35. Tegmina smoky black; dark lateral longih1dinal stripe present, especially on females; base of inner face of hind femur reddish brown; chirp rate slow with 2-4 pulses per chirp; habitat, shores of Lake Lagunita Gryllus VII - Tegmina with yellowish brown overcast, never smoky black; lat- eral stripe undifferentiated; entire inner face of hind femur VOLUME 35, NUMBER 1 103

black; chirp rate slow with 3- 6 pulses per chirp; habitat, cracks in grazed fields Grylus VIII 36. Burrowing, hind legs not saltatory; body fonn cylindrical; abdomen banded with dark brown and light brown (Jerusalem crickets, Stenopelmatidae, Stenopelmatus) 37 - Hind legs saltatory; body form hump-backed ; abdomen concolor- ous, speckled or with dark markings, never banded as above (camel crickets, Raphidophoridae) 39 37. (Adults) Overall coloration very dark, lighter b ands of abdomen hardly distinguished from darker Stenopelnwtus intermedius Davis & Smith - Overall coloration lighter; banding of abdomen distinct 38 38. Spurs of hind tibia cylindrical, apices not excavate Stenopelnwtus longispina Brunner - Spurs of hind tibia with apices excavate, spatulate Stenopelmatus fuscus (Haldeman) 39. Arboreal; dorsum of abdomen with dark markings laterally; sternum yellow; male cerci modified claspers Gammarotettix bilobatus (Thomas) - Ground-dwelling; dorsum of abdomen speckled or concolorous; sternum straw brown or whitish; male cerci not modified for clasping 40 40. Overall coloration very dark brown, yellow stripe on entire dorsum of ; hind legs of male not modified; small; found in groups under rocks, boards, leaves Pristoceuthophilus sp. (undescribed) - Overall coloration shaw brown or grayish; male hind legs heavy, tibiae frequently undulant; larger; often found singly or in small groups of two or three individuals 41 41. Coloration straw brown or yellow brown; male subgenital plate deeply cleft but without distinct styli Ceuthopilus caUfornianus Scudder - Coloration greyish or greyish brown, "salt a nd pepper"; male sub- genital plate with distinct styli Pri.stoceuthophilus mannoratus Rehn 42. Forelegs modified for grasping prey; small grey ground-dwelling forms (gray ground mantis) Litaneut-ria minor Scudder - Combination of characters not as above 43 43. ' iVingless; soft-bodied; found on chaparral trees and shrubs; male abdomen with unique unpaired dextral process at apex (chaparral stick insect) Timema caUfomicum Scudder - Males fully winged, females appearing apterous; not especially soft-bodied ; under leaves, wood, rocks in chaparral community (chaparral roach) Pa-rcohlatta americana Scudder 104 TI-lE WASMANN JOURNAL OF BIOLOGY

Species List Paratettix mexicanus (Saussure). Found as early instars only at site 14(2) on 23 July 1973.

Acrididae Catantopinae Ilesperotettix pacificus pacificus Scudder. Only two females (both green) found at site 8 in serpentine grassland. Adults by June. Melanoplus fricki Strohecker. Found in grassland areas throughout Jasper Ridge, including those in oak woodland and semimarsh areas. Usually uncommon (2-3). Adults mid-May, although most, especially those from shaded wooded areas, mature later. Present to December. 2N i5 = 23. All elements telocentric. Both 1973 individuals examined had B (supernumerary) chromosomes. Melanoplus devastator Scudder. Common in grassland areas throughout ridge, especially sites 2, 3, between 3 and 6, 6, 7, 8. Early instars seen in early May, later instars still found in mid-October. Dichromatic for hind tibiae color: 15% of females have red hind tibiae, 85'7o blue. Only 2-5% of males have red hind tibiae. Species has a reproductive dormancy. Oedaleonotus borcki (Stal). Found only on serpentine grassland 2(2); 3(2); 8(4). First instar nymphs appear early (mid-March) and, in the lab­ oratory, can become adults in less than one month. Some adults present in late July. 2N ci = 23. All elements telocentric.

Gomphocerinae Eupnigodes megocephala (McNeill). Found in serpentine grassland at sites 2(1); 8(3); 9(1). Adults by June with some still aHve in late October. 2N i5 = 23. All elements telocentric.

Oedipodinae Arphia conspersa ramona Rehn. Found at sites 3(1); 10(4); 11(1). Over­ winter in nymphal stage; adults appear in early spring. On 24 October 1973, a 4th instar female found at site 3; in the laborat01y became 5th instar on 9 December, adult 26 December. On 17 September 1974 (prior to first rains) a 3rd instar male was captured. In the laboratory it became 4th instar on 19 October, 5th instar on 3 November, adult on 21 November. An extensive mark-release-recapture program was undertaken in 1974 at site 10 with the assistance of E. French. Between 2 May and 23 May, 44 males and 7 females were marked. Individuals were most easily taken VOLUME 35, NUMBER 1 105 along the trail through the woods. Individuals were recaptured too in­ frequently for valid statistical analysis of population size. Two males, though, did move distances of 135m in 5 and 16 days respectively. Adult individuals were first caught in early April at site 10; one female was captured on 22 July at site 3. Male crepitation is infrequent but consists of a single burst. Flights by both sexes are usually short. Camnula pellucida (Scudder). At Jasper Ridge, surp1isingly restricted at sites 7 and 8, both sandstone and serpentine grassland. One individual, however, was found at site 2; also at site 19(3). First adults appear in June, last teneral individual caught on 19 July, last adults 29 September. Chimarocephala pacifica pacifica (Thomas). Found at sites 1(4); 2(4); 3(3); 5(5); 7(2); 8(5). A late winter-early spring species. Captured 4th instar female 24 October 1973; in the laboratory became 5th instar 26 October, adult 26 November. On 10 September 1974 (prior to first rains) 2nd instar male caught; in the laboratory 3rd instar 29 September, 4th instar 10 October, 5th instar 24 October, adult 7 November. r-.hles occasionally flying by late December-early January, occasional late instars into May. Male crepitation a single bmst, flight short. Females always flightless. On Jasper Ridge, individuals found in short grass on south-facing slopes ma­ ture before individuals found elsewhere. Most adults gone by May. Ninety-eight percent of males are uniformly brown, rarely with green. Fe­ males exhibit all combinations of browns, greens, and red. A large sample of adult females collected on 4 Ap1il 1973 Tevealed the following color mm-phs: green 50, brown and tan 18, red and green 11. E. French (pers. comm.) has demonstrated the ability of adults of tllis species to change integument color from green to brown and vice versa over a period of weeks to correspond to altered background coloration. Lober and Chandrashekaran (1970) commented upon the sexual and acoustical behavior in tllis subspecies. Schroeter and Hewitt (1974) discussed the cytogenetics and Arnaud and Rentz (1965) reported on parasitism by the tachinid fly Ceracia dentata (Coquillett). Six females of 140 adults (sex ratio unknown) collected on 17 April 1973 were parasitized by a total of 10 Ceracia larvae which emerged, pupated, and became adult 2 weeks later. Dissosteira pictipenn'is Bruner. Found at sites 2(2); 3(5); between 3 and 6(5); 8(2); 9(5); 10(2); 11(2); 12(2). First adults late May-early June, con­ tinuing until mid-October. Crepitation a series of bursts. Trimerotropis fontana Thomas. W e have no records from our study areas. Four males and three females caught in "Palo Alto, Cal." on 13 July 1941 by Kenneth Frick. Trimerotropis koebelei (Bruner). Generally rare; 2(1 male); 12(3). Crepi­ tation in tlus sp ecies is a single burst. 2N t = 23, all elements telocentric. 106 THE WASMANN JOURNAL OF BIOLOGY

Tri1nerotropis occidentalis (Bruner). Restlicted to serpentine grassland areas and chaparral, usually found with Dissosteira pictipennis; 2(3); 3(5); between 3 and 6(4); 6(2); 8(3); 9(4); 12(3). Adults by mid-May to early June through late November. Male crepitation is a single burst. Crepita­ tion does not occur until August. This species has a reproductive dormancy. The karyotype of this species is unique and may be of systematic value (see Rentz and Weissman, 1978). Trimerotropis pallidipennis pallicl-ipennis (Burmeister). Never a common species, although widespread; 2(1); 3(1); 9(1); 12(1). Male crepitation is a series of bursts. Adults of the first generation appear in May, those of the second generation in August- September and are usually more common than first generation. Trimerotropis pilosa McNeill. Described from a single male, collected in Palo Alto, 17 March (no year indicated). Specimen originally in Stan­ ford University Collection. This collection was subsequently transferred about 1960 to the Los Angeles County Museum of Natural History. Dr. Charles L. Hogue has kindly examined that material but has not located the holotype. We believe it to be d estroyed. McNeill (1900) described the species with "posterior tibia brown, with a pale subbasal annulus. Size small." From our knowledge of the ac1idid species of the area, this description can only refer to T. pallidipennis pallidi­ pennis. Early emerging first generation individuals of that species are com­ monly small (D.B.W., unpublished observations). We therefore synonymize T. pilosa ~!JcNeill with T. p. pallidipennis (Burmeister). Trimerotropis thalassica Bruner. A rare species, usually associated with Adenostema fasciculatum H. & A. along the border of chaparral vegetation; 3(1); 9(2); 12(2). First adults late June through mid-October. Flights by both sexes are sho1t. Male crepitation is a single burst, rarely heard, and then usually late in the year (only some 50% of males crepitate as late as mid-October). A single male hyb1id between a T. occidentalis female and a T . thalassica male was found at site 9. Fmther specifics appear elsewhere (John and Weissman, 1977).

Tettigoniidae Phaneropterinae Phaneroptera nana nana (Fieber). This European introduction can be extremely common in gardens in Palo Alto. Adults mid-July until late December. Males stridulate day and night. Platylyra californica Scudder. As adults, high in trees at site 10. Two male nymphs taken by beating Le pechinia calycina (Benth.) Epl. on 9 May 1973; matured in laboratory on 4 and 6 June; one male stlidulated 3 VOLUME 35, NUMBER l 107 days after the last molt. Grant and Rentz (1966) further discuss this monotypic genus. Scudderia furcata Brunner. Small groupings of adults can be heard throughout the Jasper Ridge area in oak woodland. A very wary species. Adults by mid-July until December. Stridulates more in late afternoon than at night.

Decticinae See Rentz and Birchim (1968) for further information on this group. Clinopleura minuta Caudell. Distributed throughout Jasper Ridge, al­ though not as common as the sympatlic Decticita brevicauda. Also found at site 19. Adults by late June; males stridulate only at night. Present until late October. Decticita brevicauda (Caudell). Found throughout Jasper Ridge in grassland situations. First instars seen in early April, adults by early May and remain until early July. Males stridulate during the day. Biology discussed by Rentz (1963b). Neduba diabolica (Scudder). A localized, but occasionally common, noc­ turnal katydid. Found at sites 5, 10, and ll associated with oak wood­ land areas. Adults by early June until October. Some males sing in mid­ late afternoon and at night. Platycleis tessellata (Charpentier). This Mediterranean species was first noted in California in 1951 and has expanded its range ever since (Rentz, 1963a). On Jasper Ridge, this macropterous species is uncommon in serpentine grasslands. It matures by mid-Jtme.

Conocepbalinae Conocephalus occidentalis (Morse). Found throughout the grasslands on Jasper Ridge in high numbers, esp ecially on wet, no1th-facing slopes. Males stridulate day and night. First instars seen late April. First adults mid-June. Adults still present late November.

Gryllidae Gtyllinae Gryllus species. Rentz and Weissman (1978) provide details concern­ ing the state of taxonomic confusion in tllis genus. VIle assign numbers to species of this genus since names cannot be used with accuracy. Gryllus I through VI will be discussed in detail in that paper; Gryllus VII and VIII are diagnosed in this paper. Of critical importance in the detetmination of species is the use of calling songs and the subsequent analysis of oscillo­ grams. Such information is not available with old museum specimens. The 108 TI-IE WASMANN JOURNAL OF BIOLOGY

Gryllus Ill fig. 18

1..--- Gryll us VI Fi g . 19

fffl ~,ij lflf H Iill ~~~ tffl ill I Gryllus VII Fi g . 2 0 25°

4ffitt ~~It Itt I INI~ ~II 4tn ~Itt mt

Gryllus VIII 1 Sec .

Figs. 18-21. Oscillograms of calling songs of Gryllus species. Fig. 18. GryZZ.us Ill, Stanford Campus; Fig. 19. Gryllus VI, Palo Alto; Fig. 20. Gryllus VII, Lake Lagunita; Fig. 21. Gryllus VIII, Palo Alto. reader is also referred to the above reference for specifics concerning the counting of teeth on the file. Gryllus III. Most widely distributed species in our study area; 15(3); 17(3); 18(2); 19(4). Tltis species lives in cracks in the ground in oak wood­ land and grasslands. It has a metallic-sounding call and a fairly rapid calling rate of some 160 cltirps per minute. The number of pulses per chirp varies from 6- 10 but is usually 7 or 8 (see Fig. 18 for oscillogram). Gryllus VI. This is the only trilling field cricket in the study area; 16(1); 18(1); 19(1). This species is never common; adults fly well. Calling song may exceed 1,000 chirps per minute-each chirp consists of 3 pulses (Fig. 19). VOLUME 35, NUMBER 1 109

Gryllus VII. Recognition characters: tegmina smoky black, yellow stiipe present laterally; basal portion of internal face of hind femur reddish brown. Geographic distribution: We have not found this species south of Palo Alto; it is very common at Lake Lagunita (Site 16) on the Stanford campus. Habitat: Males sing from cracks in the ground or from under prostrate vegetation along the banks. Seasonal occurrence: Members of the species overwinter in the nymphal stage and adults appear in the spring. Song: Gryllus VII has a very slow chirp rate, less than 100 chirps per minute. There are usually 3 pulses per chirp but this may range between \ 2 and 4 (Fig. 20).

Measurements m Gryllus VII (in mm) Males Number Length Number Length teeth body file file per rnm ±SD teeth ±SD ± SD (range) ± SD ( range) (range)

N = 15 16.6 ± 1.9 127 ± 8.0 2.9 ± 0.2 43.9 ± 2.4 (14.3- 19.5) ( 116-145) (2.5-3.3) ( 40.6--48.3)

Females Length Length body ovipositor ±SD ±SD (range) (range)

N=11 16.2 ± 1.7 11.4 ± 0.8 (13.0-18.5) ( 10.3-12.7)

Gryllus VIII. Recognition characters: similar to Gryllus VII in size, call­ ing song, and some file characters; can be separated from Gryllus VII by habitat data and, morphologically, because the inner face of hind femur is entirely black; tegmina with distinct yellow tinge. Geographic distribution: Tlus species is known only from sites 2(2); 19(4). Habitat: This species is sympatric with Gryllus III and Neonemobius eurynotus at the dry grazed grassland area above Foothill Expressway. Males sing at night and live within cracks in the ground. An extensive degree of mite parasitism in 1974 at site 19 apparently resulted in a complete absence of individuals in 1976. 110 THE WASMANN JOURNAL OF BIOLOGY

Seasonal occurrence: Adults appear by April, indicating they probably overwinter as nymphs. Song: This species has a slow calling song of around 125 chirps per minute. There are between 3 and 6 pulses per chirp (Fig. 21), although 4 or 5 is the more common condition.

Measurements in Gryllus VIII (in mm) Males Foothill Expressway Number :'-lumber Length file Length teeth body teeth fil e permm 1 ±SD ±SD ±SD ±SD (range) ( range) (range) (range) 16.2 ± 1.0 119.7 ± 9.0 2.4 ± 0.1 49.8 ± 5.6 ( 14.5-17 .0) ( 110-135) (2.2-2.5) ( 44.8-61.4)

Jasper Ridge 17.6 ± 0.1 121 2.8 43.7 ( 17.5-17.7) (117- 125) (2.6-2.9) ( 42.4-45.0)

Females Foothill Expressway Length Length body ovipositor ±SD ±SD (range) ( range) N = 13 14.6 ± 1.1 8.5 ± 0.3 ( 13.0-16.6) (8.0- 9.0)

Nemobiinae Neonemobius eu.rynotus (Rehn & Hebard), new combination. Rare at Jasper Ridge at site 1 and Stanford campus at site 15; extremely common and heavily parasitized with mites in 1974 at site 19 where thousands of males were heard trilling at night. No specimens found in 1976 at site 19 apparently due to extensive parasitism by same mite species that also depleted the numbers of Grullu.s VIII at this locality. vVe transfer this species from Nemobitts at the suggestion of Dr. V. R. Vickery, Lyman Museum, MacDonald College, Quebec, Canada. This species has not b een studied since its original description (Rehn and Hebard, 1918). The current interpretation of generic limits in this group indicates that the species belongs in Neonemobius or in a separate monotypic genus. VOLUME 35, NUMBER 1 111

Dr. Vicke1y has kindly studied the ka1yotype of this species and discovers that the male has 19 chromosomes (2N 3 = 19). All autosomes are aero­ or telocentric with an acrocentric X chromosome. The related Allonemobius has 2N 8 =17.

Oecanthidae Oecanthinae Oecanthus m·gentinus Saussure. Extremely rare in Palo Alto area. The crickets have been found only on star thistle, Centa.urea species, in disturbed fields of grasses on the Stanford campus. First adults appear in early July. Oecanthus califom icus Saussure. Chaparral inhabitant on Jasper Hidge, sings during afternoon as well as at night. First adults in early July. Found on Quercus dumosa Nutt., Quercus spp., various trees and shrubs, and grasses under these trees. Oecanthus fultoni T. ' Valker. Found on evergreen oaks and shrubs on Jasper Hidge and in Palo Alto gardens. Oecanthus quadripunctatus Beutenmuller. Extremely common on Jasper Ridge on tarweed, H emizonia species. Sings late afternoon and at night. Walker and Rentz (1967) discuss taxonomy and acoustics of this species which seems to be dwarf in the San Francisco Bay Area. Adults in early July. Oecanthus rileyi Baker. First individuals sing in late June but some males still heard in late D ecember. On oaks, Q. agrifolia Nee, Q. lobata, Q. doug­ lasii H. & A., and shrubs on Jasper Ridge and in Palo Alto gardens.

M yrmecophilidac Mymecophilus oreg011ensis Bruner. Common in ant nests of Veramessor andrei (Mayr) and Camponotus semitestareus Emery along the main b·ail on Jasper Ridge, particularly at sites 2 and 8. Apparently these inquilines emerge at night to spend considerable time on the surface of the ground.

Stenopelmatidae Stenopelmatinae A general review of the genus Stenopelrnatus is provided by Tinkham and Rentz (1969). Stenopelrnatus intermedius Davis & Smith. Present during the rainy season under rocks in areas of sandstone grassland and oak woodland. Has not been found in serpentine grassland; 7(1); 10(1). 112 THE WASMANN JOURNAL OF BIOLOGY

Stenopelmatus longispina Brunner. Single adult male collected 5 October 1974 near Jasper Ridge. Stenopelmatus fuscus (Haldeman). Subadult male and female collected in mid-August 1974 at site 19.

Rhaphidophoridae Ceuthophilinae Ceuthophilus californianus Scudder. Present in moderate numbers at site 15. Matures in mid-June. Gmnmarotettix bUobatus Thomas. Found on trunks of coast live oak, Quercus agrifolia, in early spring; 1; 5. No individuals seen since spring of 1970. Pristoceuthophilus nwrmoratus Rehn. Found at sites 4; 17; 19. In­ dividuals became adult, in laboratory, by 1 October. Pristoceuthophilus sp. near pacificus. Found at sites 14; 15. Many adults attracted to oabneal trails laid along path through redwoods in late September. Species to be described by T. H. Hubbell.

Timemidae Timema californicum Scudder. Found at sites 3; 5; 10; 11; 12. Beaten from vaxious chaparral and oak woodland vegetation including Adenostoma fasciculatum, Ceanothus cuneatus (Hook.) Nutt., C. sorediatus H. & A., Arctostaphylos spp., Qttercus durata Jeps., Q. agrifolia, Eriodictyon cali­ fomicum (H. & A.) Torr., Baccharis pilularis DC., Eriophyllum spp., and Heteromeles arbutifolia M. Roen. Adults by mid-April, many pairs found together. Between 5-10% of adults are brown, remainder green. Adults present at site lO on Q. agrifolia until late June.

Mantidae Litaneutria minor Scudder. Especially conm1on in 1974 along trail in vicinity of sites 7 and 8. Pairs frequently encountered in copula.

Blattellidae Blattellinae PaTcolJlatta americana Scudder. During the wet spring, nymphs can be found under medium to large rocks in serpentine grassland sites 2 and 3. As these areas dry through mid-April, adult females and nymphs are found at night scavenging near ant nests along the path at sites 2, 3, 8, 10. They apparently take refuge in these nests during the daytime. VOLUME 35, NUMBER 1 113

Acknowledgments \ i\Te thank Steven Weissman for field assistance; E1ika French for sharing her unpublished data with us; Hal Mooney for providing the photograph used in Fig. 1; John Thomas for plant identifications; R. Snelling for ant identifications; V. R. Vickmy for his cytological determination and information regarding Neonemobius; and \i\Terner Lober for the Grylltts oscillograms. D.B.W. was supported by NIH Training Grant GM00365.

Literature Cited Arnaud, P. H., Jr., and D. C. Rentz. 1965. Ceracia dentata, a parasite of Chimaro­ cephala pacifica pacifica in California. (Diptera: Tachinidae and Orthoptera: Acrididae). Pan-Pacific Entomologist 41:204-206. E hrlich, P. R. 1965. The population biology of the butterfly, Euphydryas editlw. II. The structure of the Jasper Ridge colony. Evolution 19:327-336. Grant, H. J., Jr., and D. C. Rentz. 1966. The katydid genus Platylyra (Orthoptera: ). Pan-Pacifi c Entomologist 42:81-88. John, B., and D . B. Weissman. 1977. Cytogenetic components of reproductive isola­ tion in Trimerotropis thalassica and T. occidentalis. Chromosoma 60:187-203. Lober, W., and M. K. Chandrashekaran. 1970. Acoustical and se:-..ual behavior in the grasshopper Chimarocephala pacifica pacifica (Oedipodinae). Entomologia Ex­ penmentalis et Applicata 13 :71-84. Md\eill, J. 1900. The Orthopteran genus Trimerotropis. Psyche 9:27-36. Rehn, J. A. G., and M. Hebard. 1918. A new species of the genus Nemobius from California (Orthoptera; Gryllidae; Gryllinae). Entomological News 29: 102-105. Rentz, D. C. 1963a. Additional records of Platycleis tessellata (Charpentier) in Cali­ fornia with biological notes. Pan-Pacific E ntomologist 39:252-254. Rentz, D. C. 1963b. Biological observations on the genus Decticita (Orthoptera: Tettigoniidae). Wasmann Journal of Biology 21 :91- 94. Rentz, D. C., and J. D. Birchim. 1968. Revisionary studies in the Nearctic Decticinae. Memoirs of the Pacific Coast Entomological Society 3:1-173. Rentz, D. C., and D. B. Weissman. 1978. Faunal affinities, systematics and bionomics of the Orthoptcra of the California Channel Islands. University of California Press, Berkeley (submitted for publication). Schroeter, G. L., and G. M. Hewitt. 1974. The effects of supemume•·ary chromatin in three species of grasshoppers. Canadian Journal of Genetics and Cytology 16:285-296. Tinkham, E. R., and D. C. Rentz. 1969. Notes on the bionomics and distribution of the genus Stenopelmatus in central California with the description of a new species (Orthoptera : Gryllacrididae). Pan-Pacific Entomologist 45:4-14. Walker, T . J., and D. C. Hentz. 1967. Host and calling song of dwarf Oecanthus quadriptmctatus BcutenmUller (Orthoptera: Gryllidae). Pan-Pacific Entomologist 43: 326--327.

(DBW) Department of Biological Sciences, Stanford University, Stanford, California 94305 and Associate, Department of Entomology, California Academy of Sciences, San Francisco, California 94118; (DCFR) Depart- 114 THE WASMANN JOURNAL OF BIOLOGY ment of Entomology, California Academy of Sciences, San Francisco, California 94118. Address reprint requests to David B. Weissman, UCIMC, Bldg. 53 Room 103, 101 City Drive South, Orange, California 92668.

Note added in proof.-H. F. Strechecker, W. W. Middlekauff, and D. C. Rentz (1968, Bulletin of the California Insect Survey 10:1-177) record Arphia conspersa hehrensi Saussure from Palo Alto, 7 June 1922. \Ve have been unable to verify this record.