The Leafmining Moths of the Genus Cameraria Associated with Fagaceae in California (Lepidoptera: Gracillariidae) PAUL A. OPLER and DONALD R. DAVIS SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 333 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." 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Smithsonian Contributions to Zoology, number 333, 58 pages, 131 figures, 2 tables, 9 maps, 1981.?The 17 species of the moth genus Cameraria that feed on Fagaceae in California are treated. A description of the major morpholog- ical features and biologies of each species is included, accompanied by appropriate distributional maps and illustrations of diagnostic characters. Fourteen new species are described, including Cameraria anomala, C. diabloensis, C. jacintoensis, C. lobatiella, C. marinensis, C. mendocinensis, C. pentekes, C. sadleri- anella, C. sempervirensella, C. serpentinensis, C. shenaniganensis, C. temblorensis, C. tildeni, and C. walsinghami. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavemosa (Linnaeus). Library of Congress Cataloging in Publication Data Opler, Paul A The leafmining moths of the genus Cameraria associated with Fagaceae in California (Lepi- doptera, Gracillariidae). (Smithsonian contributions to zoology ; no. 333) Bibliography: p. 1. Cameraria?Classification. 2. Cameraria?Host plants. 3. Fagaceae?Diseases and pests. 4. Insects?Classification. 5. Insects?California?Classification. I. Davis, Donald Ray, joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contribu- tions to zoology ; no. 333. QL1.S54 no. 333 [QL561.G7] 591s [595.78'1] 80-26743 Contents Page Introduction 1 Ecology 2 Guild Ecology 2 Interhost Adaptation 2 Evolution 4 Biogeographic Considerations 4 Host Switching 5 Cameraria Chapman 5 Key to the Species of Cameraria Chapman 6 The guttifinitella Species Group 8 Cameraria lobatiella, new species 9 Cameraria mendocinensis, new species 11 Cameraria sadlerianella, new species 11 The hamadryadella Species Group 12 Cameraria pentekes, new species 13 The agrifoliella Species Group 15 The agrifoliella Subgroup 15 Cameraria agrifoliella (Braun) 15 Cameraria anomala, new species 18 Cameraria wislizeniella Opler 19 Cameraria temblorensis, new species 21 The diabloensis Subgroup 22 Cameraria diabloensis, new species 22 Cameraria shenaniganensis, new species 24 The mediodorsella Subgroup 25 Cameraria mediodorsella (Braun) 25 Cameraria jacintoensis, new species 27 Cameraria serpentinensis, new species 28 The walsinghami Subgroup 29 Cameraria marinensis, new species 29 Cameraria walsinghami, new species 31 The sempervirensella Subgroup 32 Cameraria sempervirensella, new species 32 Cameraria tildeni, new species 34 Literature Cited 36 Figures 18-131 37 Maps 1-9 49 in The Leafmining Moths of the Genus Cameraria Associated with Fagaceae in California (Lepidoptera: Gracillariidae) Paul A. Opler and Donald R. Davis Introduction The leafmining genus Cameraria is believed to be indigenous to the Nearctic region. Before this report, it was represented by 38 species primarily restricted to the eastern deciduous forests of the United States. Intensive field investigations on the Microlepidoptera feeding on the several spe- cies of Fagaceae in California (Opler, 1974a, b) have resulted in the discovery of 14 undescribed species. Prior to this field work, only two oak- feeding Cameraria were known from California, C. agrifoliella (Braun) and C. mediodorsella (Braun), while another, C. wislizeniella Opler, was de- scribed in the course of the above work (Opler, 1971). This study was partially assisted by a grant from the Smithsonian Research Foundation to D. R. Davis in support of his work on the North American leafmining Lepidoptera. The field in- vestigations were supported by NSF grants GB4014 and GB6813X, with J. A. Powell as Paul A. Opler, Office of Endangered Species, Fish and Wildlife Service, U.S. Department of the Interior, Washington, D.C. 20240. Donald R. Davis, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. principal investigator. In addition to the above assistance, we wish to acknowledge the assistance of Ms. Biruta Akerbergs and Ms. Ann Szymkow- icz for preparing the genitalia illustrations. The habitus drawings were by Mr. Vichai Malikul. We wish also to acknowledge the generous assist- ance of J. A. Powell, University of California, Berkeley, for providing much of the material used in this study. Collection of specimens were by P. A. Opler and/or J. A. Powell unless otherwise stated. The deposition of specimens has been indicated as follows: BM(NH) British Museum (Natural History), London, England CAS California Academy of Sciences, San Francisco, California GD Gerfried Deschka, Steyr, Austria LACM Los Angeles County Museum, Los Angeles, California UCB University of California, Berkeley, California UCD University of California, Davis, Cali- fornia 2USNM former United States National Mu- seum, collections in the National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. Ecology An understanding of the ecological interactions partaken by the Cameraria species is critical to an understanding of their speciation and evolution- ary history. These interactions may be examined in two ways. First, we may examine their role within the communities (or guilds) of leaf-miners to which they belong, and, secondly, we may inspect the ecological similarities and differences between species of Cameraria that feed on different oaks. GUILD ECOLOGY.?The assemblage of miners that feed on a single host has been referred to as a "guild" (Opler, 1974a, b), since these species all utilize the same part of their host in the same way (see Root, 1967). Guilds of leaf-miners are rich on Californian oaks or their relatives, and the number of species found on a given host throughout its distribution ranges from 2 to 18. The number of miners found on a host has been shown to be a function of area of host occupation (Opler, 1974b). Californian oak leaf-miners belong to 15 genera distributed among nine families, and are primarily Lepidop- tera. Species belonging to most of these genera feed on Quercus agrifolia, and their life histories have been described in detail (Opler, 1974a). On most hosts, no more than a single species from each genus is present, and in these cases each species fills a generalized niche. For example, the Lithocarpus densiflora miner guild is composed of five species: a Cameraria, a Phyllonorycter (= Lithocolletis of authors), a Caloptilia, a Stigmella, and a Bucculatrix. Cameraria makes an upper epi- dermal blotch mine, and Phyllonorycter makes a lower epidermal blotch mine. Caloptilia makes a small, lateral, full-depth blotch on very young leaves and then skeletonizes the upper surface from within a folded shelter; Stigmella makes a SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY full-depth serpentine mine; and Bucculatrix makes a small, full-depth, linear mine near the midrib or a lateral vein, later leaving the mine to skele- tonize the lower leaf surface. Other genera have analogous stylized feeding patterns, which may have resulted from competition through evolu- tionary time. The "upper epidermal blotch niche" filled by Cameraria seems to be a basic one in the assembly of leaf-miner guilds, since only the two rarest Californian oaks, Quercus dunnii and Q tomentella, do not harbor at least one Cameraria (Opler, 1974b), and since Californian oaks with genera represented by more than one species always include Cameraria among their polyspecific genera. INTERHOST ADAPTATION.?The miners, includ- ing Cameraria, which feed on evergreen or decid- uous Fagaceae in California exhibit two strikingly different adaptive modes (Opler, 1979). It has been shown that, on the average, Cameraria that feed on evergreen Fagaceae are larger, have longer larval feeding periods, have relatively low population densities, and usually have fewer an- nual generations in comparison to those that feed on deciduous Fagaceae (Table 1; Opler, 1979). In addition to the traits mentioned above there are three additional concomitant contrasts which can be made: the degree of host specificity is greater for Cameraria that feed on evergreen Fagaceae; larval feeding periods encompass the winter months for species with evergreen hosts; and no more than one evergreen-restricted Cameraria is ever found at a single locality, even though several may be found throughout the range of the host (Table 1). Opler (1979) homologized these two constella- tions of adaptations with the concepts of r- and K-selection as proposed by MacArthur and Wil- son (1967) and as elaborated by Pianka (1970). According to this theory, species which are r- selected are small, have good colonizing ability, have high maximum reproductive rates (r), and have frequent changes in population level, seldom maximizing their resources near carrying capacity (K). In contrast, K-selected species are larger, are poor colonists, but good competitors, have lower NUMBER 333 TABLE 1.?Host relationships of California Cameraria Host Lithocarpus densiflora var. echinoides Chrysolepis sempervirens chrysophylla Quercus (Protobalanus) chrysolepis var. nana Quercus (Erythrobalanus) agrifolia wislizenii var. frutescens kelloggii Quercus (Lepidobalanus) lobata sadleriana garryana douglasii X alvordiana ( = douglasii X turbin- ella ssp. califor- nica) dumosa dumosa X engelmanii dumosa X turbinella ssp. califomica durata guttifinitella species group lobatiella, new species lobatiella, new species sadlerianella, new species mendocinensis, new species lobatiella, new species hamadryadella agrifoliella species group subgroup agrifoliella (Braun) anomala, new species wislizeniella Opler anomala, new species wislizeniella Opler wislizeniella Opler pentekes, new species pentekes, new species temblorensis, new species temblorensis, new species temblorensis, new species agrifoliella species group diabloensis mediodorsella walsinghami semperviren- subgroup subgroup subgroup J?/& subgroup marinensis, new species walsinghami, new species sempervirensella, new species tildeni new species diabloensis, new species new species diabloensis, new species mediodorsella (Braun) mediodorsella (Braun) mediodorsella (Braun) new species jacintoensis, new species serpentinensis, new species jacintoensis, new species serpentinensis, new species SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY reproductive rates, and maintain relatively con- stant population sizes near carrying capacity. It is hypothesized that evergreen oaks have higher levels of chemical defense and are more predictable resources than are deciduous oaks. In any event, Comer aria that feed on evergreen Fa- gaceae seem to be K-selected relative to those species that feed on deciduous hosts. A prediction that may be made on the above postulate is that evergreen-feeding Cameraria should oviposit fewer, relatively larger eggs than their deciduous host- associated counterparts (see also Connor et al., 1980). The observation that more than one evergreen- feeding Cameraria are never found in a single locality is not a necessary correlate of r-K theory and is novel. This implies that the evergreen- feeding Cameraria are relatively strong competi- tors, and that niche breadths may be broader for these species. In contrast, two or more deciduous- associated Cameraria often occur sympatrically, illustrating a possible lower competitive ability with resultant narrower niche breadths and greater species packing. Evolution BIOGEOGRAPHIC CONSIDERATIONS.?It has been shown previously that the number of miners found on different Californian Fagaceae is a func- tion of the distributional areas occupied by their hosts (Opler, 1974b). Even when one considers Cameraria alone this regression is still significant (S ?? 0.01 A0"66, where S ? number of Cameraria per host, and A = distributional area occupied by host). As has been discussed previously this correlation is an evolutionary one, but one that is maintained in current ecological time (Opler, 1974b). As was pointed out in the ecological discussion above, the Fagaceae with more than one Cameraria fall into either of two categories. These differing patterns of distributional diversity have been referred to by ecologists as alpha and beta diversity (Whittaker, 1965). Alpha diversity is the relative species richness or diversity at single localities or habitats and may be referred to as within-habitat diversity. On the other hand, beta diversity is the measure of increased species rich- ness or diversity when one considers more than one locality or habitat. Into one category, alpha diversity, fall those Cameraria that share a single deciduous host. More than one species of these moths often share a single host at a single locality, examples being Cameraria lobatiella and C. mediodorsella at Vaca- ville, Solano County, on Quercus lobata; and Ca- meraria lobatiella and C. pentekes near Hanford, Kings County, on Quercus lobata. More than one Cameraria were found on Quercus kelloggii, and the same is expected for Quercus garryana. It is assumed that some difference in seasonal timing or other form of host partitioning has evolved that allows these moths to coexist. In the case of sympatric occurrences on Quercus lobata, one member of the congeneric pair is always Cameraria lobatiella, a moth that frequently forms communal mines. Some partitioning of host range is seen by Cameraria that feed on Quercus lobata and Q. garry- ana. The latter, whose range extends over the greatest range of latitude, hosts three Cameraria that never occur together, while the former has two species, C. mediodorsella and C. pentekes, which have not been found sympatrically. Thus some evidence of beta diversity does exist for deciduous oak-associated Cameraria. A further observation on host-sharing by decid- uous oak Cameraria is that, with one exception, all species sharing a host belong to different species groups. For example, the three Cameraria that feed on Quercus lobata belong to the guttifinitella, agrifol- iella, and hamadryadella groups. Two of the four Cameraria that feed on Quercus garryana belong to the guttifinitella group, and are therefore an excep- tion, although it should be noted that they do not occur together. In contrast, all Cameraria that share a single evergreen host never occur sympatrically, and are therefore examples of beta diversity. In addition, all but one such example of host sharing are drawn from the same species group, and some are examples of obvious "sister species." The pairs of Cameraria that share Lithocarpus densiflora and Quer- NUMBER 333 cus chrysolepis are the most obvious of such sister species, and speciation has obviously resulted from divergence subsequent to geographic isola- tion of the same host's populations. In the case of the two Lithocarpus-keding Cameraria, one is re- stricted to the host variety echinoides, which is found only in the Siskiyou Mountains, and one of the Quercus chrysolepis-feeding Cameraria is lim- ited to Coast Range and Transverse Range loca- tions, which were isolated islands during the Pli- ocene, and to which the host variety nana is restricted. Quercus agrifolia hosts three Cameraria, all be- longing to the agrifoliella subgroup. Cameraria wis- lizeniella and C. anomala also occur widely on a different but closely related host, Q. wislizenii. As has been discussed in the species treatment, the Cameraria wislizeniella populations that feed on Quercus agrifolia exhibit different life history fea- tures from those that feed on its more usual host, and may represent a "host race" or weakly differ- entiated sibling species. HOST SWITCHING.?The events that led to many ultimate speciations of Californian oak Ca- meraria probably began with sympatric host trans- fers followed by isolation (Opler, 1974b). This is apparent when one examines the geographic dis- tribution and host composition of several Camer- aria species subgroups (Maps 1-9). One case, previously discussed by Opler (1974a), is that of the agrifoliella species group. The moths feed on disparate Fagaceae, Lithocarpus densiflora, Chryso- lepis sempervirens, C. chrysophylla, and Quercus agri- folia. In addition other species in the group, Ca- meraria gaultheriella, C. nemoris, and C. species, feed on Gaultheria shallon, Vaccinium ovatum (both Erica- ceae), and Myrica califomica (Myricaceae), respec- tively. The hosts share geographic commonality as all occur in northern coastal California (except Chrysolepis sempervirens) and all probably origi- nated with other mesic elements of the Arcto- tertiary geoflora (Wolfe, 1969). The majority of hosts for members of the mediodorsella species sub- group are most often found in chaparral or similar xeric situations. The group's locus is now in the southern Coast and Transverse Ranges. The lo- batiella species subgroup's hosts are centered in the northern Coast Ranges of northern California and southern Oregon. Evidence of this mode of switching and subse- quent speciation is provided by examples of oc- casional emergence of Cameraria from the "wrong host" at localities where a normal host also occurs. Examples are the one Cameraria lobatiella reared from Quercus douglasii, and Cameraria agrifoliella and C. mediodorsella reared from Quercus suber, intro- duced cork oak. The rate of changeover and adaptation is most rapid for moths colonizing deciduous oaks. This can be shown by two kinds of evidence. First, the Cameraria of deciduous oaks are less host specific, the most extreme of which are two California examples, Cameraria lobatiella and C. mediodorsella, which include both Lepidobalanus "white" and Erythrobalanus "red" oaks in their host range. A second kind of evidence is that the only Cameraria with clear affinities external to the Pacific Coast states are species that feed on deciduous oaks, while, in contrast, the most unusual Californian endemic Cameraria are those found on evergreen Fagaceae. Members of the hamadryadella and gut- tifinitella species groups feed on deciduous oaks, with the exception of Cameraria sadlerianella. Cameraria Chapman Cameraria Chapman, 1902:141. [Type of the genus: Lilhocol- letis guttifinitdla Clemens, 1859.] DIAGNOSIS.?Adults small, 5-12 mm in wing- span, usually red brown to orange (occasionally gray white) with white transverse bands and black patches on forewing; white bands (if dis- tinct) typically dark margined externally. Egg round and flat, transparent, deposited singly (rarely in multiples) on upper leaf surface. Larvae hypermetamorphic, forming blotch mines on the upper surface of host leaves, with one or more creases being formed in the upper epidermis over the mine at maturity. Body form flattened throughout most of larval life, becoming cylin- drical at final instar. Pupa without cremaster; SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY pupation occurring under a flat circular cocoon within the mine. The vast majority of species are restricted to a single host. ADULT.?Head: Vestiture of frons consisting of appressed, lustrous white scales; vertex with erect scales, usually white medially and orange brown laterally. Antennae nearly as long as forewing; flagellimeres white basally, black dis- tally; maxillary palpus reduced, less than 0.5 the length of apical segment of labial palpus, 2-seg- mented, segments only slightly longer than broad, subglobular. Labial palpus moderately long, ap- proximately 1.2X-1.5X diameter of eye; vestiture entirely white, or white anteriorly and black pos- teriorly. Thorax: Covered with slightly raised scales, dorsum usually white medially, orange brown laterally; tegulae orange brown or white, or or- ange brown outwardly edged with white in- wardly; pleuron and venter covered with ap- pressed, white scales. Forewing (Figure 33) 8- veined; Rl and R4 absent; Ml absent, M2 fused with 3; CuA single branched; dorsal scales non- iridescent, usually orange brown, but occasionally gray (white and black scales intermixed), with transverse white bars or fasciae often edged out- wardly with black; ventral vestiture black with smaller areas of tan. Hind wing 4-veined, Sc + Rl extremely short, terminating near base of costa; M and CuA each single branched; 1A vestigial; both surfaces pale to dark gray. Foreleg with epiphysis absent; femur white or black, or black dorsally and white ventrally; tibia usually black dorsally, white ventrally; tarsus usually black and white banded; midleg with femur usu- ally white; tibia white with lesser amounts of black, tarsus usually white with narrow black bands; hind leg with femur white, occasionally with a small black patch; tibia predominantly white with black or brown patches; tarsus white with black bands. Abdomen: Dorsum dark gray, paler at posterior (male), pale or dark gray (female), pleuron and venter white. Eighth sternite highly modified in male and usually extending ventrally beneath genitalia (as in Phyllonorycter); caudal apex either entire and rounded to subacute, or variously bifid or bilobed. Male Genitalia: Uncus absent. Tegumen par- tially membranous, reduced to a relatively nar- row ring dorsally. Vinculum often relatively well developed, either V- or Y-shaped, typically with a prominent median process (i.e., vincular pro- cess) extending posteriorly. Valvae symmetrical, without spines, slender, usually attenuated in western species or with cucullus moderately en- larged. Aedeagus relatively simple, base often flared, gradually tapering to slender apex; cornuti absent although short exogenous spines some- times present along distal half of shaft. Female Genitalia: Two pairs of apophyses pres- ent. Lamella ante vaginal is variously developed but usually extended (in western species) poste- riorly beyond ostium and with apex either entire or most often bifid. An accessory bursa usually present and arising between middle and posterior fourth of ductus bursae; corpus bursae spherical, with walls membranous except for typically two asymmetrical signa; signa less commonly single or absent. Key to the Species of Cameraria Chapman Male genitalia with costal margin of valva relatively straight (Figure 34). Female genitalia with signa absent; lamella antevaginalis (seventh sternite) simple, not deeply lobed nor divided (Figure 99). (guttifinitella group) 2 Male genitalia with costal margin of valva strongly arched or sinuate. Female genitalia with a pair of ovoid signa; lamella antevaginalis deeply lobed (Figure 130) or divided (Figure 105) 4 NUMBER 333 2. Length of forewing more than 4 mm. Larva on Quercus sadleriana Cameraria sadlerianella, new species Length of forewing less than 4 mm. Larva on either Quercus garryona or Q. lobata 3 3. Forewing relatively narrow; length/width ratio 5.0 or less. Larva on Quercus lobata Cameraria lobatiella, new species Forewing relatively broad, length/width ratio greater than 5.0. Larva on Q. garryana Cameraria mendocinensis, new species 4. Ground color mostly white with irregular golden brown bands (Figure 19). Male genitalia with vincular process greatly expanded, 5-pointed (Figure 43). Female genitalia with lamella antevaginalis deeply divided (Figure 105). {hamadryadella group) . Cameraria pentekes, new species Ground color of forewing orange brown to golden brown, banded with white. Male genitalia with vincular process often elongate but not expanded (Figures 47, 51). Female genitalia with a median, sometimes slightly cleft lobe arising from caudal margin of lamella antevaginalis (Figure 130). {agrifoliella group) 5 5. Male with eighth sternite approximately as long as broad; caudal lobes slender, reduced in size (Figures 75, 81). Aedeagus sinuate (Figures 76, 82). {mediodorsella subgroup) 6 Length of eighth sternite of male more than 1.5X the width; caudal lobes more developed and broad. Aedeagus relatively straight 8 6. Forewing with medial white fascia reduced and incomplete (Figure 28). Male genitalia with vinculum reduced, saccus slender (Figure 80). Caudal lobes of eighth sternite extremely reduced (Figure 81) Cameraria serpentinensis, new species Forewing with medial fascia extending across wing. Male genitalia with vinculum and saccus broad (Figure 74). Caudal lobes of eighth sternite more prominent (Figure 75) 7 7. Larva feeding on deciduous oaks and diapausing as a pupa during winter Cameraria mediodorsella Larva feeding on evergreen oaks, no pupal diapause Cameraria jacintoensis, new species 8. Male genitalia with digitate lobe at outer third of valva (Figure 91). Larva feeding on Chrysolepis. {sempervirensella subgroup) 9 Male genitalia without digitate lobes. Larva on Quercus or Lithocarpus . 10 9. Dorsum of thorax mostly white. Forewing with basal longitudinal streak; basal fascia absent (Figure 31) Cameraria sempervirensella, new species Dorsum golden brown. Forewing without basal streak; basal fascia pres- ent, broken (Figure 32) Cameraria tildeni, new species 10. Male with eighth sternite possessing a pair of slender caudal lobes turned 90? dorsad (Figure 62); valva with a small, ventral, keel-like lobe at subapex of cucullus (Figure 59) Cameraria temblorensis, new species 8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Male with eighth stemite variously modified but not as above; valva without subapical ventral lobe 11 11. Male with eighth stemite possessing two pairs of caudal lobes, the largest pair turned 90? dorsad (Figure 50); valva as in Figure 47, with a slightly enlarged setigerous apex Cameraria agrifoliella Male with eighth stemite bilobed, not as above; valva not as in Figure 47 12 12. Male with eighth stemite possessing a pair of moderately large, rounded, caudal lobes directed dorsad at a 45? angle (Figure 58); valva with a short apical process extending from cucullus as in Figure 55 Cameraria wislizeniella Male with eighth stemite not as in Figure 58; valva without subapical process 13 13. Male with eighth stemite possessing greatly enlarged caudal lobes sepa- rated more than one-third the length of sternite (Figure 54); valva with a large subapical lobe extending more than one-fourth the length of entire valva (Figure 51) Cameraria anomala, new species Male with caudal lobes of eighth sternite separated less than one-fourth the length of sternite; valva without large lobe as in Figure 51 . . . . 14 14. Forewings with antemedial fascia completely traversing wing. Male with a broadly rounded, ventral lobe at distal fourth of valva. Lamella antevaginalis (seventh sternite) of female with slender median cleft (Figure 127). (walsinghomi subgroup) 15 Forewings with antemedial fascia incomplete, stopping at middle of wing or less. Male without subapical lobes on valva. Lamella antevaginalis without cleft, instead produced into a median lobe (Figure 115). {diabloensis subgroup) 16 15. Eighth sternite of male strongly arched ventrad (Figure 90); valva with apex slightly expanded (Figure 87). Larva feeding on Lithocarpus densi- flora var. echinoides Cameraria walsinghami, new species Eighth sternite not strongly arched (Figure 86); valva with slender apex (Figure 83). Larva feeding on Lithocarpus densiflora Cameraria marinensis, new species 16. Costal margin of forewing with white scaling, fascia indistinct (Figure 25). Valva of male with a slight setigerous swelling at outer fourth (Figure 70) Cameraria shenaniganensis, new species Costal margin of forewing same as ground color, golden brown; fascia distinct on costal half of wing (Figure 24). Valva with setigerous swelling at outer third (Figure 63) Cameraria diabloensis, new species The guttifinitella Species Group In California, the lobatiella subgroup consists of three sibling species, each limited primarily to a The group of three species centered around C. single host Quercus, and all with nearly identical lobatiella is actually a branch of the much larger, maculation. The male genitalia of the three spe- predominantly eastern guttifinitella species group. cies are also very similar in form and resemble NUMBER 333 the genitalia of most of the eastern species of Cameraria (including the type of the genus) in possessing relatively simple unmodified valvae and eighth sternites. In this respect, the male genitalia of the lobatiella subgroup differ markedly from all other California Cameraria, which typi- cally possess sinuate valvae and more modified eighth sternites. The female genitalia are characteristic in pos- sessing two pairs of abdominal apophyses, a com- pletely membranous corpus bursae with no signa, and the ostium flush with the abdominal wall. Cameraria lobatiella, new species FIGURES 1, 2, 18, 34-36, 98, 99; MAP 1 DIAGNOSIS.?The maculation and genital mor- phology of this species closely resembles that of the other members of the lobatiella group, C. men- docinensis and C. sadlerianella. Cameraria lobatiella differs in possessing relatively short, broad wings (see Table 2) as well as in certain biological features. The larvae are gregarious in the mines (solitary in the other two species) and feed on Quercus lobata, Q. douglasii, and Q. kelloggii. Whereas the other two species in this group are single brooded, Cameraria lobatiella may undergo as many as three or four broods. MALE.?Length of forewing: 2.8-3.8 mm. Head: Labial palpus white. Antenna almost as long as forewing; scape and flagellar segments dark brown dorsally, white ventrally. Front white. Vertex brown anteriorly, white mixed with brown posteriorly. Thorax: Pronotal scaling brown with white laterally and anteriorly; tegula brown and narrow TABLE 2.?Wing measurements of the guttifinitella species group of California Cameraria C. lobatiella C. mendocinensis C. sadlerianella