taxonID	type	description	language	source
1E1F87DDFF9EFFD89939297A9AECFDA8.taxon	description	Summary of Natural History: Synergus (Hymenoptera: Cynipidae: Synergini) are usually professed to be inquilines, but are perhaps more accurately described as gallers of galls (Askew 1961). Synergus induce additional growth in existing galls, including the formation of larval chambers, and their developing young feed on the tissue of the gall (Evans 1965). Though gall inducing Synergus have been documented in Japan (Abe et al. 2011; Ide et al. 2018), gall induction in Synergus is a derived habit (Ide et al. 2018) and not known from the Nearctic. In some galls, the presence of Synergus is fatal to the developing gall inducer, but in other cases food may be sufficient such that both may emerge (Pénzes et al. 2012). In some galls, Synergus develop and emerge as adults within a matter of weeks, while others can take one or even two years to emerge (Evans 1965; Busbee 2018; Ward et al. 2020). Multiple species of Synergus can be associated with the same gall type (Askew 1961; Pénzes et al. 2012; Bird et al. 2013; Forbes et al. 2016; Weinersmith et al. 2020), while other galls have no known Synergus associates despite large collection efforts (e. g., Joseph et al. 2011). There has been some previous suggestion that two other genera of cynipid inquilines (Ceroptres and Euceroptres) of Nearctic gall wasps may not co-occur with Synergus (Brookfield 1972), but curated rearing records (e. g., Krombein et al. 1979) and our own data presented here show that this is not universally true. Relationship to galler phylogeny: Synergus wasps were reared from gall types across most of the Nearctic gall wasp phylogeny (Ward et al. 2022), with some exceptions. In only two cases were Synergus reared from gall types produced by gallers in the large clade that includes genera Melikaiella Pujade-Villar, Loxaulus Mayr, and most of the Neuroterus Hartig (Fig. S 1; gallers # 2 - 15 in Fig. 1 a). Both of these Synergus / Neuroterus associations were from Pacific coast galls. The reduced apparent association of Synergus with gall inducers in this clade might reflect that the Synergus association with oak gall wasps originated in the clade represented by the lower two-thirds of the tree. However, this hypothesized relationship is belied by records of Synergus being associated with the topmost clade of Palearctic gall wasps in fiugre 1 a (though these a) b) 1 2 3 4 5 8 6 7 11 10 12 9 14 13 15 16 17 18 19 20 21 22 23 24 25 26 28 29 27 c) 303534313332 Q 44 38 36 37403839 L 11 19 41 424745444346 V 48 49 50 51 52 53 d) 56 54 57 55 35 19 58 59 (66) (26) 60 61 62 63 66 65 67 64 4 2 68 69 (12) (4) 70 71 72 73 74 76 77 78 75 3 0 80 79 (10) (4) 81 82 83 e) f) could represent secondary colonizations). Ultimately, assessment of coevolutionary relationships of Synergus with oak gall wasps requires a phylogeny of the Holarctic Synergus. Biogeography and oak tree section: Synergus were reared from galls in all three North American oak floristic regions and from galls on trees across all three sampled oak sections (Fig. 1 b c). Of the three sections, Synergus were least often reared from galls on section Lobatae. Tree organ and gall size: Synergus were reared from galls developing on leaves, stems, buds, acorns, and petioles (Fig. 1 d). Among organs from which we sampled galls, only flower galls did not have apparent Synergus associates. Synergus in our collections were most commonly reared from medium sized galls (61 %) and least commonly reared from small galls (38 % of galls smaller than 5 mm) (Fig. 1 e). Though these differences are not large, they comport with observations of Palearctic Andricus Hartig galls which suggest that small bud and catkin galls were less likely to host Synergus (Stone et al. 1995). Alternatively, reduced association of Synergus with small bud and catkin galls could be related to their earlier temporal occurrence: 25 of the 27 putative Synergus species in Ward et al. (2020) were reared from galls developing in June or later. However, we again raise the caveat that small galls may desiccate in the lab causing associated insects to die before emergence and leading to apparent non-associations. Co-occurrence with other natural enemies: When Synergus were present, two other putative inquiline genera were significantly less likely to be present: Euceroptres (P = 0.005) and Ceroptres (P = 0.0026). Thus while our data disagree with the suggestion that Synergus and Ceroptres / Euceroptres entirely displace one another (Brookfield 1972), they do appear to co-occur less often than expected. This pattern could be due to competitive exclusion, but also or instead be an indirect result of differential adaptation to dimensions of gall hosts. Notably, different Synergus species do not apparently competitively exclude one another, with as many as five species having been reared from the same host gall (Pénzes et al. 2012). To the extent that more closely related species are expected to compete more closely when sharing the same habitat (Miller 1967; Denno et al. 1995), differential adaptation to some dimensions of the gall environment seems the more attractive hypothesis. Additional notes: Our record of four Synergus wasps reared from galls of Andricus quercuscalifornicus appear to be the first ever, despite much attention having been paid to this particularly large and common Pacific coast gall and its natural enemies (Joseph et al. 2011). Other efforts to collect and rear insects from large numbers of potential hosts often turn up uncommon associations (Yee 2008; Yee and Goughnour 2008). Given that host shifts have often been implicated in the origins of parasitic insect diversity (Diehl and Bush 1984; Drés and Mallet 2002; Forbes et al. 2017), evidence of insects occasionally being reared from unexpected hosts suggest that variation in host recognition syndromes may result in insects often “ testing ” new potential host plants.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF9CFFD699392A7A9F62FD08.taxon	description	Summary of Natural History: The genus Ceroptres are oak-associated putative inquilines (or kleptoparasitic gall-modifiers; Ronquist 1994), but the biology of members of this genus is not as well studied as many Synergus, therefore their role requires further investigation. Originally in the same tribe as Synergus, they have been moved to their own tribe, the Ceroptresini, reflecting their likely independent evolution of inquilinism (Nylander 2004; Ács et al. 2010). Ceroptres have a Holarctic distribution with most named species from the Nearctic (Pénzes et al. 2012; Lobato-Vila and Pujade-Villar 2019), though it is probable that more species remain to be described from the Eastern Palearctic (Wang et al. 2012). Note that our “ Ceroptres ” does not distinguish between Ceroptres sensu stricto and the recently described and currently monotypic genus Buffingtonella (Lobato-Vila and Pujade-Villar 2019). Relationship to galler phylogeny: Ceroptres were associated with gall inducers across the Nearctic gall wasp phylogeny (Ward et al. 2022), and are also known from Palearctic galls, including the most basal on the tree (Fig. 2 a). Ceroptres were not reared from some large gall wasp clades, including 1) the Amphibolips Reinhard galls, 2) a clade of Andricus and Callirhytis Förster bud and leaf galls, and 3) a clade of primarily Pacific coast galls (though no Ceroptres were reared from any Pacific coast galls). Ceroptres were also never reared from any of the more than 13,000 Belonocnema Mayr galls collected from across the southeastern U. S. A. Biogeography and oak tree section: Ceroptres were reared much more commonly from galls collected in the Eastern half of the United States than in the Southwest or the Pacific coast (Fig. 2 b). We reared no Ceroptres from any Pacific coast gall types, though at least two species are known from California (McCracken and Egbert 1922). Galls from all three focal oak sections produced Ceroptres, with the smallest proportion from Section Quercus (Fig. 2 c). Ceroptres are also known from oaks in section Cerris in the Palearctic and section Protobalanus in California (Lobato-Vila and Pujade-Villar 2019). Tree organ and gall size: Ceroptres emerged from galls on leaves, stems, buds, acorns and petioles, but not from flower galls (Fig. 2 d). A greater fraction of large (> 20 mm) gall types produced Ceroptres than small or medium sized galls (Fig. 2 e). a) b) 1 2 3 4 6 8 5 7 10 11 12 9 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 29 28 c) 303233343531 Q 27 55 36 37403839 L 13 17 41 424744434546 V 48 49 50 51 53 52 d) 56 57 55 54 25 10 58 59 (66) (26) 60 61 62 63 67 64 66 65 3 1 68 69 (12) (4) 70 71 72 73 74 78 77 75 76 6 0 79 80 (10) (4) 81 82 83 e) f) Co-occurrence with other natural enemies: Ceroptres had more significant correlations (four) with other gall associates than any other genus (Fig. 2 f). Two correlations were positive: Ceroptres co-occurred more often than expected with another inquiline genus, Euceroptres (see below), and with the parasitoid genus Euderus. Euderus have been suggested to specialize on galls without external spines or hairs (Ward et al. 2019), so it is possible that some species of Ceroptres have similarly restricted host ranges. Significantly less likely to co-occur with Ceroptres were Synergus (P = 0.0026) and Torymus (P = 0.0003) parasitoids (see discussion in those sections).	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF92FFD49A9D2ADA9FA3FEE8.taxon	description	Summary of Natural History: The biology of Euceroptres is not well studied and it is not known whether members of this genus function as inquilines, parasitoids, or hyperparasitoids. Only four species of Euceroptres are previously described from galls of five species of gall wasps (Buffington and Liljeblad 2008). All known hosts are oak galls. The apparently small number of species and hosts for Euceroptres, coupled with the possibility that each species may have a limited range of hosts has led to the speculation that these are the surviving members of a previously more species-rich genus (Buffington and Liljeblad 2008). Relationship to galler phylogeny: Unlike the two genera of cynipoid associates of oak galls treated above, there are no published records of Euceroptres from the Palearctic (Buffington and Liljeblad 2008), though one unpublished record from Serbia is mentioned in Buffington et al. (2020). Most Nearctic gallers in our collections were not found to have associations with Euceroptres, though the nine that mapped onto the galler phylogeny were widely scattered across the tree (Fig. 3 a). Biogeography and oak tree section: All six species reared in this effort were from galls collected in the Eastern North American floristic region (Fig. 3 b). However, two species (Euceroptres maritimus Weld on Callirhytis quercussuttoni (Bassett) and Euceroptres montanus Weld on Disholandricus truckeensis (Ashmead )) were previously reared from California and Oregon. Three of the six gall hosts in our collections were collected from trees in oak section Quercus, while the remaining three were from trees in section Lobatae (Fig. 3 c). Previous collections were from galls found on oaks in sections Quercus and Lobatae as well as Protobalanus (Buffington and Liljeblad 2008; Manos and Hipp 2021). If Euceroptres are host specific as has been previously suggested (Buffington and Liljeblad 2008), our failure to rear them from western collections may well be a function of our collections not having included specific hosts. Tree organ and gall size: All Euceroptres reared in this study were from galls on leaves or on petioles (Fig. 3 d). The host gall farthest from the leaf proper was Callirhytis scitula Bassett, a woody gall that occurs at the intersection between the petiole and stem. The two Euceroptres species known from the Pacific coast, however, are both associated with stem galls (Buffington and Liljeblad 2008). All six hosts among our collections were classified as either medium or large, with no rearings from any galls smaller than 5 mm (Fig. 3 e). Co-occurrence with other natural enemies: Euceroptres were significantly more likely to co-occur with Ceroptres (P = 0.008) and with the parasitoid genus Euderus (P = 0.015) (Fig. 3 f). All six gall types from which we reared Euceroptres also had Ceroptres emerge and only one gall type from which we reared Euceroptres failed to yield Euderus (Andricus foliaformis Gillette). In contrast, Euceroptres were significantly negatively correlated with Synergus inquilines (P = 0.005) and in fact the two genera were never reared from the same gall types. Additional Notes: The small number of rearing records for Euceroptres, both in the present study and historically, precludes definitive statements about this genus and its association with oak gall types. On the other hand, we now have initial information about which gall types do, and more importantly, apparently do not host Euceroptres. All six gall hosts identified here, and the few additional previously published host records (Buffington and Liljeblad 2008), are multichambered, integral galls (i. e., non-detachable) and larger than 0.5 cm, suggesting that one or more of these characters may be important with respect to host range for Euceroptres. However, many other galls with these same characters do not appear to be Euceroptres hosts, so while these characters may be necessary, it is not clear that they are sufficient for Euceroptres attack and development. It may be that Euceroptres requires the presence of another gall associate either because it is a parasite of wasps in one of these genera or because it is an inquiline or gall inducer that first requires, e. g., Ceroptres to make a gall within the gall (which would make Euceroptres a galler of galls-within-galls). Though we generally did not key our collections to species, the small number of collections and the availability of the key to the four species produced by Buffington and Liljeblad (2008) allowed us to make an exception for Euceroptres. Not surprisingly, given the growing recognition of cryptic diversity and higher than previously-recognised host specificity among parasitic wasps (Forbes et al. 2009; Smith et al. 2011; Condon et al. 2014; Hood et al. 2015; Sheikh et al. 2022), none of the wasps we reared matched the descriptions of any of the four named Nearctic species. All samples had 10 flagellomeres and well developed micropores on their abdominal tergites, combinations not found in Buffington and Liljeblad (2008). None of the cynipid hosts producing the gall types from which we reared Euceroptres overlapped with previous rearing records, save for Euceroptres whartoni Buffington & Liljeblad in a) b) 1 2 3 4 8 7 5 6 11 12 10 9 14 13 15 16 17 18 19 20 21 22 23 24 25 26 28 27 29 c) 303233343531 Q 3 79 36 37403839 L 3 27 41 424447464543 V 0 10 48 49 50 52 53 51 d) 57 54 55 56 3 0 59 58 (66) (26) 60 61 62 63 64 66 65 67 0 0 68 69 (12) (4) 70 71 72 73 74 78 76 75 77 3 0 79 80 (10) (4) 81 82 83 e) f) galls of Andricus quercuspetiolicola (Bassett). However, Buffington and Liljeblad (2008) list this host record as dubious. Our collections to date may therefore include one or more undescribed species.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF90FFD49AEA29BA9BCAFBC8.taxon	description	Summary of Natural History: In comparison to other genera of parasitoids associated with oak galls, wasps in the genus Ormyrus have been described as generally more restricted to galls on oaks. For example, Torymus and Eurytoma both have oak gall associated species but attack many other hosts as well. All but two of the known Nearctic Ormyrus species attack galls on oaks-the exceptions attack Cynipid gall wasps on roses and Pteromalid gall inducers on blueberries (Hanson 1992). Though 16 species were described in the most recent revision of Nearctic Ormyrus (Hanson 1992), more recent genetic and ecological data suggest that these wasps are considerably more species rich and ecologically specialized than previously supposed (Sheikh et al. 2022).	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF90FFD49AEA29BA9BCAFBC8.taxon	description	Biogeography and oak tree section: Ormyrus were reared from galls collected in all three floristic regions of North America (Fig. 4 b) and from 40 % or more of gall types monitored in each of the three oak tree sections surveyed (Fig. 4 c). Tree organ and gall size: Ormyrus were associated with galls on all tree organs represented in our collections (Fig. 4 d) and from galls of all sizes, with a slight bias towards larger galls (Fig. 4 e). Previous rearing records also show that Ormyrus can use root galls produced by cynipids (Hanson 1992). Co-occurrence with other natural enemies: We found only one significant correlation between Ormyrus and another insect genus: a positive correlation with Eurytoma (Fig. 4 f; P = 0.0033). This could represent a shared affinity for galls with similar characteristics or a currently unknown trophic association between Ormyrus and Eurytoma wasps (i. e., one is a parasite of the other). Notes: As a genus, Ormyrus are ubiquitous in their phylogenetic, geographic, taxonomic, and ecological relationships with oak gall wasps. Though individual species are often apparently specialized on a small number of gall types (Sheikh et al. 2022), they may be able to use any of several insect species within a gall as their host, or act as inquilines, or some combination thereof. Understanding host breadths and life histories of individual Ormyrus species alongside their phylogeny will be critical to understanding how and under what circumstances oak-associated Ormyrus have diversified.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF90FFD1990F2C9A9EADFCC8.taxon	description	Summary of Natural History: Torymus wasps are primarily Holarctic in their distribution and there are more than 300 named species, though not all of them attack oak gall wasps (Grissell 1995). Hosts are almost always insects in a concealed location, but from diverse orders. In the Nearctic, two species groups contain all of the oak gall-associated species. The fullawayi species group consists primarily of species that attack insects in cynipid oak galls (Grissell 1976). Species in the tubicola group have a more diverse range of hosts, but several specialize on oak galls or on galls on other plant hosts (Grissell 1976). Most female Torymus have a long ovipositor (in some species, the ovipositor is more than the length of their body) that helps them attack hosts in concealed and otherwise protected locations. There is some evidence for apparent intraspecific variation in ovipositor length in some species, both between generations and within the same generation (Eady 1958; Askew 1965), which may result in a larger than expected host range for any given species. In the Palearctic, oak gall associated species range between one and 41 host records (Askew et al. 2013). The Nearctic species Torymus tubicola (Osten Sacken) is also described as a particularly generalist species, with more than 30 named oak gall hosts across the United States (Grissell 1976; Noyes 2022). However, as for Ormyrus and Synergus some of the apparently generalist Torymus species in both the Palearctic (Kaartinen et al. 2010) and Nearctic may harbor cryptic species. Relationship to galler phylogeny: Torymus wasps have been reared from gallers across the Nearctic oak gall wasp phylogeny (Ward et al. 2022), including from all four Palearctic lineages (Fig. 5 a). Like Ormyrus, the genus houses essentially ubiquitous parasitoids in the oak gall system. a) b) 1 2 3 4 7 8 5 6 11 12 10 9 13 14 15 16 17 18 19 20 21 22 23 24 25 26 29 28 27 c) 303233313534 Q 45 37 36 37403839 L 13 17 41 424644454347 V 48 49 50 53 51 52 d) 57 56 54 55 41 12 59 58 (66) (26) 60 61 62 63 67 64 65 66 3 3 68 69 (12) (4) 70 71 72 73 74 78 77 76 75 6 1 80 79 (10) (4) 81 82 83 e) f) g) Biogeography and oak tree section: We reared Torymus from galls in all three floristic regions (Fig. 5 b) and from all three oak sections (Fig. 5 c). Torymus are also known from galls on oaks in section Protobalanus in California (Grissell 1976). Tree organ and gall size: Torymus wasps were found in association with galls on all six organ types (Fig. 5 d). They emerged more often from large and medium sized galls than from small (> 5 mm) galls (Fig. 5 e). Though our collections were exclusive to aboveground galls, Torymus have also been reared from root galls (Forbes et al. 2016). Co-occurrence with other natural enemies: Torymus were reared from the same gall types producing Eurytoma wasps significantly more often than predicted (P = 0.0200). Conversely, Torymus were significantly less likely to be reared from the same gall types as Ceroptres wasps (Fig. 5 f; P = 0.0003). Previous work has shown that Torymus wasps can emerge from the same individual galls as other insects, though these other insects are usually either known or suspected inquilines (e. g., Synergus, Brasema), or suspected parasites of inquilines (e. g., Allorhogas Gahan) (Hall 2001). Additional Notes: The implication from the current Torymus taxonomic organization (Grissell 1976) and from molecular phylogeny of family Torymidae (Janšta et al. 2018) is that oak galls have been colonized two or more times, such that gall-associated Torymus are para- or polyphyletic. Any future phylogenetic assessment of Torymus coevolution with oak galls should therefore endeavor to include non-gall associated taxa.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF95FFD19A922B9A9A57FDE8.taxon	description	Summary of Natural History: Sycophila are known as parasitoids of endophytic insects, including gall wasps (Balduf 1932; Askew et al. 2006 2013; Gómez et al. 2013). Some species are known from just a single host (e. g., Sycophila marylandica (Girault) (Balduf 1932), while the Palearctic species Sycophila buguttata (Swederus) has 80 recorded hosts (Askew et al. 2013). Recent molecular work has shown considerable cryptic diversity and more limited host ranges among the Nearctic species, (Zhang et al. 2022). Relationship to galler phylogeny: Sycophila in our collections were broadly associated with almost all clades in the Nearctic gall wasp phylogeny (Ward et al. 2022), including all Palearctic clades (Fig. 6 a). Two clades from which no Sycophila were reared were a mixture of cluster galls on leaves and stems, early spring bud galls, and small unilocular leaf galls. Sycophila appear to be reared more often from large galls (Fig. 6 e; Hall 2001; Zhang et al. 2022) such that this apparent absence may reflect a general favoring of larger galls, but could also or instead be related to phenology or a bias in survivorship from smaller galls when using our rearing methods. Biogeography and oak tree section: Sycophila were reared from galls on oaks across all three floristic regions and in all three oak sections (Fig. 6 b c). Nine of 10 galls reared from live oaks (section Virentes) were host to Sycophila. Tree organ and gall size: We reared Sycophila from all surveyed host tree organs (Fig. 6 d). At least at this genus-level resolution, they were reared from a larger fraction (74 %) of large galls than from medium (59 %) or small (26 %) galls (Fig. 6 e). PCoA analyses of Sycophila from these and other collections also suggest that wasps in the genus generally favor medium and large galls (Zhang et al. 2022). Co-occurrence with other natural enemies: Sycophila did not co-occur significantly in a positive or negative direction with any of the other seven genera analyzed (Fig. 6 f).	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF95FFCF99372ABA9D8CFE2B.taxon	description	Summary of Natural History: Eurytoma in the Nearctic comprise a large (> 80 species) group of mostly parasitic wasps associated with a large diversity of insects across several orders (Bugbee 1967). At least 10 Nearctic species have been reared from oak galls, but their direct hosts are often uncertain, and indeed some Palearctic species are confirmed to attack nongalling inquilines (Redfern and Askew 1998). Some Eurytoma are also phytophagous, although no exclusive phytophages are known from oak galls (Bugbee 1967). Though some Eurytoma are apparently endoparasites (Redfern and Askew 1998), Eurytoma in other Cynipid galls, including oak galls in the Palearctic, are uniformly ectoparasites, with species in some galls feeding on the gall organ once the primary host insect has been exhausted (Gómez et al. 2011). Reported host ranges of oak gall-associated Eurytoma vary from a single species to more than 75 hosts (Bugbee 1967; Gómez et al. 2011; Askew et al. 2013). However, the diversity of the Nearctic fauna has yet to be interrogated genetically, and those in the Palearctic only marginally so. One particularly generalist-appearing species in the Palearctic, Eurytoma brunniventris Ratzeburg, has shown some evidence of genetic structure at the COI locus, with specimens reared from five species of oak gall wasps sorting genetically by tree host section (Ács et al. 2010). Until an integrative assessment of species limits can be performed for Nearctic Eurytoma, interpretation of their host ranges will likely remain limited. Relationship to galler phylogeny: Eurytoma were or have previously been reared from most oak galls represented in the Nearctic gall wasp phylogeny (Ward et al. 2022) (Fig. 7 a). If they appear sparse anywhere on the phylogeny, it is among the Neuroterus part of the tree (Fig. S 1; tips 4 – 17), many of which were collected as small flower or leaf galls. Biogeography and oak tree section: Eurytoma were reared from galls across all three floristic regions (Fig. 7 b) and from 48 – 64 % of gall types collected from each oak section (Fig. 7 c). Tree organ and gall size: Eurytoma were reared from galls on all oak organs studied (Fig. 7 d) though were more often reared from larger galls (84 %) than from small galls (29 %; Fig. 7 e). Co-occurrence with other natural enemies: Eurytoma were significantly more likely to be present when Ormyrus (P = 0.0033) or Torymus (P = 0.0200) wasps were also present (Fig. 7 f). This may indicate a) b) 1 2 3 4 6 5 7 8 12 11 10 9 13 14 15 16 17 18 19 20 21 22 23 24 25 26 29 28 27 c) 303431323533 Q 42 40 36 37383940 L 13 17 41 424447464543 V 48 49 50 52 51 53 d) 57 54 55 56 30 18 58 59 (66) (26) 60 61 62 63 66 67 65 64 6 2 69 68 (12) (4) 70 71 72 73 74 76 77 78 75 6 2 80 79 (10) (4) 81 82 83 overlap in the types of gall morphologies favored by each parasitoid genus. Additional Notes: Like Ormyrus and Torymus, Eurytoma are near-ubiquitous in their association with oak galls: we reared them from 64 (52.5 %) of the 122 gall types that had five or more insect specimens emerge (Table S 1). Also like these other genera, Eurytoma may be more species-rich than they currently appear, with each species more specialized on particular dimensions of gall environments (Zhang et al. 2014). A thorough integrative analysis of this group will be necessary to address questions about their evolution, ecology, and taxonomy.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF8AFFCB9AF1291A9F65FC08.taxon	description	Summary of Natural History: Euderus is a moderately large genus, with> 75 species described worldwide. Where hosts are known, they are usually pupae in concealed habitats (e. g., leaf mines, inside fruits and stems) (Yoshimoto 1971). More rarely, Euderus attack insects in galls, with two oak gall associated species — Euderus crawfordi Peck and Euderus set Egan, Weinersmith & Forbes-known from the Nearctic (Yoshimoto 1971; Egan et al. 2017). Euderus set has been specifically studied as a behavioral manipulator of its host gall wasps-wasps with E. set infections chew significantly smaller exit holes in their galls than those that are not infected and then do not leave the gall but instead plug the exit hole with their head. Euderus set then consumes the body of the host wasp and leaves the gall by chewing a hole through its host’s head (Weinersmith et al. 2017; Ward et al. 2019). Though this behavior has only been studied in detail for E. set, evidence of “ head plugs ” has been found in museum collections of Southwestern U. S. Bassettia Ashmead galls (Egan et al. 2017). Relationship to galler phylogeny: We reared or found records of Euderus in association with nine gall types in the Nearctic gall wasp phylogeny (Ward et al. 2022). Seven of these (indicated by “ s ” in Fig. 8 a) have been previously identified as E. set (Ward et al. 2019). Euderus crawfordi (“ c ” in Fig. 8 a) has been reared from the Palearctic species Plagiotrochus suberi Weld, but only from galls in its introduced range in the Nearctic suggesting this is a derived host association. Euderus crawfordi is also known from the Nearctic Kokkocynips coxii (Bassett), which is not on the Ward et al. (2022) tree, but we have indicated its approximate location near its congener, Kokkocynips imbricariae (Ashmead) (“ c ” with two asterisks). Our collections also produced what appears to be a new species of Euderus associated with galls of the sexual generation of Neuroterus washingtonensis on the Pacific coast (“ n ” in Fig. Xa). The morphology of these wasps did not match that of E. set, nor any species described in Yoshimoto (1971). Two other Euderus reared from galls in the southwestern U. S. were not examined morphologically and their hosts were not among those on the Ward et al. (2022) tree. Biogeography and oak tree section: Euderus set was reared from seven gall types, all in Eastern North America but across all three oak sections (Fig. 8 b c). Euderus were reared from two types of gall in the Mexican and Central American floristic region (both on section Quercus oaks), and from one gall type in the Californian floristic province (also section Quercus). Tree organ and gall size: Gall hosts were only on stems, leaves, or petioles (Fig. 8 d). Euderus attacked a greater proportion of large (> 2 cm) galls (0.26) than medium galls (0.06), and no Euderus were reared from galls smaller than 5 mm (Fig. 8 e). Co-occurrence with other natural enemies: Euderus were found to occur alongside Ceroptres (P = 0.015) and Euceroptres (P = 0.001) significantly more often than expected (Fig. 8 f). This co-occurrence is unlikely to be because Euderus are using wasps in either of these two genera as their primary hosts - the particular biology of Euderus set causes the host to be visible from outside of the gall and the gall inducer has been shown to be the host in cases where this has been investigated (Weinersmith et al. 2017; Ward et al. 2019). Other hypotheses for the significant co-occurrence are that wasps in these three genera tend to attack galls with similar features, and / or that they all tend to attack galls in a similar temporal window (Ward et al. 2019). Additional Notes: Because we are primarily working at the level of genus or above and patterns of specialization are more likely to manifest at the species level we have not sought to test hypotheses about insect specialization on gall morphology in this paper. However, since we have Euderus rearing records to the level of species, here we can compare features of their associated galls. Previous work has noted that Euderus set has only been reared from integral galls that lack external defensive structures, perhaps because Euderus appear to attack late-stage pupae and have relatively short ovipositors (Ward et al. 2019). This pattern of host use-medium to large, integral (non-detachable) swellings, often with> 1 cells-appears to apply to E. crawfordi and to the putative new Euderus species reared from Neuroterus washingtonensis as well (Fig. 8 a; Table S 1). Other Common Affiliates - Superfamily and Family level IDs Eulophidae (Hymenoptera: Chalcidoidea: Eulophidae) We reared 21,232 (mean = 259, range = 1 – 8,655) eulophid wasps from 82 gall types. These counts exclude wasps in subfamily Entiinae (Euderus), which were treated separately above. Previously documented Eulophid associates of oak galls include representatives of the Entedoninae (Chrysochari s Förster, Eprhopalotus Girault, Horismenus Walker, Pediobius Walker), Eulophinae (Aulogymnus Förster, Cirrospilus Westwood, Pnigalio Schrank, Sympiesis Förster), and Tetrastichinae (Aprostocetus Westwood, Baryscapus Förster, Galeopsomyia Girault, Minotetrastichus Kostjukov, Pentastichus Ashmead, Quadrastichus Gilrault, Tetrastichus Walker) (Askew et al. 2013; Noyes 2022). Relatively little is known about the ecology of eulophid gall associates, including their trophic roles, host ranges, and details of their life cycles. Representatives of many eulophid genera and subfamilies are known as obligatory or facultative hyperparasitoids (Schauff et al. 1997), and eulophid larvae in some oak galls can be gregarious (Redfern and a) b) 1 c * 2 3 s 4 s 7865 1210119 sn 13 14 15 16 17 18 19 20 21 22 23 24 s 25 26 29 28 27 c) 303132343335 c ** Q 5 77 36 37394038 L 3 27 41 424443454647 s V 48 49 s 50 s 5351 52 d) 54575655 3 2 5958 (66) (26) 60 61 62 63666465670 0 6869 (12) (4) 70 71 72 73 7478757776 4 0 8079 (10) (4) 81 82 83 e) f) g) Askew 1998). Several different eulophid species have previously been reared from the same gall type (e. g., Askew 1961; Eliason and Potter 2000; Bird et al. 2013), but they may or may not be attacking the same host (s) in these communities. Future integrative taxonomic work that includes host relationships will be invaluable in understanding this important group of gall associates. Eupelmidae (Hymenoptera: Chalcidoidea: Eupelmidae) We reared 10,620 (mean = 190; range = 1 – 3,802) eupelmid wasps from 56 gall types (Table S 1). The Eupelmidae most commonly associated with oak galls are genera Brasema or Eupelmus Dalman, and it is likely that all or most of our collections are from one of these genera (though we note that Anastatus gemmaii (Ashmead) was originally recorded from Callirhytis quercusgemmaria Ashmead (Burks 1967 )). Many Brasema have been mistakenly classified as Eupelmus, and sexual dimorphism in the family makes it challenging to separate males (Gibson 2011), such that we felt it was premature to split the two genera here.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF8AFFCB9AF1291A9F65FC08.taxon	description	We reared a combined 229 (mean = 11.5, range = 1 – 80) ichneumonid and braconid wasps from 20 gall types. All but one of the ichneumonoid collections were reared from medium-sized galls or larger (> 0.5 cm) on stems, leaves, or petioles (Table S 1). Though we did not key all of these wasps to genus, the majority of these collections were wasps in the genus Allohorgas (Braconidae: Doryctinae). Allohorgas has previously been reared from several North American galls (Eliason and Potter 2000; Forbes et al. 2016; Weinersmith et al. 2020; Joele et al. 2021). Braconid wasps from subfamily Cheloninae were reared from Disholcaspis quercusmamma (Walsh & Riley) and Callirhytis frequens (Gillette) galls. Ichneumonidae were reared from Neuroterus washingtonensis, Disholcaspis quercusglobulus (Fitch), Disholcaspis quercusmamma, and Neuroterus saltaorius (Edwards). The ecology of the oak gall-associated wasps in superfamily Ichneumonoidea is still generally unknown. Recent work on genus Allorhogas shows it to be particularly labile with respect to its feeding habits, with some species acting as parasites, others as phytophagous inquilines, still others as seed predators, and some as gall formers themselves (Moreira et al. 2017; Samacá-Sáenz et al. 2020 2022). In oak galls, Allorhogas and other ichneumonid wasps may be attacking moth or beetle inquilines: Joseph et al. (2011) reported Bassus nucicola Musebeck (Braconidae) and an unidentified ichneumonid wasp, both thought to be parasitizing Cydia latiferreana Walsingham in galls of Andricus quercuscalifornicus. Cheloninae are also primarily known as parasitoids of Lepidoptera (Wharton et al. 1997). Pteromalidae (Hymenoptera: Chalcidoidea: Pteromalidae) We reared 7,545 (mean = 148; range = 1 – 2,385) pteromalid wasps from 51 different gall types (Table S 1). Our pteromalid collections were reared from galls in all three floristic regions and from all three oak tree sections. Existing keys, which acknowledge their own difficulty and incompleteness (e. g., Bouček and Heydon 1997), were insufficient for separating many of these wasps to the genus level within this large and polyphyletic family, so they will be addressed elsewhere in the future through integrative taxonomic approaches. The Western Palearctic fauna of oak gall-associated Pteromalidae includes at least nine genera (Arthrolytus Thomson, Cecidostiba Thomson, Caenacis Förster, Cyrtoptyx Delucchi, Elatoides Nikol'skaya, Hobbya Delucchi, Kaleva Graham, Mesopolobus Westwood, and Ormocerus Walker) (Noyes 2022). At least three of these (Arthrolytus, Cecidiostiba, and Ormocerus) are also known from Nearctic oak galls. Additional Nearctic gall-associated pteromalids include Acaenacis Girault, Amphidocius Dzhanokmen, Anisopteromalus Ruschka, Guolina Heydon, Lariophagus Crawford, and Pteromalus Swederus. Where the ecology of oak gall-associated pteromalids has been studied, they are ectoparasites of a variety of gall inhabitants. Some attack the gall wasp itself, while others parasitize inquilines or other parasitoids (Askew 1961). Mesopolobus are the best studied of the Palearctic oak gall-associated pteromalids and feed on several different gall inhabitants, and at various life stages, including adults (Askew 1961). As many as five different species from this genus have been reared from the same gall type (Redfern and Askew 1998). Experiments on host searching suggest that short range searches for galls by Mesopolobus wasps may rely more on visual cues than on olfaction (Askew 1961), such that host tree and / or gall morphology may be axes of specialization to explore for these and other pteromalids. Occasional associates (Hymenoptera) Bethylidae (Hymenoptera: Chrysidoidea: Bethylidae) We reared 37 (mean = 5, range 1 – 16) bethylid wasps from seven gall types. A previous study found a wasp in bethylid genus Goniozus Förster in a gall of Belonocnema fossoria Weld (Forbes et al. 2016), and other Goniozus have been reared from moth inquilines in oak galls (Fouts 1942). The four gall types with the most bethylid wasps in our collections (Andricus quercuspetiolicola and three species of Belonocnema) also produced moths, indicating a possible trophic connection.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF8FFFCA9AF42BBB9A44FA28.taxon	description	We reared 2,385 (mean = 596, range 1 – 2,307) wasps in genus Bootanomyia Girault from four gall types, all leaf galls on Quercus garryana on the Pacific coast. While several species from this genus are common parasites of oak galls wasps in the Palearctic (Doğanlar 2012; Askew et al. 2013), only one informal record exists of the genus in North America (photos # 2020534 - 6 on BugGuide. org) and these are the first records we can find of Bootanomyia associated with oak gall wasps in the Nearctic. Though we did not key all of our collections to species, we keyed one to Bootanomyia dorsalis (Fabricius) using Doğanlar (2012). Molecular analysis of B. dorsalis in the western Palearctic found evidence for host-associated genetic differentiation among wasps reared from oak galls on trees in different oak subgenera (Nicholls et al. 2018). Whether our collections represent an introduced population of B. dorsalis, another (or several) host-associated populations, or a combination of these two remains to be seen. Ceraphronoidea (Hymenoptera) We reared 24 ceraphronoid wasps (mean = 8, range 2 – 11) from three different Belonocnema leaf galls. These were extremely uncommon, accounting for <0.15 % of all associates reared from either gall type. The biology of most ceraphronoids is generally poorly known (Johnson and Musetti 2004), but some have been reared from galls of cecidomyiid midges (Loiácono and Margaría 2002). Chalcididae (Hymenoptera: Chalcidoidea) We reared just two chalcidid wasps, one from each of two gall collections: Belonocnema fossoria and Neuroterus washingtonensis. Most Chalcididae are parasites of Lepidoptera or are sometimes hyperparasites of other Hymenoptera (though usually still in lepidopteran systems; Boucek and Halstead 1997). Our B. fossoria collections did have associated Lepidoptera. While we did not officially record lepidopterans from our N. washingtonensis collections, we observed larvae and moths emerging from these fleshy galls. Some chalcidids are known from Lepidoptera-induced galls (e. g., Prinsloo 1984), so these records may represent rare oviposition “ mistakes. ” Crabronidae or Pemphredonidae (Hymenoptera: Apoidea) We r e a r e d t h r e e a p o i d w a s p s f r o m g a l l s of Andricus wheeleri Beutenmüller collected in Arizona and two more from galls of Disholcaspis quercusmamma collected in Minnesota. Some Apoid wasps create larval cells in hollow spaces associated with plants (Ashmead 1894; Blommers 2008) and so these were likely occupying a gall emptied of its original inhabitants. Encyrtidae (Hymenoptera: Chalcidoidea) We reared 22 encyrtid wasps (mean = 3.6, range 1 – 16) from six gall types. Encyrtidae are usually known as endoparasitoids and often are hyperparasitic on other hymenoptera (Noyes et al. 1997). The rarity of these wasps among our collections leads us to believe that these were non-specific attacks on other gall associates. Eumeninae (Hymenoptera: Vespoidea: Vespidae) We reared nine mason wasps (mean = 3, range 1 – 5) from three gall types. All three galls (Disholcaspis quercusmamma, Andricus quercuscalifornicus, and Disholcaspis quercusglobulus) were medium to large stem galls that are often retained on oak branches even after gall wasps and other insects have exited. While we did not identify the mason wasps in A. quercuscalifornicus, those in the two Disholcaspis galls were Bramble mason wasps (Ancistrocerus adiabatus (de Saussure )). Ancistrocerus adiabatus create larval cells of mud in existing cavities and provision cells with moth caterpillars. Bramble mason wasps are known to use abandoned homes of other insects, including galleries of cerambycid beetles, nests of other wasps, and empty galls (Gosling 1978; Holm 2021). Joseph et al. (2011) previously recorded an unidentified species of vespid wasp in galls of Andricus quercuscalifornicus, and suggested it may be acting as a facultative predator. Formicidae (Hymenoptera: Formicoidea) We found 70 ants (mean = 14, range 1 – 28) in association with five of our collections: Tapinoma Foerster in Amphibolips confluenta (Harris) galls, Camponotus Mayr (a queen), Temnothorax Mayr, and Tetramorium Mayr in Callirhytis quercuscornigera (Osten-Sacken) galls, Camponotus (workers and a queen), Temnothorax (workers and a queen), and Crematogaster Lund in Disholcaspis quercusmamma galls, Temnothorax on Andricus quercusstrobilanus (Osten-Sacken), Brachymyrmex patagonicus Mayr and Crematogaster ashmeadi Mayr in Bassettia pallida Ashmead (previously reported in Weinersmith et al. 2020). All ants recovered here are almost certainly colonists of older galls that had already been hollowed out by their original inhabitants. Previous studies have reported other ant species in other galls, e. g., Camponotus nearcticus Emery and Lasius alienus (Foerster) in galls of C. quercuscornigera (Eliason and Potter 2000), and seven different ant genera in galls of Disholcaspis cinerosa (Bassett) (Wheeler and Longino 1988). Besides living inside oak galls, some ant species are known to act as mutualists, feeding on honeydew produced by some gall types while actively defending those galls from parasites and predators (Washburn 1984; Fernandes et al. 1999). We have observed ants tending Disholcaspis eldoradensis (Beutenmuller) galls (which produces honeydew) in the Pacific northwest (KMP, pers. obs.) and Crematogaster ants both living inside and tending galls of D. cinerosa in the southeastern U. S. (SPE, pers. obs.). Platygastridae (Hymenoptera: Platygastroidea) We reared 913 platygastrids (mean = 91.3, range 1 – 867) from 10 gall types. The vast majority (867) of these were reared from Neuroterus washingtonensis galls on the Pacific coast but other sporadic collections came from Eastern galls. Weinersmith et al. (2020) previously reported three platygastrid genera (Telenomus Haliday, Calotelea Westwood, and Synopeas Förster) all from the same gall type. Platygastridae are usually egg parasites and Synopeas are known to parasitize cecidomyiid midges (Abram et al. 2012). Midges were reared from six of these 10 gall types (Table S 1), such that these rearing records might not reflect direct associations with the gall wasp or its parasites. Trichogrammatidae (Hymenoptera: Chalcidoidea) We reared 64 trichogrammatid wasps (mean = 21.3, range 1 – 62) from three gall types (62 from Neuroterus vesicula (Bassett), one each from two other gall types). Trichogrammatidae are parasites of insect eggs (Pinto 1997). A preliminary identification of these wasps keyed them to genus Poropoea Förster, known for parasitizing eggs of leaf-rolling weevils. It seems likely that these represent accidental collections of insect eggs and not formal gall associates. Occasional associates – other insect orders and non-insect arthropods.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF8EFFC999772DDB9F61FD8B.taxon	description	Cecidomyiinae (Diptera: Cecidomyiidae) We reared 169 cecidomyiid midges (mean = 7.3, range 1 – 70) from 23 gall types. Though one possibility is that some small number of cecidomyiid galls were collected alongside oak galls, Redfern and Askew (1998) suggest that cecidoymiids may sometimes be part of the successional fauna of cynipid oak galls, using the gall tissue after the original inhabitants have emerged. Previous records of cecidomyiids reared in cynipid oak gall studies include Lasioptera Meigen and Lestodiplosis Keiffer from C. quercuscornigera (Eliason and Potter 2000) and unidentified species reared from B. kinseyi and B. pallida (Forbes et al. 2016; Weinersmith et al. 2020).	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF8DFFC99AF42B5B9FCAFAA8.taxon	description	We reared 12 pachygastrinids from the same galls as the Lonchaea flies above. These specimens keyed to genus Eidalimus Kertesz (McAlpine et al. 1981). Aphididae (Hemiptera: Aphidoidea) We found seven aphids (mean = 2.3, range 1 – 3) in association with three gall types. Two were on leaf galls, the other on a stem gall. We did not key these beyond family. We assume these aphids were feeding on or next to galls and that their collection was coincidental to the presence of the gall.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF8DFFC99AF42A3A9CB9FCA8.taxon	description	We reared 30 Lonchaeinae (lance flies) from galls of C. quercuscornigera. These specimens keyed to genus Lonchaea Fallén (McAlpine et al. 1987). Lance flies are often associated with fallen trees and burrows of bark beetles (McAlpine et al. 1987; Marshall 2012).	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
1E1F87DDFF8DFFC89AF42D5B9A43FD88.taxon	description	We found four-minute pirate bugs (genus Orius Wolff) - one adult and three nymphs - across three gall types. Orius can be both predaceous and herbivorous. We suspect this was another non-specific gall association. Psyllidae (Hemiptera: Psylloidea) We found 251 psyllids associated with two galls: 250 from Callirhytis quercuspunctata (Bassett) and one from Andricus incertus Bassett. All 250 collected from C. quercuspunctata were collected from the same city (St. Louis, MO), and 249 of these were from the same collection. Psyllids can be gall formers themselves, but here are likely sap feeders and their collection alongside galls may have been due to generalist feeding or entirely coincidental. Rhyparochromidae (Hemiptera: Lygaeoidea) We found one rhyparochromid (dirt-colored seed bugs) in a collection of Disholcaspis mellifica Weld gall from California. These are seed-feeders, and this collection was likely coincidental. Lepidoptera We reared 355 moths (mean = 19.7; range 1 – 140) from 18 different gall types. We did not key all moths to family and some of the larger galls collected by the KP / DJ team (Table S 1; lettered lab codes) were observed to have moths emerge but these insects were not tallied. Moths reared from other galls have often been confirmed as (Joseph et al. 2011) or assumed to be (Forbes et al. 2016) inquilines feeding on gall tissue. Seventeen of the 18 gall hosts of Lepidoptera in our collections were large or medium galls, and most with thick outer walls which would provide ample food for a larger inquiline species. Moths in the families Gelechiidae, Pyralidae, Sesiidae, and Tortricidae have been previously reared from Nearctic galls (Eliason and Potter 2000; Joseph et al. 2011; Forbes et al. 2016). Most moths in our collections were small moths (10 – 20 mm wingspan), but we also reared the clearwing moths Synanthedon scitula Harris and Synanthedon decepiens Edwards from some Callirhytis galls. These larger (up to 30 mm) moths mimic vespid wasps and bees and their larvae feed on wood of living trees. Lacewings (Neuroptera: Chrysopoidea: Chrysopidae) Eleven lacewings (mean = 1.4, range 1 – 3) were found in association with eight gall types. Of these, ten were nymphs and one was an adult. The adult was associated with a collection of Disholcaspis quercusmamma bullet galls and may have emerged from a pupsae inside a hollowed-out gall, or may have just been sheltering inside a gall as an adult. Lacewing larvae and adults are generalist predators and may have been coincidental collections. However, on live oaks (section Virentes), lacewings have been observed to lay eggs on the asexual fuzzy leaf gall induced by Andricus quercuslanigera (SPE, pers. obs). It is not known whether this is coincident to the presence of the gall or if proximity to the gall increases lacewing survivorship in some way. Psocomorpha (Psocodea) We reared 318 barklice (mean: 18.7, range 1 – 156) from 17 gall types. Barklice in gall systems have previously been regarded as late-stage inquilines or successional associates (Joseph et al. 2011) and have been observed to enter empty galls through the exit holes of other gall inhabitants (Weinersmith et al. 2020). Thysanoptera We found 62 thrips (mean = 2.2, range 1 – 10) in association with 28 gall types. We did not key these specimens beyond the level of order. Though some thrips can be gall inducers or kleptoparasites of other thrips-induced galls (Crespi and Abbot 1999), our thrips collections may have been occupying old hollow galls (e. g., Redfern and Askew 1998). They also may not have been specifically associating with galls but rather collected accidentally while feeding externally on oak tissue. Acari (Arachnida) We found 135 mites (mean = 7.1, range 1 – 50) in association with 19 gall types. Eliason and Potter (2000) documented mites in families Oribatidae, Phytoseiidae, and Acaridae on the surface of C. quercuscornigera galls or sheltering in crevices. Histiogaster robustus Woodring (Acaridae) was also found to be apparently phoretic on C. quercuscornigera and its Synergus inquilines (Eliason and Potter 2000). In our collections, we also found mites were attached to other gall associates, especially ants and beetles. Araneae (Arachnida) We found 41 spiders (mean = 5.9, range 1 – 18) in association with collections of seven different gall types. All galls were 5 mm or larger. Spiders were likely collected as transients on external gall material, though conceivably may have been inside empty gall cavities of woody stem galls such as An. quercuscalifornicus, D. quercusglobulus, and D. quercusmammma. Eliason and Potter (2000) previously recorded spiders in the families Araneidae, Linyphiidae, Philodromidae, Salticidae, and Theridiidae on galls of C. quercuscornigera, and also observed spiders eating gall wasps that had emerged from galls. Pseudoscorpiones (Arachnida) A single pseudoscorpion was found in association with a Callirhytis quercusbatatoides gall collected in Florida. Redfern and Askew (1998) mention pseudoscorpions as successional species in galls. Chilopoda We found one centipede in a gall of Amphibolips quercusinanis. Like the Forficula earwigs associated with the same galls, this animal was likely using the gall as a transient shelter.	en	Ward, Anna K. G., Busbee, Robert W., Chen, Rachel A., Davis, Charles K., Driscoe, Amanda L., Egan, Scott P., Goldberg, Bailey A. R., Hood, Glen Ray, Jones, Dylan G., Kranz, Adam J., Meadely-Dunphy, Shannon A., Milks, Alyson K., Ott, James R., Prior, Kirsten M., Sheikh, Sofia I., Shzu, Shih-An, Weinersmith, Kelly L., Zhang, Linyi, Zhang, Y. Miles, Forbes, Andrew A. (2022): The Arthropod Associates of 155 North American Cynipid Oak Galls. Zoological Studies 61 (57): 1-30, DOI: 10.6620/ZS.2022.61-57, URL: http://dx.doi.org/10.5281/zenodo.8055962
