Birds from the Far East in Central Europe: a test of the reverse migration hypothesis

Every year birdwatchers throughout Europe record the arrival of a considerable number of vagrants from the eastern Palaearctic. We analysed the occurrence of such vagrants in Central Europe of the genera Phylloscopus, Turdus and Zoothera in Central Europe. Our results revealed that the occurrence of a species depended on total population size, but not on body size. Furthermore, the occurrence of species in Central Europe increased with distance between the breeding range and wintering range, but the distance between the breeding range and Europe had no effect. These results indicate that the migratory restlessness of species determines whether an East Asian species arrives in Europe. Overall, our data support the hypothesis that the migratory program determines vagrancy.     

Energy and distribution in subterranean rodents: sympatry between two species of the genus Ctenomys

The low basal metabolic rate (BMR) observed in subterranean rodents, compared to that of surface-dwelling species of comparable size, has been proposed to be an adaptation to underground life. Two main hypotheses have been proposed to explain this finding, the cost of burrowing and the thermal stress. The former states that the low BMR is due to the high cost of extending the tunnel system whereas the other relates it to the possibility of overheating in burrows where evaporative and convective heat exchange are restricted. Additionally, both hypotheses related the energetics of subterranean rodent with spatial distribution. The genus Ctenomys is an excellent model to evaluate the cost of burrowing or thermal stress, since they are widely distributed, with members differing markedly in body mass. The aim of this study was to assess digging and basal energetics in two Ctenomys species that live in sympatry in a coastal grassland, but differ in their microspatial distribution by soil preference. We used the obtained energetic data to test both energy-distribution hypotheses. We measured BMR and digging metabolic rate (DMR) through open flow respirometry in two species exposed to soft and hard soils. In brief, DMR in Ctenomys talarum (100-170 g), as in Ctenomys australis (250-600 g), was unaffected by soil hardness. Within thermoneutral zone of each species, DMR/RMR quotient was lower in the smaller species. Our data did not support the thermal stress hypothesis, but the cost of burrowing hypothesis was not rejected. Other alternative hypotheses are proposed to explain the distribution of C. talarum and C. australis.     

Examination of the immune responses of males and workers of the leaf-cutting ant Acromyrmex echinatior and the effect of infection

Summary. Parasites represent significant challenges to social insects. The high density, interaction rate and relatedness of individuals within colonies are all predicted to make social insect colonies particularly vulnerable to parasites. To cope with this pressure, social insects have evolved a number of defence mechanisms. These include the immune response, which, aside from in bumblebees, has been relatively little studied in social insects. Here we compare the immune responses of males and workers of the leaf-cutting ant Acromyrmex echinatior and examine the effect upon immunocompetence of prior exposure to a virulent parasite. Males have a far lower immune response than workers, suggesting either haploid susceptibility or reduced investment in immunity by the short-lived males. There was also significantly less variation in the immune response of males than of workers, which may be due to leaf-cutting ant workers being more variable in age or more genetically diverse within colonies. When exposed to the entomopathogenic fungus Metarhizium, workers expressed a substantially reduced immune response 96 h after infection, suggesting that the immune system was either depleted by having to respond to the Metarhizium infection or was depressed by the parasite. The results suggest that the immune response is a costly and limited process, but further experiments are needed to distinguish between the alternative explanations for the effects observed.Received 3 August 2004; revised 3 February 2005; accepted 2 March 2005.     

Invasive plants and their escape from root herbivory: a worldwide comparison of the root-feeding nematode communities of the dune grass <Emphasis Type="Italic">Ammophila arenaria</Emphasis> in natural and introduced ranges

Invasive plants generally have fewer aboveground pathogens and viruses in their introduced range than in their natural range, and they also have fewer pathogens than do similar plant species native to the introduced range. However, although plant abundance is strongly controlled by root herbivores and soil pathogens, there is very little knowledge on how invasive plants escape from belowground enemies. We therefore investigated if the general pattern for aboveground pathogens also applies to root-feeding nematodes and used the natural foredune grass Ammophila arenariaas a model. In the late 1800s, the European A. arenariawas introduced into southeast Australia (Tasmania), New Zealand, South Africa, and the west coast of the USA to be used for sand stabilization. In most of these regions, it has become a threat to native vegetation, because its excessive capacity to stabilize wind-blown sand has changed the geomorphology of coastal dunes. In stable dunes of most introduced regions, A. arenaria is more abundant and persists longer than in stabilized dunes of the natural range. We collected soil and root samples and used additional literature data to quantify the taxon richness of root-feeding nematodes on A.␣arenaria in its natural range and collected samples from the four major regions where it has been introduced. In most introduced regions A. arenaria did not have fewer root-feeding nematode taxa than the average number in its natural range, and native plant species did not have more nematode taxa than the introduced species. However, in the introduced range native plants had more feeding-specialist nematode taxa than A. arenaria and major feeding specialists (the sedentary endoparasitic cyst and root knot nematodes) were not found on A. arenaria in the southern hemisphere. We conclude that invasiveness of A. arenaria correlates with escape from feeding specialist nematodes, so that the pattern of escape from root-feeding nematodes is more alike escape from aboveground insect herbivores than escape from aboveground pathogens and viruses. In the natural range of A. arenaria, the number of specialist-feeding nematode taxa declines towards the margins. Growth experiments are needed to determine the relationship between nematode taxon diversity, abundance, and invasiveness of A. arenaria.     

Distribution and ecology of Demospongiae from the circalittoral of the islands of the Aegean Sea (Eastern Mediterranean)

Size-selective predation by fish is often considered to be a primary driver of seasonal declines in large-bodied Daphnia populations. However, large Daphnia commonly exhibit midsummer extinctions in ponds lacking planktivorous fish. A number of empirical and theoretical studies suggest that resource competition and its interaction with nutrient enrichment may determine variable dominance by large Daphnia. Low resource levels may favor competitive dominance by small-bodied taxa while large Daphnia may be favored under high resource conditions or following a nutrient/productivity pulse. Nutrient enrichment may also influence the strength of invertebrate predation on Daphnia by affecting how long vulnerable juveniles are exposed to predation. We investigated these hypotheses using an in situ mesocosm experiment in a permanent fishless pond that exhibited seasonal losses of Daphnia pulex. To explore the effects of nutrient enrichment, Daphnia plus a diverse assemblage of small-bodied zooplankton were exposed to three levels of enrichment (low, medium, and high). To explore the interaction between nutrient enrichment and invertebrate predation, we crossed the presence/absence of Notonecta undulata with low and high nutrient manipulations. We found no evidence of competitive reversals or shifts in dominance among nutrient levels, Daphnia performed poorly regardless of enrichment. This may have been due to shifts in algal composition to dominance by large filamentous green algae. Notonecta had significant negative effects on Daphnia alone, but no interaction with nutrient enrichment was detected. These results suggest that Daphnia are not invariably superior resource competitors compared to small taxa. Though predators can have negative effects, their presence is not necessary to explain poor Daphnia performance. Rather, abiotic conditions and/or resource-based effects are probably of greater importance.     

Univariate and multivariate analysis on phenotypic divergence in Phleum

Summary An analysis of phenotypic and genetic variability was performed on Phleum ambiguum populations obtained by vegetative propagation and grown in different environments. The investigation on a single character and the canonical analysis on several plant traits indicate that, while genetic variability (h²) is essentially constant, phenotypic variance and covariance are affected by different environments and successive clonations. The observed changes seem not to be reversible when plants, transferred to a different environment, are returned to the original one. Within-clones variability seems to be affected by environmental conditions without being directly related to them. The results reported seem to be in good agreement with the phenotypic divergence hypothesis.     

Size-assortative pairing in the lotic amphipod <Emphasis Type="Italic">Gammarus zaddachi</Emphasis>, an examination of hypotheses and the influence of current speed

Three main hypotheses have been put forward to explain size-assortative pairing in gammarid amphipods: microhabitat separation, sexual selection and loading constraint. In order to determine which hypothesis best explains this phenomenon in the estuarine species Gammarus zaddachi, I first measured the body lengths and dry weights of precopula pairs collected from two field sites with substantially different current speeds. Second, I performed three laboratory experiments in order to estimate the importance of the following processes: (1) male choice; (2) male–male competition and (3) male–female acceptability. The loading constraint hypothesis seemed best supported by the data in that field-collected male G. zaddachi size correlated well with female size in precopula pairing in both fast and slow flowing water. In the laboratory, males preferred females of their same size group (large versus small), and ‘won’ them in the male–male competition experiments. Size-assortative pairing is thus likely a consequence of the loading constraints imposed upon these males by virtue of them having to carry and manoeuvre their partners through flowing water, while attempting to maintain station in an optimal microhabitat. Males may therefore forego the largest, most fecund females, in favour of a practicable payload (small male–large female pairings were rare). However, there seems to be a lower limit to this selection, indicated by the high degree of cannibalism on small females by large males.     

Impact of fish predation on coexisting Daphnia taxa: a partial test of the temporal hybrid superiority hypothesis

Fish predation was tested as a factor mediating the coexistence of Daphnia taxa in the shallow, hypertrophic Lake Blankaart. Naturally co-occurring populations of D. galeata and the hybrid D. galeata x cucullata were subjected to different levels of fish predation in in situ enclosures. In control enclosures without fish, the largest taxon D. galeata rapidly became dominant over the intermediate sized D. galeata x cucullata, mainly as a result of higher birth rates. In enclosures with fish, population densities of D. galeata dropped relative to D. galeata x cucullata, due to higher mortality rates. These results are in concordance with the `temporal hybrid superiority hypothesis', and can be explained by a higher vulnerability of the large and more conspicuous D. galeata to the size selective predation exerted by visually hunting planktivorous fishes. After approximately one month, however, population growth rates of D. galeata and D. galeata x cucullata in the enclosures with fish converged, due to a relative reduction in the mortality rate of D. galeata. This suggests that, in the presence of fish, D. galeata may co-exist with hybrids due to a decrease in its relative vulnerability to visual predation with time. Indeed, both D. galeata and the hybrid showed strong reductions in adult body size in the enclosures with fish, but this size reduction tended to be stronger in D. galeata than in D. galeata x cucullata. In addition, turbidity increased in the enclosures with fish and may additionally have reduced the relative advantage of D. galeata x cucullata with regard to mortality caused by visual predation.     

Australopithecus afarensis and the single species hypothesis

Ferguson (1989) has recently argued that the variability seen in the fossils assigned toA. afarensis is far more than expected for a single hominid species, and therefore proposes they represent multiple taxa. In particular, he utilizes data on variation in dental metrics and in premolar morphology in support of this hypothesis. A re-evaluation of these data finds the above conclusion to be unwarranted. Variation in dental metrics providesno basis for separating this sample into multiple taxa, regardless of the analog that is used (i.e. modern primate species or fossil hominid species). Additionally, data on P₃ morphology indicate that thepattern of variation seen in the Laetoli/Hadar sample is comparable to the sexual variation seenwithin a single hominoid species. Overall, the balance of the evidence at present indicates that the fossils from Laetoli and Hadar represent a single hominid species,A. afarensis.     

Ectoparasitism and the role of green nesting material in the European starling

Summary The use of green nesting material is widespred among birds. Recent evidence suggests that birds use secondary chemicals contained in green plants to control ectoparasites. We manipulated green nesting material and ectoparasites of European starlings (Sturnus vulgaris) to test two hypotheses: (1) ectoparasites adversely affect prefledging survival and morphometrics or postfledging survival, and (2) green nesting material ameliorates the effects of ectoparasites. We recorded fat score, numbers of scabs, tarsal length, body mass, and hematocrit level on each nestling 17 days after hatching. We also fitted each nestling with unique patagial tags and resighted the starlings for 6–8 weeks after fledging to estimate survival and sighting rates. Nests devoid of green nesting material and dusted with the insecticide, carbaryl, had fewer high ectoparasite infestations, and nestlings had significantly lower scab scores, and significantly higher body masses than nestlings in undusted boxes. However, there was no difference in postfledging survival between birds from carbaryl-treated and undusted nests. There also was no difference in prefledging survival and morphometrics or postfledging survival between nestlings from boxes with and without green nesting material. These results do not support the hypothesis that starlings use green nesting material to control nest ectoparasites. We suggest an alternative hypothesis; green nesting material is used for mate selection or pairbonding in the starling.     

施用甲醇刺激矮牵牛的生长效应与其代谢关系初探

大量研究表明施用甲醇能够促进多种植物的生长,在甲醇刺激植物生长的机理中,支持碳源假说的证据最多。该研究通过考察矮牵牛甲醇代谢与甲醇刺激其生长的相关性,对碳源假说进行验证。结果表明:(1)在MS固体培养基上添加2和6mmol/L CH3OH均可促进矮牵牛的生长和叶绿素含量增加,但2mmol/L CH3OH(低浓度)效果好于6mmol/L(高浓度),而且添加6mmol/L CH3OH会诱发较强的氧化胁迫。(2)进一步用13 C-NMR分析矮牵牛对不同浓度13 CH3OH的代谢作用发现,6mmol/L 13 CH3OH处理矮牵牛中[U-13 C]Fruc和[U-13 C]Gluc的生成量显著大于2mmol/L 13 CH3OH处理,即来自甲醇的碳源在代谢过程中虽被卡尔文循环同化为糖类物质,但这部分碳源对甲醇刺激矮牵牛的生长贡献不大。这些证据表明CH3OH代谢与其刺激矮牵牛生长的效果没有关联性,该实验结果不支持碳源假说。     

Spread of plant pathogens and insect vectors at the northern range margin of cypress in Italy

The Mediterranean cypress (Cupressus sempervirens) is a multi-purpose tree widely used in the Mediterranean region. An anthropogenic range expansion of cypress has taken place at the northern margin of the range in Italy in recent decades, driven by ornamental planting in spite of climatic constraints imposed by low winter temperature. The expansion has created new habitats for pathogens and pests, which strongly limit tree survival in the historical (core) part of the range. Based on the enemy release hypothesis, we predicted that damage should be lower in the expansion area. By comparing tree and seed cone damage by pathogens and pests in core and expansion areas of Trentino, a district in the southern Alps, we showed that tree damage was significantly higher in the core area. Seed cones of C. sempervirens are intensively colonized by an aggressive and specific pathogen (the canker fungus Seiridium cardinale, Coelomycetes), associated with seed insect vectors Megastigmus wachtli (Hymenoptera Torymidae) and Orsillus maculatus (Heteroptera Lygaeidae). In contrast, we observed lower tree damage in the expansion area, where a non-aggressive fungus (Pestalotiopsis funerea, Coelomycetes) was more frequently associated with the same insect vectors. Our results indicate that both insect species have a great potential to reach the range margin, representing a continuous threat of the arrival of fungal pathogens to trees planted at extreme sites. Global warming may accelerate this process since both insects and fungi profit from increased temperature. In the future, cypress planted at the range margin may then face similar pest and pathogen threats as in the historical range.     

Scale-dependent mechanisms in the population dynamics of an insect herbivore

A multiscale approach has lead to significant advances in the understanding of species population dynamics. The scale-dependent nature of population processes has been particularly clearly illustrated for insect herbivores. However, one of the most well-studied insect herbivores, the galling sawfly Euura lasiolepis, has to date been examined almost exclusively at fine spatial scales. The preference-performance, plant vigour and larval survival hypotheses are well supported by this species. Here, we test these hypotheses at a spatial scale larger than that previously considered, i.e. across a landscape in northern Arizona represented by an altitudinal gradient encompassing a series of drainages. We also develop a qualitative model for understanding the population dynamics of E. lasiolepis based on patterns of survival and mortality found in this study and previous ones. Gall density was highly variable across the altitudinal gradient, not explained by host plant variables, and thus a poor surrogate fot population abundance. These findings for the first time fail to support the plant vigour and preference hierarchy hypotheses for E. lasiolepis. Dispersal limitation most likely explains the lack of support for these hypotheses at this scale. By contrast, sawfly survival, gall abortion, parasitism and larval mortality were well explained by host plant quality variables and altitude. The larval survival hypothesis was well supported and is thus comparatively scale-invariant. A qualitative model developed here highlighted the importance of both willow water status and disturbance in determining host plant quality, as well as an apparent trade off between shoot length and plant moisture status in determining vital rates across the altitudinal gradient. This study thus demonstrated for the first time the scale-dependent nature of mechanisms underlying the population dynamics E. lasiolepis, and identified the interaction between parasitism and altitude as a novel mechanism underlying spatial patterns in the survival and mortality patterns of this species.     

Embryogenesis inDrosophila: Can a single mechanism explain the developmental segregation of germ-line and somatic cells and their subsequent determination?

Summary A hypothesis is presented which explains the segregation of germ cells from somatic cells, and the subsequent determination of both cell types with a single mechanism. This hypothesis is in part based on that of Meinhardt (1977) and can be summarized as follows: In the newly fertilized egg, the action of a sink in the pole plasm leads to the formation of an anterior-posterior gradient of an inhibitor. The concentration of this inhibitor in the posterior 20% of the egg is below that needed to repress synthesis of an activator. When, during the nuclear division stage, nuclei enter this posterior region, synthesis of the activator begins. As the activator is autocatalytic, this leads to the formation of a peak of activator in this region; and since the activator also catalyses the synthesis of the inhibitor, a peak of inhibitor is formed in the same place. The inhibitor then diffuses anteriorly through the periplasm, forming a posterior-anterior gradient. The presence of this inhibitor in the periplasm causes the nuclei that enter the periplasm to form blastoderm cells and to take up particular segmental states appropriate to their position, while those that remain in the yolk-containing plasm develop into vitellophages. The action of the sink in the pole plasm is postulated to result in the formation of the pole cells, and subsequently to direct some of these into forming cells of the germ-line.Dr. Ilan Deak had submitted this paper shortly before his untimely death. The editors decided to publish his thoughts as expressed by him. However, with the consent of his friends and collaborators, some paragraphs reviewing earlier data were cut out, and deviating opinions were added as footnotes     

温度对中华倒刺鲃不同生理生态性能的影响:驯化有益假说的验证

驯化有益假说(Beneficial acclimation hypothesis)认为生物表型的适应性变化会增强其在诱导这些变化产生的环境中的生理机能或适合度。然而,由于动物不同生理生态性能对环境驯化的响应可能不一致,那么,测试表型性状的选择对驯化有益假说的验证就尤为关键。为此,整合表征动物生存适合度的不同生理生态性能并探究其对环境驯化的响应模式就十分必要。以我国长江中上游广泛分布的中华倒刺鲃(Spinibarbus sinensis)为对象,考察了驯化温度(18℃、28℃)和测试温度(18℃、28℃)及其交互作用对该物种有氧运动能力和无氧运动能力的影响,为驯化有益假说等相关假说的验证提供参考。研究发现,中华倒刺鲃不同生理生态性能对温度驯化的响应存在差异:(1)驯化温度对表征中华倒刺鲃无氧运动能力的快速启动游泳无显著影响(除最大加速度外)(P>0.05),研究数据倾向于支持无益假说(No-advantage hypothesis);(2)驯化温度对表征中华倒刺鲃有氧运动能力的临界游泳速度(Critical swimming speed,Ucrit)和最大代谢率(Maximum metabolic rate, MMR)影响显著(P<0.05),18℃驯化-18℃测试下的Ucrit和MMR均优于28℃驯化-18℃测试下的Ucrit和MMR,结果部分支持驯化有益假说和冷有益假说(Cooler is better hypothesis);(3)驯化温度、测试温度、游泳速度对中华倒刺鲃的运动代谢率(Active metabolic rate,MO2)和单位距离能量消耗(The energetic cost of transport, COT)影响显著(P<0.05)。值得关注的是,当游泳速度小于30 cm/s时,驯化温度对MO2和COT无影响,结果支持无益假说;而当游泳速度大于30 cm/s时,在特定的流速下经过28℃驯化的中华倒刺鲃无论在28℃还是18℃的测试环境下MO2和COT均较低,结果倾向于支持热有益假说(Warmer is better hypothesis)。研究结果提示:驯化有益假说并不具有普遍性,热驯化相关假说的验证不仅受表型性状选择的影响,而且还与测试的环境选择压力有关。     

Copulatory wounds in the monandrous ant species <Emphasis Type="Italic">Formica japonica</Emphasis> (Hymenoptera,Formicidae)

Males of several insect species inflict wounds on female genitalia during copulation, but the significance of such copulatory wounds for males is not clear. I compared the genitalia of virgin and mated Formica japonica females and for the first time report the occurrence of copulatory wounds in this monandrous ant species. All inseminated females examined had two types of melanized patches, indicating wound repair, and the serrated penis valves and sharp-pointed volsellar digitus of male genitalia were the likely instruments of these wounds. Physically damaging mating in monandrous species supports the view that copulatory wounds do not necessarily contribute to postcopulatory fitness gains for males via advantages in sperm competition or cryptic female choice. Received 10 September 2007; revised 15 October 2007; accepted 16 October 2007.     

小型哺乳动物种群周期性波动的外因调节假说

对小型哺乳动物种群数量周期性波动的外因调节假说进行介绍 ,概述了食物假说、捕食假说和复合因子假说的主要内容和研究进展。在少数生存环境严酷的小型哺乳动物种群中 ,食物假说能解释它们的周期性数量波动现象 ,可能作为调节因子起作用 ,但难以说明低数量期的确切机制 ,对于大多数小型哺乳动物而言 ,它更可能作为限制因子。捕食假说解释了北欧芬诺斯坎底亚地区某些种群的周期性波动 ,尤其是捕食的间接效应已引起许多学者的关注 ,但也有不支持该假说的证据。对于复合因子假说 ,近年颇受学者重视 ,其中验证食物和捕食交互及累加作用的实验证据较多 ,有的研究还包括气候、种间竞争、空间或社会行为等因素。有关复合因子的实验研究 ,尽管工作是困难和艰巨的 ,花费也是巨大的 ,但所得结果却极有价值 ,为深入理解种群动态调节理论提供了一个合理而有效的手段     

Juvenile hormone and the ontogeny of cockroach aggression

Our previous study [Kou et al., 2008. Juvenile hormone levels are increased in winners of cockroach fights. Horm. Behav. 52, 252–260] showed that the basic principle of the challenge hypothesis (hormone levels can respond to social stimuli to modulate aggression in vertebrates] could be applied to juvenile hormone (JH) levels and aggression in the lobster cockroach Nauphoeta cinerea. In that study, 80- to 85-day-old socially naïve males were used, as fighting is much more easily initiated in these older animals than in younger males, and JH III levels in the dominant were found to be significantly increased after an encounter compared to before the encounter and were significantly higher than those in the subordinates. In N. cinerea, newly emerged males usually show no aggressiveness towards each other and aggression is only initiated after several days of close contact. To investigate the development of aggression from an early age, in the present study, newly emerged males were paired to investigate the relationship between JH levels and aggression. The results showed that injection of JH III significantly increased the probability of the young males being fight winners. In each age group in which aggression was initiated, the dominants had significantly higher JH levels than either the subordinates or the same aged non-fighters. JH injection of subordinates on the day of rank establishment had no effect on the probability of rank switch. These results indicate that, (i) in newly emerged male pairs, JH plays a decisive role in rank establishment and the fact that dominant status is significantly associated with a higher JH titer and subordinate status with a lower JH titer is consistent with the basic principle of the challenge hypothesis, and (ii) after rank establishment, the lack of effect of JH treatment on rank change is consistent with the idea of “social inertia” in vertebrates.     

Parasitoid pressure and the radiation of a gallforming group (Cecidomyiidae: Asphondylia spp.) on creosote bush (Larrea tridentata)

Summary We tested the Enemy Impact Hypothesis, which predicts that communities of one tropic level are organized by the tropic level above. In the case of gallforming insect communities, the hypothesis predicts that gall morphology will diverge, minimizing the number of parasitoids shared among species. We used the monophyletic group of gallforming cecidomyiids (Asphondylia spp.) on creosote bush (Larrea tridentata) to test this hypothesis, predicting that species with thicker gall walls should exclude species of parasitoids with shorter ovipositors and have lower levels of parasitism. Of 17 parasitoid species reared from Asphondylia galls on creosote bush, 9 accounted for over 98% of parasitism. Seven of these 9 species had ovipositors long enough to penetrate 10 of 13 gall morphs measured. There was no significant relationship between gall wall thickness and number of associated parasitoid species (r ²=0.01, P>0.05, n=13). There was no relationship between gall wall thickness and types of parasitoid species colonizing galls: parasitoids with the shortest ovipositors colonized all types of gall morphs and were dominant members of the parasitoid assemblages in galls with the thickest walls. Ultimately, there were no significant differences in percent parasitism among Asphondylia species, regardless of gall wall thickness. We found no difference in numbers of associated parasitoids or percent parasitism in galls with different textures (e.g. hairy versus smooth), different locations on the plant or different phenologies. Our results suggest that enemy impact has not influenced the diversity of this gall community. Gall wall thickness, phenology, location on the plant and surface structure do not appear to influence the distribution of parasitoid species. Other explanations are offered to account for diversity in gall morphology among these species.