Evolution of gall morphology and host-plant relationships in willow-feeding sawflies (Hymenoptera: Tenthredinidae)

Abstract.— There are over 200 species of nematine sawflies that induce galls on willows (Salix spp.). Most of the species are monoor oligophagous, and they can be separated into seven or eight different groups based on the type of gall that they induce. We studied the evolution of different gall types and host plant associations by reconstructing the phylogeny of five outgroup and 31 ingroup species using DNA sequence data from the mitochondrial cytochrome b gene. Maximum-parsimony and maximum-likelihood analyses resulted in essentially the same phylogeny with high support for important branches. The results show that: (1) the galling species probably form a monophyletic group; (2) true closed galls evolved only once, via leaf folders; (3) with the possible exception of leaf rollers, all gall type groups are mono- or paraphyletic; (4) similar gall types are closer on the phylogeny than would be expected by a random process; (5) there is an apparent evolutionary trend in galling site from the leaf edge towards the more central parts of the host plant; and (6) many willow species have been colonized several times, which excludes the possiblity of parallel cladogenesis between willows and the gallers; however, there are signs of restrictions in the evolution of host use. Many of the patterns in the evolutionary history of nematine gallers have also been observed in earlier studies on other insect gallers, indicating convergent evolution between the independent radiations.     

Evolutionary dynamics of host-plant use in a genus of leaf-mining moths

Abstract. We used nuclear 28S rDNA sequence data to estimate the phylogeny of 77 leaf-mining Phyllonorycter (Gracillariidae) moth species, including all 55 British species, feeding on 44 different plant genera. There was strong support for both the monophyly of Phyllonorycter and the placement of the genus Cameraria as its sister group. Host-plant use was mapped onto the moth phylogeny and investigated statistically in several ways. First, we show that the estimated level of cospeciation between leaf miners and their host plants is not greater than expected by chance, despite the physical intimacy of the association. Nevertheless, the pattern of host-plant use is far from random, with closely related Phyllonorycter species generally feeding on closely related plants. However, although Phyllonorycter species from a given host plant tend to form distinct clades, there is also statistical support for multiple independent colonizations of some host-plant taxa (e.g. the order Rosales and the genus Corylus ). Despite numerous host shifts, most Phyllonorycter species feed on trees and the few species that attack shrubs or herbs have mostly acquired these habits independently. There is also limited evidence that host shifts to herbs are more likely from shrubs than from trees. Similarly, most species mine the lower surface of leaves but the few upper-surface miners have each evolved the habit independently. Consequently, these shifts to new adaptive zones have not led to substantial radiations.     

Manipulation of host plant development by the gall-midge Rhabdophaga strobiloides

Abstract. 1. The gall-midge Rhabdophaga strobiloides (O.S.) forms a gall in the apical bud of actively growing willow twigs.
2. Galls were not randomly distributed among twigs. Twigs that arose towards the distal end of the branch were much more likely to be galled. Distally located twigs also grew to greater girth than more proximally located twigs.
3. Comparisons of galled twigs with normal twigs in similar locations along their branch showed that the gall causes even greater growth in twig girth than when no gall is present.
4. The hypothesis that galled twigs draw photosynthate produced elsewhere in the plant was tested in an experiment that measured the growth of galled and normal twigs. with their leaves intact, to galled and normal twigs that were manually defoliated. Defoliation caused reduced growth in normal twigs, but galled twigs grew equally well with or without their leaves. Leaf removal had no effect on gall growth.
5. Twig diameter was positively correlated with gall diameter. Call diameter was positively correlated with larval biomass.
6. Patterns of twig and gall growth suggest that the gall-midge manipulates host plant growth and development to provide resources for growth and survival. Manipulation of the host may be an important phenomenon in the evolution of parasitic organisms.     

Amoebophrya spp. from the Bloom‐forming Dinoflagellate Cochlodinium polykrikoides: Parasites not Nested in the “Amoebophrya ceratii Complex”

Members of Amoebophrya ceratii complex are known to infect a number of free‐living dinoflagellates including harmful algal bloom species. In August and October 2012, Amoebophrya infections during two bloom events of the dinoflagellate Cochlodinium polykrikoides were observed along southern coastal waters of Korea. Microscopic observations and molecular data revealed that two different Amoebophrya parasites infected the same host species. In addition, while one developed in the host's nucleus, the other in the host's cytoplasm. Phylogenetic analyses showed that the two parasites were not nested in the previously recognized “Amoebophrya ceratii complex clade”, which contained sequences of parasites infecting numerous dinoflagellate species. Instead, they branched as sister taxa to the isolate (possibly Amoebophrya) from radiolarians Hexacontium gigantheum. Our result indicates that the two Amoebophrya parasites infecting C. polykrikoides may be different species from those inside the “complex.”     

The evolution of gall parasitism accompanied by a host shift in the gall aphid, Eriosoma yangi (Homoptera: Aphidoidea)

Eriosoma aphids form leaf-roll galls on various elm species. Eriosoma yangi occurring on the Japanese elm is an obligatory "cuckoo" parasite, invading and usurping galls of other Eriosoma species. But another host race of E. yangi induces its own galls on the Chinese elm. Multivariate morphometries show that the two host races differ significantly in quantitative characters. Overwintering eggs of the parasitic form were experimentally transferred to the Chinese elm. The resultant larvae still exhibited a parasitic habit and developed into adults, which multivariate morphometries classified as the parasitic form. This form is considered different enough from the gall maker to be given a species rank. Eriosoma aphids facultatively invade other Eriosoma galls, where they sometimes deposit offspring. It is hypothesized that the parasitic form on the Japanese elm originated from a founder population which had migrated from the Chinese elm. On this new host, only those fundatrices that invade other species' galls will be able to leave offspring, because aphid fundatrices cannot gall plants other than their primary hosts. It is assumed that the parasitic form was established under intense selective pressure following a population bottleneck. This hypothesis was corroborated by field investigations at sites where the two forms occur sympatrically.     

Masking of host plant odour in the olfactory orientation of the Colorado potato beetle

The odour of potato plants Solanum tuberosum L., elicits a true odour-conditioned positive anemotaxis in the Colorado potato beetle, Leptinotarsa decemlineata Say. Blending the odour of non-host plant species, namely wild tomatoes Lycopersicon hirsutum f. glabratum C.H. Mull or cabbage Brassica oleracea L. var. gemmifera DC., with the attractive host plant odour blocks the release of upwind responses in non-experienced as well as experienced females. The neutralization of the beetle's orientation responses is obtained without repellency. It is expected that masking of host plant odour occurs often in mixed cropping systems.

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Résumé Un certain nombre d'articles rapporte une diminution du nombre des insectes phytophages spécialistes comme une conséquence à la diversification des agroécosystèmes (Altieri & Letourneau, 1982; Cromartie, 1981; Kareiva, 1983; Risch et al., 1983). Ainsi, il a été supposé que la présence de plantes non-hôtes pouvait interférer avec celle de plantes hôtes en modifiant la nature des informations chimiques parvenant aux insectes. Les modifications comportementales qui peuvent en résulter lors de l'attraction à distance sont toutefois encore peu claires. Les expérimentations présentées ici analysent les réponses individuelles du doryphore à diverses stimulations olfactives issues de plantes entières. Des femalles ont été ainsi étudiées un jour après l'émergeance en présence d'air vierge et d'air chargé d'odeurs de Solanum tuberosum, Lycopersicon hirsutum f. glabratum, Brassica oleracea var. gemmifera, ainsi que des mélanges, S. tuberosum avec L. hirsutum et S. tuberosum avec B. oleracea. A partir de l'enregistrement continu de l'activité locomotrice dans chaque condition (à l'aide du compensateur de locomotion), nous avons déterminé que L. hirsutum et B. oleracea masquaient l'odeur de la plante hôte S. tuberosum en supprimant la réponse anémotactique positive conditionnée par l'odeur de cette dernière. Les mélanges d'odeurs ainsi obtenus sont considérés comme neutres pour l'orientation à longue distance du doryphore. L'expérience préalable par le doryphore de l'ingestion de feuillage de pomme de terre ne modifie pas l'effet du camouflage obtenu. Ces résultats laissent penser qu'il est possible de limiter la découverte de la plante hôte chez les insectes, et que les méthodes de camouflages olfactives pourraient prévenir les attaques de certains ravageurs.

     

Host plant intraspecific variation determines gall traits

• Galls display a multiplicity of traits, including colours, which are driven by pigment accumulation. Their conspicuousness has attracted researchers' attention and several hypotheses have been raised. However, plants themselves vary intra-specifically, including in their pigment concentrations. As galls are a result of host tissue development, colours may be a by-product of the host's own traits, being more conspicuous simply because the sites where galls develop already have the predisposition to accumulate more pigment. Here, we call this the host variation hypothesis.
• We test this hypothesis using the system of galls induced by Palaeomystella oligophaga on Macairea radula host plant. Using spectrophotometry, we calculated the Anthocyanin Reflectance Index (ARI) of gall projections, which are responsible for their characteristic colours. We tested the influence of occupant identity (galling insect or any natural enemy), gall volume, parenchyma thickness, height from the ground, ARI of leaf, ARI of gall surface and ARI of the respective stem.
• We corroborated the host variation hypothesis since the anthocyanin content in stems and in galls' projections were positively related. Moreover, anthocyanin in galls' projections was positively related to anthocyanin in the gall surface and negatively related to gall volume and parenchyma thickness. This shows that, besides the host specificities, galls' own traits may also be responsible for pigment accumulation, influencing their colours.
• In this study, using colour as an example, we show that although galls tend to be considered complex expressions of galling insects' stimuli, their traits may be simply influenced by previous and specific attributes of the host organs.

     

BUTTERFLIES AND PLANTS: A PHYLOGENETIC STUDY

A database on host plant records from 437 ingroup taxa has been used to test a number of hypotheses on the interaction between butterflies and their host plants using phylogenetic methods (simple character optimization, concentrated changes test, and independent contrasts test). The butterfly phylogeny was assembled from various sources and host plant clades were identified according to Chase et al.'s rbcL-based phylogeny. The ancestral host plant appears to be associated within a highly derived rosid clade, including the family Fabaceae. As fossil data suggest that this clade is older than the butterflies, they must have colonized already diversified plants. Previous studies also suggest that the patterns of association in most insect-plant interactions are more shaped by host shifts, through colonization and specialization, than by cospeciation. Consequently, we have focused explicitly on the mechanisms behind host shifts. Our results confirm, in the light of new phylogenetic evidence, the pattern reported by Ehrlich and Raven that related butterflies feed on related plants. We show that host shifts have generally been more common between closely related plants than between more distantly related plants. This finding, together with the possibility of a higher tendency of recolonizing ancestral hosts, helps to explain the apparent large-scale conservation in the patterns of association between insects and their host plants, patterns which at the same time are more flexible on a more detailed level. Plant growth form was an even more conservative aspect of the interaction between butterflies and their host plants than plant phylogeny. However, this is largely explained by a higher probability of colonizations and host shifts while feeding on trees than on other growth forms.     

Ovipositional preference ofSiphoninus phillyreae and its fitness on seven host plant species

The relationship between ovipositional preference ofSiphoninus phillyreae (Haliday) (Homoptera: Aleyrodidae) and host plant suitability on seven host plant species (Citrus sinensis (L.) cv. ‘Washington’ [navel orange],Fraxinus uhdei (Wenz.) [shamel ash],Heteromeles arbutifolia Roemer [toyon],Malus domestica Mill. cv. ‘Granny Smith’, [apple],Pistacia vera L. cv. ‘Kerman’ [pistachio],Prunus persica (L.) cv. ‘O’Henry’ [peach], andPyrus communis L. cv. ‘Bartlett’ [pear]) was evaluated. Ovipositional preference ofS. phillyreae was determined by measuring egg density after adult female whitefies were given a simultaneous choice of all host plants for oviposition. Immature survival, developmental time, and adult size were examined to determine host plant suitability forS. phillyreae. All studies were performed under greenhouse conditions.S. phillyreae showed distinct ovipositional preference among host plant species. Host plant species had a significant effect on immature survival, but little or no effect on developmental time or forewing length. For four of the seven host plant species tested, there was an association between ovipositional preference and survival.     

Ant effects on three-trophic level interactions: plant, galls, and parasitoids

1. Ants have evolved mutualistic relationships with a diverse array of plant and animal species. Usually, the predatory/aggressive behaviour of ants near food sources can limit herbivore damage. 2. Galls of Disholcaspis edura on Quercus turbinella produce a secretion that is harvested by three species of ants (Formica neorufibarbis, Liometopium apiculatum, and Monomorium cyaneum) in the chaparral vegetation of Arizona, U.S.A. The study reported here provides evidence of a mutualistic relationship between these species of ants and the gall-forming wasp Disholcaspis edura. 3. An ant exclusion experiment showed that when ants tended galls, the rate of parasitism by Platygaster sp. on Disholcaspis edura was nearly halved in comparison to a treatment in which ants were excluded. 4. In the presence of ants, galls with the largest diameter suffered a lower mortality rate due to parasitoid attack than when ants were excluded. Thus, ant presence reduced the selective pressure imposed by Platygaster sp. on the galls with larger diameter.     

The evolution of inquilinism, host-plant use and mitochondrial substitution rates in Tamalia gall aphids

We used mitochondrial DNA data to infer phylogenies for 28 samples of gall-inducing Tamalia aphids from 12 host-plant species, and for 17 samples of Tamalia inquilinus, aphid 'inquilines' that obligately inhabit galls of the gall inducers and do not form their own galls. Our phylogenetic analyses indicate that the inquilines are monophyletic and closely related to their host aphids. Tamalia coweni aphids from different host plants were, with one exception, very closely related to one another. By contrast, the T. inquilinus aphids were strongly genetically differentiated among most of their host plants. Comparison of branch lengths between the T. coweni clade and the T. inquilinus clade indicates that the T. inquilinus lineage evolves 2.5-3 times faster for the cytochrome oxidase I gene. These results demonstrate that: (1) Tamalia inquilines originated from their gall-inducing hosts, (2) communal (multi-female) gall induction apparently facilitated the origin of inquilinism, (3) diversification of the inquilines has involved rapid speciation along host-plant lines, or the rapid evolution of host-plant races, and (4) the inquilines have undergone accelerated molecular evolution relative to their hosts, probably due to reduced effective population sizes. Our findings provide insight into the behavioural causes and evolutionary consequences of transitions from resource generation to resource exploitation.     

Detecting ancient codispersals and host shifts by double dating of host and parasite phylogenies: Application in proctophyllodid feather mites associated with passerine birds

Inferring cophylogeographic events requires matching the timing of these events on both host and symbiont (e.g., parasites) phylogenies because divergences of hosts and their symbionts may not temporally coincide, and host switches may occur. We investigate a large radiation of birds (Passeriformes) and their permanent symbionts, the proctophyllodid feather mites (117 species from 116 bird species; six genes, 11,468 nt aligned) using two time‐calibration strategies for mites: fossils only and host phylogeography only. Out of 10 putative cophylogeographic events 4 agree in timing for both symbiont and host events being synchronous co‐origins or codispersals; three were based on host shifts, but agree in timing being very close to the origin of modern hosts; two disagree; and one large basal mite split was seemingly independent from host phylogeography. Among these events was an ancient (21–25.3 Mya), synchronous codispersal from the Old World leading to the origin and diversifications of New World emberizoid passerids and their mites, the thraupis + quadratus species groups of Proctophyllodes. Our framework offers a more robust detection of host and symbiont cophylogeographic events (as compared to host‐symbiont reconciliation analysis and using host phylogeography for time‐calibration) and provides independent data for testing alternative hypotheses on timing of host diversification and dispersal.     

Malaria parasites (Apicomplexa,Haematozoea) and their relationships with their hosts: is there an evolutionary cost for the specialization?

Parasite–host specialization is frequently considered to be a derived state such that it represents an 'evolutionary dead end' that strongly limits further evolution. In this study, it was tested whether this theory is applicable to the relationship of malaria parasites and their vertebrate hosts. For this, we revisited Perkins and Schall (2002) analysis of the phylogenetic relationships of the malaria parasites (belonging to the genera Plasmodium , Haemoproteus and Hepatocystis ) based on the mitochondrial Cytochrome b gene sequence, and inferred, using a maximum likelihood (ML) approach, the putative ancestral vertebrate hosts. As the topology in this study presents several unresolved branches and is slightly different from that of Perkins and Schall, a Shimodaira and Hasegawa (SH; 1999) test has been performed in order to properly consider several alternative topologies. The results of this study suggest that the common ancestor of all these malaria parasites was a reptile (more specific of the order Squamata), and that the host switches from Squamata to Aves and vice versa were quite frequent along the evolution of these parasites. On the contrary, a strong evidence that the host shift from Squamata to Mammalia had occurred only once during the evolution of these organisms was found. This evidence (added to the current knowledge about the association of the malaria parasites with their vertebrate hosts) allows us to suggest, at least considering the species included in this study, that the adaptation in mammals had required a high level of specialization. Hence, the acquisition of this host class had culminated in an evolutionary dead end for the mammalian malaria parasites.     

Host plant related properties of the antennal olfactory system in the oak flea weevil, Rhynchaenus quercus. Electroantennogram study

Electroantennogram (EAG) recordings from the antennae of a highly specialized phytophagous species — the oak flea weevil, Rhynchaenus quercus L. — revealed two general properties of its antennal olfactory system: (1) a high and to some extent selective sensitivity to chemotaxonomically nonspecific compounds, namely the so-called general green leaf volatiles as well as to geraniol and linalool, (2) a lack of marked differences in the general excitation level caused by stimulation with vapours from leaves of both host plants and non-host plants. It was also observed, that the shapes of EAG's evoked by vapours from cut leaves and general green leaf volatiles differed from those elicited by corresponding concentrations of geraniol and linalool, where recovery was distinctly slower.The results of EAG tests suggest that the general green leaf volatile complex is important in host-odour perception by the oak flea weevil. A possible function of these compounds in host-plant odour discrimination is discussed.

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Zusammenfassung Elektroantennographische Aufzeichnungen an Fühlern einer hochspezialisierten phytophagen Art, des Eichenblattflohs Rhynchaenus quercus L. zeigten zwei allgemeine Eigenschaften des Geruchsystems der Antennen:Erstens ist eine hohe und teilweise selektive Empfindlichkeit auf chemotaxonomisch nicht spezifische Verbindungen, besonders auf die allgemeinen Blattgründämpfe wie auch auf Geraniol und Linalool vorhanden.Zweitens fehlen ausgeprägte Unterschiede im allgemeinen Erregungsniveau hervorgerufen durch flüchtige Stoffe von Wirtspflanzen einerseits und Nichtwirtspflanzen andererseits. Es wurde auch beobachtet, dass die Form der EAGs hervorgerufen durch zerschnittene Blätter oder allgemeine Blattgründämpfe verschieden war von denjenigen verursacht durch Geraniol und Linalool, wo die Erholung eindeutig langsamer war.Die Resultate der EAG deuten darauf hin, dass der Komplex der allgemeinen Blattgründämpfe wichtig ist bei der Wahrnehmung der Wirtspflanzengerüche durch Rhynchaenus quercus. Eine mögliche Funktion dieser Verbindungen bei der Erkennung der Wirtspflanzen wird diskutiert.

     

Genetic variation in a phylogenetic context: Responses of two specialized leaf beetles (Coleoptera: Chrysomelidae) to host plants of their congeners

This work explores the possibility that constraints on genetic variation guide host shifts and are responsible for the evolutionary conservatism of host affiliation in phytophagous insects. To this end, we used full- and half-sib breeding designs to screen two species of the North American bettle genus Ophraella for genetic variation in larval and adult feeding responses to several host plants of other species of Ophraella. All the plants are in the family Asteraceae. In O. conferta, we observed effectively no feeding response, and hence no genetic variation in response, to three of five test plant species; only those plants related to the species' natural hosts evoked genetically variable responses. In O. artemisiae, adults displayed genetic variation in response to a congener of the natural host, but not to two distantly related plants. However, significant variation among full-sib broods in larval feeding suggests the existence of nonadditive genetic variance in feeding response to all five species of test plants—although survival was very low on most of them. The results suggest that patterns of presence versus apparent absence of detectable genetic variation may be related to the chemical similarity of plants to the insects' natural hosts, but not evidently to the phylogenetic history of host affiliation within the genus. Almost all genetic correlations in responses to host plants were not significantly different from zero; the few significant correlations were positive, and negative correlations that might explain host specificity were not found. Our data do not explain why exclusive shifts to new hosts should occur, but the apparent lack of genetic variation in responses to some plants suggests that the direction of host shifts is genetically constrained.     

Pyrrolizidine alkaloids in arctiid moths (Lep.) with a discussion on host plant relationships and the role of these secondary plant substances in the Arctiidae

Pyrrolizidine alkaloids (PA) have been identified in six species of Arctiidae reared on Senecio and Crotalaria. These include senecionine, seneciphylline, integerrimine, jacobine, jacozine, jacoline, jaconine and a metabolite (C₁₅H₂₅NO₅) from Senecio , and monocrotaline, trichodesmine and crispatine from Crotalaria.
The all-red aberration of Tyria jacobaeae (var. conyi) contained much less of the metabolite than normal examples of this species. Female Spilosoma lutea reared on the same plants of S. jacobaea contained markedly more jacobine and jacoline than die males.
Host plant relationships and secondary plant substances are discussed. It is suggested that the Arctiid moths' own deterrent secretions, directed against vertebrate predators, pre-adapts them for feeding on foliage likewise protected against large herbivores by toxic secondary plant substances such as cardenolides and pyrrolizidine alkaloids. These latter substances are more toxic to vertebrate than to insect herbivores, and their dual function of deterrent and insect aphrodisiac puts a premium on their sequestration and storage once a species has achieved the initial steps, and occupied the plant niche concerned. It is further suggested that the polyphagous habits of the Arctiidae result in a more equitable distribution of die secondary plant substances within the Mullerian complex concerned, thus providing a generalized warning message for the potential vertebrate predator.     

Host plant variation in mortality of the leaf-folding sawfly on the arroyo willow

Abstract. 1. We tested the hypothesis that survival and sources of mortality of the leaf-folding sawfly ( Phyllocolpa sp.) varied significantly among host plants of the arroyo willow ( Salix lasiolepis Bent ham).
2. Survival of the leaf folder differed among field and potted willows in a common environment in two of three cases, and sources of mortality differed among plants in four of five cases.
3. Egg mortality differed among field plants but not among the potted willow plants.
4. Larval mortality and parasitism differed among field and potted willows in 2 years, and appeared to be compensatory mortality sources.
5. Leaf folder density among plants was not generally correlated with percentage of leaf folds with no egg (galls formed but no subsequent oviposition), percentage survival, or percentage mortality, indicating a general lack of density dependence.
6. The proportion of folds with no egg oviposited differed significantly among field and potted plants, and was only correlated with survival or sources of mortality in one of three years.
7. The data support the hypothesis that host plant genotype affects the interaction of the leaf folder with its natural enemies, and thus represents a three trophic level interaction.     

Evolution of host- and habitat association in the wood-feeding cockroach, Cryptocercus

Members of the cockroach genus Cryptocercus are subsocial insects that live in temperate forests and feed on decomposing logs. At present, seven species are recognized worldwide: four in the eastern USA, one in the western USA, and one each in Russia and China. Genetic variation within and among the Nearctic species has been characterized extensively in previous studies. However, whether there has been a corresponding divergence in the host and habitat association of Cryptocercus species is not known. Here, we report on differences in host and habitat association among six of the seven Cryptocercus species, estimated from field observations, elevation data, and land cover data. Our results indicated that the eastern and western USA species differ from one another in their distribution patterns, abundance, and habitat association. The eastern USA species are associated largely with deciduous forests, whereas the western USA and the Russian species are associated with evergreen forests. Thus, the eastern USA species, which are evolutionarily the most recent ones, have adapted to a different set of tree species relative to the basal species. There were also differences in the habitat association of the various species. Specifically, in the eastern USA, Cryptocercus darwini , evolutionarily the most recent species, occupied a habitat that is predominantly at low elevation [<400 m above sea level (ASL)] while all the other Nearctic species and the Russian species occupied a habitat that is at relatively higher elevations (>400 m ASL). Mapping of the above traits on a phylogenetic tree revealed that the evolutionary trend in Cryptocercus with regard to host and habitat association has been toward the utilization of low elevation habitats dominated by deciduous forests. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 163–172.     

The evolution of egg size in the brood parasitic cuckoos

We compared genera of nonparasitic cuckoos and two groups ofparasitic cuckoos: those raised together with host young ("nonejectors")and those in which the newly hatched cuckoo either ejects thehost eggs or chicks, or kills the host young ("ejectors"). Nonejectorsare similar to their hosts in body size and parasitize largerhosts than do ejectors, which parasitize hosts much smallerthan themselves. In both types of parasite, the cuckoo's eggtends to match the host eggs in size. To achieve this, nonejectorshave evolved a smaller egg for their body size than have nonparasiticcuckoos, and ejectors have evolved an even smaller egg. Amongejector cuckoo genera, larger cuckoos have larger eggs relativeto the eggs of their hosts, and the relationship between cuckooegg volume (mass of the newly-hatched cuckoo) and host egg volume(mass to be ejected) did not differ from that predicted by weight-liftingallometry. However, comparing among Cuculus cuckoo species,the allometric slope differed from the predicted, so it is notclear that egg size is related to the need to give the cuckoochick sufficient strength for ejection. Comparing the two mostspeciose ejector genera, Chrysococcyx cuckoos (smaller and parasitizedome-nesting hosts) lay eggs more similar in size to their host'seggs than do Cuculus cuckoos (larger and parasitize open cup–nestinghosts). Closer size-matching of host eggs in Chrysococcyx mayreflect the following: (1) selection to reduce adult body massto facilitate entry through small domed nest holes to lay, and(2) less need for a large egg, because longer incubation periodsin dome-nesting hosts allow the young cuckoo more time to growbefore it need eject host eggs.