Ecological implications of parasites in natural <Emphasis Type="Italic">Daphnia</Emphasis> populations

In natural host populations, parasitism is considered to be omnipresent and to play an important role in shaping host life history and population dynamics. Here, we study parasitism in natural populations of the zooplankton host Daphnia magna investigating their individual and population level effects during a 2-year field study. Our results revealed a rich and highly prevalent community of parasites, with eight endoparasite species (four microsporidia, one amoeba, two bacteria and one nematode) and six epibionts (belonging to five different taxa: Chlorophyta, Bacillariophyceae, Ciliata, Fungi and Rotifera). Several of the endoparasites were associated with a severe overall fecundity reduction of the hosts, while such effects were not seen for epibionts. In particular, infections by Pasteuria ramosa, White Fat Cell Disease and Flabelliforma magnivora were strongly associated with a reduction in overall D. magna fecundity. Across the sampling period, average population fecundity of D. magna was negatively associated with overall infection intensity and total endoparasite richness. Population density of D. magna was negatively correlated to overall endoparasite prevalence and positively correlated with epibiont richness. Finally, the reduction in host fecundity caused by different parasite species was negatively correlated to both parasite prevalence and the length of the time period during which the parasite persisted in the host population. Consistent with epidemiological models, these results indicate that parasite mediated host damages influence the population dynamics of both hosts and parasites.     

Correlated evolution of host and parasite body size: tests of Harrison's rule using birds and lice

Large-bodied species of hosts often harbor large-bodied parasites, a pattern known as Harrison's rule. Harrison's rule has been documented for a variety of animal parasites and herbivorous insects, yet the adaptive basis of the body-size correlation is poorly understood. We used phylogenetically independent methods to test for Harrison's rule across a large assemblage of bird lice (Insecta: Phthiraptera). The analysis revealed a significant relationship between louse and host size, despite considerable variation among taxa. We explored factors underlying this variation by testing Harrison's rule within two groups of feather-specialist lice that share hosts (pigeons and doves). The two groups, wing lice (Columbicola spp.) and body lice (Physconelloidinae spp.), have similar life histories, despite spending much of their time on different feather tracts. Wing lice showed strong support for Harrison's rule, whereas body lice showed no significant correlation with host size. Wing louse size was correlated with wing feather size, which was in turn correlated with overall host size. In contrast, body louse size showed no correlation with body feather size, which also was not correlated with overall host size. The reason why body lice did not fit Harrison's rule may be related to the fact that different species of body lice use different microhabitats within body feathers. More detailed measurements of body feathers may be needed to explore the precise relationship of body louse size to relevant components of feather size. Whatever the reason, Harrison's rule does not hold in body lice, possibly because selection on body size is mediated by community-level interactions between body lice.     

Phylogenomic analysis of seal lice reveals codivergence with their hosts

Lice are considered a model system for studying the process of cospeciation because they are obligate and permanent parasites and are often highly host‐specific. Among lice, species in the family Echinophthiriidae Enderlein (Anoplura) are unique in that they infest mammalian hosts with an amphibious lifestyle, i.e. pinnipeds and the river otter. There is evidence that the ancestor of this group infested the terrestrial ancestor of pinnipeds, which suggests these parasites coevolved with their hosts during the transition to marine environments. However, there has been no previous study investigating the phylogenetic relationships among sucking lice parasitizing seals and sea lions. To uncover the evolutionary history of these parasites, we obtained genomic data for Antarctophthirus microchir Trouessart and Neumann (from two hosts), Antarctophthirus carlinii Leonardi et al., Antarctophthirus lobodontis Enderlein, Antarctophthirus ogmorhini Enderlein, Lepidophthirus macrorhini Enderlein, and Proechinophthirus fluctus Ferris. From genomic sequence reads, we assembled > 1000 nuclear genes and used these data to infer a phylogenetic tree for these lice. We also used the assembled genes in combination with read‐mapping to estimate heterozygosity and effective population size from individual lice. Our analysis supports the monophyly of lice from pinnipeds and uncovers phylogenetic relationships within the group. Surprisingly, we found that A. carlinii, A. lobodontis, and A. ogmorhini have very little genetic divergence among them, whereas the divergence between different geographic representatives of A. microchir indicate that they are possibly different species. Nevertheless, our phylogeny of Echinophthiriidae suggests that these lice have consistently codiverged with their hosts with minimal host switching. Population genomic metrics indicate that louse effective population size is linked to host demographics, which further highlights the close association between pinnipeds and their lice.     

Body size and fecundity are correlated in feather lice (Phthiraptera: Ischnocera): implications for Harrison's rule

1. Harrison's rule, which predicts that large‐bodied species of hosts have large‐bodied species of parasites, has been documented in a wide diversity of parasites. 2. Harrison's rule has been most thoroughly studied in avian feather lice, which escape from host defence (preening) by hiding in the feathers. Lice that are unable to hide are selectively removed by preening. Preening selects for small lice on small hosts, which have small feathers in which to hide. 3. Preening should not, however, select for large lice on large hosts. Instead, the larger size of lice on large hosts is thought to result from a positive relationship between size and fecundity, as shown for many other insects. 4. This study tested for a size–fecundity correlation within Columbicola columbae, the host‐specific ‘wing louse’ of rock pigeons (Columba livia). 5. The results confirm a positive relationship between female body length and number of eggs laid. 6. The study thus supports a mechanism consistent with stabilising selection leading to the evolution of the Harrison's rule pattern among species of Columbicola and their hosts.     

Patterns of host–parasite associations in tropical lice and their passerine hosts in Cameroon

Coevolutionary processes that drive the patterns of host–parasite associations can be deduced through congruence analysis of their phylogenies. Feather lice and their avian hosts have previously been used as typical model systems for congruence analysis; however, such analyses are strongly biased toward nonpasserine hosts in the temperate zone. Further, in the Afrotropical region especially, cospeciation studies of lice and birds are entirely missing. This work supplements knowledge of host–parasite associations in lice using cospeciation analysis of feather lice (genus Myrsidea and the Brueelia complex) and their avian hosts in the tropical rainforests of Cameroon. Our analysis revealed a limited number of cospeciation events in both parasite groups. The parasite–host associations in both louse groups were predominantly shaped by host switching. Despite a general dissimilarity in phylogeny for the parasites and hosts, we found significant congruence in host–parasite distance matrices, mainly driven by associations between Brueelia lice and passerine species of the Waxbill (Estrildidae) family, and Myrsidea lice and their Bulbul (Pycnonotidae) host species. As such, our study supports the importance of complex biotic interactions in tropical environments.     

Lousy grouse: Comparing evolutionary patterns in Alaska galliform lice to understand host evolution and host–parasite interactions

Understanding both sides of host–parasite relationships can provide more complete insights into host and parasite biology in natural systems. For example, phylogenetic and population genetic comparisons between a group of hosts and their closely associated parasites can reveal patterns of host dispersal, interspecies interactions, and population structure that might not be evident from host data alone. These comparisons are also useful for understanding factors that drive host–parasite coevolutionary patterns (e.g., codivergence or host switching) over different periods of time. However, few studies have compared the evolutionary histories between multiple groups of parasites from the same group of hosts at a regional geographic scale. Here, we used genomic data to compare phylogenomic and population genomic patterns of Alaska ptarmigan and grouse species (Aves: Tetraoninae) and two genera of their associated feather lice: Lagopoecus and Goniodes. We used whole‐genome sequencing to obtain hundreds of genes and thousands of single‐nucleotide polymorphisms (SNPs) for the lice and double‐digest restriction‐associated DNA sequences to obtain SNPs from Alaska populations of two species of ptarmigan. We found that both genera of lice have some codivergence with their galliform hosts, but these relationships are primarily characterized by host switching and phylogenetic incongruence. Population structure was also uncorrelated between the hosts and lice. These patterns suggest that grouse, and ptarmigan in particular, share habitats and have likely had historical and ongoing dispersal within Alaska. However, the two genera of lice also have sufficient dissimilarities in the relationships with their hosts to suggest there are other factors, such as differences in louse dispersal ability, that shape the evolutionary patterns with their hosts.     

GENETIC DIFFERENTIATION AMONG CHEWING LOUSE POPULATIONS (MALLOPHAGA: TRICHODECTIDAE) IN A POCKET GOPHER CONTACT ZONE (RODENTIA: GEOMYIDAE)

Genetic variation among populations of chewing lice (Geomydoecus actuosi) was examined in relation to chromosomal and electrophoretic variation among populations of their hosts (Thomomys bottae) at a contact zone. Louse demes were characterized by low levels of genetic heterozygosity (H? = 0.039) that may result from founder effects during primary infestation of hosts, compounded by seasonal reductions in louse population size. Louse populations sampled from different hosts showed high levels of genetic structuring both within and among host localities. Microgeographic differentiation of louse populations is high (mean F_ST = 0.092) suggesting that properties of this host–parasite system promote differentiation of louse populations living on different individual hosts. Among-population differentiation in lice (F_ST = 0.240) was similar to that measured among host populations (F_ST = 0.236), suggesting a close association between gene flow in pocket gophers and gene flow in their lice.     

Divergent host phenotypes create opportunities and constraints on the distribution of two wing‐dwelling feather mites

The diversity of symbionts (commensals, mutualists or parasites) that share the same host species may depend on opportunities and constraints on host exploitation associated with host phenotype or environment. Various host traits may differently influence host accessibility and within‐host population growth of each symbiont species, or they may determine the outcome of within‐host interactions among coexisting species. In turn, phenotypic diversity of a host species may promote divergent exploitation strategies among its symbiotic organisms. We studied the distribution of two feather mite species, Proctophyllodes sylviae and Trouessartia bifurcata, among blackcaps Sylvia atricapilla wintering in southern Spain during six winters. The host population included migratory and sedentary individuals, which were unequally distributed between two habitat types (forests and shrublands). Visual mite counts showed that both mite species often coexisted on sedentary blackcaps, but were seldom found together on migratory blackcaps. Regardless of host habitat, Proctophyllodes were highly abundant and Trouessartia were scarce on migratory blackcaps, but the abundance of both mite species converged in intermediate levels on sedentary blackcaps. Coexistence may come at a cost for Proctophyllodes, whose load decreased when Trouessartia was present on the host (the opposite was not true). Proctophyllodes load was positively correlated with host wing length (wings were longer in migratory blackcaps), while Trouessartia load was positively correlated to uropygial gland size (sedentary blackcaps had bigger glands), which might render migratory and sedentary blackcaps better hosts for Proctophyllodes and Trouessartia, respectively. Our results draw a complex scenario for mite co‐existence in the same host species, where different mite species apparently take advantage of, or are constrained by, divergent host phenotypic traits. This expands our understanding of bird–mite interactions, which are usually viewed as less dynamic in relation to variation in host phenotype, and emphasizes the role of host phenotypic divergence in the diversification of symbiotic organisms.     

Dynamic effects of parasitism by an endoparasitoid wasp on the development of two host species: implications for host quality and parasitoid fitness

1. The study reported here examined growth and developmental interactions between the gregarious larval koinobiont endoparasitoid Cotesia glomerata (Hymenoptera: Braconidae) and two of its hosts that vary considerably in growth potential: Pieris rapae and the larger P. brassicae (Lepidoptera: Pieridae). At pupation, healthy larvae of P. brassicae are over twice as large, in terms of fresh body mass, as those of P. rapae. 2. Clutch size of C. glomerata was manipulated artificially, and the relationship between parasitoid burden and the maximum weight of the parasitised host (= host–parasitoid complex) was measured. In both hosts, the maximum complex weight was correlated positively with parasitoid burden. Compared with unparasitised hosts, however, the growth of P. rapae was increased at significantly lower parasitoid burdens than in P. brassicae. Emerging wasp size was correlated negatively with parasitoid burden in both host species, whereas development time was less affected. 3. After larval parasitoid egress, the weight of the host carcass increased slightly, but not significantly, with parasitoid burden, although there was a strong correlation between the proportion of host mass consumed by C. glomerata larvae during development and parasitoid burden. 4. Clutch size was generally correlated positively with instar parasitised in both hosts, and greater in P. brassicae than in P. rapae. Sex ratios were much more female biased in L1 and L2 P. rapae than in all other host classes. Adult parasitoid size was correlated inversely with host instar at parasitism, and wasps emerging from P. brassicae were larger, and completed development faster, than conspecifics emerging from P. rapae. 5. The data reveal that parasitism by C. glomerata has profound species‐specific effects on the growth of both host species. Consequently, optimality models in which host quality is often based on host size at parasitism or unparasitised growth potential may have little utility in describing the development of gregarious koinobiont endoparasitoids. The results of this investigation are discussed in relation to the potential effectiveness of gregarious koinobionts in biological control programmes.     

Mixed sex allocation strategies in a parasitoid wasp

The sex allocation strategy of the parasitoid Laelius pedatus (Hymenoptera: Bethylidae) on different-sized hosts was investigated. The wasp lays from one to five eggs, and clutch size increases with host size. On the smallest hosts, single male eggs are laid, while on slightly larger hosts single female eggs are laid. On still larger hosts, gregarious clutches are laid which nearly always consist of a single male and one or more female eggs. The sex ratio strategy of the wasp appears to be influenced by a combination of local mate competition and conditional sex expression based on host quality. Received: 6 June 1996 / Accepted: 13 October 1996     

Mancae of the parasitic cymothoid isopod, <Emphasis Type="Italic">Anilocra apogonae</Emphasis>: early life history,host-specificity,and effect on growth and survival of preferred young cardinal fishes

Juvenile parasitic cymothoid isopods (mancae) can injure or kill fishes, yet few studies have investigated their biology. While the definitive host of the adult cymothoids is usually a single host from a particular fish species, mancae may use so-called optional intermediate hosts before settling on the definitive host. Little, however, is known about these early interactions. The cymothoid isopod, Anilocra apogonae, infests the definitive host, Cheilodipterus quinquelineatus. This study examined their host preference among potential optional intermediate hosts. Their effect on the growth and mortality of the young of three apogonid fishes, including the definitive host, was investigated. The number of mancae produced per brood was positively correlated with female length. When given a choice of intermediate hosts, significantly more mancae attached to Apogon trimaculatus (Apogonidae) than to Apogon nigrofasciatus. When presented with Ap. trimaculatus and Pomacentrus amboinensis (Pomacentridae), mancae only attached to Ap. trimaculatus suggesting that mancae may show a taxonomic affiliation with preferred hosts. Mancae fed on all three apogonid species, with C. quinquelineatus being fed on earlier than Ap. trimaculatus and Ap. nigrofasciatus. Mancae feeding frequency, adjusted for fish survival, was lowest on C. quinquelineatus and highest on Ap. trimaculatus. Infested apogonids had reduced growth and increased mortality compared with uninfested fish. A. apogonae mancae can use several species of young apogonid fishes as optional intermediate hosts. Via reduced growth and increased mortality, mancae have the potential to negatively influence definitive host populations and also other young species of apogonid fishes.     

The role of body size in host specificity: reciprocal transfer experiments with feather lice

Although most parasites show at least some degree of host specificity, factors governing the evolution of specificity remain poorly understood. Many different groups of host-specific parasites show a striking correlation between parasite and host body size, suggesting that size reinforces specificity. We tested this hypothesis by measuring the relative fitness of host-specific feather lice transferred to pigeons and doves that differ in size by an order of magnitude. To test the general influence of size, we transferred unrelated groups of wing and body lice, which are specialized for different regions of the host. Lice were transferred in both directions, from a large native host species, the rock pigeon (Columba livia), to several progressively smaller hosts, and from a small native host species, the common ground dove (Columbina passerina), to several larger hosts. We measured the relative fitness (population size) of lice transferred to these novel host species after two louse generations. Neither wing lice nor body lice could survive on novel host species that were smaller in size than the native host. However, when host defense (preening behavior) was blocked, both groups survived and reproduced on all novel hosts tested. Thus, host defense interacted with host size to govern the ability of lice to establish on small hosts. Neither wing lice nor body lice could survive on larger hosts, even when preening was blocked. In summary, host size influenced the fitness of both types of feather lice, but through different mechanisms, depending on the direction of the transfer. Our results indicate that host switching is most likely between hosts of similar body size. This finding has important implications for studies of host-parasite coevolution at both the micro- and macroevolutionary scales.     

Cophylogenetic analysis of lice in the Colpocephalum complex (Phthiraptera: Amblycera)

The chewing louse genus Colpocephalum parasitizes nearly a dozen distantly related orders of birds. Such a broad host distribution is relatively unusual in lice. However, the monophyly of the genus Colpocephalum has never been tested using molecular characters. Using one nuclear and one mitochondrial gene, we inferred a phylogeny for 54 lice from the genus Colpocephalum and other morphologically similar genera. The resulting phylogeny demonstrates that Colpocephalum itself is not monophyletic. However, these data support the existence of a Colpocephalum complex within which several lineages are restricted to particular host orders. These lineages corresponded to previously described genera, some of which are morphologically distinct and currently considered subgenera. Maddison–Slatkin tests were performed on the resulting phylogeny and showed that host order, host family and biogeographic region had significant phylogenetic signal when mapped onto the Colpocephalum complex phylogeny. A PARAFIT analysis comparing the overall Colpocephalum complex phylogeny to a host phylogeny revealed significant congruence between host and parasite trees. We also compared the cophylogenetic history of Colpocephalum and their hosts to that of a second distantly related feather louse genus, Degeeriella, which also infests diurnal birds of prey. Using PARAFIT to identify individual host–parasite links that contributed to overall congruence, there was no evidence of correlated cophylogenetic patterns between these two louse groups, suggesting that their host distribution patterns have been shaped by different evolutionary processes.     

Host sympatry and body size influence parasite straggling rate in a highly connected multihost,multiparasite system

Parasite lineages commonly diverge when host lineages diverge. However, when large clades of hosts and parasites are analyzed, some cases suggest host switching as another major diversification mechanism. The first step in host switching is the appearance of a parasite on an atypical host, or “straggling.” We analyze the conditions associated with straggling events. We use five species of colonially nesting seabirds from the Galapagos Archipelago and two genera of highly specific ectoparasitic lice to examine host switching. We use both genetic and morphological identification of lice, together with measurements of spatial distribution of hosts in mixed breeding colonies, to test: (1) effects of local host community composition on straggling parasite identity; (2) effects of relative host density within a mixed colony on straggling frequency and parasite species identity; and (3) how straggling rates are influenced by the specifics of louse attachment. Finally, we determine whether there is evidence of breeding in cases where straggling adult lice were found, which may indicate a shift from straggling to the initial stages of host switching. We analyzed more than 5,000 parasite individuals and found that only ~1% of lice could be considered stragglers, with ~5% of 436 host individuals having straggling parasites. We found that the presence of the typical host and recipient host in the same locality influenced straggling. Additionally, parasites most likely to be found on alternate hosts are those that are smaller than the typical parasite of that host, implying that the ability of lice to attach to the host might limit host switching. Given that lice generally follow Harrison's rule, with larger parasites on larger hosts, parasites infecting the larger host species are less likely to successfully colonize smaller host species. Moreover, our study supports the general perception that successful colonization of a novel host is extremely rare, as we found only one nymph of a straggling species, which may indicate successful reproduction.     

Separating the Effects of Experience, Size, Egg Load, and Genotype on Host Response in Trichogramma (Hymenoptera: Trichogrammatidae)

The effects of experience, egg load, and wasp size on the response of four strains of Trichogramma nr. brassicae (Hymenoptera: Trichogrammatidae) to three economically important hosts, Heliothis punctigera (Noctuidae), Phthorimaea opercullela (Gelechiidae), and Sitotroga cereallela (Gelechiidae) were investigated. Experience effects due to rearing host and oviposition were considered in all combinations of host species. Both these effects altered response levels to low-ranked hosts but not to highly preferred hosts. Size was correlated with host response in some strains; larger females took more encounters with a host before accepting it, while egg load was not correlated with number of host encounters. However, egg load, rearing host, and size all affected acceptance of the low-ranked host S. cereallela. Females were more likely to accept this host if they were reared on S. cereallela, had a small size, and had a high egg load. Effects were consistent across strains, although overall acceptance levels differed among strains.     

Chewing Lice of Swan Geese (Anser cygnoides): New Host-Parasite Associations

Chewing lice (Phthiraptera) that parasitize the globally threatened swan goose Anser cygnoides have been long recognized since the early 19th century, but those records were probably biased towards sampling of captive or domestic geese due to the small population size and limited distribution of its wild hosts. To better understand the lice species parasitizing swan geese that are endemic to East Asia, we collected chewing lice from 14 wild geese caught at 3 lakes in northeastern Mongolia. The lice were morphologically identified as 16 Trinoton anserinum (Fabricius, 1805), 11 Ornithobius domesticus Arnold, 2005, and 1 Anaticola anseris (Linnaeus, 1758). These species are known from other geese and swans, but all of them were new to the swan goose. This result also indicates no overlap in lice species between older records and our findings from wild birds. Thus, ectoparasites collected from domestic or captive animals may provide biased information on the occurrence, prevalence, host selection, and host-ectoparasite interactions from those on wild hosts.     

Changes in morphological traits of the cabbage aphid (<Emphasis Type="Italic">Brevicoryne brassicae</Emphasis>) associated with the use of different host plants

The genetic and morphological differentiation of insect populations in relation to the use of different host plants is an important phenomenon that predates ecological specialisation and speciation in sympatric conditions. In this study, we describe the morphological variation of populations of Brevicoryne brassicae (Homoptera: Aphididae) associated with two host species, Brassica oleracea and Brassica campestris, which occur sympatrically in the highlands of Chiapas, Mexico. The study is aimed at obtaining evidence regarding phenotypic differentiation induced by, or associated with, the use of distinct but closely related host species. Seven morphological characters were measured in 696 wingless aphids collected from plants of the two host species at four localities. Morphological variation was summarised through principal components analysis (PCA). Sixty-two percent of morphological variation was explained by the first two PCs. The first component (PC1) was related to the general size of appendages, and PC2 was interpreted as the relationship between body size (body and leg size) and antenna length. Aphids growing on B. campestris were bigger than those collected from B. oleraceae. Significant differences between hosts were detected for PC1, whereas a significant effect of locality, host, and the interaction locality × host was detected for PC2. These results indicate that the average phenotype of B. brassicae individuals inhabiting different host-plant species differs as a consequence of the contrasting feeding environments the host species provide.     

One-year survey of a single Micronesian reef reveals extraordinarily rich diversity of <Emphasis Type="Italic">Symbiodinium</Emphasis> types in soritid foraminifera

Recent molecular studies of symbiotic dinoflagellates (genus Symbiodinium) from a wide array of invertebrate hosts have revealed exceptional fine-scale symbiont diversity whose distribution among hosts, regions and environments exhibits significant biogeographic, ecological and evolutionary patterns. Here, similar molecular approaches using the internal transcribed spacer-2 (ITS-2) region were applied to investigate cryptic diversity in Symbiodinium inhabiting soritid foraminifera. Approximately 1,000 soritid specimens were collected and examined during a 12-month period over a 40 m depth gradient from a single reef in Guam, Micronesia. Out of 61 ITS-2 types distinguished, 46 were novel. Most types found are specific for soritid hosts, except for three types (C1, C15 and C19) that are common in metazoan hosts. The distribution of these symbionts was compared with the phylotype of their foraminiferal hosts, based on soritid small subunit ribosomal DNA sequences, and three new phylotypes of soritid hosts were identified based on these sequences. Phylogenetic analyses of 645 host-symbiont pairings revealed that most Symbiodinium types associated specifically with a particular foraminiferal host genus or species, and that the genetic diversity of these symbiont types was positively correlated with the genetic diversity found within each of the three host genera. Compared to previous molecular studies of Symbiodinium from other locations worldwide, the diversity reported here is exceptional and suggests that Micronesian coral reefs are home to a remarkably large Symbiodinium assemblage.     

A hitchhiker’s guide to parasite transmission: The phoretic behaviour of feather lice

Transmission to new hosts is a fundamental challenge for parasites. Some species meet this challenge by hitchhiking on other, more mobile parasite species, a behaviour known as phoresis. For example, feather-feeding lice that parasitise birds disperse to new hosts by hitchhiking on parasitic louse flies, which fly between individual birds. Oddly, however, some species of feather lice do not engage in phoresis. For example, although Rock Pigeon (Columba livia) “wing” lice (Columbicola columbae) frequently move to new hosts phoretically on louse flies (Pseudolynchia canariensis), Rock Pigeon “body” lice (Campanulotes compar) do not. This difference in phoretic behaviour is puzzling because the two species of lice have very similar life cycles and are equally dependent on transmission to new hosts. We conducted a series of experiments designed to compare the orientation, locomotion and attachment capabilities of these two species of lice, in relation to louse flies. We show that wing lice use fly activity as a cue in orientation and locomotion, whereas body lice do not. We also show that wing lice are more capable of remaining attached to active flies that are walking, grooming or flying. The superior phoretic ability of wing lice may be related to morphological adaptations for life on wing feathers, compared to body feathers.