03341nas a2200145 4500008004100000245006700041210006700108260000900175300001400184490000600198520287300204100001903077700002503096856007403121 2007 eng d00aComparative tests of ectoparasite species richness in seabirds0 aComparative tests of ectoparasite species richness in seabirds c2007 a227 - 2480 v73 a

Background: The diversity of parasites attacking a host varies substantially among different hostspecies. Understanding the factors that explain these patterns of parasite diversity is critical to identifying the ecological principles underlying biodiversity. Seabirds (Charadriiformes, Pelecaniformes and Procellariiformes) and their ectoparasitic lice (Insecta: Phthiraptera) are ideal model groups in which to study correlates of parasite species richness. We evaluated the relative importance of morphological (body size, body weight, wingspan, bill length), life-history (longevity, clutch size), ecological (population size, geographical range) and behavioural (diving versus nondiving) variables as predictors of louse diversity on 413 seabird hosts species. Diversity was measured at the level of louse suborder, genus, and species, and uneven sampling of hosts was controlled for using literature citations as a proxy for sampling effort. Results: The only variable consistently correlated with louse diversity was host population size and to a lesser extent geographic range. Other variables such as clutch size, longevity, morphological and behavioural variables including body mass showed inconsistent patterns dependent on the method of analysis. Background: The diversity of parasites attacking a host varies substantially among different host species. Understanding the factors that explain these patterns of parasite diversity is critical to identifying the ecological principles underlying biodiversity. Seabirds (Charadriiformes, Pelecaniformes and Procellariiformes) and their ectoparasitic lice (Insecta: Phthiraptera) are ideal model groups in which to study correlates of parasite species richness. We evaluated the relative importance of morphological (body size, body weight, wingspan, bill length), life-history (longevity, clutch size), ecological (population size, geographical range) and behavioural (diving versus nondiving) variables as predictors of louse diversity on 413 seabird hosts species. Diversity was measured at the level of louse suborder, genus, and species, and uneven sampling of hosts was controlled for using literature citations as a proxy for sampling effort. Results: The only variable consistently correlated with louse diversity was host population size and to a lesser extent geographic range. Other variables such as clutch size, longevity, morphological and behavioural variables including body mass showed inconsistent patterns dependent on the method of analysis. Conclusion: The comparative analysis presented herein is (to our knowledge) the first to test correlates of parasite species richness in seabirds. We believe that the comparative data and phylogeny provide a valuable framework for testing future evolutionary hypotheses relating to the diversity and distribution of parasites on seabirds.

1 aHughes, Joseph1 aPage, Roderic, D. M. uhttp://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-7-22701827nas a2200169 4500008004100000245008900041210006900130260000900199300001600208490000700224520125900231100002401490700002501514700002001539700002501559856007301584 2007 eng d00aLack of host-dependent genetic structure in ectoparasites of Calonectris shearwaters0 aLack of hostdependent genetic structure in ectoparasites of Calo c2007 a5204 - 52150 v163 a

We compared patterns of mitochondrial DNA (mtDNA) differentiation in three hostspecific lice (Halipeurus abnormis, Austromenopon echinatum and Saemundssonia peusi) and one generalist flea (Xenopsylla gratiosa), parasitizing 22 colonies of Cory’s and Cape Verde shearwater (Calonectris). The shearwater hosts show distinct phylogeographic structure corresponding to the three taxa Calonectris d. diomedea, C. d. borealis, and C. edwardsii. The host-specific lice appeared undifferentiated among the three Calonectris taxa, whereas the more generalist flea displayed significant levels of population differentiation. Neithergenetic distances among host populations, nor their spatial distribution explained the patterns of genetic variability observed in the ectoparasites. The lack of differentiation among lice is unexpected, given that previous work has found evidence of cospeciation between procellariiform seabirds and their lice, and lice typically have an elevated rate of mtDNA evolution with respect to their hosts. Our results suggest that either rates of evolution in seabird lice are not always as high as previously thought, or that the magnitude of movement of lice between seabird hosts has been substantially underestimated.

1 aGómez-Díaz, Elena1 aGonzález-Solís, J.1 aPeinado, M., A.1 aPage, Roderic, D. M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/47260.pdf00494nas a2200121 4500008004100000245013200041210006900173260000900242300001600251490000700267100002500274856007300299 2000 eng d00aComparative analysis of secondary structure of insect mitochondrial small subunit ribosomal RNA using maximum weighted matching0 aComparative analysis of secondary structure of insect mitochondr c2000 a3839 - 38450 v281 aPage, Roderic, D. M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/47167.pdf02283nas a2200301 4500008004100000245011500041210006900156260000900225300001400234490000700248520139400255653001601649653001601665653001701681653000801698653000901706653001201715653001901727653001701746653001501763653001301778100001901791700002001810700002301830700002301853700002501876856008001901 2007 eng d00aMultiple cophylogenetic analyses reveal frequent cospeciation between Pelecaniform birds and Pectinopygus lice0 aMultiple cophylogenetic analyses reveal frequent cospeciation be c2007 a232 - 2510 v563 a

Lice in the genus Pectinopygus parasitize a single order of birds (Pelecaniformes). To examine the degree of congruence between the phylogenies of 17 Pectinopygus species and their pelecaniform hosts, sequences from mitochondrial 12S rRNA, 16S rRNA, COI, and nuclear wingless and EF1-α genes (2290 nucleotides) and frommitochondrial 12S rRNA, COI, and ATPases 8 and 6 genes (1755 nucleotides) were obtained for the lice and the birds, respectively. Louse data partitions were analyzed for evidence of incongruence and evidence of long-branch attraction prior to cophylogenetic analyses. Host-parasite coevolution was studied by different methods: TreeFitter, TreeMap, ParaFit, likelihood-ratio test, data-based parsimony method, and correlation of coalescence times. All methods agree that there has been extensive cospeciation in this host-parasite system, but the results are sensitive to the selection of different phylogenetic hypotheses and analytical methods for evaluating cospeciation. Perfect congruence between phylogenies is not found in this association, probably as a result of occasional host switching by the lice. Errors due to phylogenetic reconstruction methods, incorrect or incomplete taxon sampling, or to different loci undergoing different evolutionary histories cannot be rejected, thus emphasizing the need for improved cophylogenetic methodologies.

10aCoalescence10acoevolution10acospeciation10aILD10alice10aParaFit10aPelecaniformes10aPhthiraptera10aTreeFitter10aTreeMap.1 aHughes, Joseph1 aKennedy, Martyn1 aJohnson, Kevin, P.1 aPalma, Ricardo, L.1 aPage, Roderic, D. M. uhttps://phthiraptera.myspecies.info/sites/phthiraptera.info/files/46850.pdf00553nas a2200157 4500008004100000245005600041210005600097260003800153300001400191100002100205700002400226700002700250700002000277700002500297856007300322 2003 eng d00aCophylogeny between pocket gophers and chewing lice0 aCophylogeny between pocket gophers and chewing lice bUniversity of Chicago Pressc2003 a195 - 2201 aHafner, Mark, S.1 aDemastes, James, W.1 aSpradling, Theresa, A.1 aReed, David, L.1 aPage, Roderic, D. M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/46177.pdf00619nas a2200145 4500008004100000245009100041210006900132260005700201300001300258100002500271700002400296700001600320700001500336856012200351 2003 eng d00aA statistical perspective for reconstructing the history of host-parasite associations0 astatistical perspective for reconstructing the history of hostpa aChicago, IllinoisbUniversity of Chicago Pressc2003 a93 - 1191 aPage, Roderic, D. M.1 aHuelsenbeck, J., P.1 aRannala, B.1 aLarget, B. uhttps://phthiraptera.myspecies.info/content/statistical-perspective-reconstructing-history-host-parasite-associations02763nas a2200337 4500008004100000245009300041210006900134260000900203300001400212490000700226520170000233653001201933653001901945653002501964653003301989653001102022653002602033653002002059653001402079653001502093653003702108653004502145653001402190100002302204700002802227700002302255700002702278700002502305700002202330856007302352 2003 eng d00aDramatically elevated rate of mitochondrial substitution in lice (Insecta: Phthiraptera)0 aDramatically elevated rate of mitochondrial substitution in lice c2003 a231 - 2420 v263 a

Few estimates of relative substitution rates, and the underlying mutation rates, exist between mitochondrial and nuclear genes in insects. Previous estimates for insects indicate a 2-9 times faster substitution rate in mitochondrial genes relative to nuclear genes. Here we use novel methods for estimating relative rates of substitution, which incorporate multiple substitutions, and apply these methods to a group of insects (lice, Order: Phthiraptera). First, we use a modification of copath analysis (branch length regression) to construct independent comparisons of rates, consisting of each branch in a phylogenetic tree. The branch length comparisons use maximum likelihood models to correct for multiple substitution. In addition, we estimate codon-specific rates under maximum likelihood for the different genes and compare these values. Estimates of the relative synonymous substitution rates between a mitochondrial (COI) and nuclear (EF-1alpha) gene in lice indicate a relative rate of several 100 to 1. This rapid relative mitochondrial rate (>100 times) is at least an order of magnitude faster than previous estimates for any group of organisms. Comparisons using the same methods for another group of insects (aphids) reveals that this extreme relative rate estimate is not simply attributable to the methods we used, because estimates from aphids are substantially lower. Taxon sampling affects the relative rate estimate, with comparisons involving more closely related taxa resulting in a higher estimate. Relative rate estimates also increase with model complexity, indicating that methods accounting for more multiple substitution estimate higher relative rates.

10aanimals10aBird parasites10aEvolution, Molecular10aLice/classification/genetics10amammal10aMitochondria/genetics10aModels, Genetic10aPhylogeny10apolymerase10aResearch Support, Non-U.S. Gov't10aResearch Support, U.S. Gov't, Non-P.H.S.10avariation1 aJohnson, Kevin, P.1 aCruickshank, Robert, H.1 aAdams, Richard, J.1 aSmith, Vincent, Stuart1 aPage, Roderic, D. M.1 aClayton, Dale, H. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/44823.pdf03421nam a2200121 4500008004100000245006000041210005700101260005500158300001500213520293800228100002503166856010803191 2003 eng d00aTangled trees. Phylogeny, cospeciation, and coevolution0 aTangled trees Phylogeny cospeciation and coevolution aChicago, LondonbUniversity of Chicago Pressc2003 ax + 350 pp3 a

In recent years, the use of molecular data to build phylogenetic trees and sophisticated computer-aided techniques to analyze them have led to a revolution in the study of cospeciation. Tangled Trees provides an up-to-date review and synthesis of current knowledge about phylogeny, cospeciation, and coevolution. The opening chapters present various methodological and theoretical approaches, ranging from the well-known parsimony approach to "jungles" and Bayesian statistical models. Then a series of empirical chapters discusses detailed studies of cospeciation involving vertebrate hosts and their parasites, including nematodes, viruses, and lice. Tangled Trees will be welcomed by researchers in a wide variety of fields, from parasitology and ecology to systematics and evolutionary biology.

Contributors: Sarah Al-Tamimi, Michael A. Charleston, Dale H. Clayton, James W. Demastes, Russell D. Gray, Mark S. Hafner, John P. Huelsenbeck, J.-P. Hugot, Kevin P. Johnson, Peter Kabat, Bret Larget, Joanne Martin, Yannis Michalakis, Roderic D. M. Page, Ricardo L. Palma, Adrian M. Paterson, Susan L. Perkins, Andy Purvis, Bruce Rannala, David L. Reed, Fredrik Ronquist, Theresa A. Spradling, Jason Taylor, Michael Tristem

Content:

1. Introduction
Roderic D. M. Page
2. Parsimony analysis of coevolving species associations
Fredrik Ronquist
3. Lizards, Malaria, and Jungles in the Caribbean
Michael Charleston and Susan L. Perkins
4. A Statistical Perspective for Reconstructing the History of Host-Parasite Associations
John P. Huelsenbeck, Bruce Rannala, and Bret Larget
5. Population Genetics and Cospeciation: From Process to Pattern
Bruce Rannala and Yannis Michalakis
6. New evidence for hystricognath rodent monophyly from the phylogeny of their pinworms
J.-P. Hugot
7. Cospeciation and horizontal transmission rates in the murine leukaemia-related retroviruses
Joanne Martin, Peter Kabat, and Michael Tristem
8. Cophylogeny between pocket gophers and chewing lice
Mark S. Hafner, James W. Demastes, Theresa A. Spradling, and David L. Reed
9. The Effects of Spatial and Temporal Scale on analyses of cophylogeny
James W. Demastes, Theresa A. Spradling, and Mark S. Hafner
10. Have mammals and their chewing lice diversified in parallel?
Jason Taylor and Andy Purvis
11. Coevolutionary history of ecological replicates: Comparing phylogenies of wing and body lice to Columbiform hosts
Kevin P. Johnson and Dale H. Clayton
12. Drowning on arrival, missing the boat and x-events: how likely are sorting events?
Adrian M. Paterson, Ricardo L. Palma, Russell D. Gray
13. The ecological basis of coevolutionary history
Dale H. Clayton, Sarah Al-Tamimi and Kevin P. Johnson

1 aPage, Roderic, D. M. uhttps://www.researchgate.net/publication/280881048_Tangled_Trees_Phylogeny_Cospeciation_and_Coevolution00741nas a2200157 4500008004100000245008800041210006900129260005500198300001400253520008600267100002500353700002500378700002300403700002200426856013500448 2003 eng d00aDrowning on arrival, missing the boat, and x-events: how likely are sorting events?0 aDrowning on arrival missing the boat and xevents how likely are  aChicago, LondonbUniversity of Chicago Pressc2003 a287 - 3093 a

Full version and a slightly condensed version of the paper are attached.

1 aPage, Roderic, D. M.1 aPaterson, Adrian, M.1 aPalma, Ricardo, L.1 aGray, Russell, D. uhttps://www.researchgate.net/publication/228701470_Drowning_on_arrival_missing_the_boat_and_x-events_how_likely_are_sorting_events00566nas a2200133 4500008004100000245011800041210006900159260004700228300001400275100002500289700002300314700002200337856007300359 2003 eng d00aCoevolutionary history of ecological replicates: comparing phylogenies of wing and body lice to Columbiform hosts0 aCoevolutionary history of ecological replicates comparing phylog aChicagobUniversity of Chicago Pressc2003 a262 - 2861 aPage, Roderic, D. M.1 aJohnson, Kevin, P.1 aClayton, Dale, H. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/41862.pdf00507nas a2200145 4500008004100000245005100041210004700092260004700139300001400186100002500200700002200225700001800247700002300265856007300288 2003 eng d00aThe ecological basis of coevolutionary history0 aecological basis of coevolutionary history aChicagobUniversity of Chicago Pressc2003 a310 - 3411 aPage, Roderic, D. M.1 aClayton, Dale, H.1 aAl-Tamimi, S.1 aJohnson, Kevin, P. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/41863.pdf02279nas a2200169 4500008004100000245007100041210006900112260000900181300001200190490000700202520173400209100002301943700002301966700002501989700002202014856007302036 2003 eng d00aWhen do parasites fail to speciate in response to host speciation?0 aWhen do parasites fail to speciate in response to host speciatio c2003 a37 - 470 v523 a

Abstract.— Cospeciation generally increases the similarity between host and parasite phylogenies. Incongruence between host and parasite phylogenies has previously been explained in terms of host switching, sorting, and duplication events. Here, we describe an additional process, failure of the parasite to speciate in response to host speciation, that may be im- portant in some host–parasite systems. Failure to speciate is likely to occur when gene flow among parasite populations is much higher than that of their hosts. We reconstructed trees from mitochondrial and nuclear DNA sequences for pigeons and doves (Aves: Columbiformes) and their feather lice in the genus Columbicola (Insecta: Phthiraptera). Although com- parisons of the trees from each group revealed a significant amount of cospeciation, there was also a significant degree of incongruence. Cophylogenetic analyses generally indicated that host switching may be an important process in the history of this host–parasite association. Using terminal sister taxon comparisons, we also identified three apparent cases where the host has speciated but the associated parasite has not. In two of these cases of failure to speciate, these comparisons involve allopatric sister taxa of hosts whose lice also occur on hosts sympatric with both of the allopatric sisters. These additional hosts for generalist lice may promote gene flow with lice on the allopatric sister species. Relative rate comparisons for the mitochondrial cytochrome oxidase I gene indicate that molecular substitution occurs about 11 times faster in lice than in their avian hosts. [Coevolution; Columbiformes; cospeciation; lice; molecular phylogeny; Phthiraptera.]

1 aJohnson, Kevin, P.1 aAdams, Richard, J.1 aPage, Roderic, D. M.1 aClayton, Dale, H. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/41850.pdf02114nas a2200193 4500008004100000245006300041210006300104260000900167300001400176490000700190520153200197653001901729653001601748653000901764100002701773700002501800700002301825856007201848 2004 eng d00aData incongruence and the problem of avian louse phylogeny0 aData incongruence and the problem of avian louse phylogeny c2004 a239 - 2590 v333 a

Recent studies based on different types of data (i.e. morphological and molecular) have supported conflicting phylogenies for the genera of avian feather lice (Ischnocera: Phthiraptera). We analyse new and published data from morphology and from mitochondrial (12S rRNA and COI) and nuclear (EF1-α) genes to explore the sources of this incongruence and explain these conflicts. Character convergence, multiple substitutions at high divergences, and ancient radiation over a short period of time have contributed to the problem of resolving louse phylogeny with the data currently available. We show that apparent incongruence between the molecular datasets is largely attributable to rate variation and nonstationarity of base composition. In contrast, highly significant character incongruence leads to topological incongruence between the molecular and morphological data. We consider ways in which biases in the sequence data could be misleading, using several maximum likelihood models and LogDet corrections. The hierarchical structure of the data is explored using likelihood mapping and SplitsTree methods. Ultimately, we concede there is strong discordance between the molecular and morphological data and apply the conditional combination approach in this case. We conclude that higher level phylogenetic relationships within avian Ischnocera remain extremely problematic. However, consensus between datasets is beginning to converge on a stable phylogeny for avian lice, at and below the familial rank.

10aclassification10acoevolution10amtax1 aSmith, Vincent, Stuart1 aPage, Roderic, D. M.1 aJohnson, Kevin, P. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/6699.pdf01578nas a2200265 4500008004100000245012100041210006900162260000900231300001400240490000700254520074900261653001701010653000901027653001701036653000801053653001301061100002501074700002801099700001901127700002401146700002001170700002301190700002701213856007201240 2004 eng d00aPhylogeny of "Philoceanus complex" seabird lice (Phthiraptera: Ischnocera) inferred from mitochondrial DNA sequences0 aPhylogeny of Philoceanus complex seabird lice Phthiraptera Ischn c2004 a633 - 6520 v303 a

The Philoceanus complex is a large assemblage of lice that parasitise procellariiform seabirds (petrels, albatrosses, and their relatives). We obtained mitochondrial 12S rRNA and cytochrome oxidase I DNA sequences from 39 species from diverse hosts and localities. Resolution of deeper relationships between genera was limited, however there is evidence for two major clades, one hosted by albatrosses, the other by petrels. Based on our results, the genera hosted by albatrosses are excellent candidates for detailed analysis of cospeciation. Our results also suggest that a previous estimate of a 5-fold difference in the relative rate of sequence evolution in lice and their avian hosts is an artefact of limited taxonomic sampling.

10acospeciation10alice10aPhthiraptera10aRNA10aseabirds1 aPage, Roderic, D. M.1 aCruickshank, Robert, H.1 aDickens, Megan1 aFurness, Robert, W.1 aKennedy, Martyn1 aPalma, Ricardo, L.1 aSmith, Vincent, Stuart uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/6675.pdf01607nas a2200241 4500008004100000245009600041210006900137260000900206300001400215490000700229520088200236653000901118653001201127653001401139653002301153653000801176653002401184653000901208100002501217700002801242700002301270856007201293 2002 eng d00aLouse (Insecta: Phthiraptera) mitochondrial 12S rRNA secondary structure is highly variable0 aLouse Insecta Phthiraptera mitochondrial 12S rRNA secondary stru c2002 a361 - 3690 v113 a

Lice are ectoparasitic insects hosted by birds and mammals. Mitochondrial 12S rRNA sequences obtained from lice show considerable length variation and are very difficult to align. We show that the louse12S rRNA domain III secondary structure displays considerable variation compared to other insects, in both the shape and number of stems and loops. Phylogenetic trees constructed from tree edit distances between louse 12S rRNA structures do not closely resemble trees constructed from sequence data, suggesting that at least some of this structural variation has arisen independently in different louse lineages. Taken together with previous work on mitochondrial gene order and elevated rates of substitution in louse mitochondrial sequences, the structural variation in louse 12S rRNA confirms the highly distinctive nature of molecular evolution in these insects.

10alice10ametrics10amolecular10amultiple alignment10aRNA10asecondary structure10atree1 aPage, Roderic, D. M.1 aCruickshank, Robert, H.1 aJohnson, Kevin, P. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/6680.pdf01416nas a2200133 4500008004100000245013000041210006900171260000900240300001200249490000700261520091900268100002501187856007001212 1991 eng d00aClocks, clades, and cospeciation: comparing rates of evolution and timing of cospeciation events in host-parasite assemblages0 aClocks clades and cospeciation comparing rates of evolution and  c1991 a77 - 840 v403 a

A recently described test for a consistent pattern of timing of cospeciation between host and parasite taxa uses the correlation between observed genetic distance matrices for the hosts and parasites. In this paper I argue that this method can be improved by comparing trees rather than distance matrices. In the special case of a molecular clock, the test compares cluster heights in dendrograms for the hosts and the parasites. Because tree topology alone can cause a significant correlation between the cluster heights, a randomization test is described that allows for this autocorrelation. The degree of autocorrelation is a function of tree topology, increasing as the trees become more unbalanced. Balanced trees offer more sensitive tests of hypotheses concerning timing of cospeciation. [Bootstrap; cladistics; cospeciation; dendrograms; molecular clock; randomization tests; rates of evolution.]

1 aPage, Roderic, D. M. uhttps://www.jstor.org/stable/2992256?seq=1#page_scan_tab_contents02224nas a2200289 4500008004100000245004800041210004700089260000900136300001400145490000700159520146100166653001701627653001601644653001501660653001401675653001101689653001501700653001301715653001801728653001401746653001101760653001901771100002501790700002201815700002501837856007201862 1996 eng d00aLice and cospeciation: a response to Barker0 aLice and cospeciation a response to Barker c1996 a213 - 2180 v263 a

"The student who intends working on the Mallophaga should take warning that he will be tried almost beyond endurance by the paradoxes and complexities which beset his subject but he will also find, in the dual and inter-related aspect of insect and bird, an infinite fascination." (Rothschild & Clay, 1952: pp. 156-157). The study of host louse coevolution will benefit greatly from the phylogenetic perspective offered by recent advantages in molecular systematics. However, in order to make best use of phylogenies we need to appreciate the complexities of the possible relations between host and parasite phylogeny. At the same time, the very complexity of louse-host systems has a potentially useful consequence; the presence of multiple lineages of lice on the same hosts allow for replicated tests of coevolutionary hypotheses. For example, if a number of louse clades infest the same host clade but some lice show more cospeciation than others, we might ask whether there are features of louse biology that correlate with this difference in host tracking fidelity. It may further be possible to ascertain the relative importance of these features in ecological time through controlled transfer experiments. By beginning to appreciate "the paradoxes and complexities" of host-louse evolution, lice may offer us not only "infinite fascination" but also a chance to address important questions in coevolution.

10aassociations10acoevolution10acongruence10aEvolution10agopher10aMallophaga10aparasite10aPhilopteridae10aPhylogeny10apocket10arock wallabies1 aPage, Roderic, D. M.1 aClayton, Dale, H.1 aPaterson, Adrian, M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/2977.pdf02159nas a2200469 4500008004100000245014100041210006900182260000900251300001400260490000700274520073700281653001301018653001601031653001701047653001001064653001401074653001701088653000801105653001401113653001601127653001401143653001101157653001101168653001801179653000901197653001301206653001501219653001401234653001601248653001401264653001101278653001301289653001501302653001801317653001801335653001301353653000901366653001001375653001901385100002501404856026001429 1996 eng d00aTemporal congruence revisited: comparison of mitochondrial DNA sequence divergence in cospeciating pocket gophers and their chewing lice0 aTemporal congruence revisited comparison of mitochondrial DNA se c1996 a151 - 1670 v453 a

Molecular phylogenies can be used to test hypotheses of cospeciation between hosts and parasites by comparing both cladistic relationships and branch lengths. Molecular data can also help discriminate between competing reconstructions of the history of the host-parasite association. Methods for comparing sequence divergence in hosts and parasites are described and applied to data for pocket gophers and their chewing lice. The hypothesis of cospeciation between these two clades is strongly supported. The lengths of homologous branches in the gopher and louse phylogenies are positively correlated, but there is little support for the hypothesis that lice are evolving an order of magnitude faster than are their hosts.

10aanalysis10aassemblages10aChewing lice10aclock10acomponent10acospeciation10adna10aEvolution10ageomydoecus10aGeomyidae10aGeomys10agopher10ahost parasite10alice10alineages10aMallophaga10amolecular10aOrthogeomys10aPhylogeny10apocket10asequence10aspeciation10asubstitutions10athomomydoecus10athomomys10atree10atrees10atrichodectidae1 aPage, Roderic, D. M. uhttp://oup.silverchair-cdn.com/oup/backfile/Content_public/Journal/sysbio/45/2/10.1093/sysbio/45.2.151/2/45-2-151.pdf?Expires=1485178998&Signature=Ot51qtPL0yKCXIoYGaXyDHFhW32zncDUkaJP357f8CU5td~btGv15S2jmYH8DOK8nSGL1a66mMoOQflCg02Fs47sv~qKstzYpYsm6kbvwqnA00984nas a2200229 4500008004100000245004200041210004100083260000900124300001400133490000700147520038500154653001600539653001700555653001500572653001500587653001700602653001400619653001400633653001000647100002500657856007200682 1993 eng d00aParasites, phylogeny and cospeciation0 aParasites phylogeny and cospeciation c1993 a499 - 5060 v233 a

Host-parasite assemblages offer exciting possibilities for the comparative study of rates of speciation and evolution in different organisms. The basis for such studies must be a phylogenetic analysis of the host-parasite association. Some recent developments in this field are briefly reviewed and illustrated using phylogenies for pocket gophers and their chewing lice.

10aassemblages10aBiogeography10acladistics10acongruence10acospeciation10aEvolution10aPhylogeny10atrees1 aPage, Roderic, D. M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/2753.pdf02175nas a2200337 4500008004100000245008100041210006900122260000900191300001400200490000700214520127200221653001301493653000901506653001601515653001601531653001601547653001401563653000901577653001601586653001701602653001401619653001301633653001401646653000901660653001101669100001801680700002101698700002501719700002001744856007301764 2000 eng d00aComparative body size relationships in pocket gophers and their chewing lice0 aComparative body size relationships in pocket gophers and their  c2000 a239 - 2490 v703 a

In this paper, we use the method of independent contrasts to study body size relationships between pocket gophers and their chewing lice, a host-parasite system in which both host and parasite pilylogenies are well studied. The evolution of body size of chewing lice appears to be dependent only on the body size of their hosts, which confirms the 1991 findings of Harvey and Keymer. We show that there is a positive relationship between body size and hair-shaft diameter in pocket gophers, and that there is also a positive relationship between body size and head-groove width in chewing lice. Finally, we show a positive relationship between gopher hair-shaft diameter and louse head-groove width. We postulate that changes in body size of chewing lice are driven by a mechanical relationship between the parasite's head-groove dimension and the diameter of the hairs of its host. Louse species living on larger host species may be larger simply because their hosts have thicker hairs, which requires that the lice have a wider head groove. Our study of gopher hair-shaft diameter and louse head-groove dimensions suggest that there is a 'lock-and-key' relationship between these two anatomical features. (C) 2000 The Linnean Society of London.

10aanalysis10abody10acoevolution10acomparative10acovariation10aEvolution10ahost10aindependent10alife history10aNematodes10aparasite10aPhylogeny10asize10atraits1 aMorand, Serge1 aHafner, Mark, S.1 aPage, Roderic, D. M.1 aReed, David, L. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/37965.pdf00493nas a2200121 4500008004100000245006500041210006400106260004100170100002500211700001500236700001500251856010500266 2003 eng d00aHave mammals and their chewing lice diversified in parallel?0 aHave mammals and their chewing lice diversified in parallel aChicagobUniversity of Chicagoc20031 aPage, Roderic, D. M.1 aTaylor, J.1 aPurvis, A. uhttps://phthiraptera.myspecies.info/content/have-mammals-and-their-chewing-lice-diversified-parallel00999nas a2200145 4500008004100000245005400041210005300095260000900148300001400157490000700171520052600178100002000704700002500724856010400749 2000 eng d00aRadCon: phylogenetic tree comparison and consesus0 aRadCon phylogenetic tree comparison and consesus c2000 a486 - 4870 v163 a

RadCon is a Macintosh program for manipulating and analysing phylogenetic trees. The program can determine the Cladistic Information Content of individual trees, the stability of leaves across a set of bootstrap trees, produce the strict basic Reduced Cladistic Consensus profile of a set of trees and convert a set of trees into its matrix representation for supertree construction. Availability: The program is free and available at http://taxonomy.zoology.gla.ac.uk/ approximately jthorley/radcon/radcon.html.

1 aThorley, J., L.1 aPage, Roderic, D. M. uhttps://www.researchgate.net/publication/12448127_RadCon_Phylogenetic_tree_comparison_and_consensus01924nas a2200301 4500008004100000245009300041210006900134260004200203300001400245520102800259653001401287653001301301653000901314653001601323653002501339653001901364653001401383653001801397653001401415653000901429100002501438700002101463700001901484700002101503700001801524700001201542856006801554 1996 eng d00aMolecular phylogenies and host-parasite cospeciation: gophers and lice as a model system0 aMolecular phylogenies and hostparasite cospeciation gophers and  aOxfordbOxford University Pressc1996 a255 - 2703 a

Recent methodological advances permit a rigorous comparison of phylogenetic trees parasites for hosts and their parasites to determine the extent to which these groups have cospeciated through evolutionary time. In cases where significant levels of cospeciation are indicated, comparison of amounts of evolutionary change  that have accumulated along analogous branches in the host and parasite trees provides a direct assessment of relative rates of evolution in the two groups. For such a comparison to be meaningful, the features compared in the hosts and parasites should be genetically based, evolutionary homologous, and should evolve in a roughly time-dependent fashion within each group. Nucleotide sequences encoding homologous genes in hosts and parasites are an ideal source of data for comparative studies of evotionary rates. Recent studies of pocket gophers and their lice are used to illustrate the variety of questions that can be addressed through phylogenetic study of host-parasite systems.

10aEvolution10agenetics10alice10aMastication10anucleotide sequences10aParasite hosts10aparasites10aPhylogenetics10aPhylogeny10aTaxa1 aPage, Roderic, D. M.1 aHafner, Mark, S.1 aHarvey, P., H.1 aBrown, A., J. L.1 aSmith, J., M.1 aNee, S. uhttps://www.jstor.org/stable/56126?seq=1#page_scan_tab_contents02513nas a2200277 4500008004100000245013100041210006900172260000900241300001400250490000700264520161000271653001501881653002801896653000901924653002601933653002501959653001701984653001402001100002802015700002302043700002702066700002302093700002202116700002502138856007202163 2001 eng d00aPhylogenetic analysis of partial sequences of Elongation Factor 1alpha identifies major groups of lice (Insecta: Phthiraptera)0 aPhylogenetic analysis of partial sequences of Elongation Factor  c2001 a202 - 2150 v193 a

As a first attempt to use molecular data to resolve the relationships between the four suborders of lice and within the suborder Ischnocera, we sequenced a 347-bp fragment of the elongation factor 1 gene of 127 lice (Insecta: Phthiraptera) as well as outgroup taxa from the order Psocoptera. A number of well-supported monophyletic groups were found but the relationships among many of these groups could not be resolved. While it is probable that multiple substitutions at high divergences and ancient radiation over a short period of time have contributed to the problem, we attribute most of this lack of resolution to the high ratio of taxa to characters. Nevertheless, the sequence data unequivocally support a number of important relationships that are at variance with the conclusions of morphological taxonomy. These include the sister group relationship of Chelopistes and Oxylipeurus, two lice occupying different ecological niches on the same host, which have previously been assigned to different families. These results provide evidence in support of the hypothesis that lice have speciated in situ on the host in response to niche specialization and that this has given rise to convergent morphologies in the lice of different host groups which share similar ecological niches. We discuss our attempts to overcome the limitations of this large data set, including the use of leaf stability analysis, a new method for analyzing the stability of taxa in a phylogenetic tree, and examine a number of hypotheses of relationships based on both traditional taxonomy and host associations.

10aIschnocera10aleaf stability analysis10alice10amolecular systematics10aniche specialization10aPhthiraptera10aPhylogeny1 aCruickshank, Robert, H.1 aJohnson, Kevin, P.1 aSmith, Vincent, Stuart1 aAdams, Richard, J.1 aClayton, Dale, H.1 aPage, Roderic, D. M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/1204.pdf00440nam a2200121 4500008004100000245004900041210004800090260003800138300000800176100002500184700001900209856009000228 1998 eng d00aMolecular Evolution. A phylogenetic Approach0 aMolecular Evolution A phylogenetic Approach aCambridgebUniversity Pressc1998 a3461 aPage, Roderic, D. M.1 aHolmes, E., C. uhttps://phthiraptera.myspecies.info/content/molecular-evolution-phylogenetic-approach01227nas a2200145 4500008004100000245006200041210006100103260000900164300001400173490000700187520072900194100002500923700002800948856010500976 1998 eng d00aTrees within trees: phylogeny and historical associations0 aTrees within trees phylogeny and historical associations c1998 a356 - 3590 v133 a

The association between two or more lineages over evolutionary time is a recurrent theme spanning several different fields within biology, from molecular evolution to coevolution and biogeography. In each `historical association', one lineage is associated with another, and can be thought of as tracking the other over evolutionary time with a greater or lesser degree of fidelity. Examples include genes tracking organisms, parasites tracking hosts and organisms tracking geological and geographical changes. Parallels among these problems raise the tantalizing prospect that each is a special case of a more general problem, and that a single analytical tool can be applied to all three kinds of association.

1 aPage, Roderic, D. M.1 aCharleston, Michael, A. uhttps://phthiraptera.myspecies.info/content/trees-within-trees-phylogeny-and-historical-associations01953nas a2200265 4500008004100000245011400041210006900155260000900224300001600233490000700249520116100256653000901417653001401426653002301440653001401463653000901477653001401486653001601500653000901516100002501525700002201550700001801572700002501590856007201615 1996 eng d00aDoes behavior reflect phylogeny in cave swiftlets (Aves: Apodidae)? A test using cytochrome b mtDNA sequences0 aDoes behavior reflect phylogeny in cave swiftlets Aves Apodidae  c1996 a7091 - 70960 v933 a

[INCOMPLETE: Add alternate journal title] Swiftlets are small insectivorous birds, many of which nest in caves and are known to echolocate. Due to a lack of distinguishing morphological characters, the taxonomy of swiftlets is primarily based on the presence or absence of echolocating ability, together with nest characters. To test the reliability of these behavioral characters, we constructed an independent phylogeny using cytochrome b mitochondrial DNA sequences from swiftlets and their relatives. This phylogeny is broadly consistent with the higher classification of swifts but does not support the monophyly of swiftlets. Echolocating swiftlets (Aerodramus) and the nonecholocating ''giant swiftlet'' (Hydrochous gigas) group together, but the remaining nonecholocating swiftlets belonging to Collocalia are not sister taxa to these swiftlets. While echolocation may be a synapomorphy of Aerodramus (perhaps secondarily lost in Hydrochous), no character of Aerodramus nests showed a statistically significant fit to the molecular phylogeny, indicating that nest characters are not phylogenetically reliable in this group.

10abird10aEvolution10amaximum likelihood10amolecular10anest10astructure10asystematics10atree1 aLee, Patricia, L. M.1 aClayton, Dale, H.1 aGriffiths, R.1 aPage, Roderic, D. M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0767.pdf01212nas a2200145 4500008004100000245006000041210006000101260000900161300001200170490000700182520074100189100002500930700002800955856008300983 1997 eng d00aReconciled trees and incongruent gene and species trees0 aReconciled trees and incongruent gene and species trees c1997 a57 - 700 v373 a

We present a method for visualising and quantifying the relationship between a pair of gene and species trees that constructs a third tree termed a reconciled tree. Given a gene tree and a species tree the reconciled tree represents the history of the gene tree embedded within the species tree. The reconciled tree is constructed from one or more subtrees of the species tree, and contains the gene tree as a subtree. The relationship between the gene tree and the species tree can be expressed in terns of the number of gene duplications and gene losses required to construct the reconciled tree. This number can be used as an optimality criterion for selecting the species tree that best accounts for the observed gene tree.

1 aPage, Roderic, D. M.1 aCharleston, Michael, A. uhttps://pdfs.semanticscholar.org/be60/40228f09daddc2001254850f4b54936dc0c0.pdf00436nas a2200121 4500008004100000245008000041210006900121260000900190300001400199490000700213100002500220856006900245 1990 eng d00aTemporal congruence and cladistic analysis of biogeography and cospeciation0 aTemporal congruence and cladistic analysis of biogeography and c c1990 a205 - 2260 v391 aPage, Roderic, D. M. uhttp://phthiraptera.info/sites/phthiraptera.info/files/38841.pdf01249nas a2200133 4500008004100000245008200041210006900123260000900192300001400201490000700215520079800222100002501020856007001045 1988 eng d00aQuantitive cladistic biogeography: constructing and comparing area cladograms0 aQuantitive cladistic biogeography constructing and comparing are c1988 a254 - 2700 v373 a

This paper describes methods for constructing and comparing area cladograms. Three different interpretations (Assumptions 0, 1 and 2) of widespread taxa and redundant distributions are outlined. Algorithms for Assumptions 1 and 2 are presented, and a measure of relative parsimony for area cladograms is discussed. By using quantitative measures of the similarity between two or more area cladograms it is possible to test statistically hypotheses of biogeographic congruence. The algorithms described in this paper have been implemented in a computer program available from the author. Rosen's (1978, 1979) and Cracraft's (1986) data are reanalyzed using this software. [Area cladograms; biogeography; cladistics; comparing trees; consensus methods; vicariance biogeography.]

1 aPage, Roderic, D. M. uhttps://www.jstor.org/stable/2992372?seq=1#page_scan_tab_contents01570nas a2200181 4500008004100000245008300041210006900124260000900193300001400202490000600216520098000222100002501202700002501227700001901252700002301271700002201294856007201316 1998 eng d00aA different tempo of mitochondrial evolution in birds and their parasitic lice0 adifferent tempo of mitochondrial evolution in birds and their pa c1998 a276 - 2930 v93 a

A phylogeny for the lice (Insecta: Phthiraptera: genus Dennyus) parasitic on swiftlets (Aves: Collocalliinae) was constructed based on mitochondrial cytochrome b DNA sequences. This phylogeny is congruent with previous phenetic analyses of morphometric data for the lice. Comparison with a prevoius obtained phylogeny for the hosts indicates some degree of cospeciation. These cospeciation rates are used to compare relative rates of evolution in the birds and their lice for the same segment of cytochrome b gene. Cytochrome b is evolving 2-3 times more rapidly in the lice than the bird, and louse cytochrome b is highly divergent compared to most other insects. Although generation time has been suggested as an explanation for the disparity in evolutionary rates between lice and their hosts, we suggest that the small effective population sizes of lice coupled with founder events occuring during transmission to new host individuals may be an important factor.

1 aPage, Roderic, D. M.1 aLee, Patricia, L. M.1 aBecher, Anette1 aGriffiths, Richard1 aClayton, Dale, H. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0155.pdf01316nas a2200145 4500008004100000245009700041210006900138260000900207300001400216490000600230520080900236100002501045700002801070856007201098 1997 eng d00aFrom gene to organismal phylogeny: reconciled trees and the gene tree / species tree problem0 aFrom gene to organismal phylogeny reconciled trees and the gene  c1997 a231 - 2400 v73 a

The process of gene duplication, loss and lineage sorting can result in incongruence between the phylogenies of genes and those of species. This incongruence complicates the task of infering the latter from the former. We describe the use of reconciled trees to reconstruct the history of a gene tree with respect to a species tree. Reconciled trees allow the history of the gene tree to be visualized and also quantify the relationship between the two trees. The cost of a reconciled tree is the total number of duplications and gene losses required to reconcile a gene tree with its species tree. We describe the use of heuristic searches to find the species tree that yields the reconciled tree with the lowest cost. This method can be used to infer species trees from one or more gene trees.

1 aPage, Roderic, D. M.1 aCharleston, Michael, A. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0157.pdf01923nas a2200181 4500008004100000245016600041210006900207260000900276300001400285490000700299520127100306653000901577653000801586100002201594700002101616700002501637856007901662 1996 eng d00aRevision of Dennyus (Collodennyus) lice (Phthiraptera: Menoponidae) from swiftlets, with descriptions of new taxa and a comparison of host-parasite relationships0 aRevision of Dennyus Collodennyus lice Phthiraptera Menoponidae f c1996 a179 - 2040 v213 a

Lice of the subgenus Dennyus(Collodennyus) are host specific, perminant parasites of swiftslets (Aves: Apodidae). As a prelude to a test of the hypothesis that these lice have cospeciated with their hosts, we revise the taxonomy of the subgenus, redescribing the seven previously recognised species, and adding thirteen new species and three new subspecies. All twenty three of these louse taxa are found on swiftlets (Apodiformes: Apodidae), with four from hosts of the genusCollocalia, eighteen from Aerodramus, and one from Hydrochous. Successful identification is associated in most cases with females; males are only tenuously seperable. A complete host parasite list for the subgenus Collodennyus is provided, as well as a key for the identification of these taxa. Limited morphological variation within the subgenus has prevented ready extraction of descrete characters for cladistic analysis. In the absence of such characters, a cluster analysis of female and male characters is presented. Comparison of a dendrogram for Dennyus (Collodennyus) with a molecular phylogeny for the swiftlet hosts suggests that the history of the swiftlet-louse association has been complex, including episodes of host switching and independent speciation by the lice.

10amtax10aPHP1 aClayton, Dale, H.1 aPrice, Roger, D.1 aPage, Roderic, D. M. uhttps://phthiraptera.myspecies.info/sites/phthiraptera.info/files/0143.pdf00974nas a2200193 4500008004100000245006600041210006300107260000900170300001200179490000700191520037900198653001700577653001500594653001500609653001400624653001700638100002500655856010000680 1993 eng d00aGenes, organisms, and areas: the problem of multiple lineages0 aGenes organisms and areas the problem of multiple lineages c1993 a77 - 840 v423 a

The presence of multiple species lineages can cause problems in biogeographic and cospeciation studies analogous to the problems gene paralogy can cause in molecular systematics. The differential survival of these lineages, coupled with inadequate sampling, can distort estimates of area relationships in biogeography or host-parasite relationships in parasitology.

10aBiogeography10acladistics10acongruence10aorthology10aParasitology1 aPage, Roderic, D. M. uhttps://phthiraptera.myspecies.info/content/genes-organisms-and-areas-problem-multiple-lineages01451nas a2200193 4500008004100000245008200041210006900123260000900192300001400201490000700215520087800222653001401100653001701114653001101131653000901142653000901151100002501160856007201185 1994 eng d00aParallel phylogenies: reconstructing the history of host-parasite assemblages0 aParallel phylogenies reconstructing the history of hostparasite  c1994 a155 - 1730 v103 a

A method for reconstructing the history of a host parasite assemblage is described. This method has the advantage of making explicit the relationship between the host and the parasite trees, and allows a visually intuitive representation of that history. It also inables host switches to be incorporated as an explanation of the observed pattern of host parasite associations, without the spurious overestimates of the number of host switches that can be obtained using Brooks parsimony analysis (BPA). Reconstructions that maximise the number of cospeciation events have the greatest explanatory power and are hense prefered over reconstructions with fewer cospeciation events. A heuristic algorithm to find a single maximum reconstruction, and an exact algorithm to find all such reconstructions are presented. Two empirical applications of the method are given.

10aAlgorithm10acospeciation10agopher10alice10atree1 aPage, Roderic, D. M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0012.pdf01732nas a2200253 4500008004100000245010600041210006900147260000900216300001200225490000700237520093900244653001401183653001501197653001501212653001701227653001001244653001301254653001401267653001201281653000901293653001001302100002501312856014101337 1994 eng d00aMaps between trees and cladistic analysis of historical associations among genes, organisms and areas0 aMaps between trees and cladistic analysis of historical associat c1994 a58 - 770 v433 a

The concept of a reconciled tree arose indipendently in molecular systematics, parasitology and biogeography as a means of describing historical associations. Examples of historical associations include genes and organisms, hosts and parasitic organisms, and organisms and areas. A reconciled tree combines the tree for the host and its associate into a single summary of the historical association of the two entities under the assumption that no horizontal transmission of associates has occured. In this paper I define reconciled trees, describe an algorithm for their computation, and develop measures to quantify the degree of fit between trees, host parasite cospeciation and biogeography. The problem of incorporating horizontal transmission of associates (e.g., dispersal or host switching) is also addressed by introducing the concept of maximising the amount of codivergance (shared history) between the associates.

10abiography10acladistics10acongruence10acospeciation10agenes10amapping.10aPhylogeny10aspecies10atree10atrees1 aPage, Roderic, D. M. uhttps://phthiraptera.myspecies.info/content/maps-between-trees-and-cladistic-analysis-historical-associations-amoung-genes-organisms-and01511nas a2200145 4500008004100000245009300041210006900134260000900203300001200212490000800224520101500232100002101247700002501268856007201293 1995 eng d00aMolecular phylogenies and host-parasite cospeciation: gophers and lice as a model system0 aMolecular phylogenies and hostparasite cospeciation gophers and  c1995 a77 - 830 v3493 a

Recent methodological advances permit a rigerous comparison of phylogenetic trees for hosts and their parasites to determine the extent to which these groups have cospeciated through evolutionary time. In cases where significnat levels of cospeciation are indicated, comparisons of amounts of evolutionary change that have accumulated along analogous branches in the host and parasite trees provides a direct assessment of relative rates of evolution in the two groups. For such a comparison to be meaningful, the features compared in the hosts and parasites should be genetically based, evolutionary homologous, and should evolve in a roughly time-dependent fashion within each group. Nucleotide sequences encoding homologous genes in hosts and parasites are an ideal source of data for comparitive studies of evolutionary rates. Recent studies of pocket gophers and their lice are used to illustrate the variety of questions that can be addressed through phylogenetic study of host-parasite systems.

1 aHafner, Mark, S.1 aPage, Roderic, D. M. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0001.pdf02121nas a2200301 4500008004100000245016700041210006900208260000900277300001400286490000700300520120000307653001601507653001701523653001201540653001701552653001401569653001401583653001801597653000901615653001501624653001101639653001301650653001101663100002501674700002101699700002701720856007201747 1995 eng d00aPhylogeny of Geomydoecus and Thomomydoecus pocket gopher lice (Phthiraptera: Trichodectidae) inferred from cladistic analysis of adult and first instar morphology0 aPhylogeny of Geomydoecus and Thomomydoecus pocket gopher lice Ph c1995 a129 - 1430 v203 a

The phylogeny of all 122 species of chewing lice of the genera Geomydoecus and Thomomydoecus (Phthiraptera: Trichodectidae) hosted by pocket gophers (Rodentia: Geomyidae) is estimated by a cladistic analysis of fifty-eight morphological characters obtained from adults and first instars. The data set has considerable homoplasy, but still contains phylogenetic information. The phylogeny obtained is moderately resolved and, with some notable exceptions supports the species complexes proposed by Hellenthal and Price over the last two decades. The subgenera of G. (Thaelerius) and T. (Thomomydoecus)are both shown to be monophyletic , but the monophyly of subgenus T. (Jamespattonius) could not be confirmed, perhaps due to the lack of first instar data for one of its component species. The nominate subgenus of Geomydoecus may be monophyletic, but our cadogram was insufficiently resolved to corroberate this. Mapping the pocket gopher hosts on to the phylogeny reveals a consistant pattern of louse clades being restricted to particular genera or subgenera of gophers, but the history of the host-parasite association appears complex and will require considerable effort to resolve.

10aassemblages10aChewing lice10acomplex10acospeciation10aEvolution10aGeomyidae10ahost parasite10alice10aMallophaga10amexico10arodentia10aU.S.A.1 aPage, Roderic, D. M.1 aPrice, Roger, D.1 aHellenthal, Ronald, A. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0003.pdf