03147nas a2200265 4500008004100000022001400041245007500055210006900130260001200199300001600211490000600227520235900233653001702592653001802609653002302627653001702650653001902667653002402686100002202710700002102732700002702753700002302780700002402803856005402827 2019 eng d a2045-775800aTemporal and spatial dynamics of competitive parapatry in chewing lice0 aTemporal and spatial dynamics of competitive parapatry in chewin c06-2019 a7410 - 74240 v93 a

Abstract: We synthesize observations from 1979 to 2016 of a contact zone involving two sub‐ species of pocket gophers (Thomomys bottae connectens and T. b. opulentus) and their respective chewing lice (Geomydoecus aurei and G. centralis) along the Río Grande Valley in New Mexico, U.S.A., to test predictions about the dynamics of the zone. Historically, the natural flood cycle of the Rio Grande prevented contact between the two subspecies of pocket gophers. Flood control measures completed in the 1930s permitted contact, thus establishing the hybrid zone between the pocket gophers and the contact zone between their lice (without hybridization). Since that time, the pocket gopher hybrid zone has stabilized, whereas the northern chewing louse spe‐ cies has replaced the southern louse species at a consistent rate of ~150 m/year. The 0.2–0.8 width of the replacement zone has remained constant, reflecting the con‐ stant rate of chewing louse species turnover on a single gopher and within a local pocket gopher population. In contrast, the full width of the replacement zone (north‐ ernmost G. centralis to southernmost G. aurei) has increased annually. By employing a variety of metrics of the species replacement zone, we are better able to understand the dynamics of interactions between and among the chewing lice and their pocket gopher hosts. This research provides an opportunity to observe active species re‐ placement and resulting distributional shifts in a parasitic organism in its natural setting.

Data Availability Statement:: 1. Chewing louse microsatellite genotypes and R‐scripts are available in Dryad, https://doi.org/10.5061/dryad.9sv4q00. 2. Parasite voucher specimens and frozen tissue samples from pocket gophers are stored at −80°C at the University of Northern Iowa and at Louisiana State University. 3. Pocket gopher voucher specimens are housed at the Louisiana State University Museum of Natural Science (LSUMZ), the Museum of Southwestern Biology, University of New Mexico (MSB), Museum of Vertebrate Zoology, University of California (MVZ), or the Biodiversity Research and Teaching Collections at Texas A&M University (BRTC). 4. Current and future research directions of the team may be found at https://louselab.uni.edu/homepage.

10aChewing lice10acontact zones10adispersal distance10ahybrid zones10aPocket gophers10aspecies replacement1 aHafner, David, J.1 aHafner, Mark, S.1 aSpradling, Theresa, A.1 aLight, Jessica, E.1 aDemastes, James, W. uhttps://onlinelibrary.wiley.com/toc/20457758/9/1302348nas a2200241 4500008004100000022001300041245010500054210006900159260001200228300001500240490000700255520161100262653001701873653001701890653001601907653002301923653001601946653001401962653001901976100002301995700002102018856006702039 2007 eng d a1055790300aCophylogeny and disparate rates of evolution in sympatric lineages of chewing lice on pocket gophers0 aCophylogeny and disparate rates of evolution in sympatric lineag c12-2007 a997 - 10130 v453 a

Although molecular-based phylogenetic studies of hosts and parasites are increasingly common in the literature, no study to date has examined two congeneric lineages of parasites that live in sympatry on the same lineage of hosts. This study examines phylogenetic rela- tionships among chewing lice (Phthiraptera: Trichodectidae) of the Geomydoecus coronadoi and Geomydoecus mexicanus species com- plexes and compares these to phylogenetic patterns in their hosts (pocket gophers of the rodent family Geomyidae). Sympatry of congeneric lice provides a natural experiment to test the hypothesis that closely related lineages of parasites will respond similarly to the same host. Sequence data from the mitochondrial COI and the nuclear EF-1a genes confirm that the two louse complexes are recip- rocally monophyletic and that individual clades within each species complex parasitize a different species of pocket gopher. Phylogenetic comparisons reveal that both louse complexes show a significant pattern of cophylogeny with their hosts. Comparisons of rates of nucle- otide substitution at 4-fold degenerate sites in the COI gene indicate that both groups of lice have significantly higher basal mutation rates than their hosts. The two groups of lice have similar basal rates of mutation, but lice of the G. coronadoi complex show significantly elevated rates of nucleotide substitution at all sites. These rate differences are hypothesized to result from population-level phenomena, such as effective population size, founder effects, and drift, that influence rates of nucleotide substitution.

10aCodivergence10acospeciation10aCratogeomys10aEvolutionary rates10ageomydoecus10aPhylogeny10aPocket gophers1 aLight, Jessica, E.1 aHafner, Mark, S. uhttps://linkinghub.elsevier.com/retrieve/pii/S105579030700314402467nas a2200229 4500008004100000245010300041210006900144300001400213490000700227520173100234653001901965653001401984653002501998653001302023653002602036100002402062700002202086700002102108700002302129700002702152856005802179 2019 eng d00aLoss of genetic diversity, recovery and allele surfing in a colonizing parasite, Geomydoecus aurei0 aLoss of genetic diversity recovery and allele surfing in a colon a703 - 7200 v283 a

Understanding the genetic consequences of changes in species distributions has wide-ranging implications for predicting future outcomes of climate change, for pro- tecting threatened or endangered populations and for understanding the history that has led to current genetic patterns within species. Herein, we examine the genetic consequences of range expansion over a 25‐year period in a parasite (Geomydoecus aurei) that is in the process of expanding its geographic range via invasion of a novel host. By sampling the genetics of 1,935 G. aurei lice taken from 64 host individuals collected over this time period using 12 microsatellite markers, we test hypotheses concerning linear spatial expansion, genetic recovery time and allele surfing. We find evidence of decreasing allelic richness (AR) with increasing distance from the source population, supporting a linear, stepping stone model of spatial expansion that em- phasizes the effects of repeated bottleneck events during colonization. We provide evidence of post‐bottleneck genetic recovery, with average AR of infrapopulations increasing about 30% over the 225‐generation span of time observed directly in this study. Our estimates of recovery rate suggest, however, that recovery has plateaued and that this population may not reach genetic diversity levels of the source popula- tion without further immigration from the source population. Finally, we employ a grid-based sampling scheme in the region of ongoing population expansion and pro- vide empirical evidence for the power of allele surfing to impart genetic structure on a population, even under conditions of selective neutrality and in a place that lacks strong barriers to gene flow.

10aallele surfing10aDiversity10aexpanding population10aparasite10apopulation bottleneck1 aDemastes, James, W.1 aHafner, David, J.1 aHafner, Mark, S.1 aLight, Jessica, E.1 aSpradling, Theresa, A. uhttps://onlinelibrary.wiley.com/doi/10.1111/mec.1499703327nas a2200217 4500008004100000245010800041210006900149490000700218520263600225653001602861653001602877653001402893653001002907653003302917100001802950700002402968700002702992700002203019700002103041856004703062 2019 eng d00aHost-parasite associations of the Cratogeomys fumosus species group and their chewing lice, Geomydoecus0 aHostparasite associations of the Cratogeomys fumosus species gro0 v313 a

Chewing lice (Phthiraptera: Trichodectidae) and the pocket gophers (Rodentia: Geomyidae) that they inhabit have shared an intimate his- torical association that has made them a textbook study for cophylogeny. Herein, we examine the chewing lice found on pocket gophers of the Cratogeomys fumosus species group using DNA sequence data from the mitochondrial cytochrome c oxidase subunit I gene to determine relationships among lice for comparison to published host phylogeny. Although matrix approaches indicate a correlation between host and parasite genetic distances, cophylogenetic reconstruction methods fail to detect a pattern of widespread cophylogeny. In conclusion, this study provides an exception to the rule of host-parasite cophylogeny that could be the result of the young age of the relationships considered herein and the complex history of the Trans-Mexican Volcanic Belt.
Los piojos masticadores (Phthiraptera: Trichodectidae) y las tuzas (Rodentia: Geomyidae) que habitan, han compartido una asociación histórica íntima que los ha convertido en un estudio de libro de texto para la cofilogenia. En el presente estudio, examinamos los piojos mas- ticadores que se encuentran en las tuzas del grupo de especies de Cratogeomys fumosus utilizando los datos de secuencias de ADN del gen mitocondrial Citocromo C Oxidasa subunidad I para determinar las relaciones entre los piojos en comparación con la filogenia publicada del hospedero. Si bien los enfoques matriciales indican una correlación entre las distancias genéticas del hospedero y el parásito, los métodos de reconstrucción cofilogenéticos no detectan un patrón de cofilogenia generalizada. En conclusión, este estudio, proporciona una excepción a la regla de cofilogenia parásito-hospedero que podría ser el resultado de la corta edad de las relaciones consideradas en este documento y la compleja historia del Cinturón Volcánico Trans-Mexicano.

Authors recommended citation: Popinga, Alex; Demastes, James W.; Spradling, Theresa A.; Hafner, David J.; and Hafner, Mark S., "Host-parasite associations of the Cratogeomys fumosus species group and their chewing lice, Geomydoecus" (2019). Biology Faculty Publications. 31. https://scholarworks.uni.edu/bio_facpub/31

First published in THERYA, v.10 n.2 (2019), by the Asociacion Mexicana de Mastozoologia A.C. Vol 10 Issue 2 81-89 website URL: http://www.revistas-conacyt.unam.mx/therya/index.php/THERYA/article/view/739

10acophylogeny10aCratogeomys10aGeomyidae10amtDNA10aTrans-Mexican Volcanic Belt.1 aPopinga, Alex1 aDemastes, James, W.1 aSpradling, Theresa, A.1 aHafner, David, J.1 aHafner, Mark, S. uhttps://scholarworks.uni.edu/bio_facpub/3102333nas a2200217 4500008004100000022001400041245014500055210006900200300001400269490000700283520160700290653002701897653001601924653002301940653002001963653001301983653001401996100002302010700002102033856006102054 2008 eng d a1063-515700aCodivergence in Heteromyid Rodents (Rodentia: Heteromyidae) and Their Sucking Lice of the Genus Fahrenholzia (Phthiraptera: Anoplura)0 aCodivergence in Heteromyid Rodents Rodentia Heteromyidae and The a449 - 4650 v573 a

Abstract.—Although most studies of codivergence rely primarily on topological comparisons of host and parasite phyloge- nies, temporal assessments are necessary to determine if divergence events in host and parasite trees occurred contempora- neously. A combination of cophylogenetic analyses and comparisons of branch lengths are used in this study to understand the host-parasite association between heteromyid rodents (Rodentia: Heteromyidae) and their sucking lice of the genus Fahrenholzia (Phthiraptera: Anoplura). Cophylogenetic comparisons based on nucleotide substitutions in the mitochondrial COI gene reveal a significant, but not perfect, pattern of cophylogeny between heteromyids and their sucking lice. Regres- sion analyses show a significant functional relationship between the lengths of analogous branches in the host and parasite trees, indicating that divergence events in hosts and parasites were approximately contemporaneous. Thus, the topological similarity observed between heteromyids and their lice is the result of codivergence. These analyses also show that the COI gene in lice is evolving two to three times faster than the same gene in their hosts (similar to the results of studies of other lice and their vertebrate hosts) and that divergence events in lice occurred shortly after host divergence. We recommend that future studies of codivergence include temporal comparisons and, when possible, use the same molecular marker(s) in hosts and parasites to achieve the greatest insight into the history of the host-parasite relationship.

10aCophylogenetic methods10acophylogeny10acospeciation; host10amolecular rates10aparasite10asymbiosis1 aLight, Jessica, E.1 aHafner, Mark, S. uhttps://academic.oup.com/sysbio/article/57/3/449/166461002055nas a2200205 4500008004100000022001400041245010900055210006900164260001600233300001400249490000700263520138600270100002401656700002701680700002101707700002401728700002201752700002301774856005201797 2012 eng d a0022-339500aCophylogeny on a Fine Scale: Geomydoecus Chewing Lice and Their Pocket Gopher Hosts, Pappogeomys bulleri0 aCophylogeny on a Fine Scale Geomydoecus Chewing Lice and Their P cJan-04-2012 a262 - 2700 v983 a

Many species of pocket gophers and their ectoparasitic chewing lice have broadly congruent phylogenies, indicating a history of frequent codivergence. For a variety of reasons, phylogenies of codiverging hosts and parasites are expected to be less congruent for more recently diverged taxa. This study is the first of its scale in the pocket gopher and chewing louse system, with its focus entirely on comparisons among populations within a single species of host and 3 chewing louse species in the Geomydoecus bulleri species complex. We examined mitochondrial DNA from a total of 46 specimens of Geomydoecus lice collected from 11 populations of the pocket gopher host, Pappogeomys bulleri. We also examined nuclear DNA from a subset of these chewing lice. Louse phylogenies were compared with a published pocket gopher phylogeny. Contrary to expectations, we observed a statistically significant degree of parallel cladogenesis in these closely related hosts and their parasites. We also observed a higher rate of evolution in chewing louse lineages than in their corresponding pocket gopher hosts. In addition, we found that 1 louse species (Geomydoecus burti) may not be a valid species, that subspecies within G. bulleri are not reciprocally monophyletic, and that morphological and genetic evidence support recognition of a new species of louse, Geomydoecus pricei.

1 aDemastes, James, W.1 aSpradling, Theresa, A.1 aHafner, Mark, S.1 aSpies, Gretchen, R.1 aHafner, David, J.1 aLight, Jessica, E. uhttp://www.bioone.org/doi/abs/10.1645/GE-2904.101528nas a2200145 4500008004100000245009400041210006900135260000900204300001400213490000700227520102200234100002301256700002101279856008201300 2007 eng d00aPhylogenetics and host associations of Fahrenholzia sucking lice (Phthiraptera: Anoplura)0 aPhylogenetics and host associations of Fahrenholzia sucking lice c2007 a359 - 3700 v323 a

Mitochondrial and nuclear DNA sequence data were used to reconstruct phylogenetic relationships for eleven of the twelve currently recognized species of Fahrenholzia, lice found only on rodents of the family Heteromyidae. Field collections included twenty of the thirty-three known host associations and resulted in the discovery of four new associations. Phylogenetic analyses of the mitochondrial and nuclear datasets were in general agreement, resulting in a well-resolved Fahrenholzia phylogeny. Analyses supported the monophyly of lice parasitizing the host subfamily Heteromyinae (spiny pocket mice). Lice parasitizing the genera Chaetodipus (pocket mice) and Perognathus (silky pocket mice) each represent monophyletic lineages. Phylogenetic patterns and levels of genetic differentiation suggest that the widespread Fahrenholzia pinnata may contain several cryptic species. Cryptic species may exist also within the less widely distributed species, Fahrenholzia microcephala and Fahrenholzia reducta.

1 aLight, Jessica, E.1 aHafner, Mark, S. uhttp://onlinelibrary.wiley.com/wol1/doi/10.1111/j.1365-3113.2006.00367.x/full00553nas 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.pdf02363nas a2200265 4500008004100000020001400041245012800055210006900183260001800252300001400270490000700284520148200291653001201773653001201785653002001797653004601817653001501863653001401878653004501892653003301937653001301970100002301983700002102006856007002027 1993 eng d a0020-751900aSystematic relationships among pocket gopher chewing lice (Phthiraptera: Trichodectidae) inferred from electrophoretic data0 aSystematic relationships among pocket gopher chewing lice Phthir aENGLANDc1993 a191 - 2010 v233 a

Multilocus starch-gel electrophoresis was used to obtain electrophoretic data for maximum parsimony, phenetic, and component analysis of systematic relationships among 14 species of pocket gopher chewing lice and two outgroup species. Pocket gopher lice studied included taxa representing seven of the 25 species complexes in Geomydoecus, and two of the four species complexes of Thomomydoecus. In pairwise comparisons, 12 of the 14 pocket gopher lice were characterized by two or more fixed genetic differences, and each nominal species possessed at least one fixed difference. These electrophoretic results are consistent with previous morphological analysis of species-level taxonomy. A maximum-parsimony analysis of the electrophoretic data yielded 80 trees, each with a length of 78 steps and a consistency index of 0.83. The strict consensus tree and the UPGMA phenogram of genetic distances indicate that five species of lice from Central American pocket gophers (Orthogeomys) form a clade distinct from lice hosted by North American pocket gophers (Geomys, Thomomys, and Cratogeomys). Other louse clades receiving reasonable support in bootstrap parsimony analysis include: (Thomomydoecus wardi, Th. minor); (Geomydoecus scleritus, G. mobilensis); (G. panamensis, G. setzeri); and (G. cherriei, G. costaricensis). The systematic results of this electrophoretic study are generally consistent with the boundaries of louse species complexes described previously.

10aalleles10aanimals10aelectrophoresis10aLice Infestations/parasitology/veterinary10aMallophaga10aPhylogeny10aResearch Support, U.S. Gov't, Non-P.H.S.10aRodent Diseases/parasitology10arodentia1 aNadler, Steven, A.1 aHafner, Mark, S. uhttp://www.sciencedirect.com/science/article/pii/002075199390141K01689nas a2200157 4500008004100000245014400041210006900185260000900254300001200263490000700275520111600282653001301398100002001411700002101431856007901452 2002 eng d00aPhylogenetic analysis of bacterial communities associated with ectoparasitic chewing lice of pocket gophers: A culture-independent approach0 aPhylogenetic analysis of bacterial communities associated with e c2002 a78 - 930 v443 a

This study identifies the bacteria associated with ectoparasitic chewing lice that live in the fur of pocket gophers. Samples of chewing lice were collected from pocket gopher hosts in Florida, Missouri, New Mexico, and Costa Rica. We used a molecular sampling method whereby total community DNA was extracted from samples of chewing lice, and PCR was used to selectively amplify small-subunit rRNA genes from bacteria. This culture-independent method yielded ca. 35 distinct lineages representing eight widely divergent groups within the domain Bacteria. Phylogenetic analysis of two lineages (Acinetobacter and Staphylococcus) provides evidence that multiple species of each group are found in chewing lice. Phylogenetic analysis also demon-strated that diversication within chewing lice may be evident in both Acinetobacter and Staphylococcus. Some clones amplified from chewing louse hosts appeared to be distinct from known species of Acinetobacter and Staphylococcus. This diversication may be the result of the extreme isolation of populations of both chewing lice and their pocket gopher hosts.

10abacteria1 aReed, David, L.1 aHafner, Mark, S. uhttps://phthiraptera.myspecies.info/sites/phthiraptera.info/files/6678.pdf01812nas a2200313 4500008004100000245009500041210006900136260000900205300001400214490000700228520094600235653001701181653002301198653001701221653001401238653001401252653001101266653000901277653000901286653001401295653001301309653001701322653001101339653001601350653001901366100002001385700002101405856007201426 1997 eng d00aHost specificity of chewing lice on pocket gophers: a potential mechanism for cospeciation0 aHost specificity of chewing lice on pocket gophers a potential m c1997 a655 - 6600 v783 a

Pocket gophers (Rodentia: Geomyidae) and their ectoparasitic chewing lice (Phthiraptera Trichodectidae) have congruent phylogenies and show evidence of cospeciation. We examined a potential mechanism that could generate the observed pattern of cospeciation by testing the ability of lice to survive and reproduce on hosts other than their own. Our tests were conducted at the subspecific, specific, and generic levels relative to the natural host. Although lice established successful colonies at each level, colonization of new hosts diminished with increasing phylogenetic distance from the natural host of each louse. We suggest that the pattern of cospeciation results primarily from lack of opportunity for lice to colonize new hosts. However, in rare cases where lice disperse to new hosts, survival may be difficult on hosts that are not closely related to the natural host, which would reinforce the pattern of cospeciation.

10aChewing lice10acladistic analysis10acospeciation10aEvolution10aGeomyidae10agopher10ahost10alice10amolecular10aparasite10aPhthiraptera10apocket10aspecificity10atrichodectidae1 aReed, David, L.1 aHafner, Mark, S. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/3072.pdf01556nas a2200145 4500008004100000245009400041210006900135260000900204300001400213490000700227520104200234100002301276700002101299856009001320 1989 eng d00aGenetic differentiation in sympatric species of chewing lice (Mallophaga, Trichodectidae)0 aGenetic differentiation in sympatric species of chewing lice Mal c1989 a109 - 1130 v823 a

Starch-gel electrophoresis was used to assess levels of genetic differentiation among populations representing four species of chewing lice (Geomydoecus). These species included representatives of the subgenera Geomydoecus and Thomomydoecus that coexist on individual pocket gophers (Thomomys). Fourteen enzymatic loci were resolved from individual chewing lice. Samples of conspecific lice collected from different individual pocket gophers at a single locality sometimes showed marked differences in allele frequencies. Conspecific samples of lice collected from geographically distant host populations had levels of genetic differentiation similar to those of other conspecific insect populations. Levels of Nei's genetic identity (I) within each subgenus were typical of nonsibling species of insects. The subgenera Geomydoecus and Thomomydoecus had a mean Nei's I value of 0.332. This value is characteristic of intergeneric comparisons for insects, and it supports the elevation of these subgenera to generic rank.

1 aNadler, Steven, A.1 aHafner, Mark, S. uhttps://academic.oup.com/aesa/article-abstract/82/1/109/16747?redirectedFrom=fulltext01442nas a2200217 4500008004100000245013200041210006900173260000900242300001400251490000700265520074100272653001701013653001101030653001301041653001101054100002001065700002101085700002301106700002301129856007201152 2000 eng d00aSpatial partitioning of host habitat by chewing lice of the genera Geomydoecus and Thomomydoecus (Phthiraptera: Trichodectidae)0 aSpatial partitioning of host habitat by chewing lice of the gene c2000 a951 - 9550 v863 a

Chewing lice, Geomydoecus and Thomomydoecus, coexist on pocket gophers, Thomomys spp. We investigated the spatial distribution of the 2 genera on their hosts and explored possible mechanisms of resource partitioning by chewing lice. Chewing lice appear to partition available host resources spatially, with Geomydoecus occurring primarily on the lateral and dorsal regions of the host, and Thomomydoecus occurring primarily on the lateral and ventral regions. Although spatial partitioning of the host habitat is evident, it does not appear to be explained by hair diameter. Spatial partitioning of the host's body could be the result of some other factor, possibly temperature or humidity gradients of the host's body.

10acospeciation10agopher10aparasite10apocket1 aReed, David, L.1 aHafner, Mark, S.1 aAllen, Shannon, K.1 aSmith, Melanie, B. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/2262.pdf01746nas a2200337 4500008004100000245009200041210006900133260000900202300001500211490000700226520085200233653000901085653001401094653001401108653001101122653000901133653000901142653001601151653000901167653001301176653001701189653001401206653001101220653001301231653000901244653001901253100002001272700002101292700002301313856007201336 2000 eng d00aMammalian hair diameter as a possible mechanism for host specialization in chewing lice0 aMammalian hair diameter as a possible mechanism for host special c2000 a999 - 10070 v813 a

We examined the relationship between mammalian hair diameter and body mass at several taxonomic levels (interordinal, intrafamilial, intrageneric, and intraspecific) and showed a significant, positive allometric relationship between hair diameter and body size at all taxonomic levels examined. The allometric coefficient (alpha) ranged from 0.13 to 0.33. Within pocket gophers (Geomyidae), a significant positive relationship exists between hair diameter and rostral groove dimensions of their chewing lice, Geomydoecus, which use the rostral groove to grasp hairs of their host. Coupled with previous evidence of a strong allometric relationship between rostral groove width and louse body size, our findings suggest that hair diameter of the host is an important determinant of body size in chewing lice that parasitize pocket gophers.

10abody10aEvolution10aGeomyidae10agopher10ahair10ahost10aindependent10amass10aparasite10aPhthiraptera10aPhylogeny10apocket10arodentia10asize10atrichodectidae1 aReed, David, L.1 aHafner, Mark, S.1 aAllen, Shannon, K. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/2261.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.pdf01924nas 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_contents01980nas a2200169 4500008004100000245009500041210006900136260000900205300001200214490000700226520142900233100002101662700002101683700001901704700001801723856006901741 1987 eng d00aMacrogeographic patterns of genetic differentiation in the pocket gopher Thomomys umbrinus0 aMacrogeographic patterns of genetic differentiation in the pocke c1987 a18 - 340 v363 a

Electromorphic and chromosomal variation is analyzed in 26 populations of Tho- momys umbrinus sampled from throughout the range of the species. Interpopulation levels of genic differentiation are extreme, generally exceeding values measured between conspecific populations of most animals or plants. Two principal groups of T. umbrinus are recognized based on chromosomal evidence, one with 2n = 76 chromosomes and the other with 2n = 78. Further, the 2n = 78 group (but not the 2n = 76 group) is bisected into geographic subgroups with respect to chromosome morphology and heterochromatin position. The kind and degree of chromosomal differentiation observed suggests that the three groups may be reproductively incompatible. Allozymic evidence corroborates the above groupings, and an analysis of patterns of allele sharing suggests the absence of gene flow among the groups. A cladistic analysis of electro- morphic data indicates that the two 2n = 78 groups may be independently derived from the 2n = 76 lineage. The combined evidence supports the hypothesis that T. umbrinus is actually a composite of at least three biological species and confirms the observation that speciation in the genus Thomomys is unrelated to the level of genic differentiation between populations. [Evolu- tionary genetics; geographic variation; evolutionary concordance; pocket gophers; cladistic anal- ysis; paraphyly; speciation.]

1 aHafner, Mark, S.1 aHafner, John, C.1 aPatton, J., L.1 aSmith, M., F. uhttp://www.jstor.org/stable/2413305?seq=1#page_scan_tab_contents01241nas a2200181 4500008004100000245014000041210006900181260000900250300001400259490000700273520062100280653000900901653000800910100002100918700002700939700002100966856007200987 1985 eng d00aThe Geomydoecus (Mallophaga, Trichodectidae) from the central American pocket gophers of the subgenus Macrogeomys (Rodentia, Geomyidae)0 aGeomydoecus Mallophaga Trichodectidae from the central American  c1985 a432 - 4430 v873 a

Six species and subspecies of Geomydoecus from Orthogeomys (Macrogeomys) pocket gophers are described and illustrated: G. panamensis panamensis Price and Emerson from O. (M.) cavator (Bangs); G. p. dariensis Price and Emerson from O. (M.) dariensis (Goldman); G. setzeri Price and G. davidhafneri new species from O. (M.) underwoodi (Osgood) and O. (M.) cherriei (J. A. Allen); G. costaricensis Price and Emerson from three subspecies of O. (M.) heterodus (Peters); and G. cherriei Price from O. (M.) cherriei and O. (M.) matagalpae (J. A. Allen). Keys are provided for the identification of these six taxa.

10amtax10aPHP1 aPrice, Roger, D.1 aHellenthal, Ronald, A.1 aHafner, Mark, S. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0940.pdf01566nas a2200325 4500008004100000245008400041210006900125260000900194300001600203490000600219520065300225653001600878653001700894653001600911653001400927653000900941653001500950653002200965653001000987653001300997653001601010653000901026653002001035653001901055100002401074700002101098700002201119700002701141856007201168 1998 eng d00aPocket gophers and chewing lice: a test of the maternal transmission hypothesis0 aPocket gophers and chewing lice a test of the maternal transmiss c1998 a1065 - 10690 v73 a

The life-history traits of pocket gophers and their chewing lice suggest that there is little opportunity for transmission of parasites among pocket gophers, with the exception of transmission from mother to offspring. Herein, we test the hypothesis that lice are transmitted maternally by using an indirect approach that compares the distribution of louse populations to the distribution of mitochondrial DNA haplotypes in the pocket gophers. Comparison of the chewing louse distributions to the distribution of mtDNA haplotypes for the gophers revealed no significant concordance, and thus falsifies the maternal transmission hypothesis.

10acoevolution10acolonization10ageomydoecus10aGeomyidae10ahost10aMallophaga10amitochondrial dna10amtDNA10aparasite10apopulations10aRFLP10athomomys bottae10atrichodectidae1 aDemastes, James, W.1 aHafner, Mark, S.1 aHafner, David, J.1 aSpradling, Theresa, A. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0842.pdf01666nas a2200385 4500008004100000245011600041210006900157260000900226300001400235490000700249520060200256653001700858653001000875653001600885653002000901653001400921653001200935653001400947653001100961653001800972653001100990653000901001653001101010653001101021653001601032653001301048653001001061100002101071700002401092700002201116700002701138700002301165700002301188856006901211 1998 eng d00aAge and movement of a hybrid zone: Implications for dispersal distance in pocket gophers and their chewing lice0 aAge and movement of a hybrid zone Implications for dispersal dis c1998 a278 - 2820 v523 a

Historical flood records for the Rio Grande Valley of New Mexico suggest that a pocket gopher (Thomomys bonae) hybrid zone previously thought to be 10,000 years old may actually be closer to 50 years old. Measured zone width (defined genetically) is consistent with the hypothesis of recent contact, if we assume a reasonable dispersal distance of approximately 400 m/year for pocket gophers. A five-year study of movement of the contact zone between the two species of chewing lice that parasitize these pocket gophers also is consistent with the hypothesis of recent origin of the zone.

10aChewing lice10acline10acoevolution10adifferentiation10adispersal10agenetic10aGeomyidae10agopher10ahost parasite10ahybrid10alice10amodels10apocket10apopulations10arodentia10azones1 aHafner, Mark, S.1 aDemastes, James, W.1 aHafner, David, J.1 aSpradling, Theresa, A.1 aSudman, Philip, D.1 aNadler, Steven, A. uhttp://www.jstor.org/stable/2410946?seq=1#page_scan_tab_contents01562nas a2200193 4500008004100000245007600041210006900117260000900186300001400195490000800209520097500217653001701192653001101209653001401220653001001234100002101244700002301265856008001288 1988 eng d00aPhylogenetic trees support the coevolution of parasites and their hosts0 aPhylogenetic trees support the coevolution of parasites and thei c1988 a258 - 2590 v3323 a

The close correspondence often observed between the taxonomy of parasites and their hosts has led to Fahrenholz's rule, which postulates that parasites and their hosts speciate in synchrony. This leads to the prediction that phylogenetic trees of parasites and their hosts should be topologically identical. We report here a test of this prediction which involves the construction of phylogenetic trees for rodents and their ectoparasites using protein electrophoretic data. We find a high degree of concordance in the branching patterns of the trees which suggests that there is a history of cospeciation in this host-parasite assemblage. In several cases where the branching patterns were identical in the host and parasite phylogenies, the branch lengths were also very similar which, given the assumptions of molecular clock theory, strongly suggests that the speciation of these hosts and ectoparasites was roughly contemporaneous and causally related.

10acospeciation10agopher10aPhylogeny10atrees1 aHafner, Mark, S.1 aNadler, Steven, A. uhttps://phthiraptera.myspecies.info/sites/phthiraptera.info/files/38845.pdf00522nas a2200133 4500008004100000245012400041210006900165260000900234300001400243490000700257100002100264700002300285856008000308 1990 eng d00aCospeciation in host-parasite assemblages: comparative analysis of rates of evolution and timing of cospeciation events0 aCospeciation in hostparasite assemblages comparative analysis of c1990 a192 - 2040 v391 aHafner, Mark, S.1 aNadler, Steven, A. uhttps://phthiraptera.myspecies.info/sites/phthiraptera.info/files/38840.pdf01612nas a2200169 4500008004100000245014400041210006900185260000900254300001400263490000700277520099100284100002301275700002101298700002101319700002201340856008001362 1990 eng d00aGenetic differentiation amoung chewing louse populations (Mallophaga: Trichodectidae) in a pocket gopher contact zone (Rodentia: Geomyidae)0 aGenetic differentiation amoung chewing louse populations Malloph c1990 a942 - 9510 v443 a

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

1 aNadler, Steven, A.1 aHafner, Mark, S.1 aHafner, John, C.1 aHafner, David, J. uhttps://phthiraptera.myspecies.info/sites/phthiraptera.info/files/38810.pdf01556nas a2200205 4500008004100000245008100041210006900122260000900191300001400200490000700214520094500221653001701166653001701183653001101200653000901211653001301220100002401233700002101257856007201278 1993 eng d00aCospeciation of pocket gophers (Geomys) and their chewing lice (Geomydoecus)0 aCospeciation of pocket gophers Geomys and their chewing lice Geo c1993 a521 - 5300 v743 a

Comparison of independently derived phylogenies for pocket gophers (Geomys) and their chewing lice (Geomydoecus) from Texas and Louisiana indicates a history of widespread cospeciation in this host-parasite assemblage. Inference of cospeciation is supported by statistical comparison of genetic-distance matrices for gophers and lice based on allozyme data. Although similar, host and parasite phylogenies are not identical; inconsistencies likely result from host switching by the parasites, retention of ancestral taxa on parasites, or poorly delineated taxonomic boundries. The current disjunct distribution of Geomydoecus ewingi suggests that this chewing louse once parasitised the common ancestor of Geomys breviceps and G. attawateri. Combined protein and morphologic evidence suggests that the population of Geomydoecus ewingi hosted by Geomys breviceps breviceps in northeastern Lousiana may be a cryptic species of louse.

10aChewing lice10acospeciation10aGeomys10alice10aparasite1 aDemastes, James, W.1 aHafner, Mark, S. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0031.pdf01511nas 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.pdf01710nas a2200373 4500008004100000245007900041210006900120260000900189300001600198490000800214520068000222653000900902653002000911653001200931653001100943653001600954653001400970653001100984653001500995653002201010653001401032653001101046653001301057653001801070653001001088653001901098100002101117700002301138700002901161700002701190700002401217700002301241856007201264 1994 eng d00aDisparate rates of molecular evolution in cospeciating hosts and parasites0 aDisparate rates of molecular evolution in cospeciating hosts and c1994 a1087 - 10900 v2653 a

DNA sequences for the gene encoding mitochondrial cytochrome oxidase I in a group of rodents (pocket gophers) and their ectoparasites provide evidence for cospeciation and reveal different rates of molecular evolution in the hosts and their parasites. The overall rate of nucleotide substitution (both silent and replacement changes) is approximately three times higher in lice, and the rate of synonymous substitution (based on analysis of fourfold degenerate sites) is approximately an order of magnitude greater in lice. The difference in synonymous substitution rate between lice and gophers correlates with a difference in similar magnitude in generation times.

10abase10adifferentiation10agenetic10agenome10ageomydoecus10aGeomyidae10agopher10aMallophaga10amitochondrial dna10aPhylogeny10apocket10arodentia10asubstitutions10atrees10atrichodectidae1 aHafner, Mark, S.1 aSudman, Philip, D.1 aVillablanca, Francis, X.1 aSpradling, Theresa, A.1 aDemastes, James, W.1 aNadler, Steven, A. uhttp://www.phthiraptera.info/sites/phthiraptera.info/files/0004.pdf