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/3102363nas 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/002075199390141K01812nas 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.pdf01241nas 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.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.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