We present Procrustean Approach to Cophylogeny (PACo), a novel statistical tool to test for congruence between phylogenetic trees, or between phylogenetic distance matrices of associated taxa. Unlike previous tests, PACo evaluates the dependence of one phylogeny upon the other. This makes it especially appropriate to test the classical coevolutionary model that assumes that parasites that spend part of their life in or on their hosts track the phylogeny of their hosts. The new method does not require fully resolved phylogenies and allows for multiple host-parasite associations. PACo produces a Procrustes superimposition plot enabling a graphical assessment of the fit of the parasite phylogeny onto the host phylogeny and a goodness-of-fit statistic, whose significance is established by randomization of the host-parasite association data. The contribution of each individual host-parasite association to the global fit is measured by means of jackknife estimation of their respective squared residuals and confidence intervals associated to each host-parasite link. We carried out different simulations to evaluate the performance of PACo in terms of Type I and Type II errors with respect to two similar published tests. In most instances, PACo performed at least as well as the other tests and showed higher overall statistical power. In addition, the jackknife estimation of squared residuals enabled more elaborate validations about the nature of individual links than the ParaFitLink1 test of the program ParaFit. In order to demonstrate how it can be used in real biological situations, we applied PACo to two published studies using a script written in the public-domain statistical software R.

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Evaluating phenotypic plasticity in attachment organs of parasites can provide information on the capacity to colonize new hosts and illuminate evolutionary processes driving host specificity.We analysed the variability in shape and size of the dorsal and ventral anchors of Ligophorus cephali from Mugil cephalus by means of geometric morphometrics and multivariate statistics. We also assessed the morphological integration between anchors and between the roots and points in order to gain insight into their functional morphology. Dorsal and ventral anchors showed a similar gradient of overall shape variation, but the amount of localized changes was much higher in the former. Statistical models describing variations in shape and size revealed clear differences between anchors. The dorsal anchor/bar complex seems more mobile than the ventral one in Ligophorus, and these differences may reflect different functional roles in attachment to the gills. The lower residual variation associated with the ventral anchor models suggests a tighter control of their shape and size, perhaps because these anchors seem to be responsible for firmer attachment and their size and shape would allow more effective responses to characteristics of the microenvironment within the individual host. Despite these putative functional differences, the high level of morphological integration indicates a concerted action between anchors. In addition, we found a slight, although significant, morphological integration between roots and points in both anchors, which suggests that a large fraction of the observed phenotypic variation does not compromise the functional role of anchors as levers. Given the low level of genetic variation in our sample, it is likely that much of the morphological variation reflects host-driven plastic responses. This supports the hypothesis of monogenean specificity through host-switching and rapid speciation. The present study demonstrates the potential of geometric morphometrics to provide new and previously unexplored insights into the functional morphology of attachment and evolutionary processes of host-parasite coevolution.4

An important question in the study of phenotypic evolution is whether characters are independent of each other or behave and evolve as integrated modules. Morphological integration and modularity provide a powerful framework for the analysis of the evolution of morphological traits. We used geometric morphometrics and phylogenetically independent contrasts (PIC) to test four different modularity hypotheses in the haptoral anchors of 14 monogenean species of Ligophorus. Integration between the modular units identified was further evaluated with two-block partial least squares analysis. Roots and points represented two modules in the dorsal and ventral anchors, but modularity was not statistically supported when parasite phylogeny was accounted for, which may indicate convergent evolution related to host characteristics and gill morphology. In contrast, PIC revealed medial and lateral modules in ventral anchors only. Moreover, we found evidence for ventral and dorsal anchor pairs forming two modules, supporting the notion that they play different functional roles. Integration between all identified modules was strong. We conclude that there is modular structure in the anchors of Ligophorus spp., accounted by adaptive and phylogenetic factors acting at different levels, and ventral and dorsal anchors evolve as integrated modules with specific roles in attachment.

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Ecological investigations regarding the parasite fauna of grey mullets are scarce. The present study provides a detailed description of the helminth communities of Liza aurata in the Spanish western Mediterranean and analyzes the role of spatial, temporal, and host variables in shaping the infracommunities. In total, 204 fish were collected in 2 localities, situated ca. 290 km apart, in spring and fall of 2004 and 2005. A non-metric multidimensional scaling (NMDS) was used to visualize an ordination of the infracommunities according to their relative similarities in parasite abundances. The relationship between infracommunity composition and explanatory variables (host size, locality, year, and season of harvest) was examined by permutational analysis of variance (PERMANOVA) applied to species abundances. Permutational tests for homogeneity of multivariate dispersion were used to test the null hypothesis of no differences in dispersion among groups formed by the factors whose effects were significant in PERMANOVA. A total of 33,241 helminth parasites, belonging to 18 species, was collected, i.e., 12 species of adult digeneans (23% of the parasite specimens), 3 digeneans as metacercariae (68%), 1 acanthocephalan (2.1%), and 2 monogeneans (6.5%). An important part of this helminth fauna is specialized to grey mullets, with a sizable portion of the component community restricted to the Mediterranean and northeast Atlantic. The NMDS ordination indicated high heterogeneity among infrapopulations. However, most differences at both the component and infracommunity level were related to geographic locality. In fact, the PERMANOVA showed that, among the explanatory variables considered, sampling locality accounted for the largest share of variation. The geographical differences observed may be related to local environmental characteristics or to the limited spatial dispersal of the species forming the component community. The latter was supported by the significant portion of variation explained by a 3-way interaction term. Thus, the spatial structure of our helminth infracommunities seems to be determined by a combination of differences in local environmental conditions and the transmission ability of each species at small local and time scales.

A new monogenean species, Ligophorus yucatanensis n. sp. from the gills of the flathead mullet Mugil cephalus from the Yucatan Peninsula, Mexico, is described. The new species can be differentiated from all other species of Ligophorus by the morphology of the accessory piece of the copulatory organ. Its main lobe is cylindrical, tunnelled expanded distally, slightly bowed with a characteristic membranous opening at level of medial bifurcation of the accessory piece, forming a thick-walled bulbshaped expansion that ends in a round labium. The secondary lobe is spatulate, straight, and shorter than the main lobe. In addition, the new species can be distinguished from other species by the morphology of the haptoral ventral bar, and the distal end of the vaginal duct. Furthermore the ventral anchors are shorter than those of all other species of Ligophorus reported in the Gulf of Mexico and Caribbean Sea. In addition, the zoogeographical records of Ligophorus spp. on the M. cephalus species complex are briefly reviewed and updated.

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