Deborah A. McLennan

The nature of diversity : an evolutionary voyage of discovery

All living things on earth - from individual species to entire ecosystems - have evolved through time, and evolution is the acknowledged framework of modern biology. Yet many areas of biology have moved from a focus on evolution to much narrower perspectives. Daniel R. Brooks and Deborah A. McLennan argue that it is impossible to comprehend the nature of life on earth unless evolution - the history of organisms - is restored to a central position in research. They demonstrate how the phylogenetic approach can be integrated with ecological and behavioral studies to produce a richer and more complete picture of evolution. Clearly setting out the conceptual, methodological, and empirical foundations of their research program, Brooks and McLennan show how scientists can use it to unravel the evolutionary history of virtually any characteristic of any living thing, from behaviors to ecosystems. They illustrate and test their approach with examples drawn from a wide variety of species and habitats. The Nature of Diversity provides a powerful new tool for understanding, documenting, and preserving the worlds biodiversity. It is an essential book for biologists working in evolution, ecology, behavior, conservation, and systematics.

ECOLOGICAL FITTING AS A DETERMINANT OF THE COMMUNITY STRUCTURE OF PLATYHELMINTH PARASITES OF ANURANS

Host-parasite associations are assumed to be ecologically specialized, tightly coevolved systems driven by mutual modification in which host switching is a rare phenomenon. Ecological fitting, however, increases the probability of host switching, creating incongruences between host and parasite phylogenies, when (1) specialization on a particular host resource is a shared characteristic of distantly related parasites, and (2) the resource being tracked by the parasite is widespread among many host species. We investigated the effect of ecological fitting on structuring the platyhelminth communities of anurans from a temperate forest and grassland in the United States and tropical dry and wet forests in Mexico and Costa Rica. The six communities all exhibit similar structure in terms of the genera and families inhabiting the frogs. Parasite species richness is highly correlated with the amount of time a host spends in association with aquatic habitats, a conservative aspect of both parasite and host natural history, and determined in a proximal sense by host mobility and diet breadth. The pattern of parasite genera and families within host genera across the regions examined is consistent with the prediction that ecological fitting by phylogenetically conservative species, coupled with historical accidents of speciation and dispersal, should be evidenced as a nested-subset structure; the shared requirement for aquatic habitats of tadpoles provides a baseline assemblage to which other parasite taxa are added as a function of adult host association with aquatic habitats. We conclude that parasite communities are structured by both ecological fitting and coevolution (mutual modification), the relative influences of which are expected to vary among different communities and associations.

Phylogeny and Speciation of Felids

The phylogeny of the Felidae is reconstructed using a total evidence approach combining sequences from 12S rRNA, 16S rRNA, NADH‐5, and cytochrome b genes with morphological and karyological characters. The 1504‐character data set generated two equally parsimonious trees (CI = 0.413, 1795 steps) of which a strict consensus revealed one polytomy in the placement of the bay cat group. The tree supports several traditional groupings such as the genera Panthera and Lynx and the ocelot group of small South American felids, and it provides additional resolution of relationships within and among the major felid lineages. Combining phylogenetic, distributional, and ecological data indicates that vicariant speciation has played a relatively minor role in the diversification of the felids (approximately 26% of events), while sympatric speciation has been more important than expected on theoretical grounds (approximately 51.8% of events), although postspeciation dispersal may have blurred the boundaries between sympatric, parapatric, and peripheral isolate modes. An examination of ecological changes on the felid tree shows repeated patterns of resource partitioning in time (activity patterns), space (preferred habitat type), and food (as measured by body size) among closely related species. The rapid diversification of the cats thus appears to have been associated more with ecological than with geological separation.

Phylogeny and biodiversity: Conserving our evolutionary legacy

Historical ecological studies provide information about the origins of species in an area and the origins of traits characterizing the interactions between those species and their environment. Incorporating this evolutionary information into conservation policies will broaden the base of options for making effective decisions about the preservation of biodiversity.

INTERSPECIFIC RECOGNITION AND DISCRIMINATION BASED UPON OLFACTORY CUES IN NORTHERN SWORDTAILS

Female Xiphophorus montezumae were attracted to olfactory cues from conspecific and heterospecific (X. cortezi and X. nigrensis) males when given a choice between the stimulus and water. When given a choice between conspecific and heterospecific cues, females only demonstrated a strong preference for the conspecific stimulus when it was matched against X. nigrensis. Female X. nigrensis were attracted to olfactory cues from their close relative, X. cortezi, but did not respond to cues from the more distantly related X. montezumae. They preferred the scent of their own males to the olfactory cues of both heterospecific species. Our results indicate that X. cortezi and X. nigrensis share an apomorphic change in some aspect of their olfactory cue‐receiver system that is not shared with X. montezumae. We also uncovered an asymmetry in response based on olfactory stimuli in these fishes: X. montezumae is moderately attracted to the cue from X. nigrensis, whereas X. nigrensis does not respond to the cue from X. montezumae at all.

Integrating phylogeny and experimental ethology : from pattern to process

Phylogenetic systematics offers ethologists a method for uncovering macroevolutionary patterns of behavioral diversity, which can be used to generate predictions for experimental investigation of microevolutionary processes. I used phylogenetic methods to disentangle the contributions of various selection pressures in the evolution of sexually dimorphic nuptial coloration in gasterosteid fishes in general, and Gasterosteus aculeatus in particular. Based upon the phylogenetic relationships between breeding colors and breeding behaviors in the gasterosteids, three predictions relevant to a discussion of nuptial coloration in G. aculeatus were proposed. The origin and elaboration of nuptial coloration was (1) weakly associated with male/male interactions (intrasexual selection), (2) moderately associated with parental care (natural selection) and (3) strongly associated with male/female interactions (intersexual selection). I tested the hypothesized macroevolutionary relationships between male breeding color and behavior in two sets of experiments studying changes in male nuptial coloration across the breeding cycle and female choice based on intensity of male color. The results of these experiments at the microevolutionary level of analysis corroborated the macroevolutionary predictions and confirmed the original intuition of previous researchers: male body color intensity is an important component of female mate choice in G. aculeatus. This study demonstrates that a phylogenetic hypothesis can generate predictions that are experimentally testable and falsifiable.

Comparative Study of Adaptive Radiations with an Example Using Parasitic Flatworms (Platyhelminthes: Cercomeria)

Studies of adaptive radiations require robust phylogenies, estimates of species numbers for monophyletic groups within clades, assessments of the adaptive value of putative key innovations, and estimates of the frequency of speciation modes. Four criteria are necessary to identify an adaptive radiation within the parasitic platyhelminths: (1) a group contains significantly more species than its sister group, (2) species richness is apomorphic, (3) apomorphic traits enhance the potential for adaptively driven modes of speciation (sympatric speciation and speciation by peripheral isolation via host switching), and (4) the frequency of adaptively driven speciation modes is high within the group when compared with data from free-living groups. Only the species-rich Monogenea fulfill all four criteria. The Digenea and Eucestoda also are more species rich than their sister groups, their species richness is derived, and they possess unique characters that increase the potential for host switching to occur However, because there is not enough information to determine whether the frequency of adaptive modes of speciation is high for those groups, we cannot yet assert that their radiations have been adaptive.

Macroevolutionary patterns of morphological diversification among parasitic flatworms (Platyhelminthes : Cercomeria)

Patterns of parasite morphological diversification were investigated using a morphological data base for the parasitic platyhelminths comprising 1,459 characters analyzed using phylogenetic systematic methods. Only 10.8% of the 1,882 character transformations are losses, casting doubt on views that parasites are secondarily simplified and exhibit degenerate evolution. Chi‐squared analysis indicates that character loss in the Digenea and Monogenea occurs in proportion to total change and is disproportionately lower within the Eucestoda. In the Digenea fewer female characters and more male characters have been lost than expected by the total number of characters in that group, and more male and more nonreproductive characters have been lost in proportion to their distribution across groups. In the Monogenea fewer nonreproductive and more larval characters have been lost than expected within the group, and female character loss is high relative to other groups. In the Eucestoda fewer female and more larval characters have been lost than expected within the group, whereas loss of male and nonreproductive character is low, and loss of larval characters is high, compared to the other groups. Patterns of character loss result partially from characters that show repeated (homoplasious) loss in different groups. High consistency index and low homoplasy slope ratio values indicate that the parasitic platyhelminths show unusually low levels of homoplasy, casting doubt on views that parasite morphology is unusually adaptively plastic. Homoplasy within the monogeneans occurs in proportion to overall character change, is slightly higher than expected in the digeneans, and is much lower than expected within the eucestodes. Homoplasy occurs less often than expected in larval characters, and more often than expected in nonreproductive characters in the Digenea. Monogeneans show more homoplasy than expected for larval characters both within and among groups. Eucestodes show fewer homoplasious male and nonreproductive, and more homoplasious larval, characters than expected within the group, and higher homoplasy in larval characters and lower homoplasy in female and nonreproductive characters among groups.

Male mate choice based upon female nuptial coloration in the brook stickleback, Culaea inconstans (Kirtland)

Female brook sticklebacks develop distinct nuptial coloration following ovulation. When given a choice between a gravid, uncoloured, inter-spawning interval female and a gravid, nuptially coloured female, male brook sticklebacks from two Ontario populations (1) approached the nuptially coloured female first, (2) directed their first courtship pummel towards her, and (3) spent significantly more time with her than with the inter-spawning interval female. Males from Tooley Creek spent more of their time near the nuptially coloured female engaged in courtship behaviour and more of their time near the inter-spawning interval female engaged in aggressive behaviour. Males from Algonquin Park showed no diVerence in the intensity of courtship or aggression directed towards the two females. These results corroborate the hypothesis that female nuptial coloration plays a role in signalling courtship readiness in at least some gasterosteids. The hypothesis that males should become more discriminating in their choice of spawning partners as they increase their parental investment was tested by comparing male mate choice based upon female colour for males with no eggs, one clutch and two clutches. The results corroborated the hypothesis: compared to when they were eggless, males with two clutches (1) spent less time with the inter-spawning interval female, (2) spent more time with the nuptially coloured female, (3) spent a greater proportion of the time near the nuptially coloured female engaged in courtship behaviour, and (4) displayed a trend towards increasing levels of aggression directed towards the inter-spawning interval female with increasing parental investment. ? 1995 The Association for the Study of Animal Behaviour

CLADOGRAM VERSATILITY: EVOLUTION AND BIOGEOGRAPHY OF ACANTHUROID FISHES

A highly corroborated cladogram of acanthuroid fishes is used to explore several aspects of the biogeographic and microevolutionary events during the evolution of the group. Five events in acanthuroid evolution are documented here, which demonstrate not only the power of cladograms to provide adaptational hypotheses, but also how they can be used to frame significant questions for further research. Biogeographic analysis indicates that basal cladogenesis of the acanthuroids must have occurred prior to the Eocene (50‐55 mya), because at least the basal lineages of the Acanthuridae, one of the last acanthuroid families to have evolved, were present at that time. In addition, optimization of current distributions suggests that the ancestors of the Acanthurinae, of the Acanthuridae, and of the Acanthuroidei each had an Indo–West Pacific distribution. Subsequent dispersion and/or speciation in one or more of these lineages may have been related either to the closure of the Tethys (ca 20 mya), or to colonization from the Pacific of the Caribbean/Atlantic prior to the formation of the Panamanian isthmus. Hypotheses about adaptation in acanthuroids begin with a discussion of several cases of dietary plasticity. The importance of information from fossils is illustrated in an investigation of predorsal bone evolution. Morphologically divergent and convergent tendencies in the modification of the pelvic fins concludes the examples. The morphologically most derived genus, Ctenochaetus, seems the least active taxon evolutionarily in the characters investigated.