Determining variation in the success of New Zealand land birds

The New Zealand avifauna has changed considerably since the arrival of humans approximately 1000 years ago. The extinction of native species followed by their replacement with introduced exotic species is recognized as having greatly reduced the distinctiveness of the avifauna. Using phylogenetically independent methods, I tested whether life history and ecological correlates of extinction, introduction and persistence were significant across species and within taxa. Extinct land bird species were, on average, and independent of phylogenetic relatedness, larger-bodied than extant species. Introduction success was found to be significantly associated with increasing species body size, generation time and indices of human effort. In contrast, the geographical distribution of extant land birds was correlated with traits associated with high population growth rates (i.e. small, rapidly developing species, with high fecundity). My results suggest that selection pressures have changed significantly since the arrival of human colonists in New Zealand and that these changes have favoured different types of land birds, to the general detriment of endemic species. Comparative methods that explicitly examine historical changes are necessary to elucidate the changing roles of biological and extrinsic traits in the differential success of species in any assemblage.     

Phylogenetic perspective on ecological niche evolution in american blackbirds (Family Icteridae)

Analysis of ecological characters on phylogenetic frameworks has only recently appeared in the literature, with several studies addressing patterns of niche evolution, generally over relatively recent time frames. In the present study, we examined patterns of niche evolution for a broad radiation of American blackbird species (Family Icteridae), exploring more deeply into phylogenetic history. Within each of three major blackbird lineages, overlap of ecological niches in principal components analysis transformed environmental space varied from high to none. Comparative phylogenetic analyses of ecological niche characteristics showed a general pattern of niche conservatism over evolutionary time, with differing degrees of innovation among lineages. Although blackbird niches were evolutionarily plastic over differing periods of time, they diverged within a limited set of ecological possibilities, resulting in examples of niche convergence among extant blackbird species. Hence, an understanding of the patterns of ecological niche evolution on broad phylogenetic scales sets the stage for framing questions of evolutionary causation, historical biogeography, and ancestral ecological characteristics more appropriately.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 869–878.     

Inferring life history from ovipositor morphology in parasitoid wasps using phylogenetic regression and discriminant analysis

Our understanding of the critical shift in life history among insect parasitoids from ectoparasitism to endoparasitism is hindered by the fact that this particular life history trait is not known for many phylogenetically important taxa. One method of coping with this problem is to seek correlations between the ovipositor structure and this life history trait among taxa whose life history is known, and then to use this to infer the trait in species whose life history in unknown. In one group of parasitoid wasps, the Ichneumonoidea (Hymenoptera), we scored a total of 20 morphometric and morphological characters of the ovipositor for 41 species whose life histories are known – representing all the main clades in which there have been independent transitions to endoparasitism plus a broad range of the ectoparasitoid groups; we then used phylogenetic regression and discriminant analyses to infer the life history of four species whose life histories are unknown. To allow for the effect of phylogenetic non-independence in the discriminant analysis, we carried out analyses using different randomly chosen representatives of the endoparasitoid clades (phylogenetic regression controls for phylogeny). These two methods gave congruent results from which we conclude that Megalohelcon and Gnamptodon are endoparasitoids, and Aspilodemon and Allobracon are ectoparasitoids. We discuss the consequences of these inferences for our understanding of the evolution of endoparasitism in the Ichneumonoidea.  © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society , 2003, 139 , 213–228.     

Evidence for ecological speciation in the sister species Candidula unifasciata (Poiret, 1801) and C. rugosiuscula (Michaud, 1831) (Helicellinae, Gastropoda)

This paper studies the speciation of two land snail species in south-east France. By using two mitochondrial and one nuclear gene, the species are shown to be sister species within the larger clade of western European Candidula species. The species never occurred syntopically, but a narrow contact zone population was identified. Recent range expansions inferred from phylogeographical methods indicated that the present day distributions are not limited by the colonization capacity of the species. Analysis of environmental variables suggested distribution along an ecotone. The gastropod community is also correlated to this gradient. Morphometric shell analysis showed that the divergence between the sister species is a result of desiccation-resistant shell characters. We therefore conclude that the cause for the ecological speciation is most probably the establishment of the Mediterranean climate ≈ 3.2 mya. A model of ecological speciation with a moving ecotone is put forward.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 611–628.     

A new phylogenetic classification for the gymnophthalmid genera Cercosaura , Pantodactylus and Prionodactylus (Reptilia: Squamata)

Because of the poor state of knowledge of many of the gymnophthalmid genera, systematic revision is necessary to render the classification consistent with evolutionary history. To that end, I conducted a review of the species of three genera of the Cercosaurinae which appear to form a monophyletic group: Cercosaura , Pantodactylus , and Prionodactylus . Phylogenetic analysis of 61 morphological characters was conducted after specimens of all species were examined to evaluate the composition of each taxon. The phylogenetic reconstruction suggested that the genus Prionodactylus was paraphyletic. A new phylogenetic classification is proposed that synonymizes Pantodactylus and Prionodactylus with Cercosaura. Cercosaura is redefined to include 11 species and seven subspecies. A key is provided to distinguish among species.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 101−115.     

Are there body size implications for the success of globally introduced land birds?

Verbal models have hypothesized a relation between body size and the successful introduction of animal species. This relation is largely based on studies of intrinsic rate of increase in what have been termed "colonizing" species. From these studies it has been predicted that introduction success should be negatively correlated with body size across taxa but positively correlated within closely related taxa. T examine this relation for globally introduced land birds. Introduced land birds are. on average, larger bodied than extant land bird species. Across species, families, and higher family nodes, global introduction success is significantly related to decreasing body size. However, within taxa there is a significant positive relationship between introduction success and body size. I discuss possible explanations for the observed relations and conclude that an indirect but genuine relationship between the introduction success of land birds and their body size is currently the most plausible.     

Is the morphological disparity of lemur skulls (Primates) controlled by phylogeny and/or environmental constraints?

The cranial morphology of Lemur catta and of the five species of the genus Eulemur is investigated here by landmark identification and Procrustes superimpositions. This geometrical morphometric method makes it possible to describe pure shape differences independent of size effects, and to quantify differences between specimens. The aim of this study is to determine whether the morphological disparity of lemur skulls is constrained by the environment and/or by the group's phylogenetic history. First, Procrustes residuals are analysed by principal component analysis and the scatter-plots interpreted against the geographical distributions of taxa to determine whether morphology is correlated with geography. Then, a morphological distance tree is computed and compared with various cladograms reported in the literature to test for any correlation between morphology and phylogeny. Morphological disparity is found to be closely correlated with geographical distribution but independent of phylogeny. This confirms that the morphological disparity of lemur skulls is associated with a high degree of homoplasy, probably as a result of ecological constraints.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 577–590.     

Historical influences on genital morphology among sympatric species: gonopod evolution and reproductive isolation in the crayfish genus Orconectes (Cambaridae)

Freshwater crayfishes represent an incredibly diverse component of temperate aquatic ecosystems. The genus Orconectes (Cambaridae) comprises approximately 17% of total global crayfish diversity and is native to central and eastern North America. Using both Bayesian and parsimony-based phylogenetic hypotheses from complete cytochrome c oxidase subunit I (COI) mtDNA sequence data (1545 bp) along with information on male gonopod shape and species distribution patterns, we examined the evolution of male genital morphology and its potential influence on species co-occurrence. We found that sympatric species exhibited similar male genital morphology more often than expected based on the frequency of genital shapes within the genus. When evolutionary history was incorporated into this analysis, the result was no longer significant, suggesting that species co-occurrence and the associated genital shapes of sympatric congeners cannot be explained solely by interspecific ecological interactions. Our results also do not support the current subgeneric classifications within Orconectes or the monophyly of the genus, both of which have been based previously on male genital morphology.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 1–12.     

Evolution and determinants of host specificity in the genus Lamellodiscus (Monogenea)

The evolution and determinants of host specificity in Lamellodiscus species (Monogenea, Diplectanidae) were investigated. The 20 known Mediterranean species were studied, all parasites of fishes from the family Sparidae (Teleostei). An index of specificity, which takes into account the phylogenetic relationships of their fish host species, was defined. The link between specificity and its potential determinants was investigated in a phylogenetic context using the method of independent contrasts. Host specificity in Lamellodiscus species appeared to be highly constrained by phylogeny, but also linked to host size. Mapping specificity onto the parasite phylogenetic tree suggests that specialist species do not represent an evolutionary dead end, and that specialization is not a derived condition. It is hypothesized that the ability to be generalist or specialist in Lamellodiscus is controlled by intrinsic, phylogenetically-related characteristics, and that specialist species tend to use large hosts, which may be more predictable.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 431−443.     

Genetic structure and population differentiation of the Mediterranean pioneer spiny broom Calicotome villosa across the Strait of Gibraltar

The region around the Strait of Gibraltar is considered to be one of the most relevant 'hot spots' of biodiversity in the Mediterranean Basin due to its historical, biogeographical, and ecological features. Prominent among these is its role as a land bridge for the migration and differentiation of species during the Pleistocene, as a consequence of the lowering of sea level and climate changes associated with the Ice Ages. In the present study, we report a multilevel hierarchical investigation of the genetic diversity of Calicotome villosa , a common pioneer legume shrub, at the regional scale. The results of genetic analysis of progeny arrays are consistent with a predominantly outcrossing mating system in all the populations analysed. Geographically, a pattern of population isolation by distance was found, but the Strait accounted for only approximately 2% of the among-population genetic differentiation. Consequently, extensive historical gene flow appears to be the rule for this species in this area. According to the natural history traits of C. villosa (pollination, dispersal, and colonization ability), we hypothesize that gene flow must be strongly influenced by seed dispersal because pollen flow is very limited. Based on the history of trade and land use, cattle and human movements across the Strait must have strongly favoured seed dispersal. We review and discuss these results and compare them with those of other reported studies of genetic and phylogenetic differentiation across the Strait of Gibraltar. It is stressed that colonization ability, which depends upon seed dispersal and life form, can be a more critical factor in gene flow than pollination.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 39–51.     

The evolution of large brain size in birds is related to social,not genetic,monogamy

There are several hypotheses suggesting that social complexity, including pair bonding, is important in the evolution of increased brain size. I examined whether genetic or social monogamy was related to large brain size in birds. Recent work has indicated that the length and strength of pair bonds are associated with large brain size. I tested several hypotheses for the evolution of large brain size in 42 species of bird by including life history variables in a regression model. A test on 100 phylogenetic trees revealed no phylogenetic signal in brain size. Controlling for body size, a principal components analysis was run on the life history variables and degrees of extra‐pair paternity. The main principal component (PC1) was regressed on brain size revealing a strong, positive association. Social, but not genetic, monogamy was positively related to brain size. Large brain size is related to the selective pressures of procuring extra‐pair copulations whilst maintaining a social partnership. However, other life history variables also loaded positively and significantly on brain size. These results indicate that the evolution of large brain size in birds was driven by several important selective pressures. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 668–678.     

Seed dormancy and germination behaviour in two Euphrasia species (Orobanchaceae) occurring in the Swiss Alps

The optimal time for germination of a seed depends on environmental conditions of its habitat, the life cycle of the germinating plant, and the conditions for successful establishment, growing and reproduction. We studied the germination behaviour of the alpine annual Euphrasia minima and an alpine ecotype of E. salisburgensis in a lowland garden experiment. Seeds of both species and their hybrids germinated at constant (5 °C) and at varying temperatures (3–10 °C), and never before spring after seed ripening. Germination was spread over 3 years, which suggests that the seeds formed a persistent seed bank. The two species together with E. minima and E. minima hybrids differed significantly in the germination rate in the first and second spring.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 649–656.     

Speciation mirrors geomorphology and palaeoclimatic history in African laminate-toothed rats (Muridae: Otomyini) of the Otomys denti and Otomys lacustris species-complexes in the 'Montane Circle' of East Africa

We adopted an integrated systematic approach to delimit evolutionary species and describe phylogeographic, morphometric and ecological relationships in Otomys denti (from the Albertine Rift, Southern Rift in Malawi and the northern Eastern Arc Mountains) and Otomys lacustris (from the Southern Rift in Tanzania and Zambia, and the southern Eastern Arc Mountains). Molecular [cytochrome (cyt) b sequences, 1143 bp, N  = 18], craniometric (classical, N  = 100 and geometric, N  = 60) and ecological (Partial Least Squares regression of shape and ecogeographic variables) approaches show a profound, parallel disjunction between two groups: (1) Eastern Arc and Southern Rift (including the Malawi Rift) ( O. lacustris and Otomys denti sungae ) and (2) Albertine Rift ( Otomys denti denti and Otomys denti kempi ) taxa. Within both groups, cyt b sequences or craniometric analysis provided evidence for the differentiation of both southern and northern Eastern Arc from Southern Rift lineages (across the so-called Makambako Gap). Within the Albertine Rift ( denti – kempi ) lineage, populations from individual mountain ranges differed significantly in skull shape (but not size), but were similar genetically. Over-reliance in the past on very few morphological characters (e.g. number of molar laminae) and a polytypic species concept has obscured phylogenetic relationships and species discrimination in this group. We recognize at least three species in this group, and distinct lineages within two of these species. Each species or lineage was endemic to one of three regions: the Albertine Rift, the Malawi Rift or the Eastern Arc. Our result echo conclusions of recent studies of other mammalian and bird taxa and reflect the geomorphology and palaeoclimatic history of the region.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 913–941.     

Polyploidy in fishes: patterns and processes

Fishes are the most speciose group of vertebrates, with more than 24 000 species. They are characterized by great diversity in ecology, morphology, life history, behaviour and physiology. Here, the phylogenetic patterns of orders in which polyploidy has been recorded are considered, with special reference to patterns of species richness and hybridization: these orders include such phylogenetically diverse taxa as the Lepidosireniformes (lungfish) and the Perciformes (perch). Examples, predominantly drawn from the Cyprinidae and Salmonidae, are used to illustrate attributes of polyploidy in fishes. It is concluded (i) that polyploidy may have been of considerable importance in the evolution of fishes, and (ii) that fishes, with their diverse life histories, represent a useful model system with which to test theories relating to the origin and consequences of polyploidy that have been derived from work on plants.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 431–442.     

Thuja sutchuenensis: a rediscovered species of the Cupressaceae

After more than a century, the only conifer species listed as being extinct in the wild (EW) by IUCN-SSC has been rediscovered in the Dabashan Mountains of central China. The history, taxonomy, ecology and conservation of Thuja sutchuenensis are described, and illustrations, based on both the earliest and latest collection of botanical material of this tree, are provided. The taxonomic context of this rare species is discussed.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 139 , 305–310.     

Ecological niche differentiation in the Aphelocoma jays: a phylogenetic perspective

The Aphelocoma jays have become an important touchstone in behavioural ecology and biogeography – the corpus of studies of this genus makes it an important point of reference. Aphelocoma evolutionary history, nevertheless, has been the subject of two papers reaching opposite conclusions, even though they were based on the same allozyme data set. Herein, we present a second molecular data set – 500 bases of the ND2 gene – and analyse it cladistically to arrive at a new hypothesis of phylogenetic relationships. Recent hypotheses by other investigators of a hybrid origin of Aphelocoma populations are strongly contradicted. The ecological context within which these evolutionary processes are taking place is characterized using new tools for modelling ecological niches of species along a spectrum from humid tropical to dry temperate habitats. Evolutionary patterns of ecological niches are shown to consist of drastic departures from rate-uniformity and ecological niche conservatism.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 369–383.     

Propagule size and the relative success of exotic ungulate and bird introductions to New Zealand

We investigated factors affecting the success of 14 species of ungulates introduced to New Zealand around 1851-1926. The 11 successful species had a shorter maximum life span and were introduced in greater numbers than the three unsuccessful species. Because introduction effort was confounded with other life-history traits, we examined whether independent introductions of the same species were more likely to succeed when a greater number of individuals were introduced. For the six species with introductions that both succeeded and failed, successful introductions always involved an equal or greater number of individuals than unsuccessful introductions of the same species. For all independent introductions, there was a highly significant relationship between the number of individuals introduced and introduction success. When data for ungulate and bird introductions to New Zealand were combined, a variable categorizing species as ungulate or bird was a highly significant predictor of introduction success, after variation in introduction effort was controlled. For a given number of individuals introduced, ungulates were much more likely to succeed than birds.     

Fluvial range expansion, allopatry, and parallel evolution in a Danubian snail lineage (Neritidae: Theodoxus)

The snails Theodoxus danubialis and Theodoxus prevostianus form a single clade native to freshwaters of south-eastern Europe whose inter- and intraspecific relationships remain unresolved. The present study utilized a phylogeographical approach to clarify the relationship of these species as well as to reconstruct the evolutionary and demographic history of populations in the western portion of their range. Phylogenetic, population genetic, and nested clade analyses reveal a clade that has distributed itself upriver from a lower Danube River source population and become genetically distinct primarily through range expansion and localized allopatric divergence. Notably, this geographical pattern is replicated phylogenetically in the form of two cytochrome c oxidase subunit I (CO I) lineages that are present simultaneously in individual snails. Haplotypes from polymorphic individuals form two distinct clades, both of which show phylogenetic and nucleotide substitution patterns consistent with a mitochondrial origin, and whose common ancestor must have occurred in a lower Danube source population. Separated allopatrically from their Danubian relatives, populations of T. danubialis in northern Italy have also undergone substantial range expansion, much more recently than Danube watershed lineages. In addition to repeated patterns of range expansion, parallelism is found in T. prevostianus , which is shown to be a nonmonophyletic taxon of remarkable morphological and ecological similarity.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 603–617.     

Combining molecular, morphological and ecological data to infer species boundaries in a cryptic tropical pitviper

Few operational methods exist for delimiting species boundaries, and these usually require sampling strategies that are unrealistic for widespread organisms that occur at low densities. Here we apply molecular, morphological and ecological species delimitation criteria to a wide-ranging, fragmented group of Asian green pitvipers, the Popeia popeiorum complex. A mitochondrial DNA phylogeny for the group indicates two well-differentiated clades, corresponding mainly to northern and southern parts of its range. Strong phylogeographical structure within each clade suggests isolation in forest refugia during the Pliocene and a southward colonization of the Sunda islands during the Pleistocene. Multivariate analysis of morphological characters reveals a generally conserved pattern of geographical variation, incongruent with the recovered phylogenetic history. We compare groups delineated by mtDNA variation to morphological and ecological divisions in the complex, and discuss the implications of these for the taxonomy of the group. Discordance between species boundaries inferred from different criteria suggests that combining independent sources of data provides the most reliable estimation of species boundaries in organisms that are difficult to sample in large numbers.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 343–364.