Trophic differentiation in the phylogenetically young Cyprinodon species flock (Cyprinodontidae, Teleostei) from Laguna Chichancanab (Mexico)

Analysis of the gut contents of six syntopic Cyprinodon species from Laguna Chichancanab, Mexico, shows that the supposed basal form, C. beltrani , feeds on detritus and has the longest gut. All other species have significantly shorter guts. Whereas C. simus exclusively ingests detritus despite short gut length, all other species additionally rely on different benthic invertebrates. Each species has developed specific preferences: C. labiosus (amphipods), C. verecundus (bivalves), C. maya (ostracods and gastropods) and C. esconditus (ostracods and chironomid larvae). However, all of them ingest a minimum of c. 40% detritus. According to Schoener's index, pairwise niche overlap varies from c. 40% to over 95% in the species flock. Contrary to this, there is almost no overlap between the members of the flock and the only other native fish species, Gambusia sexradiata , which feeds nearly exclusively on terrestrial arthropods dropping on the water surface. Recently, the cichlid genus Oreochromis and the characid Astyanax fasciatus gained access to the lake. The cichlids show moderate to high niche overlap, whereas in A. fasciatus it is low. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 125–134.     

Delimiting species in the hoplonemertean genus Tetrastemma (phylum Nemertea): morphology is not concordant with phylogeny as evidenced from mtDNA sequences

The marine genus Tetrastemma contains monostiliferous hoplonemertean (phylum Nemertea) species which mostly undergo direct development with no free-swimming stages of larvae. The lack of a pelagic phase, and the fact that many benthic species lay eggs attached to the bottom substrate, are obvious restrictions on dispersal and gene flow. Nevertheless, some of the species, for example T. candidum and T. melanocephalum , are described as ubiquitous and are reported from waters all over the world. We studied genetic variation and evolutionary relationships in order to assess whether they are concordant with external characters in samples of nine morphologically distinct forms formally named as different Tetrastemma species, from different geographical localities. We estimated the phylogeny and species network based on 539 base pairs of the mitochondrial protein-coding gene cytochrome oxidase-1 (CO1) for 30 ingroup specimens. From this, we assessed the evolutionary history and phylogenetic relationships between these forms. We conclude that in most cases there was no correspondence between evolutionary lineage and morphotype. Our results thus indicate that morphological species delimitation in nemerteans may be questionable, and that this in turn may have a profound effect upon estimates of species diversity within the phylum.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 201–212.     

New data on the Cainotheriidae (Mammalia, Artiodactyla) from the early Oligocene of south-western France

The Cainotheriidae are small artiodactyls that suddenly appeared in the late Eocene of western Europe. A revision of early Oligocene cainotheriid lineages is proposed on the basis of newly dated material from the Quercy Phosphorites (south-western France). A significant diversification of the group occurred at the end of the Eocene. Few species seem to have persisted through the Eocene/Oligocene boundary, but the Cainotheriidae subsequently diversified rapidly during the early Oligocene.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 145−166.     

Two new species from Gabon show the need to reduce Commitheca to the synonymy of Pauridiantha (Rubiaceae, Pauridiantheae)

Two new Pauridiantha species (Pauridiantheae, Rubiaceae) from Gabon are described and illustrated. Pauridiantha pleiantha is characterized by a combination of entirely glabrous stems, stipules with a narrow base, and large inflorescences. Pauridiantha smetsiana is characterized by entirely glabrous stems, narrow, early deciduous stipules, leaves red-tinged when dry, and small inflorescences. The two new species, especially P. smetsiana , close the morphological gap between Commitheca and Pauridiantha in combining typical Commitheca characters, such as glabrous stems and a narrow stipular base, with Pauridiantha characters, such as eucamptodromous venation. A detailed morphological comparison (including ovary structure, seed morphology, exotesta anatomy, and pollen morphology) between the two genera is given; in conclusion, Commitheca is relegated to the synonymy of Pauridiantha . The necessary new specific combinations are provided.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 105–117.     

Ecological opportunity and levels of morphological variance within freshwater stickleback populations

The 'ecological opportunity' hypothesis predicts that when interspecific competition or predation is reduced, populations will exhibit increases in phenotypic variance as a result of colonization and adaptation to vacant or underutilized ecological niches (i.e. character release). We assessed this hypothesis by examining morphological diversity within stickleback populations in 40 undisturbed lakes from six islands off the mid-coast of British Columbia, Canada. Because larger lakes with well-developed littoral and limnetic zones will have greater trophic niche diversity than smaller lakes with only littoral zones, we predicted a positive association between lake size and variation in trophic morphology. Conversely, reduced vertebrate predation in small bog lakes allows increased variance in defensive structures without costs to fitness. Consistent with both predictions, we observed that phenotypic variance in two traits that are involved in feeding (gape width and pectoral fin length) increased with lake size while variability in defence structures (lateral plate number and dorsal spine length) was inversely related to lake size. Moreover, increased variance in defence morphology was accentuated in populations with severe armour reduction (spine loss, decreased plate overlap), another strong indicator of reduced vertebrate predation. In the majority of cases, these patterns were repeatable among islands, independent from the geographical distance between lakes, and arose from a combination of high variance within each of the sexes and increases in sexual dimorphism. These findings suggest that character release can be trait-specific and reflect the combined effects of competition, predation and habitat heterogeneity.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 297–308.     

'Shouldering' exaggerated genitalia: a unique behavioural adaptation for the retraction of the elongate intromittant organ by the male rove beetle (Aleochara tristis Gravenhorst)

Complex genitalia are ubiquitous among arthropods, but little attention has been given to the fact that the evolution of such elaborate structures may have led to biomechanical constraints that hinder their usage. In the rove beetle, Aleochara tristis, the male's intromittant organ consists of a long flagellum that is more than twice the body length. It is introduced into the spermathecal duct of the female during copulation. The flagellum apparently functions as a guiding rod for a tube growing from the spermatophore that the male deposits in the female's genital chamber. The extraordinary length of the intromittant organ poses a unique physical challenge for the male. During its retraction from the female after mating, the flagellum is under considerable tension. Any sudden release of this tension would result in the flagellum becoming severely entangled, preventing the male from mating again. In response to this novel physical challenge, males have apparently evolved a specialized behavioural adaptation that prevents entanglement after copulation. While retracting the flagellum from the female, the male secures it between a wing shoulder and the pronotum ('shouldering'), holding it taut for about one half of its length. This allows the stepwise retraction of the flagellum from the female and allows it to be retracted back into the male's body in an orderly fashion. This is, to our knowledge, the first demonstration of a behavioural adaptation that has evolved to ameliorate the biomechanical problems caused by exaggerated genital morphology.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 307–312.     

Is the last glaciation the only relevant event for the present genetic population structure of the meadow brown butterfly Maniola jurtina (Lepidoptera: Nymphalidae)?

Phylogeographical studies are available for a considerable number of European species, but few analyses exist for temperate species with very large and fairly continuous populations that are also absent from Northern Europe. Therefore, we studied the butterfly Maniola jurtina as a model for this group. The species has two major genetic lineages (mean genetic distance between lineages: 0.033; F _CT: 0.052), most probably evolving in glacial differentiation centres in the western and eastern Mediterranean. The onset of this differentiation might have been the beginning of the last glacial stage maximum some 40 kyr bp . A hybrid zone between these two lineages exists in western Central Europe. No genetic substructures have been found within the two lineages ( F _SC: 0.017) and average genetic distances are very small. Therefore, it is highly probable that postglacial expansion was of the phalanx type. There is, at most, very limited differentiation at regional and local scales. However, the genetic diversity within populations is high (means: A : 2.68; H _E: 17.2%; P : 78%), as would be predicted for such a common species. Comparison of these results with a published allozyme analysis revealed a similar phylogeographical pattern, but lower genetic diversity in the latter. Morphological patterns of wings and genitalia show similar geographical patterns as allozyme data.   © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 419–431.     

A new species of Selaginella from Hainan Island, China, with a key to the Hainan species

Selaginella hainanensis X.C.Zhang & Noot. sp. nov. , a new species from Hainan Island, South China, is described and illustrated. Diagnostic notes and a key to the species distributed in Hainan are also provided.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 323–327.     

Revision of the subgenus Udamochiras of Melaloncha bee-killing flies (Diptera: Phoridae: Metopininae)

The genus Melaloncha Brues is defined and groundplan character states established based on outgroup comparison with Phalacrotophora Enderlein and Melittophora Brues. Major groupings within Melaloncha are recognized, and two subgenera are established: Udamochiras Enderlein (type species M. colossia (Enderlein)) and Melaloncha s.s. (type species M. pulchella Brues). Subgenus Udamochiras is revised and 42 species are recognized, including the following 33 new to science: M. anaticula , M. angustifrons , M. apicula , M. aprica , M. basella , M. biseta , M. brevicarina , M. carinata , M. compressicauda , M. exigua , M. falcata , M. flavilata , M. hamata , M. hansoni , M. horologia , M. individa , M. lobata , M. maculifrons , M. parkeri , M. paxilla , M. premordica , M. rhampha , M. rhypopoda , M. rostrata , M. sinuosa , M. spatula , M. spicula , M. triangularis , M. trua , M. valeria , M. vargasi , M. villosa , M. woodi . Melaloncha simillima Borgmeier is removed from synonymy with M. piliapex Borgmeier and reinstated as a separate species; lectotypes are designated for both.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 1−42     

A new species of Lepidostrobus from the Upper Devonian of Xinjiang, China and its bearing on the phylogenetic significance of the order Isoëtales

Lepidostrobus xinjiangensis sp. nov. is described from Upper Devonian rocks of the eastern Junggar Basin, Xinjiang, north-west China. It provides new insights into the reproductive diversification and phylogenetic relationships of lycopsids in the Late Devonian. The strobilus shares more characteristics with arborescent lycopsids than any herbaceous forms, and falls within the diagnosis of Lepidostrobus . Each sporophyll consists of a deltoid pedicel and a triangular lamina. Sporophylls are horizontally inserted into the strobilus axis in low spirals. The pedicel, with lateral alations and an abaxial keel, extends distally into the upturned lamina and downturned heel, producing a peltate appearance. A single sporangium with terminal longitudinal dehiscence is a radially elongate, dorsiventrally flattened ovoid and is attached along its length to the adaxial surface of the pedicel. A column-like subarchesporial pad is found in the sporangium. A possible ligule occurs on the adaxial surface of the pedicel distal to the sporangium. The strobilus is microsporangiate, containing Lycospora -type spores with granulate ornamentation and an equatorial flange. Based on this new species, the reproductive diversification and evolutionary pattern in arborescent lycopsids from the Devonian through the Carboniferous are discussed in a phylogenetic framework. We suggest that the reproductive strategies represented by bisporangiate- and monosporangiate-strobili had proliferated by the Late Devonian, which implies that phylogenetically advanced arborescent lycopsids bearing Lepidostrobus strobili had an earlier origin than previously thought.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 55−67.     

Dispersal history of a spider (Stegodyphus lineatus) across contiguous deserts: vicariance and range expansion

Israel marks a crossroads between three continents encompassing several phytogeographical and zoogeographical zones. In this complex area, the flow of species from different biogeographical regions creates opportunities to study how geographical division and colonization routes affect current distribution and structure of resident populations of organisms associated with desert and arid environments, habitats that may have persisted throughout Pleistocene glacial periods. The present paper analyses the population history of the spider Stegodyphus lineatus in the contiguous Negev and Judean deserts in Israel using allozyme and mtDNA variation. The distinct patterns of variation indicate that Judean and Negev populations are vicariant lineages. The residence time was longer in Judea, where populations were more polymorphic for mtDNA, showed isolation by distance and were less structured than in the Negev. The Negev population, possibly linked to other Mediterranean populations of S. lineatus , consisted of two subdivisions derived from a recent eastward expansion across the central Negev watershed. Despite differences in age and level of structure, all lineages show similar dispersal processes dominated by restricted gene flow. The distribution patterns of allozyme and mtDNA markers are unrelated to geographical patterns of precipitation and vegetation. Rather, they follow large-scale topographic features, namely the water divide between Mediterranean and Afro-Syrian rift drainages and between eastern and western Negev drainages.   © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 739–754.     

Difference in the predation impact enhanced by morphological divergence between introduced fish populations

The strength of predation impact on recipient environments may vary among introduced populations due to their local adaptations to different prey. We examined whether functional diversification associated with morphological differences may be observed among the introduced populations of invasive bluegill sunfish Lepomis macrochirus (Perciformes, Centrarchidae) in Japan. The two examined populations are morphologically different, although they were recently derived from a common American source and colonized in different lakes. We performed a laboratory experiment wherein these populations were fed the benthic (chironomid larva) and the pelagic prey (daphnid zooplankton). The results revealed that a population colonizing in a shallower lake and foraging on benthic invertebrates in the wild had a greater impact on the benthic prey, whereas the other population colonizing in a deeper lake and foraging on crustacean zooplankton have consumed the pelagic prey more efficiently. A series of regression analyses showed that morphological differences among individuals were responsible for these population differences. The evidence obtained suggests that morphological adaptations by introduced bluegill populations enhance the strength of predation impact on a prey resource consumed in a relevant environment, but reduce the impact on the other prey. Thus, although the introduced Japanese populations were recently derived from a common ancestor, the predation impacts on the native prey community vary due to morphological adaptations to different prey.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 601–610.     

Molecular systematics and patterns of morphological evolution in the Centropagidae (Copepoda: Calanoida) of Argentina

Recent studies have shown the value of complementing standard taxonomy with genetic analyses to reveal cryptic diversity and to aid in the understanding of patterns of evolution. We surveyed variation in the COI mitochondrial gene in members of the three genera of centropagid copepods from the inland waters in Argentina. In general, we found a close association between molecular and morphological systematics in this group. Similar to findings for marine calanoids, genetic distances within Boeckella species were modest (< 4%), while distances among morphospecies were much larger (> 11%). Parabroteas is currently monotypic, although we detected cryptic genetic diversity, with two lineages showing 5.5% divergence. In contrast, Karukinka was not a valid genus, apparently representing an interesting and atavistic offshoot of B. poppei , a result reinforcing the value of considering both morphological and molecular evidence. Moreover, we used combined genetic and morphological information, analysed with maximum likelihood methods, to evaluate the common assumption that evolution tends to proceed via the loss of structures in crustaceans. Although analysis of other taxa and character types is required to evaluate fully the reduction hypothesis, our results suggest that structures may be gained readily as well as lost.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 279–292.     

Sensory and skeletal development and growth in relation to the duration of the embryonic and larval stages in damselfishes (Pomacentridae)

Several trends were found in comparisons of rates of growth and development of larvae of four coral-reef damselfishes ( Chromis atripectoralis, Pomacentrus amboinensis, Premnas biaculeatus, Acanthochromis polyacanthus ), which were reared under constant temperature conditions in the laboratory, and which varied in their early life stage durations (respectively, egg stage durations were 2, 4, 7, 16; larval stage durations were 25, 23, 14, 0). Parameters measured included standard length, muscle area, eye diameter, and selected stages of retinal development, olfactory development, and skeletal ossification. Rates of ossification and olfactory development were inversely related to growth rate (in length and muscle area) among most species. Rates of eye growth and retinal development were also negatively correlated among all species. These results are consistent with the concept of a trade-off between growth and development. We observed a positive relationship between egg stage duration and developmental rate, and a negative correlation between larval stage duration and developmental rate. Acanthochromis developed slower than predicted by the general trends, although retinal development was very rapid. Specific retinal stages correlated with settlement, regardless of ontogenetic rates. Olfactory development was especially rapid in the anemonefish Premnas biaculeatus , which imprints to olfactory cues as an embryo. Skeletal ossification was rapid in species with pelagic larvae, and much slower in the benthic brooder. Literature-derived data on size and age at hatching and settlement from > 40 species of tropical pomacentrids were transformed into growth and developmental rates; correlations of these literature-derived parameters were mostly consistent with our controlled four-species comparison.  © 2003 The Linnean Society of London, The Biological Journal of the Linnean Society , 2003, 80 , 187−206.     

High photobiont diversity associated with the euryoecious lichen-forming ascomycete Lecanora rupicola (Lecanoraceae, Ascomycota)

The diversity and phylogenetic position of photobionts in the widespread saxicolous, crustose lichen-forming ascomycete Lecanora rupicola s.l. is presented. The algal partners of this lichen species complex belong to diverse and unrelated lineages in the genus Trebouxia . Specimens were sampled from different habitats and geographical origins. Either whole thallus DNA extractions or minute fragments of the algal layer of the lichen thallus were subjected to polymerase chain reaction, using primers that specifically amplify internal transcribed spacer rDNA of the photobionts. No correlations between different chemical races of L. rupicola with particular lineages of Trebouxia spp. were found. Irrespective of the different algal partners, all lichen thalli abundantly developed ascomata. L. rupicola apparently maintains full fecundity with a low degree of selectivity for photobionts, which promotes the occurrence of this lichen-forming species in ample ecological situations.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 283–293.     

Empirical evaluation of phenotype–environment correlation and trait utility with allopatric and sympatric whitefish, Coregonus lavaretus (L.), populations in subarctic lakes

Adaptive phenotypic divergence of sympatric morphs in a single species may have significant evolutionary consequences. In the present study, phenotypic impacts of predator on zooplankton prey populations were compared in two northern Finnish lakes; one with an allopatric whitefish, Coregonus lavaretus (L.), population and the other with three sympatric whitefish populations. First, we examined whether there were phenotypic associations with specific niches in allopatric and sympatric whitefish. Second, trait utility (i.e. number of gillrakers) of allopatric and sympatric whitefish in utilizing a pelagic resource was explored by comparing predator avoidance of prey, prey size in environment, and prey size in predator diet. The allopatric living large sparsely rakered (LSR) whitefish morph, was a generalist using both pelagic and benthic niches. In contrast, sympatric living whitefish morphs were specialized: LSR whitefish was a littoral benthivore, small sparsely rakered whitefish was a profundal benthivore and densely rakered (DR) whitefish was a pelagic planktivore. In the lake with allopatric whitefish, zooplankton prey did not migrate vertically to avoid predation whereas, in the lake with sympatric whitefish, all important prey taxa migrated significantly. Trait utility was observed as significantly smaller size of prey in environment and predator diet in the lake with DR whitefish than in the lake with only LSR whitefish.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 561–572.     

Phylogenetic relationships between parthenogens and their sexual relatives: the possible routes to parthenogenesis in animals

In theory, parthenogenetic lineages have low evolutionary potential because they inexorably accumulate deleterious mutations and do not generate much genotypic diversity. As a result, most parthenogenetic taxa occupy the terminal nodes of phylogenetic trees. The rate and mode of development of parthenogenesis are important factors to consider when assessing its costs and benefits since they determine both the level of genetic diversity and the ecological adaptability of the resulting lineages. The origin of parthenogenesis is polyphyletic in many taxa, suggesting that genetic systems maintaining sexuality are often labile. In addition, the loss of sex may be achieved in several ways, leading to parthenogenetic lineages with distinct genetic profiles. This could then influence not only the fate of such lineages in the long term, but also the outcome of competition with their sexual counterparts in the short term. In this paper, we review the possible evolutionary routes to parthenogenesis based on a survey of the phylogenetic relationships between sexual and parthenogenetic lineages in a broad range of animals. We also examine the different mechanisms by which parthenogenetic lineages could arise, and discuss the influence of these mechanisms on both the genetic properties and the ecological life styles of the resulting lineages.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 79 , 151–163.     

Generic delimitation and a new infrageneric system in the genus Holcoglossum (Orchidaceae: Aeridinae)

Based on herbarium and field observations, we confirm the generic delimitation of Holcoglossum and present a new infrageneric system for the genus. Our work supports Holcoglossum as a natural group including H. amesianum (Rchb.f.) Christenson and H. subulifolium (Rchb.f.) Christenson. The new infrageneric system of Holcoglossum includes two subgenera: Holcoglossum subgen. Brachycentron subgen. nov. and Holcoglossum subgen. Holcoglossum , the latter comprising two new sections, Holcoglossum sect. Holcoglossum and Holcoglossum sect. Sorotylos .  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 465–468.     

Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus Pecado

The genus Labulla Simon is circumscribed in phylogenetic terms to include the species Labulla thoracica (Wider), L. flahaulti Simon and L. machadoi sp. nov. The genital anatomy of the genus is described in detail and the taxonomy of the genus is reviewed. The monophyly of Labulla is supported by numerous morphological apomorphies of the male palp and female epigynum. Based on a cladistic analysis, a new genus, Pecado gen. nov. , is erected to place Labulla impudica Denis, from Northern Africa. Lepthyphantes insularis Saito and ' Labulla ' nepula Tikader, both formerly included in Labulla , are not congeneric with the type species of Labulla .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 359–404.