Comparative body size relationships in pocket gophers and their chewing lice

In this paper, we use the method of independent contrasts to study body size relationships between pocket gophers and their chewing lice, a host-parasite system in which both host and parasite phylogcnies are well studied. The evolution of body size of chewing lice appears to be dependent only on the body size of their hosts, which confirms the 1991 findings of Harvey and Keymer. We show that there is a positive relationship between body size and hair-shaft diameter in pocket gophers, and that there is also a positive relationship between body size and head-groove width in chewing lice. Finally, we show a positive relationship between gopher hair-shaft diameter and louse head-groove width. We postulate that changes in body size of chewing lice are driven by a mechanical relationship between the parasite's head-groove dimension and the diameter of the hairs of its host. Louse species livingon larger host species may be larger simply because their hosts have thicker hairs, which requires that the lice have a wider head groove. Our study of gopher hair-shaft diameter and louse head-groove dimensions suggest that there is a 'lock-and-key' relationship between these two anatomical features.     

Faster evolutionary rates in endosymbiotic bacteria than in cospeciating insect hosts

The hypothesis of a universal molecular clock holds that divergent lineages exhibit approximately constant rates of nucleotide substitution over evolutionary time for a particular macromolecule. We compare divergences of ribosomal DNA for aphids (Insecta) and Buchnera, the maternally transmitted, endosymbiotic bacteria that have cospeciated with aphids since initially infecting them over 100 million years ago. Substitution rates average 36 times greater for Buchnera than for their aphid hosts for regions of small-subunit rDNA that are homologous for prokaryotes and eukaryotes. Aphids exhibit 18S rDNA substitution rates that are within the range observed in related insects. In contrast, 16S rDNA evolves about twice as fast in Buchnera as in related free-living bacterial lineages. Nonetheless, the difference between Buchnera and aphids is much greater, suggesting that rates may be generally higher in bacteria. This finding adds to evidence that molecular clocks are only locally rather than universally valid among taxonomic groups. It is consistent with the hypothesis that rates of sequence evolution depend on generation time.     

Impact of pocket gopher disturbance on plant species diversity in a shortgrass prairie community

Summary We examined the impact of pocket gopher disturbances on the dynamics of a shortgrass prairie community. Through their burrowing activity, pocket gophers (Thomomys bottae) cast up mounds of soil which both kill existing vegetation and create sites for colonization by competitively-inferior plant species. Three major patterns emerge from these disturbances: First, we show that 10 of the most common herbaceous perennial dicots benefit from pocket gopher disturbance; that is, a greater proportion of seedlings are found in the open space created by pocket gopher disturbance than would be expected based on the availability of disturbed habitat. Additionally, these seedlings exhibited higher growth rates than adjacent seedlings of the same species growing in undisturbed habitat. Second, we tested two predictions of the Intermediate Disturbance Hypothesis and found that species diversity was greatest for plots characterized by disturbances of intermediate age. However, we did not detect significant differences in diversity between plots characterized by intermediate and high levels of disturbance, indicating that many species are adapted to or at least tolerant of high levels of disturbance. Third, we noted that the abundance of grasses decreased with increasing disturbance, while the abundance of dicots increased with increasing disturbance.     

Survivorship of penstemon grandiflorus in an oak woodland: combined effects of fire and pocket gophers

Summary The effects of fire and pocket gophers, Geomys bursarius, on the survivorship of Penstemon grandiflorus growing in an oak woodland in Minnesota were studied from 1986 to 1990. Plants growing in sparse vegetation experienced mortality rates twice that of plants growing in dense vegetation. This difference was due partly to pocket gophers whose earth moving activities reduce the density of vegetation and bury and kill individual Penstemon plants. Laboratory feeding trials showed that gophers readily eat Penstemon, particularly the fleshy roots. An experiment involving the removal of 25–75% of the root tissue in 90 plants showed that root loss significantly reduced survivorship, suggesting that gopher herbivory might also kill plants. When gophers were experimentally excluded, plants growing in sparse vegetation exhibited significantly lower mortality rates than those growing in dense vegetation. Plants in the smallest size class exhibited reduced survivorship following a late spring burn; however, overall patterns of survivorship of plants in burned areas did not differ markedly from those in the unburned areas. A longitudenal analysis of plants with different reproductive histories revealed no survivorship cost to reproduction. Mortality rates decreased with increasing plant size. Small plants were more likely to be killed by fire and by being buried under gopher mounds. Differences in underground energy reserves of small and large plants can account for most of the survivorship patterns observed in this study. The study shows that within openings of the oak woodland, fire and gophers reduce the survival of individual Penstemon plants. Nevertheless, since both gophers and fire also serve to perpetuate suitable habitat in the woodland, Penstemon is ultimately dependent on both for its long term persistence in the landscape.     

Soil translocation by pocket gophers in a Mima moundfield

Summary We measured soil translocation due to the tunneling of valley pocket gophers (Thomomys bottae) in a Mima moundfield at Miramar Mounds National Landmark, San Diego, California, from December, 1984 through December, 1985. We placed 1-l soil plugs containing 20 11-g iron pellets into pocket gopher tunnels at locations between mound tops and points about one mound radius beyond mound edges. After about 4–10 d, sites to which the marker-containing soil had been translocated were located with a metal detector and the horizontal and vertical displacements measured. Between 1 October and 15 May (the cooler, wetter portion of the year), pocket gophers removed an average of 63% of the experimental plugs and moved an average of 38% of the markers that we recovered. From 15 May through 1 October (the hotter, drier portion of the year), only 32% of plugs were cleared and 12% of the recovered markers were moved. On average, markers that were moved were displaced 41 cm moundward and 4.9 cm upward in elevation. The intensity of moundward translocation increased with distance from the mound center. At a distance of 0.5–1.0 mound radius beyond the edge of the mound, the moundward translocation tendency averaged 71 cm. The intensity of moundward translocation was also inversely related to maximum mound height. These observations provide strong support for the fossorial rodent hypothesis of Mima mound origin, and constitute a first step in development of a mathematical model of mound formation.     

Eukaryote evolution: A view based on cytochromec sequence data

Summary We have compared the amino acid sequences of cytochromec's from 45 species of organisms representing all five kingdoms, including one species each for the Protista and Monera. We have made a phylogeny for these data by reconstructing probable ancestral sequences which generate the present descendants through a minimum number of mutations. Several trials with different data sets produced the same minimal configuration. Assuming the occurrence of no major shifts in mutation acceptance rate, we find an early differentiation between prokaryote and eukaryote stocks. Afterward the eukaryote stem gave rise first to the protozoan flagellate branch and later to the multicellular green plant branch; after this the fungi and multicellular animal stems diverged from each other. A probable ancestral sequence was estimated for each kingdom of multicellular organisms. The basic eukaryote ancestor was probably a non-photosynthetic, heterotrophic flagellate. The photosynthetic apparatus could have been a later symbiotic acquisition in the plant ancestry. The dicotyledons had differentiated into two stocks before the emergence of a monocotyledon line as did the Ascomycetes before the emergence of the Basidiomycetes. The mollusc and chordate lines may have had a common acoelomate ancestor at the divergence of the arthropod stock. The numbers of mutations on all of the branches of the phylogenetic tree were calculated as well as the numbers of mutations and repeated mutations at each amino acid position.     

Extreme rates and heterogeneity in insect DNA evolution

Summary DNA-DNA hybridization studies of insects, more specificallyDrosophila and cave crickets, have revealed interesting patterns of genome evolution that contrast markedly with what has been seen in other taxa, especially mammals and birds. Insect genomes are composed of sections of single-copy DNA with extreme variation in rates of evolutionary change. This variation is more extreme than between introns and exons; introns fall into the relatively conserved fraction of the genome. Attempts to calculate absolute rates of change inDrosophila DNA have all led to estimates some 5–10 times faster than those found in most vertebrates; this is true even for the more conservative part of the nuclear genome. Finally we point out that morphological similarity, chromosomal similarity, and/or ability to form interspecific hybrids is often associated with quite high levels of single-copy DNA divergence in insects as compared to mammals and birds.     

Confrontation of morphological and molecular data: the Praomys group (Rodentia, Murinae) as a case of adaptive convergences and morphological stasis

Phylogenetic relationships in a group of 21 African rodent species designated as the Praomys group (Murinae) were investigated using morphological characters and sequence data from the complete mitochondrial cytochrome b gene and nuclear IRBP gene fragment (840bp). The molecular results confirm the monophyly of the Praomys group, including the species Malacomys verschureni, while the other Malacomys species appear very divergent. The basal relationships within the Praomys group are poorly resolved, suggesting a rapid radiation at about 7-9 million years ago based on genetic divergence rates calibrated from the fossil record. Discrepancies between molecular and morphological results probably reflect of numerous convergences as well as variations in the rates of morphological evolution among lineages. Reconstructions of the ancestral character states suggest a savannah origin for the Praomys group, along with some morphological traits conserved by stasis in savannah taxa. At the same time, forest taxa seem to be characterized by an accelerated morphological evolution, with acquisition of convergent adaptive characters.     

Phylogeny of the avian genus Pitohui and the evolution of toxicity in birds

Bird species in the avian genus Pitohui contain potent neurotoxic alkaloids that may be used for defense. The genus comprises multiple species that are endemic to New Guinea and were presumed to belong to the family Pachycephalidae or Colluricinclidae, within the core corvoidea, an ancient Australasian radiation of crow-like birds. In order to understand the evolution of toxicity within the genus Pitohui, we sequenced three mitochondrial and two nuclear gene segments and reconstructed a phylogeny of the genus Pitohui and its putative relatives. We show that the genus Pitohui is polyphyletic, and consists of five different lineages. Using Bayesian ancestral state reconstruction, we estimate that toxicity likely evolved multiple times within this group. Furthermore, because the morphological and behavioral similarity among these poisonous birds appears to have evolved convergently, we hypothesize that this may be a possible example of Müllerian mimicry in birds. The Morningbird of Palau, Micronesia, that has often been included in the genus Pitohui, actually belongs in the genus Pachycephala and offers an intriguing case of pronounced evolution on a remote oceanic island.     

Enamel hypoplasia in sympatric chimpanzee and gorilla

The prevalence of enamel hypoplasia in 229Pan andGorilla, from the Republic of Cameroon, is described. A substantially higher proportion of gorillas (76%) than chimpanzees (58%) was affected. The incidence of enamel hypoplasia was not a function of sex, body size, or pathology. A study of tooth formation, from radiographs of a further series of immature apes, indicated that the mandibular canine bore 99% of all information about hypoplasia events. In both species a marked regularity of hypoplastic grooving with an interval of about 11.4% of canine crown height was observed. This appears to reflect a semi-annual cycle of stress which is tentatively linked to the twice-yearly rainy season. Uniform spacing of hypoplastic grooves has been observed in a variety of fossil hominids. Readily observable hypoplastic time markers in the teeth have potential for disclosing growth rates in early Hominidae. This is considered important because of the profound significance which prolonged maturation and longevity characteristic of recent human beings have for the transmission of learned behavior and social bonding.     

The structure of cytochromec and the rates of molecular evolution

Summary The x-ray structure analysis of ferricytochromec shows the reasons for the evolutionary conservatism of hydrophobic and aromatic side chains, lysines, and glycines, which had been observed from comparisons of amino acid sequences from over 30 species. It also shows that the negative character of one portion of the molecular surface is conserved, even though individual acidic side chains are not, and that positive charges are localized around two hydrophobic channels leading from the interior to the surface.The reason for the unusual evolutionary conservation of surface features in cytochromesc is probably the interaction of the molecule with two other large macromolecular complexes, its reductase and oxidase. This conservation of surface structure also explains the relatively slow rate of change of cytochromec sequences in comparison with the globins and enzymes of similar size.The rate of evolution of a protein is the rate of occurrence of mutations in the genome modified by the probability that a random change in amino acid sequence will be tolerable in a functioning protein. The observed rates of change in fibrinopeptides, the globins, cytochromec, and several enzymes are interpreted in terms of the proteins' biological roles.Contribution No. 4114 from the Norman W. Church Laboratory of Chemical Biology, California Institute of Technology, Pasadena, California 91109.     

Isolation and growth rates of methanol utilizing rhodospirillaceae

38 pure culture strains belonging to seven species of the Rhodospirillaceae were isolated from 39 methanol enrichment cultures inoculated with water and mud samples of different habitats. None of the strains exhibited doubling times shorter than 10 h in methanol-bicarbonate media.     

Marbleseeds are gromwells – Systematics and evolution of Lithospermum and allies (Boraginaceae tribe Lithospermeae) based on molecular and morphological data

Phylogenetic relationships are complex within the Lithospermeae, a large subgroup of the Boraginaceae s.str. The relationships of New World Lasiarrhenum, Macromeria, Nomosa, Onosmodium, Perittostoma, and Psilolaemus to subcosmopolitan and much larger Lithospermum have not been critically investigated in the recent past. No molecular data on the phylogeny of these genera and Lithospermum have so far been published. We investigated the relationships within Lithospermeae using three loci (nuclear ITS plus 5.8S rRNA, chloroplast trnL-F-spacer, and trnS-G-spacer) and micromorphological character traits (pollen, nutlets). Lithospermum s.l. constitutes the sistergroup of Asian Ulugbekia and is monophyletic only when its American segregates “Macromeria”, monotypic Nomosa, and Onosmodium are included. Both the African and the South American species groups of Lithospermum are monophyletic, but North American representatives are not resolved in a single clade. Morphological characters that have been considered as important for generic delimitation in the past (such as large, yellow corollas without faucal scales, particular pollen types, coarsely veined leaves, shrubby habit) have evolved in at least two only distantly related lineages within Lithospermum s.l. The reduction of American “Macromeria”, Nomosa, and Onosmodium as well as Asian Ulugbekia under Lithospermum is proposed to render the latter monophyletic. This redefined Lithospermum s.l. appears to have undergone a type of recent “island radiation” in the Americas, reflected in a morphological diversity far exceeding that found in the Old World.     

Comparative demography of sympatric populations of Lacerta vivipara and Lacerta agilis

Summary A mark-recapture study was carried out in sympatric populations of Lacerta agilis and Lacerta vivipara in the Netherlands from 1976 to 1982. In most years the age structure of both populations was pyramidal. For both species life expectation of females was higher and on average they did live longer. Hence the sex ratio for adults deviated significantly from 1.0 in favour of females. Maximum age for Lacerta vivipara was 8 years (female) and for Lacerta agilis 12 years (male). The density of both species fluctuated around 100/ha. The biomass of Lacerta agilis was twice that of Lacerta vivipara. In Lacerta vivipara the 3rd and 4th calendar year class supplied 78% of total net reproduction; in Lacerta agilis the 4th, 5th, and 6th calendar year classes supplied 68%. In both populations the population replacement rate was 2. Population turnover time was 4.83 years for Lacerta agilis and 2.81 for Lacerta vivipara. The life history strategy of the Lacerta vivipara population is compared with six other European Lacertavivipara populations.     

Relative rates of nucleotide substitution at the rbcl locus of monocotyledonous plants

Summary We subjected 35 rbcL nucleotide sequences from monocotyledonous taxa to maximum likelihood relative rate tests and estimated relative differences in rates of nucleotide substitution between groups of sequences without relying on knowledge of divergence times between taxa. Rate tests revealed that there is a hierarchy of substitution rate at the rbcL locus within the monocots. Among the taxa analyzed the grasses have the most rapid substitution rate; they are followed in rate by the Orchidales, the Liliales, the Bromeliales, and the Arecales. The overall substitution rate for the rbcL locus of grasses is over 5 times the substitution rate in the rbcL of the palms. The substitution rate at the third codon positions in the rbcL of the grasses is over 8 times the third position rate in the palms. The pattern of rate variation is consistent with the generation-time-effect hypothesis. Heterogenous rates of substitution have important implications for phylogenetic reconstruction.Offprint requests to: M.T. Clegg     

Comparative life histories and microhabitat use in three sympatric sculpins (Cottidae:Cottus) in northeastern California

Synopsis Life histories of three sculpins endemic to the Pit River system, northeastern California, are detailed.Cottus pitensis is widely distributed throughout the drainage. This species is typically found in rapidly flowing, shallow water on coarse substrate. It grows to a large size, >100 mm, and lives to 5 years. Spawning occurs annually, in early spring. Fecundity is relatively low, <320 oocytes.Cottus asperrimus is restricted to a large population in Fall River and smaller concentrations in Hat Creek and Pit River. It is a relatively small fish, <80 mm, and lives to 5 years. It occurs in deep, slow-moving, often spring-fed streams, typically on fine substrates. Spawning lasts from autumn to early spring. Males establish nests on hard substrates and guard several clutches of eggs. Fecundity ranges from 140–580 oocytes.Cottus klamathensis macrops is relatively rare in the drainage. The largest concentration is in the Hat Creek system, smaller populations exist in Fall River and Pit River. This sculpin is found on coarse substrate, often where aquatic vegetation is abundant. It is typically found in slow, relatively shallow water. It grows to >100mm and lives to 5 years. Males guard eggs during the short spawning season beginning in late winter. Fecundity in this species reaches a maximum of 650 oocytes. Life history strategies vary within the genus, yet the life histories of these sculpins are more similar to each other than they are to many of their congeners. This within-group similarity may have resulted from exposure of all three species to similar environmental pressures. This is predicted by life-history theory.     

Phylogeny of cardinalfishes (Teleostei: Gobiiformes: Apogonidae) and the evolution of visceral bioluminescence

The cardinalfishes (Apogonidae) are a diverse clade of small, mostly reef-dwelling fishes, for which a variety of morphological data have not yielded a consistent phylogeny. We use DNA sequence to hypothesize phylogenetic relationships within Apogonidae and among apogonids and other acanthomorph families, to examine patterns of evolution including the distribution of a visceral bioluminescence system. In conformance with previous studies, Apogonidae is placed in a clade with Pempheridae, Kurtidae, Leiognathidae, and Gobioidei. The apogonid genus Pseudamia is recovered outside the remainder of the family, not as sister to the superficially similar genus Gymnapogon. Species sampled from the Caribbean and Western Atlantic (Phaeoptyx, Astrapogon, and some Apogon species) form a clade, as do the larger-bodied Glossamia and Cheilodipterus. Incidence of visceral bioluminescence is found scattered throughout the phylogeny, independently for each group in which it is present. Examination of the fine structure of the visceral bioluminescence system through histology shows that light organs exhibit a range of morphologies, with some composed of complex masses of tubules (Siphamia, Pempheris, Parapriacanthus) and others lacking tubules but containing chambers formed by folds of the visceral epithelium (Acropoma, Archamia, Jaydia, and Rhabdamia). Light organs in Siphamia, Acropoma, Pempheris and Parapriacanthus are distinct from but connected to the gut; those in Archamia, Jaydia, and Rhabdamia are simply portions of the intestinal tract, and are little differentiated from the surrounding tissues. The presence or absence of symbiotic luminescent bacteria does not correlate with light organ structure; the tubular light organs of Siphamia and chambered tubes of Acropoma house bacteria, those in Pempheridae and the other Apogonidae do not.     

Generic relationships and dating of lineages in Winteraceae based on nuclear (ITS) and plastid (rpS16 and psbA-trnH) sequence data

Phylogenetic analyses of representative species from the five genera of Winteraceae (Drimys, Pseudowintera, Takhtajania, Tasmannia, and Zygogynum s.l.) were performed using ITS nuclear sequences and a combined data-set of ITS + psbA-trnH + rpS16 sequences (sampling of 30 and 15 species, respectively). Indel informativity using simple gap coding or gaps as a fifth character was examined in both data-sets. Parsimony and Bayesian analyses support the monophyly of Drimys, Tasmannia, and Zygogynum s.l., but do not support the monophyly of Belliolum, Zygogynum s.s., and Bubbia. Within Drimys, the combined data-set recovers two subclades. Divergence time estimates suggest that the splitting between Drimys and its sister clade (Pseudowintera + Zygogynum s.l.) occurred around the end of the Cretaceous; in contrast, the divergence between the two subclades within Drimys is more recent (15.5–18.5 MY) and coincides in time with the Andean uplift. Estimates suggest that the earliest divergences within Winteraceae could have predated the first events of Gondwana fragmentation.     

Ovarian development rates in the Old World screw-worm fly, Chrysomya bezziana

Rates of ovarian development in relation to temperature were determined for autogenous females of the screw-worm fly, Chrysomya bezziana. Percentage durations of the different ovarian stages (scaled 2–10) were estimated on the basis of observed lengths of the developing oocytes. Mean durations (h) of each ovarian stage was determined at 20, 25, 28 and 35°C. A model of ovarian development rate (%/d) in relation to temperature (T) is presented, the fitted curve being give by R(T)=EXP (–2.73+0.362T–0.0055T²).