Carnivores, biases and Bergmann's rule

Studies of Bergmann's rule may encompass a non-random subsample of extant homeotherms. We examined patterns of correlation between skull length and geographical latitude in 44 species of carnivores in order to test the validity of Bergmann's rule in the Carnivora. Results were then compared to those of other studies. Significant positive correlation between skull length and latitude was found in 50% of carnivore species, while significant negative correlation was found in only 11% of species. These results indicate that the occurrence of Bergmann's rule in the Carnivora is less frequent than earlier published data suggest. Publication bias is not detected in published data. Therefore, previous studies of geographical size variation might be biased in favour of species known to follow Bergmann's rule.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 579–588.     

Fluctuating asymmetry in bank vole populations (Rodentia, Arvicolinae) reflects stress caused by landscape fragmentation in the Mont-Saint-Michel Bay

In intensively farmed, reclaimed areas (polders) of Mont-St-Michel Bay, France, bank voles ( Clethrionomys glareolus ) live in fragmented hedgerows, where populations are small and dispersal rates and genetic diversity are low. These small populations are likely to have been exposed to potential environmental and/or genetic stress. The sensitivity of development to stress can be measured by fluctuating asymmetry (FA). FA was calculated for three samples from a disturbed area and one sample from an adjacent, more connected and undisturbed landscape. Size FA was estimated from 16 measurements of the skull and teeth whilst shape asymmetry was estimated from the skull alone. Bank voles in fragmented hedgerows of the disturbed area had a higher degree of FA than bank voles from the more extensive and more connected hedges of the undisturbed area. These results were confirmed by the study of shape asymmetry, body mass and centroid size of the skull. There were no differences in FA between the three disturbed area samples. We conclude that FA does not reveal differences in the development of bank voles living in isolation under different local conditions in the various parts of the disturbed area. However, FA may allow differentiation between populations from greatly contrasting landscapes.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80, 37–44.     

Mitochondrial cytochrome b sequence yields new insight into the speciation of social voles in south-west Asia

We established a cytochrome b (cyt b ) phylogeny for six species of social voles. A Bayesian approach to phylogenetic reconstruction (BI) and a maximum likelihood (ML) tree revealed a dichotomy into two major clusters, namely a Microtus guentheri cluster and a M. socialis cluster. The three main lineages that emerged within each of these two clusters were separated by the K2P divergences which are above the intraspecific variation in Microtus . All six species were also retrieved in the minimum spanning network. Within its present taxonomic scope, M. guentheri is paraphyletic and consists of two allopatric sibling species: M. guentheri (Syria, Israel) and M. hartingi (Anatolia and the Balkans). The closest relative to these two species is M. dogramacii , which is possibly a sister species to M. hartingi . The two geographic samples were identified as M. irani , one from Shiraz (Iran) and the other from Balkusan (Turkey). The cyt b sequence confirmed the specific status of M. anatolicus within the M. socialis cluster. Although five species of social voles occur within a radius of < 500 km in the north-eastern corner of the Mediterranean, small-scale sympatry is exceptional. Species richness in this region possibly originates from past fragmentation with subsequent allopatric speciation in refugial areas.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 121–128.     

The genus Maesa (Maesaceae) in the Philippines

A comprehensive taxonomic revision of the genus Maesa Forssk. (Maesaceae) occurring in the Philippines is presented for the first time. Thirteen species are recognized, including a newly described species, M. stonei , which is most similar to M. reflexa . Several previously used names are reduced to synonymy, including M. pachyphylla and M. undulata (both reduced to synonymy with M. cumingii ), and M. elmeri , M. laxa and M. ramosii (all reduced to synonymy with M. indica ). Extensive lectotypification has been necessary due to the destruction of many type specimens, particularly those of C. Mez (in B) and E. D. Merrill (in PNH). The conservation status of each species is determined using the IUCN red list categories and criteria.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 17–43.     

Body size of the red fox Vulpes vulpes in Spain: the effect of agriculture

The body size of animals is affected by several factors, including ambient temperature and food availability. Ambient temperature is often negatively related to body size (Bergmann's rule) whereas an improved diet, especially during growth, has a positive effect. Animals commensal with man commonly exploit additional food sources (e.g. garbage dumps), thereby increasing their food supply. Using museum material, we studied morphological variation in skull size (and thus body size) among Spanish red foxes. Four measurements were taken of each skull and were related to the habitat from which the foxes were collected (agricultural and non-agricultural), and to latitude as a proxy for ambient temperature. The skull size of foxes collected in agricultural areas during the late 20th Century was significantly larger than that of those from non-agricultural areas, and was negatively related to latitude, thus contradicting Bergmann's rule. We suggest that increased food availability from animal husbandry is the cause for the observed increase in skull size (and thus body size).  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 729–734.     

Variability in diapause propensity within populations of a temperate insect species: interactions between insecticide resistance genes and photoperiodism

Temperate insects generally use day length as a reliable cue for long-term seasonal changes in their environment. Significant variation in photoperiodism between and within populations is thought to be associated with genetic variation resulting from local adaptation. In this study, we investigated whether genetic variation associated with selection for insecticide resistance may be a source of divergence in the photoperiodic timing of diapause through pleiotropic interactions. Critical photoperiods for diapause induction were estimated in one susceptible and two insecticide-resistant homozygous strains of the codling moth Cydia pomonella , as well as in their reciprocal F₁ progeny. Diapause responses to naturally decreasing day length were subsequently followed in the laboratory strains and in two field populations of C. pomonella in south-eastern France. We found higher critical photoperiods for diapause induction in homozygous resistant individuals than in both homozygous susceptible and heterozygous ones. This partly explained the significantly earlier timings of diapause found in homozygous resistant individuals of both laboratory and field populations of C. pomonella under natural photoperiods, relative to those found in both homozygous susceptible and heterozygous ones. We assume that adaptive genetic changes associated with selection for insecticide resistance may generate substantial variation in the seasonal timing of diapause, an essential ecological feature of the fitness of insecticide resistance genes in this species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 341–351.     

Size, shape and structural versatility of the skull of the subterranean rodent Ctenomys (Rodentia, Caviomorpha): functional and morphological analysis

Morphological analysis of the skull of the subterranean rodent Ctenomys , a highly speciose genus which uses both claws and teeth when digging, shows that for a broad range of species size, scaling was associated with both variation and maintenance of shape. Our results show that the angle of incisor procumbency (AIP), a character largely viewed as an adaptation to digging with teeth, is highly variable. We found a non-significant relationship between AIP and basicranial axis (basioccipital + basisphenoid) length, a measure of overall skull size. Accordingly, both small and large Ctenomys species possess either high or low AIP. A significant relationship between AIP and diastema length, given the rostral allometry seen in Ctenomys , suggests that hypermorphosis to a certain extent influences AIP. However, the roots of the incisor are lateral to those of the cheek teeth and their position may thus shift freely. This observation supports the notion that skull structural design, and to a certain extent rostral allometry, underlies variation in AIP. On the other hand, the positive allometry of incisor width and thickness indicates that, in larger species, proportionately powerful incisors are able to resist greater bending forces. We found that the out-lever arm of the jaw adductor muscles scales with positive allometry against basicranial axis length. However, we found an isometric relationship between in- and out-lever arms. In this case, conservation of skull proportions, regardless of variation in size, is a feature possibly related to the maintenance of an effective tooth digging capability. Functional and ecological data are discussed when assessing the implications of size and shape variation in the skull of Ctenomys .  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 78 , 85−96.     

Clarification of the name Nothofagus alpina and a new epithet for a Nothofagus hybrid

Arguments in favour of using Nothofagus alpina (Poepp. & Endl.) Oerst. (Fagaceae) rather than N. procera Oerst. or N. nervosa (Phil.) Krasser as the correct name for rauli are presented. We also refute suggestions that N. alpina is based on hybrid material. The hybrid of N. alpina with N. obliqua (Mirb.) Blume is formally described as N . ×  dodecaphleps Mike L. Grant & E. J. Clement and a key to the deciduous taxa of Nothofagus is provided.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 447–451.     

Mitochondrial DNA in the hybrid zone between Mus musculus musculus and Mus musculus domesticus: a comparison of two transects

We studied mtDNA introgression across the contact zone between Mus musculus musculus and M. m. domesticus in two independent transects in the Czech Republic and Bavaria, Germany. A total of 1270 mice from 98 localities in the Czech transect and 456 mice from 41 localities in the Bavarian transect were examined for presence or absence of a Bam HI restriction site in the mt-Nd1 gene. Using this simple mtDNA marker, variants that belonged to the M. m. domesticus lineage (presence of restriction site) could be unequivocally distinguished from those belonging to the M. m. musculus lineage (absence of restriction site). The extent of introgression of mtDNA, three autosomal allozymes and the X chromosome was compared. The introgression of X markers was more limited than was that of the allozymes and mtDNA. In the Czech transect, the centre for the mtDNA cline was shifted about 3.6 km to the west relative to the X chromosome cline, with asymmetric introgression from M. m. musculus to M. m. domesticus . Interestingly, in the Bavarian transect, the centre of the mtDNA cline was shifted about 10.9 km to the east relative to the X chromosome cline, with asymmetric introgression from M. m. domesticus to M. m. musculus, opposite in direction to that observed in the Czech transect.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 363–378.     

The relationship between skull morphology, biting performance and foraging mode in Kalahari lacertid lizards

Lizards are a diverse clade in which one radiation consists entirely of sit-and-wait foragers and another consists of wide foragers. Lizards utilizing these two foraging modes are known to differ in diet, but little is known about how feeding morphology relates to diet and/or foraging mode. This study tested the hypothesis that skull morphology and biting performance are related to diet preference, and consequently, coevolve with foraging mode. Four species of lacertid lizard were studied because they vary in foraging mode, their phylogenetic relationships are known and they are well studied ecologically. Using an 'ecomorphological' approach, skull morphology and biting performance were quantified and mapped on to the phylogeny for the species. The results indicate that sit-and-wait species have shorter, wider skulls than the wide foraging species, and that all are significantly different in overall head shape. The sit-and-wait species had similar values for biting performance; however, clear phylogenetic patterns of covariation were not present between sit-and-wait and wide foraging species for either biting performance or skull morphology. Thus, skull morphology and performance have little influence on diet and foraging mode in these species. Instead it is likely that other factors such as seasonal prey availability and/or life history strategy shape foraging mode decisions.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 403–416.     

Systematic revision of Myristicaceae (Magnoliales) in Madagascar, with four new species of Mauloutchia

The systematics of Malagasy Myristicaceae is revised to take into account new collections made since the work by Capuron in the 1970s, as well as improved knowledge from fieldwork and two scanning electron microscopy studies by the author. Four new species are described: Mauloutchia annickiae Sauquet, M. capuronii Sauquet, M. echinocarpa Capuron ex Sauquet, and M. sambiranensis (Capuron) Sauquet. In addition, basic information is given for each remaining Malagasy species of the family (synonymy, type specimen, updated distribution and main distinctive features). According to this treatment, Malagasy Myristicaceae now consist of four endemic genera and 15 species: Brochoneura Warb. (three species), Doyleanthus Sauquet (one species), Haematodendron Capuron (one species) and Mauloutchia (Baill.) Warb. (ten species). Two identification keys to these species are provided: one based primarily on fruit characters and one based primarily on male flower and inflorescence characters. Putative phylogenetic relationships among these species are also indicated, based on a previous combined morphological and molecular study by the author.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 351–368.     

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     

Primary growth and morphological markers of interannual growth limits in Cupressaceae from Patagonia

The present study shows experimentally that primary growth of the native Patagonian Cupressaceae Austrocedrus chilensis (D. Don) Pic.-Serm et Bizz., Fitzroya cupressoides (Molina) I. M. Johnst. and Pilgerodendron uviferum (D. Don) Florin is potentially continuous. External morphological markers of interannual growth limits have been identified for the main axes of these species after several years of observation. Such limits correspond with axis portions in which internodes are relatively short and leaves have a small distal free end forming an acute angle with the axis. The causality of these results and their utility for studying the productivity, adaptation to different environments, population management and conservation of these threatened species are discussed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 146 , 285–293.     

Recent increase in body size of the American marten Martes americana in Alaska

Using museum specimens, we studied recent changes in skull size of the American marten Martes americana , in continental Alaska. In Alaska, global warming has resulted in milder winters that may contribute to an improved food supply in the wild. In the present study, we tested the hypothesis that body size of the marten had increased during the second half of the 20th century, in response to global warming. We found that skull size, and by implication body size, increased significantly during the second half of the 20th century, possibly due to an improved food supply and/or lower metabolic demands in winter. Improved food availability in winter may result from the improved nutritional conditions for prey, and/or from increased access to prey resulting from a longer snow-free season. Longitude had a significant positive effect on skull size and a significant negative effect on teeth size. In Alaska, the climate is milder along the western coast and becomes harsher inland. Hence, the milder climate was associated with larger body size providing further support for our prediction that body size of the American marten was influenced by food availability and reduced energy expenditure. The negative relationship between longitude and teeth size may indicate a trend towards a larger prey in inland marten populations, but we have no data to support or refute this hypothesis.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 701–707.     

Variation in the digging apparatus of the subterranean silvery mole-rat, Heliophobius argenteocinereus (Rodentia, Bathyergidae): the role of ecology and geography

The skull of most subterranean tooth-digging rodents is markedly affected by their digging mode. In the present study, we investigated the cranial variation in a strictly subterranean, highly specialized Afrotropical tooth-digger, Heliophobius argenteocinereus (Bathyergidae, Rodentia), using a geometric morphometric approach and evaluated the effect of different factors on size and shape differences among four populations. No evidence for sexual dimorphism was found in skull size or shape. The cranial shape variation was large and influenced mainly by the type of habitat (miombo woodland versus farmland and grassland) and the latitudinal gradient. The dorsal side of the skull appears to be more plastic and adaptable to local environments, as well as more independent of size, than the ventral side. Only the shortening of the rostrum is presumably an adaptive process independent of size that leads to an increase of efficacy of the tooth-digging apparatus in Heliophobius , whereas the increase in the in-force and the more procumbent incisors both comprise size-related changes caused by ontogenetic allometric growth.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 822–831.