Structure and composition of the courtship phenotype in the bird of paradise Parotia lawesii (Aves: Paradisaeidae)

Ethology is rooted in the idea that behavior is composed of discrete units and sub-units that can be compared among taxa in a phylogenetic framework. This means that behavior, like morphology and genes, is inherently modular. Yet, the concept of modularity is not well integrated into how we envision the behavioral components of phenotype. Understanding ethological modularity, and its implications for animal phenotype organization and evolution, requires that we construct interpretive schemes that permit us to examine it. In this study, I describe the structure and composition of a complex part of the behavioral phenotype of Parotia lawesii Ramsay, 1885--a bird of paradise (Aves: Paradisaeidae) from the forests of eastern New Guinea. I use archived voucher video clips, photographic ethograms, and phenotype ontology diagrams to describe the modular units comprising courtship at various levels of integration. Results show P. lawesii to have 15 courtship and mating behaviors (11 males, 4 females) hierarchically arranged within a complex seven-level structure. At the finest level examined, male displays are comprised of 49 modular sub-units (elements) differentially employed to form more complex modular units (phases and versions) at higher-levels of integration. With its emphasis on hierarchical modularity, this study provides an important conceptual framework for understanding courtship-related phenotypic complexity and provides a solid basis for comparative study of the genus Parotia.     

Uncovering tropical diversity: six sympatric cryptic species of Blepharoneura (Diptera: Tephritidae) in flowers of Gurania spinulosa (Cucurbitaceae) in eastern Ecuador

Diversification of phytophagous insects is often associated with changes in the use of host taxa and host parts. We focus on a group of newly discovered Neotropical tephritids in the genus Blepharoneura , and report the discovery of an extraordinary number of sympatric, morphologically cryptic species, all feeding as larvae on calyces of flowers of a single functionally dioecious and highly sexually dimorphic host species ( Gurania spinulosa ) in eastern Ecuador. Molecular analyses of the mitochondrial cytochrome oxidase-I gene from flies reared from flowers of G. spinulosa reveal six distinct haplotype groups that differ by 7.2–10.1% bp (uncorrected pairwise distances; N  = 624 bp). Haplotype groups correspond to six distinct and well-supported clades. Members of five clades specialize on the calyces of flowers of a particular sex: three clades comprise male flower specialists; two clades comprise female flower specialists; the sixth clade comprises generalists reared from male and female flowers. The six clades occupy significantly different morphological spaces defined by wing pigmentation patterns; however, diagnostic morphological characters were not discovered. Behavioural observations suggest specific courtship behaviours may play a role in maintaining reproductive isolation among sympatric species. Journal compilation  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 779–797. No claim to original US government works.     

Hybridization between two polyphagous fruit-fly species (Diptera: Tephritidae) causes sex-biased reduction in developmental stability

When hybridization modifies the genetic constitution of individuals or populations, the stability of phenotypic development may either decrease or increase, depending on the divergence in the gene systems that control the development between the hybridizing taxa, i.e. on the relative effects of outbreeding and heterosis. In genetically closely related species, strong heterotic effects are less likely to occur, and hence hybridization may be expected to cause an overall decrease in developmental stability (DS) resulting from the disruption of coadapted gene complexes. To test this hypothesis, we experimentally crossed two closely related species of Ceratitis fruit flies and compared multiple-trait fluctuating asymmetry (FA, a measure of DS) in male and female offspring between parental species and two crossbred types. All traits measured play an important role in the fanning and buzzing behaviour associated with male courtship in Ceratitis , or are located on body parts involved in this behaviour. As predicted, hybrid offspring developed more asymmetrically than offspring of either parental species – most notably in meristic traits – and the increase in FA was consistently and significantly stronger in females than in males. The fact that males buffered their development more efficiently than females is in concordance with the presumed between-sex variation in functionality, and hence cost of asymmetry, of the measured traits. Absence of a similar sex difference in DS among parental offspring is believed to result from overall weak association between DS and FA in the absence of genetic stress, resulting from the random nature of the underlying processes that trigger asymmetric development.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 579–588.     

Kleptoparasitic interactions throughout the animal kingdom and a re-evaluation, based on participant mobility, of the conditions promoting the evolution of kleptoparasitism

Understanding of the conditions encouraging the evolution of kleptoparasitism has been limited by the preponderance of attention focused on a limited number of taxa (predominantly birds). However, most animal taxa contain kleptoparasitic members. In many ways, adult birds are aberrant because most are acrobatic, capable of three-dimensional movement, and steal from hosts that are capable of similar feats. The mobility of kleptoparasites and their hosts falls along a continuum, with the acrobatic birds at one end and sessile hosts and kleptoparasites at the other. Considering sedentary systems requires amendments to the criteria previously outlined as necessary for kleptoparasitism. Kleptoparasitic interactions between agile participants are ephemeral, involve numerous hosts over the lifetime of the kleptoparasite, and usually rely on vision. Large concentrations of hosts and large, high-quality food items are important in mobile systems. By contrast, the long-term, intimate associations of sedentary interactions demand that the host not consume the kleptoparasite's offspring and that the host must either have a longer life cycle than the kleptoparasite or be located in aggregations that have staggered life cycles (so the kleptoparasite can move to a new host when the original host dies). Expanding studies of kleptoparasitism to other taxa will differentiate between the selective pressures and evolutionary responses that are universal among kleptoparasitic symbioses and those that are restricted to certain groups.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 745–762.     

A new species of Achillea (Asteraceae: Anthemideae) from south-east Peloponnisos, Greece

A new species of Achillea L., Achillea occulta Constantin. & Kalpoutz., from the summit area of Mount Koulochera in south-east Peloponnisos, Greece, is described and illustrated. It belongs to A . sect. Ptarmica and further, to a small group of taxa known under the illegitimate sectional name 'Anthemoideae'. In Greece, A. occulta has no close allies. The taxonomically related taxa, i.e. A. barrelieri (Ten.) Sch. Bip., A. mucronulata Bertol., A. oxyloba (DC.) Sch. Bip. and A. schurii Sch. Bip. are found in the Italian Peninsula, the Alps and the Carpathians. Phytogeographically, the finding of a species with such taxonomic connections in south-east Greece was unexpected. The new species grows in semi-shade, often hidden in suitable limestone rock hollows and the foot of rocks, together with several other Greek endemics. Its chromosome number of 2 n  = 18, counted in root tips, is also reported and illustrated.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 249–256.     

Genetic parsimony: a factor in the evolution of complexity, order and emergence

Two conjectures, drawn from Gregory Chaitin's Algorithmic Information Theory, are examined with respect to the relationship between an algorithm and its product; in particular his finding that, where an algorithm is minimal, its length provides a measure of the complexity of the product. Algorithmic complexity is considered from the perspective of the relationship between genotype and phenotype, which Chaitin suggests is analogous to other algorithm-product systems. The first conjecture is that the genome is a minimal set of algorithms for the phenotype. Evidence is presented for a factor, here termed 'genetic parsimony', which is thought to have helped minimize the growth of genome size during evolution. Species that depend on rapid replication, such as prokaryotes which are generally r -selected are more likely to have small genomes, while the K -strategists accumulate introns and have large genomes. The second conjecture is that genome size could provide a measure of organism complexity. A surrogate index for coding DNA is in agreement with an established phenotypic index (number of cell types), in exhibiting an evolutionary trend of increasing organism complexity over time. Evidence for genetic parsimony indicates that simplicity in coding has been selected, and is responsible for phenotypic order. It is proposed that order evolved because order in the phenotype can be encoded more economically than disorder. Thus order arises due to selection for genetic parsimony, as does the evolution of other 'emergent' properties.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 295–308.     

Canarian land snail diversity: conflict between anatomical and molecular data on the phylogenetic placement of five new species of Napaeus (Gastropoda, Pulmonata, Enidae)

Five new species of land snail (family Enidae) are described from La Gomera (Canary Islands) of which the majority, on the basis of anatomy alone, could be incorporated within a new supraspecific taxon. In addition to the morphological study of these new species, a region of the 16S mitochondrial gene is sequenced from three of the new species and a range of species of Napaeus from within its two subgenera ( Napaeinus and Napaeus ) . There is a disparity between the morphological and preliminary molecular phylogenetic data. Possible explanations for this conflict are discussed, as well as the evolutionary relationships among these different taxa, and it is suggested that this group may be an excellent model for further studies of adaptation and diversification.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 169–187.     

Cytogenetics, geographical distribution, and pollen fertility of diploid and tetraploid cytotypes of Santolina pectinata Lag. (Asteraceae: Anthemideae)

A cytogenetic study of 62 populations of Santolina pectinata in Spain shows the existence of two ploidy levels. The diploid cytotypes with 2 n  = 18 occupy the eastern Betic mountains, and the tetraploid cytotypes with 2 n  = 36 are located on the spurs of the Iberian System. The former show a much wider ecological spectrum than the latter. Mixed cytotypes were observed in two diploid populations, with one tetraploid in each, showing different karyotypes. Three trisomic individuals were detected, one in a diploid population and the other two in a tetraploid population. Also, three hypotetraploid individuals were detected in a tetraploid population. Polyploidy is shown to be spontaneous and recurrent, promoting partial sterility in the pollen. Structural chromosomal changes, principally translocations, and local speciation through autopolyploidy are the principal factors in the evolution and diversification of this species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 657–667.     

Cytological studies on some representative species of the tribe Orchideae (Orchidaceae) from China

Cytological studies were carried out on 14 taxa belonging to Amitostigma , Chusua , Galearis , Habenaria , Hemipilia , Hemipiliopsis , Herminium , Peristylus and Ponerochis , collected mostly from the south-eastern part of the Hengduan Mountain Region, south-west China. Cytological data on 11 of the taxa are reported for the first time. The interphase nuclei were either of the simple chromocentre type or intermediate between simple and complex chromocentre types. The nuclear morphology of Hemipiliopsis at interphase supports the conclusion that it is related more closely to Chusua and Ponerochis than to Habenaria . At the whole tribe level, however, the results did not indicate a clear correlation between morphological features of the interphase nuclei and phylogeny. The somatic chromosome numbers were 2 n  = 42 in ten species and 2 n  = 32, 38, 40, 64 and 72 in four species. The chromosome counts of 2 n  = 32 and 64 in Habenaria aitchsonii are rare in the genus. It is proposed that the repeated change of chromosome number from x  = 7 to x  = 8 has played an important role in the evolution of the tribe Orchideae. This change has occurred mainly in the European subtribe Orchidinae, but also in the Asian subtribe Habenariinae.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 231–238.     

Karyological studies in Tripleurospermum (Asteraceae, Anthemideae) from north-east Anatolia

Chromosome number and morphology in 14 taxa belonging to 19 populations of Tripleurospermum Sch. Bip. were studied using karyological and numerical taxonomical techniques. Data on chromosome measurements were analysed using cluster analysis. Chromosome numbers of these taxa are 2 n  = 2 x  = 18, 4 x  = 36 and 5 x  = 42–48. Seven records are new, two are not consistent with previous counts, and the remainder confirm the very limited previous data (one to three records). A new ploidy level (pentaploidy) is reported for the first time for the genus. Some correlations between ploidy levels and morphological characters are noted and several systematic and evolutionary aspects of the genus are discussed in the light of karyological data.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 146 , 427–438.     

Gagea tisoniana, a new species of Gagea Salisb. sect. Gagea (Liliaceae) from central Italy

Gagea tisoniana Peruzzi et al. sp. nov. is described and its taxonomic relationship is discussed. The new species, which appears to be endemic to central Italy, is close to G. pratensis (Pers.) Dumort., G. pusilla (F. W. Schmidt) Sweet, and other mainly poorly known [i.e. G. succedanea Griseb. et Schenk, G. transversalis Stev., and G. paczoskii (Zapal.) Grossh.] or as yet undescribed taxa from eastern Europe and the eastern Mediterranean area. The recognition of the new taxon G. tisoniana is claimed on the basis of anatomical, karyological, ecological, and phytogeographical data. Maps of the Italian distribution of G. pratensis (2 n  = 60), G. pusilla (2 n  = 24), and G. tisoniana (2 n  = 24) are presented, together with an analytical identification key. Finally, G. pratensis is recorded here for the first time in Tuscany (Monte Cetona).  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 337–347.     

Faecal case architecture in the gibbosus species group of Neochlamisus Karren, 1972 (Coleoptera: Chrysomelidae: Cryptocephalinae: Chlamisini)

Constructions composed of faeces are rare in insects, but occur in certain leaf-beetle clades. Members of the subfamily Cryptocephalinae share a complex behavioural and morphological synapomorphy, involving portable faecal cases that house the immature stages. A maternally initiated egg case is expanded and enlarged through four larval stadia, then sealed to provide a pupal chamber from which adults eventually emerge. We analyse and compare faecal-case architecture in ten taxa of the cryptocephaline genus Neochlamisus , and assess structural variation within a life cycle, between different 'host forms' of Neochlamisus bebbianae , and among species. These cases proved to be comprised primarily of faeces, with plant trichomes representing the only common secondary component. General architectural trends reflected variation in shape, faecal texture, and the incorporation and density of trichomes. Deviations of the Neochlamisus case from a simple geometrical gnomon reflect the changing body size of the animal, differential application of faeces, and shifts in the orientation of the carriage of the case. Neochlamisus cases are presumably energetically costly to produce, carry, and maintain, and some adaptive hypotheses of case evolution are proposed. Additionally, literature on case morphology in other camptosomates is reviewed.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 315–351.     

Nomenclatural changes in preparation for a World Rubiaceae Checklist

During the preparation of a World Rubiaceae Checklist , numerous unplaced taxa were encountered, including illegitimate and invalid names, and species for which generic placement is uncertain. In this contribution, 35 new combinations and 20 new names are proposed, and the names of three taxa are validated.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 115–124.     

Patterns of genetic diversity in related taxa of Antirrhinum L. assessed using allozymes

Allozyme diversity was studied within and among populations of five related taxa of Antirrhinum L. endemic to the Iberian Peninsula ( A. graniticum Rothm. ssp. graniticum , ssp. brachycalyx Sutton and ssp. ambiguum (Lange) Mateu & Segarra, A. boissieri Rothm. and A. onubensis (Fdez. Casas) Fdez. Casas). All of the studied taxa are obligate outcrossing endemic perennial herbs which form isolated populations. However, the taxa vary in range and population sizes, and are found on different soil types. The level and distribution of allozyme diversity differed widely between taxa: A. graniticum ssp. brachycalyx had the lowest level of allozyme diversity (H_T = 0.09), whilst the highest level was detected in A. boissieri (H_T = 0.25). Total variation was partitioned into within- and among-population variation. The proportion attributable to variation within populations varied from about 67% up to 84.3% and 89.5% in A. graniticum ssp. brachycalyx and A. graniticum ssp. ambiguum , respectively. Both these subspecies also showed little population divergence (G_ST = 0.10 and 0.09, respectively) and had high levels of estimated gene flow (Nm = 2.18 and 2.62, respectively). These results are discussed in relation to geographical proximity of populations and habitat continuity. Isolation by distance was not detected in any of the studied taxa. This result suggests that divergence among populations is due to random genetic drift.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 79 , 299–307.     

Heterochrony in Tingidae (Insecta: Heteroptera): paedomorphosis and/or peramorphosis?

Tingidae (Heteroptera: Insecta) exhibit cephalic tubercles that present a very diverse shape in larvae but that are much simpler in adults. A phylogeny based on adult and last instar characters showed that these tubercles evolved independently from simple to complex states in two clades, and reversed from complex to simple in some taxa. These homoplasies are analysed in the light of ontogenetic sequences and interpreted as heterochronic events. The general trend of evolution of the cephalic tubercles in Tingidae is in mosaic, and could be generally peramorphic, with some isolated cases of paedomorphosis.  Journal compilation © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 71–80. No claim to original French government works.     

Mitochondrial DNA sequences of the Afro-Arabian spiny-tailed lizards (genus Uromastyx; family Agamidae): phylogenetic analyses and evolution of gene arrangements

Approximately 1.7 kbp of mitochondrial DNA were sequenced from 29 individuals assignable to 11 Uromastyx species or subspecies and two other agamids. U. ocellata and U. ornata had an insertion between the glutamine and isoleucine tRNA genes, which could be folded into a stable stem-and-loop structure, and the insertion for U. ornata additionally retained a sequence similar to the glutamine tRNA gene. This corroborates the role of tandem duplication in reshaping mitochondrial gene arrangements, and supports the idea that the origin of light-strand replication could be relocated within mitochondrial genomes. Molecular phylogeny from different tree-building methods consistently placed African and Arabian taxa in mutually monophyletic groups, excluding U. hardwickii inhabiting India and Pakistan. Unlike previous studies based on morphology , U. macfadyeni did not cluster with morphologically similar Arabian taxa, suggesting convergent evolution to be responsible for the morphological similarities. Divergence times estimated among the Uromastyx taxa, together with geological and palaeontological evidence, suggest that the Uromastyx agamids originated from Central Asia during the Eocene and colonized Africa after its connection with Eurasia in the early Miocene. Their radiation may have been facilitated by repeated aridification of North Africa since the middle Miocene, and geological events such as the expansion of the Red Sea and the East African Rift Valley.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 247–260.     

Chromosomal evolution of the house mouse, Mus musculus domesticus, in the Aeolian Archipelago (Sicily, Italy)

We describe the chromosomal evolution of the metacentric populations of the house mouse, Mus musculus domesticus , which constitute the Robertsonian System of Aeolian Islands (Sicily, Italy). Eighty-nine specimens from all the seven islands that form the Archipelago were cytogenetically examined. The analysis shows the presence of 4 Rb races with a large number of shared metacentric chromosomes: 2 n  = 36 on Panarea, 2 n  = 34 on Alicudi, 2 n  = 26 on Lipari and Stromboli, and a different 2 n  = 26 race on Vulcano. On Salina and Filicudi, the standard karyotype was found. Polymorphism was only found in a population on Panarea Island and this population shares no metacentrics with the other races. The distribution of metacentrics among the races and the comparison between the Aeolian metacentrics and those found in the 97 previously documented metacentric populations allows us to formulate a hypothesis of chromosomal evolution for the Aeolian Robertsonian system. Six of the twelve metacentric chromosomes found in the Aeolian Islands come from localities outside the archipelago. The evolutionary model highlights how the chromosomal races originated inside the Archipelago and involve several factors, such as formation in situ of metacentrics, zonal raciation and, whole arm reciprocal translocation. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 194–202.