Comparative osteology and relationships of the Umbridae (Pisces: Salmoniformes)

The phylogenetic relationships of the five living species of umbrids are examined through a comparative osteological study based on a series of cleared-and-stained specimens of each species. The Umbridae appear to be strictly monophyletic. In addition, for 45 characters, the outgroup Esox shares one character-state with at least one umbrid species, while two or more umbrid species share the other state. Assuming that the states shared with Esox are primitive for the Umbridae, the hypothesis that Dallia is more closely related to Umbra than to Novumbra is supported by compatibility and Wagner-tree analysis of the data. Thirteen derived characters are shared by Dallia and Umbra , 20 more are shared by the three species of Umbra , and four more by U. limi and U. pygmaea. Eight characters are considered likely to have undergone reversal or parallel evolution.     

Evolution of the amphibian tympanic ear and the origin of frogs

Recent anurans plus all but the most primitive temnospondyl labyrinthodont amphibians are proposed as a monophyletic taxon, based on shared stapedial characters which are derived with respect to all other tetrapods. Within temnospondyls, the mostly Lower Permian dissorophoids are proposed as most closely related to Recent anurans, based on interpretation of the dissorophoid dorsal quadrate process and the anuran tympanic annulus as sequential steps in a character transformation series. The otic features described here reinforce the concept of the amphibian tympanic ear as a prior "invention" with no genealogical relationship to amniote tympanic ears.     

Mobile organisms in a geologically complex area: Lepidoptera in the Indo-Australian tropics

The Indo-Australian tropical archipelagos have increased in extent and proximity through the Tertiary whilst also changing their configuration. The development of new land areas between old cratonic ones involves extensive dispersals of plants and animals from the latter to the former. The vicariance approach, though suited to the study of the fragmentation of old, cratonic land areas, is of uncertain value in this situation; the representation of emergence or convergence and fusion of land areas in cladogram form presents immense problems and to attempt it may be pointless.
Distribution of Lepidoptera species largely reflects the present day geography of the area for both lowland and montane habitats. Significant discordances at a higher taxonomic level can be examined in the light of hypotheses of past geography such as those concerning the evolution of the Melanesian Arcs.
There are many advantages and some disadvantages in using Lepidoptera for such studies, involving their taxonomy, completeness of geographical coverage, range of dispersal ability and specificity to environmental conditions. Both cladistic and phenetic methods can provide basic data for biogeographic studies. The importance of an understanding of habitat preferences is stressed.
Patterns of probable recent development involving well-defined groups of very similar species are examined in the hope they will provide a yardstick whereby potentially older patterns can be assessed, and lead in time to methodology that will satisfy the stringent requirements of current thinking in the philosophy of science.     

The systematics of the Phelsuma madagascariensis species group of day geckos (Reptilia: Gekkonidae) in the Seychelles

Conventional taxonomy of the Seychelles populations of the Phelsuma madagascariensis species group has failed to illustrate the principal patterns of phenetic variation between island populations. The taxonomic history of the populations is discussed critically, and a revised nomenclature is suggested from the results of a recent comprehensive study of the forms using multivariate morphometrics. Three phenetic groups from the granitic islands are referred to as Phelsuma sundbergi sundbergi from the north western islands; P. s. ladiguensis from the north eastern islands; and P. s. longinsulae from the southern islands. Phelsuma abbotti populations from Aldabra and Assumption are considered subspecifically distinct, and not conspecific with the granitic island forms. The other coralline island populations are closely allied to the races from the granitic islands and probably represent recent colonization or introductions by man.     

A revision of the brassoline genus Selenophanes (Lepidoptera: Rhopalocera)

The Brassoline genus Selenophanes is revised: three new subspecies are described and three lectotypes designated. The nine known holotypes and two of the three lectotypes are figured. Along the Andes there is a fairly close agreement in the distribution of some of the species and subspecies of Selenophanes with the recently recognized endemic centres. East of the Andes there is no close correspondence between subspecies and endemic centres; each subspecies has, in general, a distribution across several centres. A similar distribution has been noted previously in the Brassoline genus Catoblepia.     

Systematics of the genus Cidaria Treitschke (Lepidoptera, Geometridae, Larentiinae)

The genus Cidaria Treitschke is revised. Eight species of the genus which occur widely in the Palaearctic and northern India, are recognized, of which one, Cidaria luteata sp. nov. , is described as new while two subspecies of C. fulvata (Forster) are elevated to species, C. nugata Felder stat. rev., and C. distinctata Staudinger stat. nov. C. ochripennis Prout is proposed as a junior synonym of C. ochreata Staudinger. C. deletaria Hampson is excluded from the genus. All species of Cidaria and their genitalia are described and illustrated. The cladistic analysis of these eight species of Cidaria is carried out using two data matrices, the first comprising morphological characters alone and the second morphological and distributional characters. The most parsimonious cladogram of Cidaria is selected, and its monophyly defined. The character analysis shows that some distribution characters contribute to resolving the in group node. The choice of multiple trees is discussed.     

Differential winter mortality and seasonal selection in the polymorphic ladybird Adalia bipunctata (L) in the Netherlands

Seasonal selection acting on the melanic polymorphism in the two-spot ladybird Adalia bipunctata was investigated in The Netherlands. An increase in melanic frequency over the spring-summer reproductive period was quantified. The selective advantage gained by melanics averaged 9%, but significant heterogeneity occurred between populations. Adult hibernation behaviour is described. The beetles when outdoors show a highly clumped distribution both between and within trees. The distribution of the morph classes between aggregations is random. Survivorship in a hibernating cohort (initial n= 1898) on a grid of 70 lime trees near Utrecht was monitored by making three counts over the winter of 1981–1982. Intense selection favouring each melanic morph occurred during December and January. The relative fitness of non-melanics was 0.55 (melanics =1). The discovery of dead beetles in late January (about 5% of total losses) and the absence of spatially density-dependent mortality were consistent with a climatic stress rather than selective predation. The period of selection was associated with very cold temperatures averaging up to 4°C below normal and an overall mortality of nearly 75%. There was no change in morph frequency, near normal temperatures and a lower mortality from February to early April. Examination of groups of nearby trees in late January strongly suggested that similar differential mortality had occurred except on some willows. This difference was probably due to the more protected hibernation sites available on these trees. Samples of hibernating cohorts at three other sites showed no evidence of differential mortality. Laboratory experiments with hibernating beetles found no difference in survivorship or rate of weight loss between starved non-melanics and melanics in temperature regimes with and without periods of adult activity. It is concluded that the intense winter selection on the study limes is probably exceptional. Examination of changes in morph frequency through the annual cycle suggests that at some sites the selection favouring melanics during reproduction is counterbalanced by selection against melanics in late summer or early autumn. The results are discussed in relation to mathematical models of cyclical selection and to other field studies including that of Timoféeff-Ressovsky (1940), who found large decreases in melanic frequency during hibernation in Berlin.     

Two cryptospore-bearing land plants from the Lower Devonian (Lochkovian) of the Welsh Borderland

Two examples of fragmentary, coalified plant fossils with cellular preservation and in situ spores are described using scanning and transmission electron microscopy (SEM and TEM), from a Lower Devonian (Lochkovian) locality in the Welsh Borderland. A sporophyte which produced dyads of ?Cymbohilates of. horridus is unique in that stomata are numerous on the supporting axis, and are comparable with those described from contemporary vascular plant remains. The in situ dyads possess a bilayered exospore wall, with an outer exosporal envelope present over the distal faces. A fusiform sporangium, with externally smooth epidermis, contains specimens of the tetrad Velatitetras sp. Each tetrad is encompassed by a laevigate, folded, exosporal envelope of uniform thickness, which contains a layer of regular voids. Spores within the tetrads are ultrastructurally bilayered, with a complex, digitate outer margin presumably representing spore wall ornamentation. Neither the tetrads nor dyads reported in this paper are lamellate in ultrastructure. The combination of stomata, branching and dyads in the same sporophyte holds significance for the understanding of cryptospore affinities.     

Growth patterns in the stem of the palm Archontophoenix cunninghamiana

Studies on the growth of Archontophoenix cunninghamiana in two separate populations show that this species establishes only a narrow apical meristem complex before rising above ground This complex continues to expand in girth as it rises, until at a height of about 12 m its girth becomes fixed. The lower regions of the trunk expand, more than equalling the expansion at the apex, producing at least a 20-fold increase in volume. Anatomical observations show that this expansion, termed sustained primary growth , is due to cell enlargement and cell division in the ground tissue, and is relatively uniform along the radius of the stem. When the apex reaches a height of 12 m, sustained primary growth ceases abruptly, fixing the taper of the trunk. Subsequent growth produces a cylindrical top on this tapered base.
These results are considered in the light of a review of earlier work on palm growth. The determinate nature of this type of arborescence is discussed, and brief reference is made to its evolutionary status.     

Molecular phylogeny of the genus Dahlia Cav. (Asteraceae, Heliantheae-Coreopsidinae) using sequences derived from the internal transcribed spacers of nuclear ribosomal DNA

Sequences of the internal transcribed spacers (ITS) of the 18S-26S nuclear ribosomal DNA were used to resolve phylogenetic relationships among 13 species of the genus Dahlia . Phylogenetic reconstruction from both parsimony analysis and pairwise distance data produced a single tree in which four clades could be distinguished, these broadly separating the species into the different chromosome numbers found in Dahlia ( x = 16, x = 17 and x = 18). Although species based on x = 16 were divided into two clades, there is the suggestion that this separation may be artificial and that these should be combined as a single clade. The phylogeny produced here has been discussed with reference to the existing taxonomic treatment of the genus. It currently appears that the genus Dohlia is monophyletic and it is evident that the different basic chromosome numbers withim the dysploid series in Dahlia have evolved only once. This proposed monophyly of the genus coupled with the successful production of hybrids withim the dysploid series (2 n = 32, 2 n = 34 and 2 n = 36) suggests that the species of Dahlia have evolved from a common ancestor.     

The biology of ant-like jumping spiders: intraspecific interactions of Myrmarachne lupata (Araneae, Salticidae)

Myrmarachne lupata is an ant-like salticid in which males have very large chelicerae. The display repertoire of this species is unusually large and complex for a salticid spider. Each individual male uses one of three different mating tactics depending on the female's maturity and location. With adult females outside nests type 1 courtship occurs which seems to be a form of visual communication and includes specialized movements and postures of the legs, palps and body. With adult females inside nests, males use type 2 courtship, which seems to be a form of non-visual communication and consists primarily of probing with the legs on the silk; males mate with receptive females inside the nests. With subadult females, males first use type 2 courtship then spin an adjacent silken chamber and cohabit. After she moults and matures, mating occurs inside the nest. Vacant nests of conspecific females, but not those of another sympatric salticid species, elicit courtship behaviour from males. During male-male interactions, embracing occurs with the large chelicerae spread apart. Females and subadults also display, and different displays occur in interactions depending on the sex/age classes of the spiders involved. Despite the unusual morphology of these spiders, their individual displays are similar to those of more typical salticids. During copulation males stand beside the female instead of over or on her as occurs with typical salticids.     

Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics

CARLQUIST, S., 1984. Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics. Qualitative and quantitative data, based mostly upon liquid-preserved specimens, are presented for Akebia, Roquila, Decaisnea, Holbodia, Lardizabala, Sinofranchetia and Stauntonia . Because Decaisnea is a shrub whereas the other genera are vines, anatomical differences attributable to the scandent habit can be considered. These include exceptionally wide vessels, a high proportion of vessels to tracheids (or other imperforate trdcheary elements) as seen in transection, simple perforation plates, multiseriate rays which are wide and tall, and pith which is partly or wholly sclerenchymatous. With respect to ecology, two features are discussed: spirals in narrower vessels may relate to adaptation to freezing in the species of colder areas, and crystalliferous sclereids seem adapted in morphology and position to deterrence of phytophagous insects or herbivores. The wood may provide mechanisms for maintaining conduction even if wider vessels are deactivated temporarily by formation of air embolisms. Wood and stem anatomy of Lardizabalaceae compare closely to those of Berberidaceae and of Clematis (Ranunculaceae), as well as to other families of Berberidales. Decaisnea is more primitive than these in having consistently sralariform perforation plates and in having scalariform pitting on lateral walls of vessels. A tentative listing of anatomical features which may correspond to generic limits is given.     

Functional design and evolution of the pharyngeal jaw apparatus in euteleostean fishes

Functional and structural patterns in the pharyngeal jaw apparatus of euteleostean fishes are described and analysed as a case study of the transformation of a complex biological design. The sequential acquisition of structural novelties in the pharyngeal apparatus is considered in relation to both current hypotheses of euteleostean phylogeny and patterns of pharyngeal jaw function. Several euteleostean clades are corroborated as being monophyletic, and morphologically conservative features of the pharyngeal jaw apparatus are recognized. Functional analysis, using cinematography and electromyography, reveals four distinct patterns of muscle activity during feeding in primitive euteleosts (Esox) and in derived euteleostean fishes(Perca, Micropterus, Ambloplites, Pomoxis). The initial strike, buccal manipulation, pharyngeal manipulation, and the pharyngeal transport of prey into the oesophagus all involve unique muscle activity patterns that must be distinguished in analyses of pharyngeal jaw function. During pharyngeal transport, the upper and lower pharyngeal jaws are simultaneously protracted and retracted by the action of dorsal and ventral musculoskeletal gill arch couplings. The levator externus four and retractor dorsalis muscles, anatomical antagonists, overlap for 70–90°of their activity period. Levatores externi one and two are the main protractors of the upper pharyngeal jaws in the acanthopterygian fishes studied. The major features of pharyngeal jaw movement in primitive euteleosts are retained in many derived clades in spite of a dramatic structural reorganization of the pharyngeal region. Homologous muscles have radically changed their relative activity periods while pharyngeal jaw kinematics have been modified relatively little. Patterns of transformation of activity may thus bear little direct relationship to the sequence of structural modification in the evolution of complex designs. Overall function of a structural system may be maintained, however, through co-ordinated modifications of the timing of muscle activity and anatomical reorientation of the musculoskeletal system. Deeper understanding of the principles underlying the origin and transformation of functional design in vertebrates awaits further information on the acquisition of both structural and functional novelties at successive hierarchical levels within monophyietic clades. This is suggested as a key goal of future research in functional and evolutionary morphology.     

How tetrapods feed in water: a functional analysis by paradigm

Mechanical theory is used to erect a paradigm predicting the manipulations used by carnivorous aquatic amphibians, reptiles, birds and mammals to catch, subdue, process and swallow their prey. These predictions are confirmed by observational evidence. Most aquatic predatory tetrapods use long, prehensile tooth-armed jaws as pincer jaws to snap shut onto the prey and catch and kill it, although some use the flexibility of long necks in spear fishing and some odontocetes may stun prey with sonar. Most do not have cutting or nipping dentitions as these cannot be used on prey which is freely floating. They use caniniform dentition to hold and kill prey, or in some cases crushing dentition to break open hard-shelled prey. They dismember prey by dynamic loading, snatching bites so quickly that the prey tears. They use shake feeding, shaking the prey apart from side to side above the water. If the prey is too large to lift above the water they use twist feeding, twisting pieces off. Small pieces are easily swallowed but larger pieces are held above the water and swallowed by tilting the head back in gravity feeding, or by jerking the head back and forth in incrtial feeding. Some animals use mobile jaws to pull prey back into the mouth in ratchet feeding. Filter feeding evades these problems by feeding on very small prey. The use of paradigms in functional analysis is discussed with special reference to this work. The paradigm method is shown to be the most suitable one. There has been repeated convergent and parallel evolution of adaptations to feed in water.     

Whorl overlap and the economical construction of the gastropod shell

A model is used to show how changes in the amount of overlap between successive whorls alters the ratio of the area of shell material to the volume enclosed. An optimum amount of overlap which minimizes this ratio is shown to exist.
Measurements on the shells of a number of British gastropods show that the actual amount of overlap is always greater than the optimum. It is concluded that selection for economy is being opposed by some other factor such as shell strength, shape or aperture size.     

The compounds in Aloë leaf exudates: a review

The drug Bitter Aloes has been used for several thousands of years because of its purgative properties and later several species of the plant have been cultivated for horticultural interest. In addition, many more local therapeutic properties have been recorded. Research has been largely directed towards the physiologically active components during the 50s and 60s. Extensions to this work in the last decade have been made in Germany, Japan and South Africa. Compounds have been characterized from the dried exudate prepared for drug purposes and from a few species. A number of anthraquinones have been recognized both in the free state and as glycosides. Other compounds with phenolic groups include chromones and a phenyl-pyranone. Chromatographic methods have been developed for recognizing some of these compounds and estimating the levels of those of pharmacological interest. Distribution of some of the compounds among a number of species is described showing that among some groups it has taxonomic significance. The biology of secretion and exudation from Aloë leaves remains largely unknown although fragmentary evidence points to interesting and potentially useful activity in some of the components.     

The Bornean Lithocarpus B1. section Synaedrys (Lindl.) Barnett (Fagaceae): its circumscription and description of a new species

The Bomean species of Lithocarpus B1. section Synaedrys edl.) Barnett (Fagaceae) are discussed in terms of their unifying characteristics, and keys to the species are provided. A novel morphological feature present in the genus, where the seed in the mature fruit is enclosed by a lignified and thickened receptacle, is illustrated. A new species, L palungensis Cannon & Manos, is described. It can be distinguished from L. pulcher (King) Markgr. by its submontane distribution, prominent reflexed cupular spines, present at all stages, and broadly oblong leaves with scattered open fasciculate trichomes on the abaxial surface. Foliar trichome types present within the section are also described. Distinctiveness of fruit wall and receptacle shape, as described by two-dimensional elliptic Fourier transformation, is demonstrated by principal components analysis.     

The biology of the Pe-ret' Toad, Otophryne robusta- (Microhylidae), with special consideration of its fossorial larva and systematic relationships

Adult and larval Otophryne robusta- were collected in Colombia, South America. Although this genus has been assigned to the family Microhylidae, a variety of both adult and larval features distinguish it from all other microhylids. The adult is a diurnally active leaf mimic with complex agonistic behaviours and vocalizations. It tends to walk rather than hop, and does not burrow in captivity. The larva is unique in having minute, dagger-like, keratinized teeth and a sinistral spiracle at the tip of a long tube. The mature larva lives shallowly buried in sand at the bottom of clear, shallow streams. The anatomy of the tadpole was examined for clues to how it burrows into and survives in sand, as well as to the correct phylogenetic association of the genus. Internally, the tadpole is most similar to microhylid tadpoles and has a wealth of oral features (e.g. unperforated internal nares, branchial food traps arranged in crescentic organs, ventral velum divided on the midline, glottis located rostral to the free edge of the ventral velum, large gill filters etc.) that characterize that family. Most, if not all, of the unique features of the Otophryne larva can be associated with its fossorial existence. The arrangement of the muscles suggest that it can actively raise and lower its snout during burrowing and possibly dorsiflex its head on its vertebral column. Although the larval teeth first suggest carnivory, many features indicate that maerophagy is not possible. Stomach contents include a variety of bacteria and micro-organisms, but no fragments of macroscopic animals. We conclude that the Otophryne tadpole is a microphagous suspension feeder and its teeth are more important for keeping sand grains out of its mouth than for grasping prey. Since the O. robusta larva does not burrow deeply, the tip of its spiracular tube probably extends upward above the sandy bottom and into the current. In this position pressure would be lower at the spiracle than at the mouth and a current could be passively drawn (by the Bernoulli effect) through the oral cavity for feeding and respiration. If this is correct then the Otophryne larva is the first fossorial vertebrate known to passively filter-feed. All features of the tadpole, except the keratinized teeth and sinistral spiracle, indicate an affinity of Otophryne-/i with South American microhylids. The tadpole and adult are, however, so unusual as to warrant subfamilial status+ADs- we propose the subfamily Otophryninac for this monotypic genus.     

Male secondary sexual characteristics, sexual selection, and molecular divergence in fanged ranid frogs of Southeast Asia

A group of Southeast Asian frogs of the family Ranidae are characterized by a suite of unusual secondary sexual characteristics. Males are larger than females, possess bony odontoid processes (fangs) and have enlarged heads. While all members of this clade are sexually dimorphic in this suite of characters, the degree of development of the secondary sexual characteristics in males varies widely across species. One of the most common species, Rana blythii Boulenger, has an extremely broad geographical range. This frog lacks an advertisement call in Borneo but calling has been reported in populations from peninsular Malaysia and Vietnam. This study examines the relationship between the expression of male secondary sexual characteristics, molecular divergence and number of speciation events in widely separated populations of R. blythii , and between R. blythii and other species of closely related fanged frogs belonging to the grunniens species group. Results from morphometric and sequence data indicate that (1) there is no correlation between morphological and molecular distance in these frogs and (2) there is no correlation between morphological distance and the number of speciation events separating taxa. Systematic analysis demonstrates that R. blythii , as currently recognized, is not a single species nor is it monophyletic.