Showing papers in "Biological Journal of The Linnean Society in 1992"

Plasticity of fish body shape. The effects of diet, development, family and age in two species of Geophagus (Pisces : Cichlidae)

Abstract: Fish body shape is affected by the genetic makeup of an individual as well as environmental influences, such as diet, development, growth rate and nutrition. Fishes in the family Cichlidae exhibit tremendous morphological diversity in body shape and morphology related to feeding. Certain aspects of cichlid feeding morphology have been shown to be plastic in response to different diets but plasticity in body shape has not been examined previously. Plasticity affects ecological interactions, the direction and rate of evolution, and has ramifications for characters used in systematic studies. I examined the effect of different diets: chironomid larvae (bloodworms) and brine shrimp nauplii, on body shape in two species of the Neotropical cichlid genus Geophagus which differ in the size at which young begin feeding on external food sources. The fry of G. brasiliensis, a substrate spawner, begin to feed on external food sources earlier than the fry of G. steindachneri, a mouthbrooder. I hypothesized that the difference in size at first feeding could lead to a difference in the amount of plasticity inducible in the two species. The magnitudes of changes were mostly similar, although G. brasiliensis responded to the different diets with slightly greater changes in some of the head measurements. The pattern of changes in the two species were also similar, with fish fed ferine shrimp nauplii developing longer and shallower heads and shallower bodies and tails than fish fed chironomid larvae. I also examined the consequences of considering family and age as additional factors besides diet in G. steindachneri. Considering family or age as additional factors in the analyses did not change the conclusion that different diets induce differences, albeit small ones, in body shape. I argue that morphological plasticity is dependent on behavioural flexibility and that it may enhance evolutionary morphological diversification.

The relationship between floral display size, pollen carryover and geitonogamy in Myosotis colensoi (Kirk) Macbride (Boraginaceae)

Abstract: The relative amounts of self- and cross-pollen deposited on stigmas depends on both the number of pollinator visits that occur within plants and the amount of pollen carryover. Data collected for Myosotis colensoi (Kirk) Macbride and compiled from a survey of the literature, reveal that pollen carryover is frequently very high (upwards of 80%) and this at least partially relieves some of the effects of geitonogamous pollinator movements. It is suggested that in some cases, selection for traits that confer a high rate of pollen carryover may occur. Aspects of the plant–pollinator interaction that are likely to influence pollen carryover are discussed.

Seasonal plasticity in life history traits: growth and development in Polygonia c-album (Lepidoptera: Nymphalidae)

Abstract: The potentially multivoltine comma butterfly, Polygonia c-album L., hibernates in the adult stage. The adult seasonal morph is demonstrated to be a good indicator of whether an individual has entered reproductive diapause or is developing directly to sexual maturation. This fact, and the assumption that a short development time is not equally important to all categories of individuals, was used to test predictions on variation in life-history traits among categories (morphs and sexes) and environments (temperature and photoperiod) at the level of individuals and to some extent families and populations (the univoltine Stockholm population and the partially bivoltine Oxford population). Individuals developing to adults in a short time were expected to be smaller and lighter as a result of a basic trade-off between the two traits. Development times varied in accordance with predictions, but in most cases this was due to plastic growth and development in both the larval and pupal stages rather than through variation in size or weight, i.e. size was a highly canalized trait. This suggests a relationship between plasticity and canalization and a strong potential for plasticity to shield life-history traits from selection. Individuals regulated development times also within developmental pathways, in response to photoperiods indicating the progression of the season. These and other results suggest that development times are not normaily minimized in temperate butterflies unless this is enforced by direct development and protandry. There is thus scope for a high degree of adaptive plasticity in growth- and developmental rates which may devalue the basic trade-offs assumed by life-history theory and account for inconsistencies with its predictions.

Numbers and size of sperm storage tubules and the duration of sperm storage in birds: a comparative study

Abstract: Estimates of the numbers of sperm storage tubules (SSTs) in the utero-vaginal junction of 11 bird species are presented. Numbers of SSTs varied by a factor of 40 between species, and ranged from 500 to 20000. Body mass accounted for over 50% of the variation in SST mumbers. SST length was positively correlated with the length of spermatozoa across species. The duration of sperm storage was not correlated with the number of SSTs or the volume of sperm storage tissue. However, the number of ‘active’ SSTs appears to vary between species and it was not possible to make allowance for this. Sperm storage duration was weakly, positively correlated wth clutch size, but showed a significant positive relationship with the number of days over which laying occurred. The number of SSTs was also positively correlated with the number of sperm per ejaculate. The best predictor of sperm storage duration was a multiple regression equation using the spread of laying and the length of sperm storage tubules. The duration of sperm storage in birds which remain together during the pre-laying period is such that a single insemination immediately before the start of laying could fertilize the entire clutch.

Endemism and speciation in a lowland flora from the Cape Floristic Region

Abstract: Taxonomic, edaphic and biological aspects of endemism were studied in a phanerogamous flora from the Agulhas Plain, a coastal lowland area of the Cape Floristic Region. Of the 1751 species in the flora, 23.6% were regional endemics and 5.7% were local endemics. Families which were over-represented in terms of endemics included the Ericaceae, Rutaceae, Proteaceae and Polygalaceae. Under-represented families included the Poaceae, Cyperaceae, Scrophulariaceae and Orchidaceae. Highest levels of local endemism were recorded on limestone and colluvial acid sand. Sixty-nine percent of regional endemics and 85% of local endemics were confined to a single substratum. An analysis of the frequency of biological traits associated with species with different categories of endemism enabled the establishment of a biological profile of a local endemic: a dwarf to low, non-sprouting shrub with soil stored seeds which are ant-dispersed and/or form a symbiotic relationship with microbes. It is argued that lineages with these characteristics are vulnerable to severe population reduction or even local extinction. An effect of this would be the promotion of rapid, edaphic speciation as a result of catastrophic selection. Thus, certain traits (e.g. non-sprouting) prevail or even predominate in the flora not because of any adaptive advantage but because high speciation rates of lineages which possess them, overwhelm low survival rates.

Evolution of fruit characters and dispersal modes in the tropical family Rubiaceae.

Abstract: We investigated the evolution of fruit characters, animals versus abiotic dispersal modes, life forms and geographical distribution, in the large, mostly tropical, family Rubiaceae. As a basis for our analysis we used a phylogenetic tree derived from chloroplast DNA variation. Fleshy fruits have evolved independently at least 12 times in the family. Most of these originations appear to have occurred during Eocene to Oligocene, i.e. the radiation period for some animal taxa (bird families, mammal orders) comprising most extant dispersers of Rubiaceae fruits. Changes of dispersal modes may be of more recent origin in a few cases, e.g. evolution of drupes in some lineages, and shifts from drupes to nuts. The distribution of fruit characters suggested that in several lineages animal-dispersed fruits, such as berries and drupes have remained largely unaltered since the time of origination. This is in contrast to the occurrence of winged seeds in capsules, and pterophylls, i.e. enlarged calyx lobes promoting wind dispersal of fruits, which apparently have shifted more frequently during evolution, indicating a difference in ‘phylogenetic plasticity’ between modes of animal and wind dispersal. Animal dispersal was over-represented among genera dominated by shrubs, whereas abiotic dispersal was most prevalent among herbaceous genera. Drupes were over-represented in groups with transoceanic distributions, and on islands, indicating dispersal over long distances, probably by birds. In contrast, no evidence was found to support the view that animal dispersal in general enhances long distance dispersal. We also analysed geographical patterns on the tribal level but these were too complex to yield any resolved area cladograms due to the occurrence of many widespread taxa and area redundancy.

Relict floras of Atlantic islands: patterns assessed

Abstract: St. Helena (South Atlantic Ocean) and Macaronesia have endemic plants that have been interpreted as relicts. This paper divides these relicts into three main types. It is suggested that type 1 relicts are of Miocene age. They are usually wet tropical forest trees, geographically and taxonomically highly disjunct from their nearest relatives. Type 2 relicts are of Late Miocene or Pliocene age, usually trees of tropical or subtropical seasonally dry vegetation, often with trans-African disjunction, and moderate taxonomic isolation. Type 3 relicts are of Late Pliocene or Pleistocene age, usually herbs of arid zone or temperate vegetation, with little taxonomic or geographical disjunction from their nearest relatives. Relict theory is defined, with discussion of its explanatory power to account for patterns of endemism, and more generally, taxonomic and geographical patterns of biodiversity.

The colouration of the venomous coral snakes (family Elapidae) and their mimics (families Aniliidae and Colubridae)

Abstract: The bright coloured, highly venomous coral snakes, Leptomicrurus, Micrurus and Micruroides (family Elapidae) and a series of harmless or mildly toxic mimics form an important component of the snake fauna of the Americas. Coral snake patterns are defined as any dorsal pattern found in any species of venomous coral snake and/or any dorsal pattern containing a substantial amount of red, pink or orange distributed so as to resemble that of some species of venomous coral snake. The components of coral snake colouration are described and four principal dorsal patterns are recognized: unicolour, bicolour, tricolour and quadricolour. The tricolour patterns may be further clustered based on the number of black bands or rings separating the red ones as: monads, dyads, triads, tetrads or pentads. A detailed classification of all coral snake colour patterns is presented and each pattern is illustrated. The taxonomic distribution of these patterns is surveyed for mimics and the 56 species of highly venomous coral snakes. Among the latter, the most frequent encountered patterns are tricolour monads, tricolour triads and bicolour rings, in that order. No venomous coral snakes have a tricolour dyad, tricolour tetrad or quadricolour pattern. As many as 115 species of harmless or mildly toxic species, c. 18% of all American snakes, are regarded as coral snake mimics. The colouration and behavioural traits of venomous coral snakes combine to form a significant antipredator defence of an aposematic type. The mimics in turn receive protection from predators that innately or through learning avoid coral snake colour patterns. The precise resemblances in colouration between sympatric non-coral snakes and venomous coral snakes and the concordant geographic variation between the two strongly support this view. Batesian mimicry with the highly venomous coral snakes as the models and the other forms as the mimics is the favoured explanation for this situation. It is further concluded that a number of species in the genera Elaphe, Farancia, Nerodia and Thamnophis, although having red in their colouration, should not be included in the coral snake mimic guild.

Host specificity of arboreal and free‐living insect herbivores in rain forests

Abstract: Available information about the host specificity of arboreal and free-living insect herbivores in rain forests is scarce, but suggests that polyphagy may be more widespread than previously thought. The study of the arthropod fauna associated with an overstorey tree in Australia, Argyrodendron actinophyllum (Sterculiaceae), supports this contention. A conservative 11% of herbivore species foraging within this tree appear to be specialists, whereas this proportion decreases further to 3.0–4.5% when non-herbivore species are included. Argyrodendron actinophyllum presents several features which may promote a polyphagous strategy among herbivores. These observations are discussed in the wider context of ecological and evolutionary pressures shaping feeding strategies of insect herbivores in rain forests. These selection pressures include principally plant biochemistry, predation pressure of generalist predators, and structural diversity and environmental unpredictability of rainforest environments. The strength of association between herbivores and rainforest host-trees is likely to depend on three factors, acting at both the tree and rainforest community level: chemical traits of foliage (including chemical defences and nutrient levels), young foliage availability and enemy-free space. Given the complex interactions possible among these factors, rainforest host-trees may sustain herbivore faunas ranging from highly specialized to highly generalist. SUMMARY Argyrodendron actinophyllum presents several features which may promote a polyphagous strategy among herbivores: low nutrient levels and high fibre content, apparent scarcity of qualitative defences, relatively unpredictable availability of young foliage and low predation-pressure from arboreal ants in its foliage. This situation may be different for other rainforest tree species, depending on their biological features. Without speculating on the relative importance of each factor, these observations suggest that the strength of association between communities of insect herbivores and their rainforest host-trees may depend on the following factors, acting at both the component and compound rainforest-community level: chemical traits of foliage (including chemical defences and nutrient levels), young foliage availability (including factors related to plant phenology, plant abundance and plant growth strategy) and enemy-free space. These factors, which may be interrelated (i.e. amount and type of chemical defences related to nutrient availability, host phenology related to nutrient availability and searching behaviour of generalist predators/ parasitoids related to host phenology and apparency), may generate complex sets of combinations, which are likely to differ between rainforest component communities. In these conditions, rainforest host-trees may sustain herbivore faunas ranging from highly specialized to highly generalist. These considerations demonstrate a requirement for the study of arboreal insect communities associated with host-trees of dissimilar features growing in similar rain forest types, as well as in different rainforest environments.

Canine carnassials: character displacement in the wolves, jackals and foxes of Israel

Abstract: Previous research implies that competitive character displacement in felids and mustelids of Israel is expressed by canine size. Anatomy and observed killing behaviour of canids suggest that canines in this group are less adapted for the stylized role they play in felids and mustelids. Thus we hypothesized that character displacement, if it exists in canids, should not be manifested more clearly by canine size than by other traits. Five sympatric and at least partially syntopic canids occupy Israel, while in North Africa the largest (wolf) and smallest (Blanford's fox) are absent. Sexual size dimorphism in Israeli canids is generally less than in felids and mustelids (in which we analysed each sex as a separate ‘morphospecies’), so we used mixed-sex samples to represent each species. The three largest species (wolf, golden jackal and red fox) are also represented by Middle Palaeolithic samples in Israel, and all three had larger carnassial lengths then. Carnassial lengths, canine diameters and skull lengths are all remarkable evenly spaced among the five recent species in Israel. In Egypt, no trait manifests significant equality. Despite regional fluctuations in size, the carnassial length ratios of the three smaller species (foxes) are strikingly constant (1.18–1.21) throughout the region, while the ratios for the three larger species (wolf, jackal and red fox), sympatric only in Israel, are larger (1.33–1.34). Finally, mean carnassial length of jackals is constant across North Africa, while skull length and canine diameter both increase from Algeria through Egypt. All three traits are larger in Egypt than in Israel. We tentatively ascribe the equal ratios in Israel to competitive character displacement, though this hypothesis is speculative because of numerous lacunae in knowledge of diet, killing behaviour, available resources and extent of food limitation. Furthermore, humans have greatly affected range, density and ecology of wolves and jackals in the last century. Larger sizes in the Palaeolithic may well be manifestations of Bergmann's rule. The constancy of carnassial length in North African jackals, notwithstanding a longitudinal cline in CBLs of these populations, and the constant ratio between jackal and red fox carnassial length are both consistent with a hypothesis of character release in the absence of the wolf.

The phylogeny of a mutualism: evolution and coadaptation between Trollius and its seed-parasitic pollinators

Abstract: Interactions between seed-parasitic pollinators and their hosts provide useful model systems for the analysis of evolution of mutualism and potential coevolution between plants and insects. Here I present the systematics, pollination ecology and evolution of one of these interactions. I have documented and analysed the phylogenetic and geographic associations between Trollius (Ranunculaceae: 18 spp.) and Chiastocheta (Diptera: Anthomyiidae; 17 spp.), a host-specific genus of seed-parasitic flies that pollinate their host plants to varying extent. Their interactions are usually facultative mutualisms, but in the specialized T. europaeus three fly species are obligate mutualists and a fourth species is an antagonist. The distribution patterns of fly species among Trollius species suggest that the flies evolved in associations with five highly derived Trollius species, and secondarily colonized four more primitive taxa in the parts of their ranges that overlapped with primary hosts. In general, host specificity is maintained primarily through allopatry, with colonization occurring in regions of overlap between parapatric taxa. Fly speciation has occurred in allopatry, both within and among host taxa. Cospeciation is not evident, but convergent evolution in Trollius flowers of several traits, viz. orange sepals, elongated staminodia and increased carpel number per flower, may be the result of mutualism with Chiastocheta.

Single cytotype areas in autopolyploid Plantago media L.

Abstract: The European distribution of diploid and tetraploid cytotypes of Plantago media has been studied on three different scales: Europe as a whole, the Pyrenees and the upper Rio Gallego valley (Central Pyrenees). On a European scale, the tetraploids are widely and more or less continuously distributed; the diploids are restricted and disjunct. In the Pyrenees the distributions are essentially parapatric with diploids occupying the western part and tetraploids the eastern part. The overlap zone in the Rio Gallego valley consists of a mosaic of diploid, tetraploid and mixed populations. The Pyrenean distribution is best explained by colonization by cytotypes from different origins during the late glacial period followed by secondary contact. The limited intermingling of cytotypes is explained by exclusion, probably genetic and possibly also ecological. Reduced fitness of hybrid cytotypes may account for the existence of large areas with a single cytotype (‘single cytotype areas’) in allopolyploid species in general.

Canine Teeth of Bats (Microchiroptera): Size, Shape and Role in Crack Propagation

Abstract: Upper canines in microchiropteran bats show a variety of cross-sectional shapes. A consistent feature of all species studied here is that the tooth is edged and not simply round or oval. Prominent sharp edges are posi- tioned in several directions but particularly antero-medially toward the incisors and posteriorly toward the premo- lars. These edges appear to direct the cracks made in food items to the incisors or to the premolars. A continuous cut- ting edge is apparent in the occlusal view of the palate running from tip of canine to the ectoloph of the molars. Size and shape analysis indicates that larger bats have slender, rather than stouter, canines for their height, a condition that may be attributable to the nature of the prey. Most bats take prey that have little hard substance imbedded within. The compromises in tooth shape may vary between that of a terrestrial predator with short, conical canines for process- ing endoskeletal prey to that of a small fl ying predator with long, slender, edged canines for capturing and processing exoskeletal prey. Unicuspid teeth and how they might function in food break-up have been overlooked to the litera- ture; such a study could lead to an understanding of how more complex teeth function.

Primary faunal succession in volcanic terrain: lava and cave studies on the Canary Islands

Abstract: Invertebrate communities in volcanic habitats of different ages on the islands of La Palma and El Hierro were studied using standardized trapping and searching techniques. A variety of graphical and numerical approaches were used to analyse relationships among the sites. Young, barren lava flows constitute aeolian ecosystems with a fauna of generalized detritivores and predators, especially collembolans, earwigs, thysamirans and crickets. Surface samples have many individuals and low diversity; those from caves have smaller numbers but similar taxonomic composition. Vegetated surface habitats have richer communities, with diverse herbivores and predators but largely without the pioneer ‘lavicolous’ species. Caves with high humidity and stable temperature contain mainly specialized troglobitic species, but if there are both dry and humid sections lavicoles may also be present. Divergence into distinct epigean and hypogean communities results from both abiotic and biotic processes, including erosion and plant succession. While these occur mainly on the surface they also affect caves, increasing humidity and providing insulation from variations in external environmental conditions; the process is considered as a form of ‘maturation’ of the caves. Various models of succession are considered, which might help to account for the disappearance of lavicoles from mature epigean and hypogean communities.

An ecological classification of odonate mating systems: the relative influence of natural, inter‐ and intra‐sexual selection on males

Abstract: We separate the mating systems of odonates into two main groups: non-resource and resource-based systems. These two groups comprise five classes of mating system: encounter-limited mating, free female choice, resource-limitation, resource-control and female-control. These classes are consistent with previous classifications of odonate mating systems and with the overall classification of mating systems by Emlen & Oring (1977: Science, 797: 215–223). Whereas Emlen & Oring's classification of mating systems was concerned with differences in sexual selection between mating systems, our classification of odonate mating systems also addresses the influence of inter-and intra-sexual selection on males within a mating system. Predictions about such relationships are useful in multivariate analysis of odonate lifetime reproduction success. Among most odonate mating systems, much of the sexual selection on males results from male-male competition for access to mates. Sexual selection via female choice is relatively less important or operates indirectly through females' choices of times or places to mate. We place resource-control and resource-limitation at opposite ends of a resource-defence continuum and postulate female choice will have greater influence in mating systems that are more like a resource-limitation system and less influence in mating systems that are more like resource-control. Sexual selection is likely to be weak in species that resort to encounter-limited mating where longevity is likely to contribute strongly to variation in reproductive success. Females have limited opportunity to exercise choice among males in the female-control mating system and in this system selection is most likely to operate on male characters which contribute to their efficiency in searching for and capturing mates. Predictions about the differences in the intensity of sexual selection between different odonate mating systems should be made on the basis of the variation in the number of potential fertilizations per male or even per ejaculate, rather than the number of fertilizable females per male. Very different mating systems could result in similar patterns of variation in male reproductive success.

Land snail faunas of the Napier and Oscar Ranges, Western Australia; diversity, distribution and speciation

Abstract: The snail faunas of the Napier and Oscar Ranges, Western Australia have two major components: large camaenid species which are restricted endemics, with tiny geographical distributions and a pattern of geographical replacement within the Ranges, and smaller species of wider distribution, most of which are found throughout the Ranges. Sampling of 55 similar sites, protected from fire, over the length of the Ranges reveals this pattern clearly, with y diversities being far higher for camaenids than for others, in which there is no pattern of geographical replacement. This difference is not found in temperate and subtropical snail faunas from elsewhere, nor does the camaenid pattern occur in surveys of diversity in other organisms. These results are discussed in terms of dispersal and allopatric speciation related to demonstrable environmental fluctuation. An additional finding is that within one major taxonomic group (Gastropoda), a community may be partly structured by competition and exclusion and partly by accidents of colonization and accumulation: some, but not all niches may be full.

Population systematics of Russell's viper: a multivariate study

Abstract: A multivariate analysis of the population systematics of Russell's viper, based on scalation and colour pattern characters, reveals that the populations of this viper constitute two well-defined taxa: a western form, comprising all populations from the Indian subcontinent, and an eastern form, comprising all populations from east of the Bay of Bengal. The two forms could be considered either as subspecies of one species, or as two separate species, depending on the species concept used. Within the western form, there is no clear pattern of geographic variation. Within the eastern form, the populations from the Lesser Sunda Islands are clearly divergent from the populations of mainland Asia and Java. The conventionally recognized subspecies of Vipera russelli fail to portray this pattern of geographic variation. There is no clear relationship between the pattern of geographic variation in morphology and the pattern of geographic variation in the clinical effects of the venom in human bite victims: some populations with considerable differences in venom effects are equally distinct morphologically, whereas other populations with equally strong venom differences are morphologically very similar. The distribution of Russell's viper can be attributed to Pleistocene changes in climate and sea level, coupled with the viper's ecological requirements, which appear to include a seasonally dry climate.

Morphological constraints in the digging apparatus of pocket gophers (Mammalia: Geomyidae)

Abstract: The digging apparatus of pocket gophers offers a unique opportunity to examine morphological constraints within a historical context because relationships among extant taxa are well resolved and the features enhancing digging performance are relatively well understood. Structural and functional considerations suggest that the muscles associated with tooth- and claw-digging in pocket gophers are subjected to contrasting levels of morphological constraints. To assess this hypothesis, we analysed the bones and muscles of the jaws and forelimbs in four genera comprising five species of pocket gophers. Morphometric analyses were performed on 12 osteological measurements selected to reflect overall skull size, variation in rostral shape and procumbency, differences in overall length of the forelimbs and processes relating to the function of lever systems used in claw-digging. In addition, dissections were made of the jaw, hyoid, neck and all of the forelimb muscles excluding the intrinsic muscles of the manus. Results of our morphometric analyses corroborate the recent suggestion that pocket gophers encompass a wide range of morphological variation extending from claw-diggers to tooth-diggers. Myologically, however, we found structural variation only in the forelimb muscles, some of which may be advantageous for digging. No changes in jaw, neck and hyoid muscles, other than differences in muscle mass or those concordant with differences in rostral shape, were noted. These results support our hypothesis that constrasting levels of morphological constraint exist between the jaw and forelimb muscles of pocket gophers. We present a discussion of the structural and functional constraints on jaws and forelimbs in gophers as well as an analysis of historical constraints acting on this group, and perhaps on mammals in general.

A developmental constraint on the fledging time of birds

Abstract: We examined the hypothesis that the rate of bone growth limits the minimum fledging time of birds. Previous observations in California gulls indicate that linear growth of wing bones may be the rate limiting factor in wing development. If bone growth is rate limiting, then birds with relatively long bones for their size could be expected to have longer fledging periods than birds with relatively short bones. We tested this by comparing the length of wing bones, relative to body mass, to the relative length of fledging periods among 25 families. The results support the hypothesis. A strong correlation exists between relative fledging period and relative bone length. Species which have relatively long bones for their body size tend to take longer to fly. In contrast, parameters that influence flight style and performance, such as size of the pectoralis muscle and wing loading, show little or no correlation with fledging time. The analysis also indicates that, when altricial and precocial species are considered together, bone length is more highly correlated with fledging time than is body mass or rate of increase in body mass during growth. These observations suggest that linear growth of bones does limit the growth of avian wings and that it is one of the factors that influences the fledging time of birds.

A metric for the study of evolutionary trends in the complexity of serial structures

Abstract: Little empirical work has been done to see what sort of patterns of change in morphological complexity ocrur in evolution, mainly because the complexity of whole organisms has been so hard to define and to measure. For serial structures within organisms, there are fewer difficulties; this paper introduces a set of complexity metrics that are designed especially for serial structures, and then explores some of the properties of the new metrics. Also, a principle proposed in the last century by Herbert Spenrer, and offered recently in a new form by the thermodynamic school of evolutionary thought, predicts that complexity should increase in evolution as a consequence of the accumulation of perturbations. Here, simulations in which perturbations are introduced to ideal and real series of vertebral measurements show how the complexity increase predicted by Spencer’s principle would be captured by the new metrics.

Biogeography of the octocorallian coelenterate fauna of southern Africa

Abstract: The geographical and bathymetric distribution of southern African octocorals is analysed. Of the over 200 estimated species of regional octocorals, 81 confirmed and adequately described species are studied using a radial sector method. Two primary faunal components are recognized–endemic (53.3% of the fauna by numbers of species) and Indo-Pacific (39.4%). An Atlantic component contributes only minimally (about 1.7%), while the remaining fauna is made up of cosmopolites (2.8%) and scattered species (2.8%). A subantarctic component is not evident for the present-day, although evidence for previous contact is presented. A sister-group analysis using genera as a guide to sister species, shows the biogeographic affinities for the present-day fauna as a whole to be 45% Indo-Pacific, 31% cosmopolitan, 10% endemic, 10% Atlantic and 4% southern oceans (subantarctic). Applying the same method to only those genera with endemic species shows the affinities of the present-day endemic fauna to be 27.5% Indo-Pacific, 27.5% endemic, 24% cosmopolitan, 14% Atlantic and 7% subantarctic. Clearly defined boundaries for west, south, and east coast faunas (as recognized by previous authors in describing various intertidal faunas) are found not be present with regard to the octocoral fauna (largely due to its overwhelmingly subtidal nature). Instead two primary zoogeographic provinces are recognized–the Cape Endemic Province (extending from Liideritz to Inhaca Island) and the south-western fringe of the Indo-Pacific Province from East London north-eastwards. An overlap zone between these two is recognized between East London and Inhaca Island, with the region in the vicinity of Richards Bay having an essentially evenly mixed fauna (roughly 50% Cape Endemic Province and 50% Indo-Pacific). Of the 84 octocoral genera recorded for the region, seven (or 8.3%) are endemic, and of these, five are monotypic while two are ditypic. The fauna is shown to be predominantly sublittoral (about 95% by numbers of species), the shallow sublittoral (< 100 m in depth) being the region with highest species richness. Pennatulaceans are eurybathic (intertidal to 4756 m) and clearly show a high proportion of cosmopolites (20% of presently identified species). Soft corals are stenobathic and restricted to the intertidal, continental shelf and uppermost portion of the continental slope (<500m), while gorgonians are intermediate in depth distributon (intertidal to 1200 m). No cosmopolitan alcyonaceans are presently recorded. The centre of the Cape Endemic Province is the Agulhas Bank–an extensive region of shallow continential shelf (< 200 m in depth) between Cape Town and East London. Two regions of octocoral radiation for southern Africa are postulated–the Agulhas Bank and the western Indian Ocean.

The linyphiid fauna of Eastern Africa (Araneae, Linyphiidae) - distribution patterns, diversity and endemism

Abstract: Distributions, endemism and diversity among East African linyphiids are analysed and discussed in relation to other forest organisms and the environmental history of eastern African. A total of 231 species are reported from eastern Africa, of which 14 are confined to the Afroalpine region and 114 species to moist forests. Only 12 of the latter are widely distributed. The rest are only known from one or two localities. Information on habitats and distributions of all species is tabulated. Few species are shared between East African mountains and there are no detectable gradients of species diversity between mountains. There is, however, a gradient of decreasing species diversity from high latitudes to the Equator. Vicariance patterns are demonstrated for Elgonia, Ophrynia and Callitrichia in the Eastern Arc Mountains, Tanzania—areas that also hold the highest degree of endemism (> 80% on individual mountains) among linyphiids. The many endemic species on nearby mountains suggest that intermontane dispersal (ballooning) is rare or non-existent. There is no evidence for a distinction between highland and lowland linyphiid faunas, but altitudinal segregation of single species is demonstrated. The question of the reality of highland and lowland faunas cannot be solved by studying the altitudinal distribution of single species. Phylogenetic relationships must be taken into consideration to determine where sister-groups/species are located (lowland or highland).

The origins of allometry: size and shape polymorphism in the common waterstrider, Gerris remigis Say (Heteroptera, Gerridae)

Abstract: Changes in size, whether ontogenetic or phylogenetic, tend to be associated with changes in shape. This allometry can arise through two different evolutionary mechanisms: (1) selection acting primarily on overall size may be associated with changes in shape because of physiological and mechanical constraints or differential responses of different body components; or (2) selection acting primarily on shape (on the size of specific body components) may be associated with changes in overall size because of genetic correlations, and thus correlated responses, of other body components. To assess the relative importance of these two mechanisms, shape polymorphism is examined along two axes of size dimorphism (sex and wing morphology) in the common waterstrider, Gerris remigis Say. Eight measurements were made of body and appendage components of 234 adults, from three independent populations. Univariate and multivariate analyses reveal that both sexes and wing morphs differ significantly in size and shape. Shape differentiation along the two axes of size dimorphism is found to be dissimilar, partially independent of size, and strongly correlated with the ecological specialization of the various morphs. These observations suggest that selection is acting directly on shape, and thus that allometry in this species primarily reflects shape-mediated changes in size (mechanism 2), rather than size-mediated changes in shape. The role of developmental processes in facilitating this shape differentiation is discussed.

The geographic, taxonomic and temporal distribution of determinate growth in marine gastropods

Abstract: Many marine gastropods are characterized by determinate growth, as inferred from the presence of unique terminal elaborations of the shell's aperture. Although determinate growth has evolved repeatedly in most major gastropod clades, it is especially frequent among siphonate caenogastropods. Analyses of shallow-water assemblages show that the incidence of species with determinate growth is far higher in the tropics (especially the tropical Pacific and Indian Oceans) than at higher latitudes. Compilations of fossil assemblages from warm-water environments indicate that, although determinate growth occurred in some Palaeozoic gastropods, it became widespread only in the Neogene. In some groups, terminal apertural elaborations arose in lineages whose growth was more or less continuous and indeterminate, but in others it was derived either from or was ancestral to episodic growth. The hypothesis that periodic or terminal apertural elaborations evolved as a means to dispose of calcium carbonate once growth in the spiral direction ceased is rejected in favour of functional interpretations. Among the latter, the roles of modified apertures in defence and in mate recognition are explored, but no firm conclusion regarding the latter possibility can be drawn owing to our ignorance of mate recognition in gastropods.

Genetic variability and large scale differentiation in two species of littorinid gastropods with planktotrophic development, Littorina littorea (L.) and Melarhaphe (Littorina) neritoides (L.) (Prosobranchia: Littorinacea), with notes on a mass occurrence of M. neritoides in Sweden

Abstract: A planktotrophic larval development suggests a relatively high gene flow over long distances. Both Littorina littorea and Melarhaphe neritoides have an egg capsule and planktotrophic larvae which taken together are pelagic over a period of 4–8 weeks. In the absence of differential selection one would expect low levels of genetic differentiation over large distances in these two species. In this study of allozyme variation, low levels of differentiation were found over thousands of kilometres (Norway to Spain for L. littorea and Sweden to Greece for M. neritoides). This supports the hypothesis of two species with a high dispersal potential. A second expectation from neutral theory is that effective population size is positively correlated with average levels of genetic variation within species. In light of the generally high densities of local populations of these species and the high interpopulation migration rate, both L. littorea and M. neritoides may be considered as having high Ne s. Contrary to neutral expectation, L. littorea revealed very low levels of heterozygosity over its whole European range (mean Hexp= 0.015, 15 loci), while average heterozygosity of M. neritoides (Hexp= 0.084, 11 loci) was no more than in other littorinid species. This paper also reports the occurrence of M. neritoides on the Swedish west coast in 1988, 89 and 90, and two factors which may have promoted this unusually large invasion are discussed.

When did the house mouse colonize Europe

Abstract: This paper is a reply to a recent proposal by Sage and co-workers giving an early date for the colonization of Europe by the house mouse, based on mitochondria! DNA nucleotide divergence estimates. The large discrepancy in time between the date given by these authors and our own is discussed in the light of available palaeontological and genetic data. This leads us to reject the hypothesis supported by Sage and co-workers, and suggest a much more recent date of colonization.

Development and morphology of rostral cartilages in batoid fishes (Chondrichthyes: Batoidea), with comments on homology within vertebrates

Abstract: The rostral cartilages of batoid fishes were examined to elucidate their development, morphology and homology. Comparison of a variety of rostral cartilages among elasmobranchs with other groups of vertebrates shows that rostral cartilages originate embryologically from the trabecula and/or lamina orbitonasalis. Because different morphogenetic patterns of the derivatives of the two embryonic cartilages give rise to a wide variety of forms of rostral cartilages even within elasmobranchs, and because morphogenesis involves complex interactions among participating structures in the ethmo-orbital area, we put forward conceptual and empirical discussions to elucidate the homology of the rostral cartilages in batoid fishes. With six assumptions given in this study and based on recent discussions of biological and historical homology, our discussions centre on: (1) recognition of complex interactions of participating biological entities in development and evolution; (2) elucidation of a set of interacting biological and evolutionary factors to define a given morphological structure; (3) assessment of causal explanations for similarities or differences between homologous structures by determining genetic, epigenetic and evolutionary factors. Examples of conceptual approaches are given to make the approaches testable. Although a paucity of knowledge of rostral cartilage formation is the major obstacle to thorough analysis of the conceptual framework, several tentative conclusions are made on the homology of rostral cartilages that will hopefully attract more research on development and evolution in vertebrate morphology. These are: (1) the rostral cartilage in each group of vertebrates examined can be defined by both developmentally associated and adult structural attributes, yet such data do not allow us to assess homology of a variety of forms of rostral cartilages at higher taxonomic categories; (2) the entire rostral cartilage in elasmobranchs is formed by the contribution of the embryonic trabecula and lamina orbitonasalis. The status of the development and homology of the rostral cartilage in holocephalans remains uncertain; (3) there is no simple picture of evolution of rostral cartilages among three putative monophyletic assemblages of elasmobranchs, galeomorphs, squaloids (possibly plus Squatina, Chlamydoselachus and hexanchoids as the orbitostylic group) and batoid fishes. It is highly likely that rostral cartilages in each subgroup or subgroups of these assemblages may be of phylogenetic significance but that it may not serve as a basis to unite these assemblages into much higher assemblages; (4) the tripodal rostral cartilage is unique in form in the group including some carcharhinoid and lamnoid sharks. The status of the analogous tripodal cartilage in some squaloids remains uncertain. The unfused tripodal cartilage of the electric ray Narke is interpreted as developmentally equivalent to, but not homologous with, the unfused or fused ones in the sharks; (5) the rostral cartilage in the electric ray Torpedo is uniquely formed because of its embryonic origin solely from the ventro-medial part of the lamina orbitonasalis, but it is regarded as homologous with the rostral cartilages which are formed by the trabecula and other components of the lamina orbitonasalis in other batoid fishes; (6) the cornu trabecula contributes to the formation of the ventral stem of the rostral cartilage at least in elasmobranchs, especially to a particular set of rostral cartilages, i.e. the tripodal rostral cartilage in the shark Scyliorhinus and dorso-ventrally flattened rostral shaft in the narcinidid electric rays; (7) there is a unique form of a rostral shaft with rostral appendix in skates and probably guitarfishes; (8) there is no rostral cartilage in adult benthic stingrays, pelagic stingrays Dasyatis violacea and Myliobatidae, although it is present in embryonic stages; (9) there is a unique form of the rostral cartilage as a rostral projection from the dorso-lateral part of the lamina orbitonasalis in pelagic stingrays Rhinopteridae and Mobulidae, which together with part of the pectoral fins, forms a pair of cephalic fins; (10) different developmental mechanisms may be responsible for the absence or loss of rostral cartilages in different groups, i.e. absence of the cartilage derived from the medial area of the trabecula in Torpedo vs absence of the rostral cartilage in benthic stingrays; (11) the rostral cartilages in some placental mammals (cetaceans and sirenians) arise only from the medial area of the trabecula because monotreme and placental mammals do not form the trabecula cranii; (12) some actinopterygians and sacropterygians possess a rostral cartilage which originates only from the medial area of the trabecula. One scombroid group, including Sardini and Thunnini, Scomberomorus, Acanthocybium, Istiophoridae and Xiphias, possesses a unique larval beak composed of the rostral cartilage, ethmoid cartilage and premaxillar bone. The development and homology of other rostral cartilages remain to be further elucidated; (13) urodeles possess a medial rostral process whose anlage is probably developmentally equivalent to that in batoid fishes but the occurrence in urodeles is either atavistic or unique (autapomorphic); (14) the upper jaw of tadpoles is unique in possessing the suprarostral cartilage; the anlage of the cartilage is probably developmentally equivalent to the outgrowth of the cornu trabecula in batoid fishes.

Phylogenetic Relationships and Rates of Allozyme Evolution among the Lineages of Sceloporine Sand Lizards

Abstract: Parsimony analysis of characters derived from an electrophoretic survey of allozyme variation in the sceloporine sand lizards indicates that Uma is outside of a clade formed by the rest of the sand lizards and that Cophosaurus and Holbrookia share a more recent common ancestor with one another than either does with Callisaurus. Previous electrophoretic studies used phenetic clustering based on genetic distance data to assess relationships among these taxa. The resulting dendrograms were used to argue that Holbrookia is the sister group of all other sand lizards and that Callisaurus and Cophosaurus are sister taxa. When reanalysed using parsimony methods, the data from these previous studies are found to support the conclusions of the present study, namely, that Uma rather than Holbrookia is the sister group of all other sand lizards and that Cophosaurus is the sister taxon of Holbrookia rather than of Callisaurus. Relative rate tests indicate that the incongruencies between branching diagrams derived from phenetic clustering of genetic distances versus those derived from parsimony analysis of electrophoretic characters are attributable to increased rates of protein evolution in the Holbrookia lineage.

Reversed plumage ontogeny in a female hummingbird: implications for the evolution of iridescent colours and sexual dichromatism

Abstract: In polygynous birds, bright plumage is typically more extensive in the sexually competitive males and develops at or after sexual maturity. These patterns, coupled with the importance of male plumage in sexual displays, fostered the traditional hypothesis that bright plumages and sexual dichromatism develop through the actions of sexual selection on males. This view remains problematic for hummingbirds, all of which are polygynous, because their bright iridescent plumages are also important non-sexual signals associated with dominance at floral nectar sources. Here I show that female amethyst-throated sunangels [Heliangelus amethysticollis (d'Orbigny & Lafresnaye)], moult from an immature plumage with an iridescent gorget to an adult plumage with a non-iridescent gorget. This ‘reversed’ ontogeny contradicts the notion that iridescent plumage has a sexual function because sexual selection in polygynous birds should be lowest among non-reproductive immature females. Moreover, loss of iridescent plumage in adult females indicates that adult sexual dichromatism in H. amethysticollis is due in large part to changes in female ontogeny. I suggest that both the ontogeny and sexual dichromatism evolved in response to competition for nectar.