Cephalopod consciousness on “their own terms”: Who are “they”?

The ecological, morphological, and physiological diversity of species in the Hawaiian silversword alliance is exceptional. The 28 species, which belong to the endemic genera Argyroxiphium, Dubautia, and Wilkesia, have a wide variety of geographical distributions and elevational ranges within the archipelago. They grow in habitats as varied as dry scrub and woodland, wet scrub and forest, cinder and lava, and bog. Ecological diversity is also evident among sympatric species. At a site of sympatry on the island of Hawaii, for example, D. ciliolata and D. scabra are restricted to different lava flows, even though individuals of the two species may grow within a meter of one another. The 28 species have growth forms as varied as rosette shrubs, shrubs, trees, and lianas. They have a wide range of leaf sizes and shapes, with the Dubautia species exhibiting significant variation in leaf turgor maintenance capacities. Morphological and physiological diversity is also evident among sympatric species. At a site of sympatry on the island of Maui, for example, A. sandwicense and D. menziesii exhibit different suites of morphological and physiological traits enabling them to cope with the severe environmental conditions. The patterns of diversity and the genomic relationships among the 28 species suggest that a variety of factors may have played important roles in their adaptive radiation. The Hawaiian silversword alliance is a premier example of adaptive radiation in plants (Carlquist, 1980; Carr et al., 1989). The alliance includes 28 species in three endemic genera: Argyroxiphium, Dubautia, and Wlilkesia (Carr, 1985). The species grow in a wide range of habitats and have a wide variety of growth forms. They are also closely related, as evidenced by the high frequency of spontaneous interspecific and intergeneric hybrids in nature, coupled with the ease of production of artificial hybrids in the laboratory (Carr & Kyhos, 1981, 1986). The detailed analysis of the hybrids and parental taxa using cytogenetic, electrophoretic, and molecular approaches has provided compelling evidence that the silversword alliance is a genetically cohesive group whose origin and diversification probably trace to a single colonizing ancestor (Baldwin et al., 1988; Carr et al., 1989). Our objective in this review is to provide insight into the ecological, morphological, and physiological diversity of species in the silversword alliance. With respect to ecological diversity, we compare the geographical distributions, habitats, and elevational ranges of the 28 species, then analyze the local distributions of several sympatric Dubautia species. With respect to morphological and physiological diversity, we compare the growth forms, leaf sizes, and leaf shapes of the 28 species, then examine the turgor maintenance capacities of the Dubautia species and the water and temperature balances of two sympatric Argyroxiphium and Dubautia species. Our primary theme is that the ' This paper is dedicated to Dr. Sherwin Carlquist in honor of his pioneering research on the Hawaiian silversword alliance. The research was supported by NSF Grant DEB-82064 11 and a gift from the Atlantic Richfield Foundation to the senior author. We thank Lani Stemmermann for invaluable companionship, botanical insight, and technical assistance in the field. We also thank the staff of Haleakala National Park, particularly R. Nagata and L. Loope, for critical logistical support, and J. Canfield and N. Friedman for generous photographic assistance. 2 Department of Ecology & Evolutionary Biology, University of Arizona, Tucson, Arizona 85721, U.S.A. < Department of Botany, University of Hawaii, Honolulu, Hawaii 96822, U.S.A. 4 Department of Plant & Soil Sciences, University of Maine, Orono, Maine 04469, U.S.A. Department of Botany, University of California, Davis, California 95616, U.S.A. ANN. MISSOURI BOT. GARD. 77: 64-72. 1990. This content downloaded from 152.1.161.146 on Thu, 27 Mar 2014 19:39:59 PM All use subject to JSTOR Terms and Conditions Volume 77, Number 1 Robichaux et al. 65 1990 Adaptive Radiation of Hawaiian Silversword Alliance

AimMorphological and taxonomic diversity are intuitive measures of biological diversity. Previous studies have shown discordance between these measures at large spatial and temporal scales, but the implications of this pattern for the underlying processes are not understood. Using oceanic archipelagos as spatial units, we examine potential links between the morphological and taxonomic diversity of their land snail faunas in a biogeographical framework.LocationEleven major oceanic archipelagos.MethodsFor each archipelago, we assembled lists of indigenous land snail species, classified by family and genus, with shell height and width for each species (1723 species in total). We used biogeographic and climatic variables as potential predictors of diversity patterns. We employed regression analyses to evaluate (1) whether morphological diversity scales with taxonomic diversity at the species, genus or family level, and (2) whether morphological and taxonomic diversity correlate similarly with biogeographic/climatic factors. We also assessed which taxonomic level contributes most to morphological variation within archipelagos.ResultsMorphological diversity across archipelagos was strongly related to genus but not species richness. Within archipelagos, morphological variation reflected differences among genera and families but not species. Species richness was best explained by archipelago area, but morphological diversity was not significantly related to any of the physical features of archipelagos.Main conclusionsAcross archipelagos, species richness and morphological diversity of land snail faunas are decoupled. The relationship between species richness and the available ecological space (captured mainly by area) indicates the prevalence of niche‐based processes while, for morphological diversity, the strong conservatism of morphology at the genus level suggests the presence of diversification‐based limits. Assuming genera effectively reflect diversification, our findings indicate that morphological space on oceanic archipelagos depends primarily on the number of evolutionary units that have colonized and/or diversified through time.

Exceptional ecological niche diversity, clear waters and unique divergent selection pressures have often been invoked to explain high morphological and genetic diversity of taxa within ancient lakes. However, it is possible that in some ancient lake taxa high diversity has arisen because these historically stable environments have allowed accumulation of lineages over evolutionary timescales, a process impossible in neighbouring aquatic habitats undergoing desiccation and reflooding. Here we examined the evolution of a unique morphologically diverse assemblage of thiarid gastropods belonging to the Melanoides polymorpha'complex' in Lake Malawi. Using mitochondrial DNA sequences, we found this Lake Malawi complex was not monophyletic, instead sharing common ancestry with Melanoides anomala and Melanoides mweruensis from the Congo Basin. Fossil calibrations of molecular divergence placed the origins of this complex to within the last 4 million years. Nuclear amplified fragment length polymorphism markers revealed sympatric M. polymorpha morphs to be strongly genetically differentiated lineages, and males were absent from our samples indicating that reproduction is predominantly parthenogenetic. These results imply the presence of Lake Malawi as a standing water body over the last million years or more has facilitated accumulation of clonal morphological diversity, a process that has not taken place in more transient freshwater habitats. As such, the historical stability of aquatic environments may have been critical in determining present spatial distributions of biodiversity.

ABSTRACTBermudagrasses (Cynodon spp.) are genetically and morphologically diverse warm‐season species that are valuable for turf and forage. Diverse accessions of Cynodon are preserved in various germplasm banks in Asia, Europe, and the United States. However, the extent of genetic and ecological diversity within Australia has not been previously explored and characterized. To address this issue, a broad‐scale collection of Cynodon germplasm across Australia was recently conducted. A core collection was generated from this resource to optimize its utilization in breeding and research. Genetic diversity of 690 Cynodon accessions was characterized using 16 expressed sequence tag (EST)‐simple sequence repeat (SSR) markers. The mean number of alleles generated was 7.44 per marker. Genetic data was combined with passport, ploidy level, morphological, and experimental performance data to generate a core collection using a directed stratified sampling method. The core collection comprised 13% of the source germplasm and captured 96% of the allelic diversity. Redundancy within the core collection was avoided with all accessions representing unique genotypes. This core collection will facilitate the future use of Australian germplasm in Cynodon breeding, genomics, conservation, taxonomy, and phylogeographical research.

BackgroundBiological diversity is a hot topic in current research, especially its observed decrease in modern times. Investigations of past ecosystems offer additional insights to help better understand the processes underlying biodiversity. The Cretaceous period is of special interest in this context, especially with respect to arthropods. During that period, representatives of many modern lineages appeared for the first time, while representatives of more ancient groups also co-occurred. At the same time, side branches of radiating groups with ‘experimental morphologies’ emerged that seemed to go extinct shortly afterwards. However, larval forms, with their morphological diversity, are largely neglected in such studies, but may provide important insights into morphological and ecological diversity and its changes in the past.ResultsWe present here a new fossil insectan larva, a larval lacewing, in Cretaceous amber, exhibiting a rather unusual, ‘experimental’ morphology. The specimen possesses extremely large (in relation to body size) mandibulo-maxillary piercing stylets. Additionally, the labial palps are very long and are subdivided into numerous elements, overall appearing antenniform. In other aspects, the larva resembles many other neuropteran-type larvae.ConclusionsWe provide a comparison that includes quantitative aspects of different types of neuropteran larvae to emphasise the exceptionality of the new larva, and discuss its possible relationships to known lineages of Neuroptera; possible interpretations are closer relationships to Dilaridae or Osmylidae. In any case, several of the observed characters must have evolved convergently. With this new find, we expand the known morphological diversity of neuropterans in the Cretaceous fauna.

Amphipods from the genus Niphargus represent an important part of the Western Palearctic subterranean fauna. The genus is morphologically diverse, comprising several distinct ecomorphs bound to microhabitats in the subterranean environment. The most impressive among them are “lake giants,” a series of massive, large‐bodied species. These range from morphologically distinct to morphologically cryptic taxa. We analysed the taxonomic structure of the Niphargus arbiter–Niphargus salonitanus species complex, belonging to “lake giants” from the Dinaric Karst (West Balkans), and assessed their phylogenetic, morphological and ecological diversity. Multilocus phylogeny suggested that the complex is monophyletic and nested within other cave lake ecomorphs. Unilocus and multilocus coalescence species delimitations indicated that the complex totals nine species. These species substantially overlap in morphology and cannot be unambiguously told apart without the use of molecular markers. An analysis of splitting events within a palaeogeological context, and modelling of environmental characteristics on the phylogeny unveiled a complex history of diversification. Part of this diversification might have been influenced by ecological divergence along the altitudinal gradient reaching from the Adriatic coast to inland Dinaric mountain chains and Poljes. Other splits coincide with the marine regression–transgression cycles during Pliocene. We describe Niphargus alpheus sp. n., Niphargus anchialinus sp. n., Niphargus antipodes sp. n., Niphargus arethusa sp. n., Niphargus doli sp. n., Niphargus fjakae sp. n. and Niphargus pincinovae sp. n., and by doing so hope to prompt their further research.

A new genus and species of fossil bat is described from New Zealand’s only pre-Pleistocene Cenozoic terrestrial fauna, the early Miocene St Bathans Fauna of Central Otago, South Island. Bayesian total evidence phylogenetic analysis places this new Southern Hemisphere taxon among the burrowing bats (mystacinids) of New Zealand and Australia, although its lower dentition also resembles Africa’s endemic sucker-footed bats (myzopodids). As the first new bat genus to be added to New Zealand’s fauna in more than 150 years, it provides new insight into the original diversity of chiropterans in Australasia. It also underscores the significant decline in morphological diversity that has taken place in the highly distinctive, semi-terrestrial bat family Mystacinidae since the Miocene. This bat was relatively large, with an estimated body mass of ~40 g, and its dentition suggests it had an omnivorous diet. Its striking dental autapomorphies, including development of a large hypocone, signal a shift of diet compared with other mystacinids, and may provide evidence of an adaptive radiation in feeding strategy in this group of noctilionoid bats.

This study develops the random phylogenies rate test (RAPRATE), a likelihood method that simulates morphological evolution along randomly generated phylogenies, and uses it to determine whether a considerable difference in morphological diversity between two sister clades of South American fishes should be taken as evidence of differing rates of morphological change or lineage turnover. Despite identical ages of origin, similar species richness, and sympatric geographic distributions, the morphological and ecological diversity of the superfamily Anostomoidea exceeds that of the Curimatoidea. The test shows with 90% confidence (using variance among species as the measure of morphological diversity) or 99% confidence (using volume of occupied morphospace) that the rate of morphological change per unit time in the Anostomoidea likely exceeded that of the Curimatoidea. Variation in the rate of lineage turnover (speciation and extinction rates) is not found to affect greatly the morphological diversity of simulated clades and is not a likely explanation of the observed difference in morphological diversity in this case study. Though a 17% or greater delay in the onset of diversification in the Curimatoidea remains a possible alternative explanation of unequal morphological diversification, further simulations suggest that two clades drawn from the possible treespace of the Anostomoidea and Curimatoidea will rarely differ so greatly in the onset of diversification. Several uniquely derived morphological and ecological features of the Anostomoidea and Curimatoidea may have accelerated or decelerated their rate of morphological change, including a marked lengthening of the quadrate that may have relaxed structural constraints on the evolution of the anostomoid jaw.

Biodiversity encompasses multiple facets, among which taxonomic, functional and phylogenetic aspects are the most often considered. Understanding how those diversity facets are distributed and what are their determinants has become a central concern in the current context of biodiversity crisis, but such multi‐faceted measures over large geographical areas are still pending. Here, we measured the congruence between the biogeographical patterns of freshwater fish morphological, ecological and phylogenetic diversity across Europe and identified the natural and anthropogenic drivers shaping those patterns. Based on freshwater fish occurrence records in 290 European river catchments, we computed richness and evenness for morphological, ecological and phylogenetic diversity using standardized effect sizes for each diversity index. We then used linear models including climatic, geo‐morphological, biotic and human‐related factors to determine the key drivers shaping freshwater fish biodiversity patterns across Europe. We found a weak spatial congruence between facets of diversity. Patterns of diversity were mainly driven by elevation range, climatic seasonality and species richness while other factors played a minor role. Finally, we found that non‐native species introductions significantly affected diversity patterns and influenced the effects of some environmental drivers. Morphological, ecological and phylogenetic diversity constitute complementary facets of fish diversity rather than surrogates, testifying that they deserve to be considered altogether to properly assess biodiversity. Although the same environmental and anthropogenic factors overall explained those diversity facets, their relative influence varied. In the current context of global change, non‐native species introductions may also lead to important reshuffling of assemblages resulting in profound changes of diversity patterns.

The correlated evolution of traits may be a principal factor in morphological evolution, but it is typically studied in genetic or developmental systems. Most studies examining phenotypic trait correlations, through analysis of morphological integration, consider only few taxa, with limited ability to test hypotheses of the influence of trait integration on morphological variation and diversity. The few comparative studies in less inclusive groups have yielded varying relationships of integration to the key factors of phylogeny and diet. In this paper, I present analyses of cranial morphological integration in 30 species from the mammalian order Carnivora, spanning eight extant families and a wide range of ecological and morphological diversity. Fifty-five cranial landmarks were captured through three-dimensional digitization of 15-22 specimens for each species. Using a node-based phylogenetic distance matrix, a significant correlation was found between similarity in patterns of integration and phylogenetic relatedness within Felidae (cats) and Canidae (dogs), but not within more inclusive clades, when size-related variation was removed. When size was included, significant correlations were found across all Caniformia, Musteloidea, Mustelidae, and Felidae. There was a significant correlation between phylogeny and morphological integration only within the higher-level clade Feliformia (cats, civets, mongooses, and hyaenas) when a branch-length-based phylogenetic distance matrix was analyzed, with and without size. In contrast, diet was significantly correlated with similarity in morphological integration in arctoid carnivorans (bears, raccoons, and weasels), but had no significant relationship with integration in feliforms or canids. These results support the proposition that evolutionary history is correlated with cranial integration across large clades, although in some smaller clades diet also exerts significant influence on the correlated evolution of traits.

The correlated evolution of traits may be a principal factor in morphological evolution, but it is typically studied in genetic or developmental systems. Most studies examining phenotypic trait correlations, through analysis of morphological integration, consider only few taxa, with limited ability to test hypotheses of the influence of trait integration on morphological variation and diversity. The few comparative studies in less inclusive groups have yielded varying relationships of integration to the key factors of phylogeny and diet. In this paper, I present analyses of cranial morphological integration in 30 species from the mammalian order Carnivora, spanning eight extant families and a wide range of ecological and morphological diversity. Fifty-five cranial landmarks were captured through threedimensional digitization of 15–22 specimens for each species. Using a node-based phylogenetic distance matrix, a significant correlation was found between similarity in patterns of integration and phylogenetic relatedness within Felidae (cats) and Canidae (dogs), but not within more inclusive clades, when size-related variation was removed. When size was included, significant correlations were found across all Caniformia, Musteloidea, Mustelidae, and Felidae. There was a significant correlation between phylogeny and morphological integration only within the higherlevel clade Feliformia (cats, civets, mongooses, and hyaenas) when a branch-length-based phylogenetic distance matrix was analyzed, with and without size. In contrast, diet was significantly correlated with similarity in morphological integration in arctoid carnivorans (bears, raccoons, and weasels), but had no significant relationship with integration in feliforms or canids. These results support the proposition that evolutionary history is correlated with cranial integration acrosslarge clades, although in some smaller clades diet also exerts significant influence on the correlated evolution of traits.

Aerva javanica is one of Egypt’s most important traditional medicinal plants used as antidiarrheal and anthelmintic medicine and recently as an anticancer agent. In this study, variations among ten populations of Aerva javanica in different sites in the Eastern Desert of Egypt were analyzed based on morphological and ecological attributes and molecular variation expressed by Inter-Simple Sequence Repeat (ISSR) markers. Morphological diversity was higher for populations in the Wadi El-Markh and Bir Abbady regions than others. The polymorphism revealed by ten ISSR primers was 79.4% among populations. Distance trees created using the results obtained from soil variables, morphological characterizations, and molecular data showed that the highest similarity was 0.974 between Populations 8 and 9, while the lowest similarity was 0.715 between Population 1 and Population 3 regions. In conclusion, the obtained data are important to design a plan for sustainable conservation of Aerva javanica as an important medicinal plant having a wide interspecific genetic variability within various populations.

Disparity, or morphological diversity, is often quantified by evolutionary biologists investigating the macroevolutionary history of clades over geological timescales. Disparity is typically quantified using proxies for morphology, such as measurements, discrete anatomical characters, or geometric morphometrics. If different proxies produce differing results, then the accurate quantification of disparity in deep time may be problematic. However, despite this, few studies have attempted to examine disparity of a single clade using multiple morphological proxies. Here, as a case study for this question, we examine the disparity of the volant Mesozoic fossil reptile clade Pterosauria, an intensively studied group that achieved substantial morphological, ecological and taxonomic diversity during their 145+ million-year evolutionary history. We characterize broadscale patterns of cranial morphological disparity for pterosaurs for the first time using landmark-based geometric morphometrics and make comparisons to calculations of pterosaur disparity based on alternative metrics. Landmark-based disparity calculations suggest that monofenestratan pterosaurs were more diverse cranially than basal non-monofenestratan pterosaurs (at least when the aberrant anurognathids are excluded), and that peak cranial disparity may have occurred in the Early Cretaceous, relatively late in pterosaur evolution. Significantly, our cranial disparity results are broadly congruent with those based on whole skeleton discrete character and limb proportion data sets, indicating that these divergent approaches document a consistent pattern of pterosaur morphological evolution. Therefore, pterosaurs provide an exemplar case demonstrating that different proxies for morphological form can converge on the same disparity signal, which is encouraging because often only one such proxy is available for extinct clades represented by fossils. Furthermore, mapping phylogeny into cranial morphospace demonstrates that pterosaur cranial morphology is significantly correlated with, and potentially constrained by, phylogenetic relationships.

From the ecologist’s perspective, centrarchid fishes are widely recognized as a model system for investigating the role of phenotypic variation in shaping ecological patterns. To the ichthyologist, this group is considered among the most morphologically and ecologically diverse of North America’s freshwater ichthyofauna. This chapter is intended to bring these perspectives together, highlighting the contributions of studies linking resource use patterns to morphology in order to make sense of the ecological, functional, and morphological diversity exhibited within the Centrarchidae. We review literature on feeding and on locomotion. Historically, the diversity represented within this radiation helped inspire the development of ecomorphology, a research perspective that investigates hypothesized associations between organismal design and ecology. Working independently, Werner (1974, 1977) and Keast (1978, 1985; Keast and Webb 1967) were among the first to point out a general association between head and body form and resource use in centrarchid species. Using bluegill sunfish (Lepomis macrochirus), green sunfish (Lepomis cyanellus), and largemouth bass (Micropterus salmoides) to represent the range of ecological and morphological diversity in centrarchids, Werner and coworkers developed the first mechanistic insights into the implications of variation in body and head morphology. The diversity of form and feeding habits represented by bluegill, largemouth bass, black crappie (Pomoxis nigromaculatus), and rock bass (Ambloplites rupestris) motivated Keast’s proposal that different suites of morphological features confer varying prey capture and habitat use capabilities on these species and that these differences underlie the capacity for these species to coexist in sympatry. The rationale for ecomorphology research can be seen in both Werner’s and Keast’s work: an organism’s morphology affects its capacity to perform an ecologically relevant task, and this performance capacity affects the resources available for its use. This research program was made more explicit (Werner 1977; Mittelbach 1984; Wainwright 1996) by emphasizing that researchers’ ability to explain ecological phenomena through organismal design requires focus on characters whose performance consequences are predictable. This stipulation established a primary role for functional morphology research, which investigates the morphological basis of performance variation. Moreover, the ecomorphology research perspective led to widespread recognition that the choice of an appropriate performance measure is vital to the success of studies that seek to understand the relationship between morphology and resource use. Performance variables range from proximate measures that focus on the mechanical capacities of isolated functional units, such as maximum pharyngeal jaw bite force, to more integrative measures that involve multiple functional units, like prey handling time, which is influenced by the fish’s ability to capture and process prey. In either case, the performance measure should have predictable consequences for resource use. This is not a trivial issue, as the link between any given performance measure and resource use is more frequently assumed than demonstrated. Nevertheless, studies involving centrarchid fishes provide some of the best examples of the ecomorphology research program carried out to completion. The morphological diversity of centrarchid fishes ranges between the forms exhibited by the predominant ecomorphs: piscivore/crayfish predator, zooplanktivore, molluscivore, and insectivore, which possess combinations of head and body characters that are associated with different patterns of resource use. Although these ecomorphs are named according to trophic habits, they are generally associated with habitat use patterns as well. Here, we highlight a set of morphological characters that have well-known consequences for performance and resource use. We focus on mouth gape, degree

The question of grouping students into homogeneous and heterogeneous groups is not new, but it does not find an unambiguous answer in the literature, especially in mathematics. In this paper, we address the question of whether grouping students into homogeneous and heterogeneous groups in mathematics improves their knowledge. The quasi-experiment involved 126 8-grade (i.e., 13-14 years old) Slovenian primary school students, who were divided into two equal groups: the control group worked in homogeneous groups, and the experimental worked in heterogeneous groups. The results of the post-test show that the students from the experimental group had better results in mathematics, which indicates that heterogeneous groups should be preferred in mathematics. Lesson observations have identified differences in teacher behavior: educators working in homogeneous groups tend to give students with lower competencies tasks from lower taxonomic levels, and teachers favor abler groups. Such differences have not been observed among teachers working in heterogeneous groups. The implications for educators are also discussed.