Floral morphology and evolution in Anisophylleaceae

TOBE, H. & RAVEN, P. H., 1988. Floral morphology and evolution in Anisophylleaceae. The four genera of Anisophylleaceae ( Anisophyllea, Combretocarpus, Poga , and Polygonanthus ) are very uniform in their floral structures. Characteristic floral features of the family are: flowers small (except for the female flowers of Polygonanthus ), merism nearly fixed (i.e. 3- or 4-mery), petals deeply incised (except in Polygonanthus ), ovary inferior and multi-loculed, ovules few (one or two) per carpel, styles separate, intra- and interstaminal nectariferous tissues present, and floral vasculature simple. Comparisons with related groups support the distinctiveness of Anisophylleaceae, and suggest a close affinity with both Rhizophoraceae and the Myrtales. The presence of incised petals in both groups suggests an especially close relationship between Anisophylleaceae and Rhizophoraceae, while new evidence from comparative floral morphology suggests that Anisophylleaceae occupy an intermediate position between Rhizophoraceae and Myrtales. Within the Anisophylleaceae, Poga and Polygonanthus share several synapomorphies in floral structure, while Combretocarpus is the most divergent genus in the family and is more distantly related to Poga and Polygonanthus . It is uncertain whether Anisophyllea is more closely related to Poga and Polygonanthus or Combretocarpus , because the evidence from comparative floral morphology conflicts with that from embryology; more data from other kinds of characters are needed to resolve this issue.     

Principal directions of the evolution of Monimotrochida

Phylogenetic relations among the main groups of Monimotrochida are considered. The principal directions of monimotrochid evolution were defined by comparative investigations of mastax morphology (SEM), basic body structures, and general biology. On the basis of these results we propose a revision of previous rotifer taxonomy. We suggest to place the Monimotrochida in the order Protoramida divided into two suborders Flosculariina and Conochilina.     

Comparative evolution of flower and fruit morphology

Angiosperm diversification has resulted in a vast array of plant morphologies. Only recently has it been appreciated that diversification might have proceeded quite differently for the two key diagnostic structures of this clade, flowers and fruits. These structures are hypothesized to have experienced different selective pressures via their interactions with animals in dispersal mutualisms, resulting in a greater amount of morphological diversification in animal-pollinated flowers than in animal-dispersed fruits. I tested this idea using size and colour traits for the flowers and fruits of 472 species occurring in three floras (St John, Hawaii and the Great Plains). Phylogenetically controlled analyses of nearest-neighbour distances in multidimensional trait space matched the predicted pattern: in each of the three floras, flowers were more divergent from one another than were fruits. In addition, the spacing of species clusters differed for flowers versus fruits in the flora of St John, with clusters in flower space more divergent than those in fruit space. The results are consistent with the idea that a major driver of angiosperm diversification has been stronger selection for divergent floral morphology than for divergent fruit morphology, although genetic, physiological and ecological constraints may also play a role.     

Gravity and the evolution of cardiopulmonary morphology in snakes

Physiological investigations of snakes have established the importance of heart position and pulmonary structure in contexts of gravity effects on blood circulation. Here we investigate morphological correlates of cardiopulmonary physiology in contexts related to ecology, behavior and evolution. We analyze data for heart position and length of vascular lung in 154 species of snakes that exhibit a broad range of characteristic behaviors and habitat associations. We construct a composite phylogeny for these species, and we codify gravitational stress according to species habitat and behavior. We use conventional regression and phylogenetically independent contrasts to evaluate whether trait diversity is correlated with gravitational habitat related to evolutionary transitions within the composite tree topology. We demonstrate that snake species living in arboreal habitats, or which express strongly climbing behaviors, possess relatively short blood columns between the heart and the head, as well as relatively short vascular lungs, compared to terrestrial species. Aquatic species, which experience little or no gravity stress in water, show the reverse — significantly longer heart-head distance and longer vascular lungs. These phylogenetic differences complement the results of physiological studies and are reflected in multiple habitat transitions during the evolutionary histories of these snake lineages, providing strong evidence that heart-to-head distance and length of vascular lung are co-adaptive cardiopulmonary features of snakes.     

Functional morphology and evolution of tail autotomy in salamanders

Basal tail constriction occurs in about two-thirds of the species of plethodontid salamanders. The constriction, which marks the site of tail autotomy, is a result of a reduction in length and diameter of the first caudal segment. Gross and microscopic anatomical studies reveal that many structural specializations are associated with basal constriction, and these are considered in detail. Areas of weakness in the skin at the posterior end of the first caudal segment, at the attachment of the musculature to the intermyotomal septum at the anterior end of the same segment, and between the last caudosacral and first caudal vertebrae precisely define the route of tail breakage. During autotomy the entire tail is shed, and a cylinder of skin one segment long closes over the wound at the end of the body. It is suggested that specializations described in this paper have evolved independently in three different groups of salamanders. Experiments and field observations reveal that, contrary to expectations, frequency of tail breakage is less in species with apparent provisions for tail autotomy than in less specialized species. The tail is a very important, highly functional organ in salamanders and it is suggested that selection has been for behavioral and structural adaptations for control of tail loss, rather than for tail loss per se.     

Comparative morphology and evolution of cheek pouches in rodents

There are two types of cheek pouches in extant rodents. Internal cheek pouches are evaginations of the oral cavity deep to M. platysma and M. sphincter colli profundus, and have evolved independently in some species of the superfamilies Sciuroidea and Muroidea. External, furlined cheek pouches open lateral to and separate from the oral cavity, (also deep to M. platysma and M. sphincter colli profundus), and occur in all species of the families Geomyidae and Heteromyidae. The presence of external, furlined cheek pouches is a synapomorphy for the superfamily Geomyoidea. The posterior retractor muscle of the pouch is derived from facial musculature in sciurids, from trapezius musculature in cricetids, and from both facial and trapezius muscle groups in the Geomyoidea. Differences also exist in the musculature associated with the pouch opening. In the Sciuridae and Cricetidae, the M. buccinatorius muscle group acts as a sphincter to control the size of the pouch opening. In the Geomyoidea, the size of the opening is controlled by the M. orbicularis sacculi in concert with a slip of the M. platysma myoides. Thin sections and scanning electron micrographs of the pouch tissue reveal the presence of dermal papillae in Phodopus sungorus but not in a close relative, Mesocricetus auratus. All members of the subfamily Cricetinae have a peninsula of highly folded tissue projecting anteriorly from the posteromedial pouch wall. This folding allows for expansion of the pouch walls when food is stored in the pouch.     

Functional morphology and evolution of the cystoid Echinosphaerites

Echinosphaerites , known from the Lower and Middle Ordovician, has branched biserial brachioles. Such features are so far unknown in the Rhombifera. Echinosphaerites had a skeletal meshwork like that of other blastozoan echinoderms, with a fine outer mesh layer and an inner coarse mesh layer. During evolution the number and location of brachioles, including the pattern of brachiole branching, changed by increase in the number of brachioles and increased complexity of the branching pattern. The exothecal pore structures increased in the complexity of patterns of tangential canals. The pattern of skeletal growth in Echinosphaerites is discussed. The Echinosphaeritidae and Caryocystitidae families are closely related and show parallel development.     

Forelimb skeletal morphology and flight mode evolution in pelecaniform birds

The total length and mid-shaft diameters of wing elements of 50 species of pelecaniform birds were examined to investigate how forelimb skeletal morphology varies with body size and flight mode within this group. Pelecaniforms were assigned to flight mode categories based on primary habitual behaviors (soar, flap–glide, continuous flap). Allometric and discriminant function analyses were conducted on wing element variables in both historical (using independent contrasts) and ahistorical contexts. Results of this study indicate that when phylogenetic relationships are taken into account, only the length of the ulna scales with positive allometry, whereas all other variables exhibit isometry. These results differ from the ahistorical allometric analysis. Discriminant function analysis (DFA) significantly separated the flight mode groups (Wilk's λ=0.002, p<0.00001), with only six individuals from two species (out of n=284) misclassified. Results of historical canonical variates analysis supported the ahistorical DFA and identified two carpometacarpal (CMC) variables as important for separating the flight mode groups: dorsoventral CMC diameter and total CMC length. The carpometacarpus is that portion of the forelimb skeleton that serves as the attachment point for the primary flight feathers, and thus, that portion of the airfoil surface that mediates detailed flight control. Its morphology, more than any other element, reflects differences in flight mode in pelecaniforms. Results of this study indicate that, in pelecaniforms, wing bones generally exhibit isometry (with the exception of the ulna) and do possess specific morphologies reflective of the demands associated with different types of aerial locomotor specialization.     

Sexual selection and the evolution of sperm morphology in sharks

Post‐copulatory sexual selection, and sperm competition in particular, is a powerful selective force shaping the evolution of sperm morphology. Although mounting evidence suggests that post‐copulatory sexual selection influences the evolution of sperm morphology among species, recent evidence also suggests that sperm competition influences variation in sperm morphology at the intraspecific level. However, contradictory empirical results and limited taxonomic scope have led to difficulty in assessing the generality of sperm morphological responses to variation in the strength of sperm competition. Here, we use phylogenetically controlled analyses to explore the effects of sperm competition on sperm morphology and variance in sharks, a basal vertebrate group characterized by wide variation in rates of multiple mating by females, and consequently sperm competition risk. Our analyses reveal that shark species experiencing greater levels of sperm competition produce sperm with longer flagella and that sperm flagellum length is less variable in species under higher sperm competition risk. In contrast, neither the length of the sperm head and midpiece nor variation in sperm head and midpiece length was associated with sperm competition risk. Our findings demonstrate that selection influences both the inter‐ and intraspecific variation in sperm morphology and suggest that the flagellum is an important target of sexual selection in sharks. These findings provide important insight into patterns of selection on the ejaculate in a basal vertebrate lineage.     

Sperm competition and the evolution of gamete morphology in frogs

Despite detailed knowledge of the ultrastructure of spermatozoa, there is a paucity of information on the selective pressures that influence sperm form and function. Theoretical models for both internal and external fertilizers predict that sperm competition could favour the evolution of longer sperm. Empirical tests of the external-fertilization model have been restricted to just one group, the fishes, and these tests have proved equivocal. We investigated how sperm competition affects sperm morphology in externally fertilizing myobatrachid frogs. We also examined selection acting on egg size, and covariation between sperm and egg morphology. Species were ranked according to probability of group spawning and hence risk of sperm competition. Body size, testis size and oviposition environment may also influence gamete traits and were included in our analyses. After controlling for phylogenetic relationships between the species examined, we found that an increased risk of sperm competition was associated with increased sperm head and tail lengths. Path analysis showed that sperm competition had its greatest direct effect on sperm tail length, as might be expected under selection resulting from competitive fertilization. Sperm competition did not influence egg size. Oviposition location had a strong influence on egg size and a weak influence on sperm length, with terrestrial spawners having larger gametes than aquatic spawners. Our analysis revealed significant correlated evolution between egg morphology and sperm morphology. These data provide a conclusive demonstration that sperm competition selects for increased sperm length in frogs, and evidence for evolutionary covariance between aspects of male and female gamete morphology.     

Labyrinth morphology and the evolution of low-frequency phonoreception in elasmobranchs

Labyrinth morphology in extant elasmobranchs (neoselachians: sharks, skates and rays) and several extinct chondrichthyans ranging in age from Pliocene to Devonian is investigated using high-resolution computed tomography (CT scanning) and digital reconstitution techniques. The elasmobranch labyrinth is highly specialized toward low-frequency semi-directional sound detection (LFSDP), optimally around 100 Hz. Several features associated with LFSDP in neoselachians also occur in Mesozoic hybodonts (e.g., Egertonodus, Tribodus) and in some incertae sedis extinct sharks (Acronemus, Tristychius), but are absent in osteichthyans, extant and fossil holocephalans, and certain Paleozoic chondrichthyans (ctenacanths, symmoriiforms, Pucapampella). Thus, LFSDP is regarded as an evolutionary novelty of elasmobranchs that arose some time after their divergence from chimaeroids. The suite of characters associated with LFSDP was probably acquired progressively, some characters being more widely distributed among fossil chondrichthyans than others. LFSDP evolved only within chondrichthyans whose otico-occipital fissure became secondarily closed during ontogeny.     

The evolution of larval morphology and swimming performance in ascidians

The complexity of organismal function challenges our ability to understand the evolution of animal locomotion. To meet this challenge, we used a combination of biomechanics, phylogenetic comparative analyses, and theoretical morphology to examine evolutionary changes in body shape and how those changes affected swimming performance in ascidian larvae. Results of phylogenetic comparative analyses suggest that coloniality evolved at least three times among ascidians and that colonial species have a convergent larval morphology characterized by a large trunk volume and shorter tail length in proportion to the trunk. To explore the functional significance of this evolutionary change, we first verified the accuracy of a mathematical model of swimming biomechanics in a solitary (C. intestinalis) and a colonial (D. occidentalis) species and then ran numerous simulations of the model that varied in tail length and trunk volume. The results of these simulations were used to construct landscapes of speed and cost of transport predictions within a trunk volume/tail length morphospace. Our results suggest that the reduction of proportionate tail length in colonial species resulted in improved energetic economy of swimming. The increase in the size of larvae with the origin of coloniality facilitated faster swimming with negligible energetic cost, but may have required a reduction in adult fecundity. Therefore, the evolution of ascidians appears to be influenced by a trade-off between the fecundity of the adult stage and the swimming performance of larvae.     

Floral nectary morphology and evolution in Pedicularis (Orobanchaceae)

Intricate associations between floral morphology and pollinator foraging behaviour are common. In this context, the presence and form of floral nectaries can play a crucial role in driving floral evolution and diversity in flowering plants. However, the reconstruction of the ancestral state of nectary form is often hampered by a lack of anatomical studies and well‐resolved phylogenetic trees. Here, we studied 39 differentially pollinated Pedicularis spp., a genus with pronounced interspecific variation in colour, shape and size of the corolla. Anatomical and scanning electron microscopy observations revealed two nectary forms [bulged (N = 27) or elongated (N = 5)] or the absence of nectaries (N = 7). In a phylogenetic context, our data suggest that: (1) the bulged nectary should be the ancestral state; (2) nectaries were independently lost in some beaked species; and (3) elongated nectaries evolved independently in some clades of beakless species. Phylogenetic path analysis showed that nectary presence is indirectly correlated with beak length/pollinator behaviour through an intermediate factor, nectar production. No significant correlation was found between nectary type and nectar production, beak length or pollinator behaviour. Some beaked species had nectary structures, although they did not produce nectar. The nectary in beaked species may be a vestigial structure retained during a recent rapid radiation of Pedicularis, especially in the Himalaya–Hengduan Mountains of south‐western China. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 592–607.     

被子植物性系统的多样性、生态功能及分布规律

性系统是被子植物繁育系统的核心, 决定着植物种群的遗传特征、进化方向与速度, 在种群动态、群落结构及生态系统的构建与维持中具有重要意义。本文回顾了被子植物性系统的发展历程及研究方向, 总结了近30年基于性系统研究的前沿科学问题, 包括性系统的多样性和进化、与其他功能性状的生态关联、沿环境梯度的分布格局及变化规律、与群落物种共存机制和群落动态的关系及其对干扰的响应。尽管有关植物性系统的研究已经延伸到生态学领域的诸多方面, 有力地推动了各方面的发展, 但仍有很多值得关注和需要着重研究的方向和问题。本文对未来基于植物性系统的研究方向等提出了展望, 并指出, 在当前全球气候变化背景下, 性系统可作为重要的功能性状应用于指导生物多样性保护和生态系统管理政策的制定。     

Repeated evolution of exaggerated dewlaps and other throat morphology in lizards

The existence of elaborate ornamental structures in males is often assumed to reflect the outcome of female mate choice for showy males. However, female mate choice appears weak in many iguanian lizards, but males still exhibit an array of ornament‐like structures around the throat. We performed a phylogenetic comparative study to assess whether these structures have originated in response to male–male competition or the need for improved signal efficiency in visually difficult environments. We found little evidence for the influence of male–male competition. Instead, forest species were more likely to exhibit colourful throat appendages than species living in open habitats, suggesting selection for signal efficiency. On at least three independent occasions, throat ornamentation has become further elaborated into a large, conspicuously coloured moving dewlap. Although the function of the dewlap is convergent, the underlying hyoid apparatus has evolved very differently, revealing the same adaptive outcome has been achieved through multiple evolutionary trajectories. More generally, our findings highlight that extravagant, ornament‐like morphology can evolve in males without the direct influence of female mate choice and that failure to consider alternative hypotheses for the evolution of these structures can obscure the true origins of signal diversity among closely related taxa.