Musculature of the head and cervical region in several palaearctic species from the superfamilies Cossoidea,Zygaenoidea, and Sesioidea (Lepidoptera,Cossiformes)

Musculature of the head and cervical region in eight lepidopteran species from the families Cossidae, Sesiidae, Limacodidae, and Zygaenidae (Cossiformes) was studied. Besides, musculature of the anterior cervical region in Brachodes appendiculata Esper (Brachodidae) is discussed. The data on musculature are analyzed and compared with the authors’ earlier results on a relatively primitive ditrysian moth from the family Tineidae (Korzeev, 2004). The absence of the labro-epipharyngeal musculature in Cossiformes is confirmed and the presence of a new, indirectly acting secondary labral extensors CMx.5 in Zygaenidae is revealed. These secondary extensors are strongly developed in Papilionomorpha. Cossiformes and Tineidae have been shown to possess several common plesiomorphic traits in the structure of labrum, while the dilatators of salivarium within Cossiformes are reduced in number from two pairs to a single pair. The present study shows that the musculature of the cervical region has remained relatively stable in the evolution of Lepidoptera despite the loss of some muscles in different taxa. The groundplan of the head musculature is reconstructed for Glossata, Papilionomorpha, Cossiformes, and the “higher” Lepidoptera from the Geometriformes-Noctuiformes complex of superfamilies. In addition, the groundplan of the musculature of the cervical region is depicted for Myoglossata, Papilionomorpha, Cossiformes, and the Geometriformes-Noctuiformes complex of superfamilies.     

Extremely miniaturised and highly complex: The thoracic morphology of the first instar larva of Mengenilla chobauti (Insecta,Strepsiptera)

Thoracic structures of the extremely small first instar larva of the strepsipteran species Mengenilla chobauti (ca. 200 μm) were examined, described and reconstructed 3-dimensionally. The focus is on the skeletomuscular system. The characters were compared to conditions found in other insect larvae of very small (Ptiliidae) or large (Dytiscus) size (both Coleoptera) and features of “triungulin” larvae, first instar larvae of Rhipiphoridae, Meloidae (both Coleoptera), and Mantispidae (Neuroptera).The specific lifestyle and the extreme degree of miniaturisation result in numerous thoracic modifications. Many sclerites of the exo- and endoskeleton are reduced. Cervical sclerites, pleural ridges, furcae and spinae are absent. Most of the longitudinal muscles are connected within the thorax, and a pair of ventral longitudinal muscles is present in the pleural region of the meso- and metathorax. This results in a high intersegmental flexibility. Due to the size reduction and the correlated shift of the brain to the thorax, with 94 identified muscles the thoracic musculature appears highly compact. Compared to larger larvae the number of both the individual muscles and the muscle bundles are distinctly reduced. The thorax of the first instar larvae displays many additional strepsipteran autapomorphies. At least partly due to the highly specialised condition, potential synapomorphies with other groups were not found.     

Observations and comments on the reliability of muscle reconstruction in fossil vertebrates

In Canis and Ursus the largest proportion of attachments of muscles of the shoulder and brachium on the scapula and humerus is direct; fewer attachments are aponeurotic or tendinous. In both genera most attachments can be associated with superficial osteological features (scars or delimitable surfaces); attachments that lack such features are direct. Most aponeurotic attachments are associated with rugose scarring whereas tendinous attachments are often associated with smooth surfaces. Although most attachments can be associated with osteological features the areal extent of attachment is often not inferrable from the bone. The inference of muscle size or functional significance from osteological features is problematic. The amount of myological information that can be deciphered from the osteology in Canis and Ursus is greater than that reported for particular members of other vertebrate groups which suggests that there may be differences in the degree to which muscles can be reconstructed from superficial osteology alone. Nonetheless, even in mammals such as the Carnivora, detailed muscular reconstructions in extinct taxa cannot be achieved without reference to the musculature of extant relatives. Such reconstructions rely on assumptions, that often have not been adequately tested, regarding the similarity of musculature in closely related taxa. This testing and well corroborated hypotheses of phylogenetic relationship are essential for the evaluation of the accuracy of reconstructions of the musculature in fossil vertebrates.     

Patterns of musculature as taxonomic characters for the Turbellaria Acoela

While turbellarians are generally assumed to have body-wall musculature consisting routinely of longitudinal, circular, and diagonal fibers, members of the Acoela examined by a fluorescence-microscopy technique specific for actin showed more complicated and distinctive arrangements of muscles, giving promise for better delimiting taxa within this taxonomically difficult order. Certain globose or tear-drop-shaped worms such as Convoluta pulchra and species of Pseudaphanostoma, Mecynostomum, and Otocelis, showed a complex pattern in which muscles longitudinal in the anterior half of the body arc diagonally across the posterior half; complex brushes of parenchymal muscles that cross at the level of the statocyst and arc postero-laterally also characterize these groups. The more elongate acoel Paratomella sp. was found to have musculature dominated by strictly longitudinal fibers and with relatively weak circular fibers and few fibers running diagonally to the body axis, yet the elongate mecynostomid Paedomecynostomum bruneum showed a crossing of antero-longitudinal fibers similar to that seen in the more globose Mecynostomum sp. A distinctive looping of muscles around the mouth is seen in P. bruneum and the Anaperidae. Such similarities and differences in pattern of musculature promise to provide easily recognizable characters for taxonomy of the Acoela at levels ranging from species to family. This revised version was published online in July 2006 with corrections to the Cover Date.     

Comparative analysis of the intrinsic leg musculature of the American cockroach,Periplaneta americana (L.)

The American cockroach has a total of 368 muscles inserting on the post-coxal segments of its legs. By using a narrow morphological definition for delimiting individual muscles, it is shown (i) that the protrochanteral musculatures (23 muscles/leg) differ from the essentially identical meso- and metatrochanteral musculatures (24 and 26 muscles/leg) in number and disposition of extensors and in having a completely different flexor composition, and (ii) that the musculatures of the more distal segments of the legs are completely serially homologous, there being 2 muscles for moving each femur, 23 for each tibia, 7 for each first tarsomere, and 5 for each of the paired pretarsal claws. In all six legs, the trochanteral and tibial musculatures each contain single slender muscles that may be acting proprioceptively to measure the angular displacements between, respectively, the coxas and trochanters, and the femurs and tibias. Neurological and phylogenetic considerations are used to demonstrate why a narrow morphological definition should be employed, and why the widely used functional definition of Snodgrass ('35) is not only fallacious on evolutionary grounds, but also leads to making erroneous conclusions regarding the manner in which insect musculature is controlled by the insect central nervous system. Finally, it is hypothesized that the physiological limitations imposed by having an open circulatory system and the problems inherent in the neural control of large muscles may have been major evolutionary factors in forcing insects to use many slender muscles to control their body movements.     

长白山苔原带昆虫群落组成与时间动态

【目的】苔原是一种极地景观类型,本研究调查了长白山苔原带昆虫的物种组成与时间动态,以期为苔原带昆虫保护以及昆虫与苔原植物协同进化研究提供基础依据。【方法】2005-2007年每年6-9月,在长白山北坡、西坡苔原带利用网捕、灯诱、巴氏罐法采集昆虫标本。【结果】2005-2007年从长白山苔原带共获得昆虫标本4 634头,隶属于11目105科550种,其中鳞翅目、鞘翅目、膜翅目、双翅目的种类和个体数量较为丰富,这4个类群的物种数与个体数的最高峰都出现在7月份。对应分析显示,鳞翅目昆虫对7月份,鞘翅目昆虫对8和9月,膜翅目和双翅目昆虫对6月的适应性较强。苔原带7月份昆虫的物种数(382种)、个体数(2 571个个体)和多样性指数(4.673)都最高,物种数和个体数在9月份最低,仅22种265个个体。不同月份昆虫的物种数与个体数之间呈显著正相关(R=0.992)。不同月份间昆虫的相似性低(相似性系数＜0.20)。【结论】长白山苔原带昆虫多样性较低,7月为昆虫的活跃高峰期,9月昆虫的个体数和物种数均较少。鳞翅目昆虫的物种最丰富,对7月份气侯的适应性相对较强,鞘翅目对季节变化的适应能力强于其他类群,在维持苔原带生态平衡中起着重要作用。     

Walking on inclines: how do desert ants monitor slope and step length

Background

In order to increase the weak database concerning the organogenesis of Acoela – a clade regarded by many as the earliest extant offshoot of Bilateria and thus of particular interest for studies concerning the evolution of animal bodyplans – we analyzed the development of the musculature of Symsagittifera roscoffensis using F-actin labelling, confocal laserscanning microscopy, and 3D reconstruction software.

Results

At 40% of development between egg deposition and hatching short subepidermal fibres form. Muscle fibre development in the anterior body half precedes myogenesis in the posterior half. At 42% of development a grid of outer circular and inner longitudinal muscles is present in the bodywall. New circular muscles either branch off from present fibres or form adjacent to existing ones. The number of circular muscles is higher than that of the longitudinal muscles throughout all life cycle stages. Diagonal, circular and longitudinal muscles are initially rare but their number increases with time. The ventral side bears U-shaped muscles around the mouth, which in addition is surrounded by a sphincter muscle. With the exception of the region of the statocyst, dorsoventral muscles are present along the entire body of juveniles and adults, while adults additionally exhibit radially oriented internal muscles in the anterior tip. Outer diagonal muscles are present at the dorsal anterior tip of the adult. In adult animals, the male gonopore with its associated sexual organs expresses distinct muscles. No specific statocyst muscles were found. The muscle mantles of the needle-shaped sagittocysts are situated along the lateral edges of the animal and in the posterior end close to the male gonopore. In both juveniles and adults, non-muscular filaments, which stain positively for F-actin, are associated with certain sensory cells outside the bodywall musculature.

Conclusion

Compared to the myoanatomy of other acoel taxa, Symsagittifera roscoffensis shows a very complex musculature. Although data on presumably basal acoel clades are still scarce, the information currently available suggests an elaborated musculature with longitudinal, circular and U-shaped muscles as being part of the ancestral acoel bodyplan, thus increasing the possibility that Urbilateria likewise had a relatively complicated muscular ground pattern.     

Extrinsic Snout Musculature in Afrotheria and Lipotyphla

As currently recognized, the mammalian order Lipotyphla contains six extant families: Chrysochloridae, Erinaceidae, Solenodontidae, Soricidae, Talpidae, and Tenrecidae. Although most mammalogists have accepted this taxon, the morphological support for Lipotyphla is relatively weak, and recent phylogenetic studies using molecular data have concluded that it is not monophyletic. Instead, these molecular studies place chrysochlorids and tenrecids in the proposed clade Afrotheria, together with aardvarks, elephants, elephant shrews, hyraxes, and sirenians. Despite strong molecular support, Afrotheria has received little or no morphological support. It was recently suggested that a mobile snout might be a morphological feature uniting afrotherians. To test this proposal, I dissected the extrinsic snout musculature in an assortment of lipotyphlan and afrotherian mammals. These muscles provide support for Lipotyphla but not for Afrotheria. The snout is moved by different muscles in different afrotherian taxa, suggesting that the mobile snout is not homologous across different afrotherian lineages. In contrast, lipotyphlans have a distinctive set of five snout muscles moving the snout tip that appears to be unique to these six families. In addition, in soricids and talpids, four of the five snout muscles originate posterior to the zygomatic arch, supporting sister-taxon status for these two lineages. Although the extrinsic snout musculature does not support Afrotheria as presently proposed, it is consistent with an Afrotheria that does not include chrysochlorids and tenrecids.     

Variation in the trunk musculature of caecilians (Amphibia: Gymnophiona)

Variation in the trunk musculature of 28 species of caecilians. representing 24 of the 33 genera and all five families. is summarized. All forms examined have the same muscles in similar positions. Existing variation largely conforms to the current classification of the group. and some variation may be attributable to different modes of locomotion. such as burrowing versus swimming. Caecilian trunk musculature is more similar to that of salamanders than to that of frogs. but the similarity is probably syrnplesiomorphous. Trunk musculature so far has provided no clues to lissamphibian relationships.     

Musculature in sipunculan worms: ontogeny and ancestral states

SUMMARY Molecular phylogenetics suggests that the Sipuncula fall into the Annelida, although they are morphologically very distinct and lack segmentation. To understand the evolutionary transformations from the annelid to the sipunculan body plan, it is important to reconstruct the ancestral states within the respective clades at all life history stages. Here we reconstruct the ancestral states for the head/introvert retractor muscles and the body wall musculature in the Sipuncula using Bayesian statistics. In addition, we describe the ontogenetic transformations of the two muscle systems in four sipunculan species with different developmental modes, using F-actin staining with fluorescent-labeled phalloidin in conjunction with confocal laser scanning microscopy. All four species, which have smooth body wall musculature and less than the full set of four introvert retractor muscles as adults, go through developmental stages with four retractor muscles that are eventually reduced to a lower number in the adult. The circular and sometimes the longitudinal body wall musculature are split into bands that later transform into a smooth sheath. Our ancestral state reconstructions suggest with nearly 100% probability that the ancestral sipunculan had four introvert retractor muscles, longitudinal body wall musculature in bands and circular body wall musculature arranged as a smooth sheath. Species with crawling larvae have more strongly developed body wall musculature than those with swimming larvae. To interpret our findings in the context of annelid evolution, a more solid phylogenetic framework is needed for the entire group and more data on ontogenetic transformations of annelid musculature are desirable.     

Effects of stream size on taxa richness and other commonly used benthic bioassessment metrics

Benthic macroinvertebrate samples were collected from natural substrates in disturbed and undisturbed South Carolina upper coastal plain streams to determine if taxa richness and other bioassessment metrics were significantly related to stream size as predicted by the River Continuum Concept (RCC). Linear, quadratic, and lognormal regression models indicated that stream width was positively related to total number of taxa; number of Ephemeroptera, Plecoptera, and Trichoptera (EPT) taxa; and total number of organisms. Linear regression showed that the expected number of taxa at undisturbed sites ranged from 35 in 2.0 m wide streams to 64 in 16.0 m streams. Comparable values were 8–20 for EPT and 109–261 for number of organisms. Stream width was inversely related to biotic index values indicating a decrease in average organism tolerance with increasing stream size. ANCOVA showed that the effects of stream size were similar for disturbed and undisturbed sites. Rank correlations and multidimensional scaling (MDS) showed that Lepidoptera and Trichoptera were more abundant in larger streams and Annelida in smaller streams. Stream size related changes in benthic macroinvertebrate community composition are often ignored in bioassessment protocols; however, failure to adjust metrics for stream size can lead to erroneous conclusions. Adjustments are possible by analyzing regression residuals stripped of stream size related variance, dividing the area beneath the maximum taxa richness line into equal size units for metric scoring, or scaling metrics based on predicted reference values. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

The lantern of Aristotle: organization of its coelom and origin of its muscles (Echinodermata,Echinoida)

Summary Comparative ultrastructural analyses of the muscles that work the lantern of Aristotle support the opinion that the muscles in question are myoepithelially organized or derivatives of myoepithelia. They are part of the epithelium of the peripharyngeal cavity (=lantern coelom). The coelom epithelium may become multiplelayered in certain regions and is composed of (1) a layer of muscle cells that vary in number and size, (2) nerve cells and their processes that are interspersed between the muscle layer and (3) monociliated adluminal cells that build a continuous cell lining and completely cover the muscle layer. According to their complexity, the lantern muscles exhibit consecutive stages of myoepithelial variations and may finally simulate subepithelial musculature. The results of this study support the hypothesis of a histological development of subepithelial musculature from simple myoepithelia, although both epithelial and mesenchymal musculature may occur in the Echinodermata. Detailed knowledge of the organization of the lantern's coelom space was a prerequisite for the present study. In contrast to previous examinations the lantern coelom is not a continuous space, but is subdivided into several cavities that are partially completely separated from each other. On the one hand, this subdivision is probably caused by the sophisticated arrangement of the lantern's ossicles and on the other by the septa that give rise to muscles that fulfill different functions. lanter's ossicles and on the other by the septa that give     

The musculature of Squatinella rostrum (Milne, 1886) (Rotifera: Lepadellidae) as revealed by confocal laser scanning microscopy with additional new data on its trophi and overall morphology

Wilts, E.F., Wulfken, D., Ahlrichs, W.H. and Martínez Arbizu, P. 2012. The musculature of Squatinella rostrum (Milne, 1886) (Rotifera: Lepadellidae) as revealed by confocal laser scanning microscopy with additional new data on its trophi and overall morphology.—Acta Zoologica (Stockholm) 93 : 14–27. The monogonont rotifer Squatinella rostrum was investigated with light, scanning electron and confocal laser scanning microscopy to reveal new morphological data on its inner and outer anatomy. In total, the visualized somatic musculature displays five paired longitudinal muscles (musculi longitudinales I–V) and nine circular muscles (musculi circulares I–IX). Compared to other species, S. rostrum is characterized by the absence of several longitudinal and circular muscles (e.g. musculus longitudinalis capitis, corona sphincter and pars coronalis). A reconstruction of the mastax musculature revealed a total number of seven paired and two unpaired mastax muscles. Possibly homologous somatic and mastax muscles in other, thus far investigated rotifers are discussed. Moreover, we provide a phylogenetic evaluation of the revealed morphological characters and suggest possible autapomorphic characters supporting Squatinella and Lepadellidae. Finally, we refer to some striking similarities in the morphology, ecology and way of movement of Squatinella and Bryceella that may indicate a closer relationship of both taxa.     

Evolution of the skeleton and musculature of the male genitalia in the family lycaenidae (Lepidoptera): II. Infratribe Polyommatina Swainson, 1827

Skeleton and musculature of valvae were examined in 19 species from the section Polyommatus (Lepidoptera, Lycaenidae) revised by Bálint and Johnson (1997). Functional organization of male genitalia in the section Polyommatus was shown to be relatively constant. Significant interspecific variation of the skeleton and muscles was found only in the structure of valvae. During their evolution, the valvae became elongated, the costal torsion and longitudinal medial fold appeared on their medial wall, the subcostal groove was formed, and the intravalval muscle differentiated into two muscles forming a chiasma. This muscular chiasma was formed at least twice in the evolution of the group, in Polyommatus Latreille, 1804 + Aricia [Reichenbach], 1819 and in Plebeius Kluk, 1802. The genus Chilades Moore, [1881] is the least advanced as concerns the functional organization of the valvae. The genera Polyommatus and Aricia are the most advanced and constitute a sister pair. The intermediate position is occupied by Plebeius Kluk, 1802 and probably Madeleinea Bálint, 1993. The genus Albulina Tutt, 1809 is a compound taxon, its species occupying an intermediate position between Polyommatus and Aricia on the one hand, and Plebeius on the other hand. With a sole exception, they are characterized by a plesiomorphic state of the valval musculature that does not allow their mutual phylogenetic relations to be evaluated consistently. The only examined species of the genus Agriades Hübner, [1819], A. glandon (De Prunner, 1798), also possesses undifferentiated musculature, but according to the characters of the vaginal area, its females are close to the branch Polyommatus + Aricia.     

Musculature of two bdelloid rotifers, Adineta ricciae and Macrotrachela quadricornifera: organization in a functional and evolutionary perspective

Organization of muscles in microinvertebrates has often been studied to answer functional questions and understand phylogenetic relationships among taxa. In this study, the musculature of two bdelloid species, Adineta ricciae and Macrotrachela quadricornifera , was illustrated, and their organization was compared with other rotifer taxa to generate possible hypotheses of evolutionary relationships among Rotifera. The two species share a common habitat but differ from each other in feeding and locomotion. A. ricciae feeds on the biofilm by scraping it, is unable to swim, and slides on the head cilia using the foot to propel over the substratum. M. quadricornifera feeds by filtration, can swim, and advances by looping in a leech-like motion. Their musculature, stained with TRITC-phalloidin, was observed using confocal laser scanning microscopy. Major differences between the two species were observed in the muscles of head and foot, possibly reflecting differences in their life style. Muscles of the trunk were similarly arranged: circular muscles surrounded longitudinal bands, which were inserted at different points on the body wall. In both bdelloids, circular muscles of the trunk were incomplete ventrally, a condition also present in Seison and in soft-bodied monogononts from benthos. Within rotifers, circular muscles in the form of complete rings are present in acanthocephalans and in soft-bodied planktonic monogononts but are absent in loricate monogononts, which generally possess dorsoventral bands. The diversity of muscle organization among rotifers was interpreted and discussed.     

A quantitative analysis of the Eutherian orbit: correlations with masticatory apparatus

The mammalian orbit, or eye-socket, is a highly plastic region of the skull. It comprises between seven and nine bones, all of which vary widely in their contribution to this region among the different mammalian orders and families. It is hypothesised that the structure of the mammalian orbit is principally influenced by the forces generated by the jaw-closing musculature. In order to quantify the orbit, fourteen linear, angular and area measurements were taken from 84 species of placental mammals using a Microscribe-3D digitiser. The results were then analysed using principal components analysis. The results of the multivariate analysis on untransformed data showed a clear division of the mammalian taxa into temporalis-dominant forms and masseter-dominant forms. This correlation between orbital structure and masticatory musculature was reinforced by results from the size-corrected data, which showed a separation of the taxa into the three specialised feeding types proposed by Turnbull (1970): i.e. 'carnivore-shear', 'ungulate-grinding' and 'rodent-gnawing'. Moreover, within the rodents there was a clear distinction between species in which the masseter is highly developed and those in which the temporalis has more prominence. These results were reinforced by analysis of variance which showed significant differences in the relative orbital areas of certain bones between temporalis-dominant and masseter-dominant taxa. Subsequent cluster analysis suggested that most of the variables could be grouped into three assemblages: those associated with the length of the rostrum; those associated with the width of the skull; and those associated with the relative size of the orbit and the shape of the face. However, the relative area of the palatine bone showed weak correlations with the other variables and did not fit into any group. Overall the relative area of the palatine was most closely correlated with feeding type, and this measure that appeared to be most strongly associated with the arrangement of the masticatory musculature. These results give a strong indication that, although orbital structure is in part determined by the relative size and orientation of the orbits, the forces generated by the muscles of mastication also have a large effect.     

Sexual dimorphism in snake tail length: sexual selection,natural selection,or morphological constraint?

Male snakes typically have longer tails relative to body length than females, but the extent of this dimorphism varies among species. Three hypotheses have been suggested to explain tail dimorphism. The Morphological Constraint Hypothesis proposes that males have relatively longer tails to accommodate hemipenes and retractor muscles. The Female Reproductive Output Hypothesis proposes that females have relatively shorter tails as a secondary result of natural selection for increased reproductive capacity. The Male Mating Ability Hypothesis proposes that sexual selection favours relatively longer tails in males during courtship. These hypotheses make different predictions about the relationships among tail length, body size, male reproductive morphology, female reproductive output, mode of reproduction, and male mating behaviour among and within taxa. Predictions were tested using published data for 56 genera in the family Colubridae and original data for the water snake, Nerodia sipedon. Tail length dimorphism was more male-biased in tam having relatively short tails (r=–0.52, P < 0.001), hemipenes and retractor muscles occupied a greater proportion of the tail in taxa having relatively short tails (r=– 0.71, P < 0.00l and r=– 0.66, P = 0.001, respectively), and tail length dimorphism was more male-biased in taxa in which body size dimorphism was more female-biased (r=– 0.60, P < 0.001). These results support both the Morphological Constraint Hypotheses and the Female Reproductive Output Hypothesis. However, tests of other predictions, including those regarding patterns within N. sipedon , failed to support any of the three hypotheses. Comparisons among taxa suggest several species in which further tests of these hypotheses would be especially appropriate.     

Three-dimensional reconstruction of the naupliar musculature and a scanning electron microscopy atlas of nauplius development of Balanus improvisus (Crustacea: Cirripedia: Thoracica)

An atlas of the naupliar development of the cirripede Balanus improvisus Darwin, 1854 using scanning electron microscopy (SEM) is provided. Existing spikes on the hindbody increase in number with each moult and are an applicable character for identification of the different nauplius stages, as is the setation pattern of the first antennae. The naupliar musculature of B. improvisus was stained with phalloidin to visualise F-actin, followed by analysis using confocal laser scanning microscopy (CLSM) with subsequent application of 3D imaging software. The larval musculature is already fully established in the first nauplius stage and remains largely unchanged during all the six nauplius stages. The musculature associated with the feeding apparatus is highly elaborated and the labrum possesses lateral muscles and distal F-actin-positive structures. The alimentary tract is entirely surrounded by circular muscles. The extrinsic limb musculature comprises muscles originating from the dorsal and the ventral sides of the head shield, respectively. The hindbody shows very prominent postero-lateral muscles that insert on the dorso-lateral side of the head shield and bend towards ventro-posterior. We conclude that the key features of the naupliar gross anatomy and muscular architecture of B. improvisus are important characters for phylogenetic inferences if analysed in a comparative evolutionary framework.     

Evolution and homologies of primate and modern human hand and forearm muscles, with notes on thumb movements and tool use

In this paper, we explore how the results of a primate-wide higher-level phylogenetic analysis of muscle characters can improve our understanding of the evolution and homologies of the forearm and hand muscles of modern humans. Contrary to what is often suggested in the literature, none of the forearm and hand muscle structures usually present in modern humans are autapomorphic. All are found in one or more extant non-human primate taxa. What is unique is the particular combination of muscles. However, more muscles go to the thumb in modern humans than in almost all other primates, reinforcing the hypothesis that focal thumb movements probably played an important role in human evolution. What makes the modern human thumb myology special within the primate clade is not so much its intrinsic musculature but two extrinsic muscles, extensor pollicis brevis and flexor pollicis longus, that are otherwise only found in hylobatids. It is likely that these two forearm muscles play different functional roles in hylobatids and modern humans. In the former, the thumb is separated from elongated digits by a deep cleft and there is no pulp-to-pulp opposition, whereas modern humans exhibit powerful thumb flexion and greater manipulative abilities, such as those involved in the manufacture and use of tools. The functional and evolutionary significance of a third peculiar structure, the intrinsic hand structure that is often called the ‘interosseous volaris primus of Henle’ (and which we suggest is referred to as the musculus adductor pollicis accessorius) is still obscure. The presence of distinct contrahentes digitorum and intermetacarpales in adult chimpanzees is likely the result of prolonged or delayed development of the hand musculature of these apes. In relation to these structures, extant chimpanzees are more neotenic than modern humans.     

Interrelation between the flower structure and composition of the pollinator groups for Dipsacaceae and Asteraceae with externally similar anthodia

The competitive relations between members of phylogenetically distant plant families Asteraceae (Centaurea and Cirsium) and Dipsacaceae (Knautia and Succisa) with purple anthodia, sharing a common wide range of pollen vectors and competing for them, were studied. The composition of pollen vectors is somewhat different in different plant species. Only bumble-bees, the most effective pollinators, were observed visiting every studied plants species. Syrphidae flies, Lepidoptera, Coleoptera, and some other insects were also observed in different proportions. The principal importance for pollination of the corolla tube size, correlating with the size of insect mouthparts, and the additional importance of particular traits of the inflorescence are confirmed. Convergent similarity of the aspect of anthodia in two species of different families is shown to be based on different structural and functional features. Insect pollinators are the factor of anthodia convergence. The plant species studied are divided into the following three groups, according to the proportion of bumble-bees among pollen vectors and to the range of species-specific pollinators; species coadapted to one pollinator taxon; species coadapted to two or three pollinator taxa; and species coadapted to many pollinator taxa. Asteraceae species in general (with the exception for Cirsium arvense) are characterized by constant contacts with a narrower range of pollinators than Dipsacaceae species (and Cirsium arvense), characterized by wider range of pollinators. Among flowering plants with similar anthodia, the tighter structural coadaptations of Asteraceae with their effective pollinators provide their greater competitive ability as compared to Dipsacaceae.