Intra‐specific host preference for Littoraria filosa by the dipteran parasitoid Sarcophaga megafilosia: the consequences of attacking individuals outside the preferred size range

1. The marine snail, Littoraria filosa, lives on the leaves and trunks of mangroves. Populations of L. filosa are annual when its dipteran parasitoid Sarcophaga megafilosia is present, but perennial in its absence. 2. The size distribution of naturally occurring parasitised L. filosa suggests S. megafilosia has a distinct preference for snails with shells ≥ 10 mm long but, paradoxically, most individuals ≥ 18 mm long disappear from annual populations in late spring/early summer without appearing in the parasitised cohort. 3. When S. megafilosia was offered an unrestricted size range of L. filosa, only individuals >10 mm long were attacked and those from 19 to 20 mm long were preferred. 4. In no‐choice experiments, L. filosa <10 mm long were parasitised but pupation did not occur. For L. filosa >23 mm long, most attacked snails succeeded in ejecting the parasitoid larva but often lost their grip on the wall or ceiling of the cage and fell. Littoraria filosa is unlikely to survive such dislodgement from mangrove trees. 5. These results can explain the different size distributions of live and parasitised snails in the field, including the disappearance of large snails that do not subsequently appear in the parasitised cohort. However, this does not explain the preference by S. megafilosia for L. filosa around 19–20 mm for which more than 50% of attacks failed. This preference may be also determined by increased parasitoid fecundity and longevity as snail size increases. 6. If small differences in host size affect parasitoid fitness, it may help explain why attempts at biological control using known parasitoids of an exotic species are often ineffective or unsuccessful.     

Epipharynx shape as a tool to reveal differentiation patterns between insect sister species: insights from Onthophagus taurus and O. illyricus (Insecta: Coleoptera: Scarabaeidae)

We evaluated the potential of the adult epipharynx to reveal interspecific differentiation patterns in closely related Onthophagus species. Although easy to analyse, this structure has received little attention in Scarabaeidae taxonomy, probably because a qualitative inspection of its shape often does not yield differences between congeneric species. The polyphenic sister species Onthophagus taurus Schreber, 1759 and O. illyricus Scopoli, 1763 were chosen as a case study. They are extremely similar, to the extent that in some cases they cannot be identified unambiguously without the help of biomolecular analysis. In this study, a combination of linear measurements and a landmark-based approach was employed to quantify inter- and intraspecific shape variation in the epipharynx of specimens sampled at the same study site. Our results showed that the epipharynx is a monomorphic structure: its shape does not vary as a function of sex or male phenotype. In males, epipharynx shape does not change with head shape or horn length. The close proximity of the epipharynx to the horns and the synchronous developmental patterns of these two structures suggest that a developmental trade-off may act between them. Despite these predisposing conditions, however, our results suggest that epipharynx size is not subject to costs associated with horn development, and that the trait is highly canalised. Surprisingly, when using geometric morphometrics the epipharynx appears to be a better tool than genitalia for discriminating between the two sister species.     

Reproductive bionomics and life history traits of three gammaridean amphipods, Cymadusa filosa Savigny, Ampithoe laxipodus Appadoo and Myers and Mallacoota schellenbergi Ledoyer from the tropical Indian Ocean (Mauritius)

The reproductive bionomics and life history traits of two corophiid amphipods (Ampithoe laxipodus, Cymadusa filosa) and one melitid (Mallacoota schellenbergi) were studied in Mauritius (Indian Ocean) for the period March 1999 to February 2000. Results on the population structure, monthly size class variations, sex ratio, female reproductive states and fecundity are presented. The study demonstrates multivoltinism and continuous reproduction in the three species. Increase in number of juveniles was observed in warmer months for C. filosa and A. laxipodus. Sexual maturity was attained at smaller sizes in warmer months in the three species. Linear relationship on body length and number of eggs in brood pouch are presented. Size-independent analysis of egg number revealed a decrease in number of eggs in cooler months. Sex ratio is male skewed in M. schellenbergi and female skewed in C. filosa and A. laxipodus. Some of the plausible explanations for the reproductive strategies adopted by these three species in a tropical system are discussed.     

Environment and space as drivers of variation in skull shape in two widely distributed South‐American Tayassuidae,Pecari tajacu and Tayassu pecari (Mammalia: Cetartiodactyla)

The influence of the environment on the geographical variation of morphological traits has been recognized in a number of taxa. Pecari tajacu and Tayassu pecari are ideal models to investigate intraspecific geographic variation in skull because of their wide and heterogeneous geographical distribution in South America. We used geometric morphometric procedures to examine the geographical variation in skull shape of 294 adult specimens of these species from 134 localities. We quantified to what extent skull shape variation was explained by environment, skull size and geographical space using variation partitioning analysis. We detected a strong pattern of geographic variation for P. tajacu skull shape, but not for T. pecari. The environment seems to be the major selective force that drives skull shape variation in both species. Nevertheless, other spatially structured processes (e.g. genetic drift, gene flow) might also have affected variation in the skull shape of the more widespread species P. tajacu. Allometric relationships might reflect the biomechanical constraints that are thought to be strong enough to limit size‐related changes in T. pecari skull shape.     

The effects of two parasitoids on the life history and metapopulation structure of the intertidal snail Littoraria filosa in different-sized patches of mangrove forest

We describe differences in life history of the intertidal arboreal snail, Littoraria filosa, among patches of mangroves ranging in size from isolated trees to large stands several square kilometres in area. Recruitment of L. filosa occurred from mid spring (October) to early winter (June), recruits grew rapidly and copulating adults were found during the following September–April. Populations within large patches of forest were annuals; all or most individuals died between October–January (spring–midsummer). In contrast, those in smaller peripheral patches were more likely to survive over the summer but survival differed among patches and years. These differences in life history were caused by a parasitoid fly (genus Sarcophaga) that attacked L. filosa 10 mm and longer and was present in all large patches, but absent from, or rare, in smaller peripheral patches. Experimental introductions to isolated trees confirmed that the fly could kill L. filosa. Another sarcophagid parasitoid that attacked L. filosa from 4 to less than 10 mm long was also found in every patch. The combined effects of these parasitoids appear to determine the metapopulation structure of L. filosa. Most adults in large patches were killed by the larger fly during early summer. Summer recruits were often killed by the smaller fly within a month of settlement and when this happened effective recruitment of L. filosa was reduced to autumn. The planktotrophic larval stage of L. filosa lasts less than 1 month, so the source of autumn recruits to all patches must have been adults that survived the early summer, most of which were in small patches or on isolated trees. Consequently these ”peripheral sources” are likely to be important for persistence of the metapopulation of L. filosa. The results of this study demonstrate that metapopulation structure may be determined by complex interactions and that common models cannot be assumed to apply in all habitats. Received: 15 September 1999 / Accepted: 31 January 2000     

Males Resemble Females: Re-Evaluating Sexual Dimorphism in Protoceratops andrewsi (Neoceratopsia,Protoceratopsidae)

BackgroundProtoceratops andrewsi (Neoceratopsia, Protoceratopsidae) is a well-known dinosaur from the Upper Cretaceous of Mongolia. Some previous workers hypothesized sexual dimorphism in the cranial shape of this taxon, using qualitative and quantitative observations. In particular, width and height of the frill as well as the development of a nasal horn have been hypothesized as potentially sexually dimorphic.Conclusions/SignificanceSexual dimorphism within Protoceratops andrewsi is not strongly supported by our results, as previously proposed by several authors. Anatomical traits such as height and width of the frill, and skull size thus may not be sexually dimorphic. Based on PCA for a data set focusing on the rostrum and associated ANOVA results, nasal horn height is the only feature with potential dimorphism. As a whole, most purported dimorphic variation is probably primarily the result of ontogenetic cranial shape changes as well as intraspecific cranial variation independent of sex.     

Morphological variation in vanishing Mexican desert fishes of the genus Characodon (Goodeidae)

This study investigated a rapidly vanishing group of fishes in the genus Characodon (including Characodon lateralis and Characodon audax) from the upper Rio Mezquital drainage in Mexico. Using specimens from museum collections, morphological variation was assessed to quantify body shape differentiation among historic (i.e. extirpated and extant) collection sites. In both sexes, body shape (particularly head shape, the proportion of the caudal peduncle and the position and size of dorsal and anal fins) varied significantly among populations and species. Variation among collection sites could at least partially be attributed to geography, as the presence of distinct hydrographic units and a major waterfall coincided with major body shape differences. These results are discussed in the light of previously published molecular genetic analyses, as they have direct implications for taxonomic problems and the need for conservation measures for these endangered fishes.     

Geometric craniometric analysis of sexual dimorphism and ontogenetic variation: A case study based on two geographically disparate species, Aethomys ineptus from southern Africa and Arvicanthis niloticus from Sudan (Rodentia: Muridae)

Non-geographic morphometric variation, particularly at the level of sexual dimorphism and ontogenetic (age-related) variation, has been documented in rodents, and useful for establishing whether to analyse sexes separately or together, and for selecting adult specimens for subsequent data recording and analysis. However, such studies have largely been based on traditional morphometric analyses of linear measurements that mainly focus on overall size, rather than shape-related morphometric variation. Unit-free, landmark/outline-based geometric morphometric analyses are considered to offer a more appropriate tool for assessing shape-related morphometric variation. In this study, we used geometric cranial morphometric analysis to assess the nature and extent of sexual dimorphism and age variation within the Tete veld rat, Aethomys ineptus (Thomas and Wroughton, 1908) from southern Africa and the African Nile rat, Arvicanthis niloticus (Desmarest, 1822) from Sudan. The results obtained were in turn compared with previously published results based on independent geometric and traditional cranial morphometric data from the same sampled populations examined in the present study. While our geometric morphometric results detected statistically significant sexual dimorphism in cranial shape within Ar. niloticus only, previously published results based on traditional morphometric data failed to detect significant sexual dimorphism within this species. However, similar to previously published traditional morphometric data, our geometric morphometric results detected statistically significant age-related variation in cranial shape and size within both Ae. ineptus and Ar. niloticus, with individuals of age classes 5 and 6 being considered to represent adult specimens. Our results highlight the importance of carefully evaluating both size- and shape-related non-geographic morphometric variation prior to the analysis of geographic variation and the delineation of species. Erroneous conclusions of non-geographic variation may have implications in the interpretation of geographic and evolutionary processes that may be responsible for morphological differences at both the inter- and intra-specific levels.     

New species of Peliosanthes,Rohdea and Tupistra (Asparagaceae) from Laos and Vietnam

We describe and illustrate four new species, Peliosanthes choriandra, P. tatianae and Tupistra orlovii from central to northern Laos, and Rohdea filosa from northern Vietnam. These are all very local in distribution and endemic to the respective countries. We also report new localities and their ecological conditions for five other species of Peliosanthes (P. argenteostriata, P. hirsuta, P. irinae, P. micrantha and P. nivea) recently described from Laos and/or Vietnam. Furthermore, Peliosanthes nivea is recorded as new to the flora of Laos.     

Morphological assessment of the Cambrian trilobites Oryctocephalus indicus (Reed 1910) from China and Oryctocephalus ‘reticulatus’ (Lermontova 1940) from Siberia

Oryctocephalus indicus is a widespread trilobite species described from the lower–middle Cambrian (provisional Cambrian Stage 5) of south China, North Korea, India, USA and possibly North Greenland. Principal component analysis (PCA) and thin‐plate splines (TPSs) based on a pooled sample of 359 specimens from four sections in the Guizhou Province, south China, demonstrated that the shape of this species is very constant, and only a small difference is detected in specimens from deeper part of the basin. Most of the variation is taphonomically controlled, and only the specimens from Sanwan Section show a modest natural difference due to a slightly different depositional environment. A comparison between the Chinese specimens and Oryctocephalus ‘reticulatus’ from the Molodo Section, Siberia, has been carried out. PCA and TPS show that O. ‘reticulatus occupies the same morphospace of O. indicus. However, the canonical variates analysis and the analysis of similarities detect small differences between the Siberian and Chinese populations, indicating a minor geographical variation caused by different environmental settings. The ontogenic variation is also described from early meraspids to holaspids. The most substantial changes take place during the first period of the early meraspid development, in which the glabella changes from a bell‐like shape to a cylindrical shape. The ocular ridge changes from poorly defined in early meraspid to well developed in late merapids. The results indicate that O. ‘reticulatus’ is a junior synonym of O. indicus, suggesting that the base O. indicus of Siberia can be correlated with the O. indicus Zone of south China.     

Seasonal variations of opisthaptoral hard parts of Gyrodactylus salaris Malmberg, 1957 (Monogenea: Gyrodactylidae) on parr of Atlantic salmon Salmo salar L. in the River Batnfjordselva,Norway

Seasonal variations in size and shape of the marginal hooks, the anchors and the ventral bar of the opisthaptor of Gyrodactylus salaris Malmberg, 1957 were studied. The G. salaris specimens were collected from parr of Atlantic salmon Salmo salar L. in the River Batnfjordselva, north-western Norway. Samples were taken at roughly monthly intervals during a two-year period. The marginal hooks, the anchors and the ventral bar showed considerable seasonal variation in size, but varied only slightly in shape. The variation in 12 of the 14 characters measured showed a significant regression to the variation in the water temperature. The total length of the marginal hooks of G. salaris can be considerably longer than the previously reported maximum of 40 m for the species.     

Shape Variation in the Dermatocranium of the Greater Short-Horned Lizard <Emphasis Type="Italic">Phrynosoma hernandesi</Emphasis> (Reptilia: Squamata: Phrynosomatidae)

Dermatocranial shape and horn morphology display great disparity among the species of Phrynosoma. Ontogenetic change in dermatocranial shape in a series of 79 specimens of the short-horned Phrynosoma hernandesi (54F: 25M) was examined using geometric morphometric techniques. A multivariate ANCOVA of Procrustes residuals with sex as a factor and ln(centroid size) as the covariate indicated sexual shape dimorphism. Separate multivariate regressions of Procrustes residuals on ln(centroid size) for each sex indicated that allometry accounts for ~52–54% of the total sample shape variance. Comparisons of ontogenetic shape change between sexes indicate that sexual shape dimorphism is minimal and of uncertain biological significance. Groupings of multivariate regression coefficients by magnitude and sign suggest that allometric integration of the dermatocranium is not uniform over the dermatocranium. Principal component analysis of the landmark configurations corrected for sex and allometry yields a first principal component which describes shape variance concentrated in the posterolateral and posterior regions of the dermatocranium, and again is indicative of non-uniform shape variation over the dermatocranium. Our findings for P. hernandesi indicate that the adult shape of the dermatocranium may contribute to a passive defence against predation. We hypothesize that the complexity in dermatocranial shape demonstrated here for P. hernandesi indicates parcellation of shape variance, which may contribute to explanations of the pronounced dermatocranial disparity exhibited by the species of Phrynosoma.     

New data about the functional morphology of the chaetiferous leech‐like annelids Acanthobdella peledina (Grube, 1851) and Paracanthobdella livanowi (Epshtein, 1966) (Clitellata,Acanthobdellida)

Approximately 130 individuals of Acanthobdella peledina and 100 individuals of Paracanthobdella livanowi were studied. Morphometric measurements were taken to explore the body form. The digestive and the reproductive systems of leech‐like annelids were analyzed for the first time in such a high number of specimens. Observation of A. peledina and P. livanowi revealed crucial differences in the reproductive system of the analyzed taxa, mostly regarding variation in the shape of the testisacs and the length of the ovisacs. The results of a digestive system analysis suggest that blood‐sucking parasites of the order Acanthobdellida may also lead a predatory lifestyle. The presented findings support the taxonomic division of the order Acanthobdellida into the families Acanthobdellidae and Paracanthobdellidae. Multiple similarities between Acanthobdellida and Hirudinida were also discussed. J. Morphol. 275:528–539, 2014. © 2013 Wiley Periodicals, Inc.     

Morphological and molecular concordance of Rhynchozoon clades (Bryozoa, Cheilostomata) from Alaska

Abstract. Alpha‐level taxonomy in the bryozoan order Cheilostomata relies almost exclusively on hard‐part morphology. Geographical, ecophenotypic, and intracolony variation often make it difficult to distinguish intra‐ from interspecific variation and to recognize taxonomically informative characters. DNA sequences provide a source of data independent of morphology by which to gauge the relative reliability of various morphological characters for taxonomy. We present a case study involving a limited number of specimens of Rhynchozoon sp. from Ketchikan, Alaska to show the utility of DNA data in identifying genetic lineages for subsequent morphological analysis. The study illustrates that the use of genetic data need not involve massive, broad‐scale phylogenetic studies to address problems in invertebrate α‐level taxonomy. Phylogeny reconstruction with a 430‐bp fragment of the 16S mitochondrial ribosomal RNA gene showed two moderately diverged clades, here termed Rhynchozoon clades A and B, separated by an average genetic distance of 2.38% (K2P+Γ). Comparison of voucher specimens by scanning electron microscopy showed two congruent, morphologically distinct forms (forms A and B, respectively) distinguishable by a polythetic suite of characters including degree of frontal costation, range of spine number, number of beads on the primary orifice, number of areolar pores, and peri‐orificial sculpturing. Orifice shape and ovicell form proved not to be good diagnostic characters. The status of the two forms as biological species is unclear, although maintenance of distinct suites of morphological characters in the two mitochondrial lineages suggests they may be reproductively isolated from one another. For Rhynchozoon form B, which tends to have a highly costate frontal wall, we suggest a resurrection of the name Rhynchozoon tumulosum, which had been previously synonymized with R. rostratum. Rhynchozoon form A may be conspecific with “Rhynchozoon sp. A” previously reported from Washington state.     

Cranial shape varies along an elevation gradient in Gambel's white‐footed mouse (Peromyscus maniculatus gambelii) in the Grinnell Resurvey Yosemite transect

Environmental variation over a species's range creates differing pressures to which organisms must adjust in order to survive. Taxa can respond to these pressures at population and individual levels, leading to localized phenotypic differentiation. Assessing the spatial distribution of phenotypic variation can illuminate how dramatically varying environmental factors shape phenotypes and may forecast a taxon's ability to adapt should conditions change. We characterized morphological variation along a transect sampled in the Grinnell Resurvey project to determine whether Gambel's white‐footed mouse (Peromyscus maniculatus gambelii), a generalist taxon inhabiting the full elevational range of habitats in Yosemite National Park and surrounding areas, has responded morphologically to variation in its environment. We quantified variation in modern P. m. gambelii cranial shape using 2D generalized Procrustes analysis and Euclidean distance matrix‐based geometric morphometrics. We performed multivariate regression of shape coordinates on elevation to test for environmental influences on shape within the principal geographic dimension of change along the transect. We observe a statistically significant correlation with shape on elevation for occlusal and lateral views of the cranium, explaining a small percentage of the overall variation in shape. Modern P. m. gambelii crania show a pattern of flexion in which the angle of the cranial base decreases at higher elevations. Results of EDMA parallel these findings, but highlight additional areas of the cranium that vary with elevation. Collectively, the patterns of variation detected suggest a biological response to the environment that warrants further study. This work lays the foundation for comparison with morphological data from historical specimens, which can address evolutionary scenarios generated from our findings, and for investigation of other taxa included in the resurvey project. J. Morphol. 271:897–909, 2010. © 2010 Wiley‐Liss, Inc.     

The family Myriotrochidae Théel (Echinodermata: Holothurioidea) in the deep northeast Atlantic Ocean

Recent sampling in the Rockall Trough, Porcupine Seabight and Porcupine Abyssal Plain, in the NE Atlantic, has yielded 200 specimens of apodous holothurians belonging to seven species of the family Myriotrochidae Théel from depths between 1000 and 4310 m. These include the type species of a new genus and two new species of existing genera. Parvotrochus belyaevi gen. et sp. nov . is described from some minute specimens from the Rockall Trough that possess both exceptionally small wheel-like deposits and numerous, large curved rods in the body wall and tentacles. Myriotrochus clarki sp. Nov . has affinities to M. vitreus (M. Sars) but shows differences in the shape of the plates of the calcareous ring and in having larger, frequently abnormally formed, wheels. Siniotrochus myriodontus sp. Nov . is similar to the type species of this genus, S. phoxus, Pawson, but differs in the arrangement of the teeth on the wheels. A single small specimen of Prototrochus Belyaev et Mironov from the Whittard Canyon, northern Bay of Biscay, is similar to P. minutus (Östergren), a species known only from the Sea of Japan, but differs in some characteristics of the wheels and tentacles. A new subspecies, P. zenkevitchi rockallensis subsp. nov. , is proposed for a form of P. zenkevitchi (Belyaev) that was the most common myriotrochid encountered. P. zenkevitchi was known previously only from a few localities in the Pacific and S Atlantic deep-sea trenches. Similarly, two species previously known only from the NE Pacific, Myriotrochus bathybius H. L. Clark and M. giganteus H. L. Clark, are also recorded from the N Atlantic for the first time. The synonymy of M. giganteus and M. sp. ex gr. macquariensis-giganteus Belyaev et Mironov is proposed. The greatest number both of specimens and species came from the areas most intensively sampled. The present records show that several species have world-wide distributions. The wide geographic separation of many records is the result of both poor sampling effort in the deep sea, particularly with fine-meshed gear, and the difficulties in sampling infaunal animals in the deep sea. Box core samples in the Rockall Trough suggest that the myriotrochids are more common in this area than would be supposed from epibenthic sledge data. Hence, myriotrochids may be more prevalent in the deep sea than previously thought.     

Morphometric convergence among European sand gobies in freshwater (Gobiiformes: Gobionellidae)

The five genera of sand gobies inhabit the seas and freshwaters of Europe and western Asia and occupy habitats ranging from fully marine to exclusively freshwater. In this study, we use geometric morphometrics to quantify body shape among sand gobies, in order to investigate how shape has evolved and how it is related to habitat. We also compare body shape between preserved museum specimens and fresh specimens, to determine whether or not fixation and storage in ethanol introduce detectable bias. We confirm that the fixed specimens exhibit significant shape changes as compared to fresh specimens, and so, we perform the bulk of our analyses exclusively on fixed specimens. We find that Economidichthys, Orsinigobius, and Pomatoschistus occupy distinct regions of morphospace. Knipowitschia and Ninnigobius have intermediate forms that overlap with Pomatoschistus and Orsinigobius, but not Economidichthys. This pattern is also in rough accordance with their habitats: Pomatoschistus is fully marine, Economidichthys fully freshwater, and the others fresh with some brackish tolerance. We augment a recent phylogeny of sand gobies with data for P. quagga and interpret morphometric shape change on that tree. We then evaluate convergence in form among disparate lineages of freshwater species by constructing a phylomorphospace and applying pattern‐based (convevol) measures of convergence. We find that freshwater taxa occupy a mostly separate region of morphospace from marine taxa and exhibit significant convergence in form. Freshwater taxa are characterized by relatively larger heads and stockier bodies than their marine relatives, potentially due to a common pattern of heterochronic size reduction.