Trypanoxyuris (Trypanoxyuris) minutus (Schneider, 1866) among Alouatta guariba clamitans (Cabrera, 1940) in the state of Rio de Janeiro,Brazil

This study reports on infection by Trypanoxyuris (Trypanoxyuris) minutus among Alouatta guariba clamitans through biometry on adults and eggs. A total of 58 adult helminths were recovered from the small segment of the animal's large intestine. This study is the first report of this parasite among A. guariba clamitans in Rio de Janeiro.     

A remarkable new genus of Atalophlebiinae (Ephemeroptera: Leptophlebiidae) from the Neotropics

An unusual species of Leptophlebiidae is described based on males, females, and nymphs from Rio de Janeiro State, Brazil. As a consequence of its distinct characteristics on all stages, a new genus is established. The new genus can be distinguished from other South American Leptophlebiid genera mostly by: Adults: vein MA of fore wings asymmetrical; hind wings with costal projection well developed, Sc ending at cross vein near costal projection; tarsal claws dissimilar, one blunt other acute; projections of styliger plate forming two well developed lobes with rounded apex, ventrally obstructing view of the penes; penes fused on basal half, each lobe with a ventral furrow and a long and slender spine directed anteriorly. Nymphs: Head prognathous, wider than labrum; labrum with prominent median emargination, with three subtle crenulations; body flattened; hind wings pads present; tarsal claws with over 20 denticles, subapical denticle much larger than remaining denticles; gills long and narrow, present on abdominal segments I-VII; posterolateral projections present on abdominal segments VIII-IX. Phylogenetic analyses conducted based on a previously published data matrix that included other South America leptophlebiid genera placed Poranga nessimiani gen. nov. et sp. nov. as sister to Bessierus + Perissophlebioides. In most analyses this clade was recovered within the Farrodes complex. Nymphs of the new taxa are particularly similar to Bessierus, whereas adults share the very acute costal projection on hind wings seen in non-dipterous members of the Farrodes complex.     

Pseudascarophis brasiliensis sp. nov. (Nematoda: Cystidicolidae) parasitic in the Bermuda chub Kyphosus sectatrix (Perciformes: Kyphosidae) from southeastern Brazil

A new species of Pseudascarophis (Nematoda: Cystidicolidae) found in the stomach of Kyphosus sectatrix (Linnaeus) (Kyphosidae), off Rio de Janeiro, Brazil, is described. The new species can be differentiated from the other congeners by the presence of lateral alae, distinct but inconspicuous cephalic papillae at the anterior end, three pairs of precloacal and one pair of adcloacal papillae in males, egg morphology and morphometry of glandular oesophagus and spicules. Pseudascarophis tropica is transferred to Ascarophis as Ascarophis tropica (Solov''eva) comb. n. due to its ambiguous diagnosis.     

Long-term study of zooplankton in the estuarine system of Ribeira Bay,near a power plant (Rio de Janeiro,Brazil)

The structure of the zooplankton community in an estuary adjacent to the Admiral álvaro Alberto Nuclear Power Plant at Angra dos Reis, state of Rio de Janeiro, Brazil was studied from 2001 through 2005. At that time, the power plant had been operating for 20 years. The results were compared with a previous study in 1991–1993. The zooplankton was sampled 4 times a year, in vertical hauls using a 150 μm net, at two fixed points near the plant’s intake and discharge sites. Temperature, salinity, and chlorophyll a were measured. The water of Ribeira Bay is warm, with salinities typical of Coastal Water and more saline waters. Zooplankton density at the discharge site (Mean: 245,434 ind m⁻³, SD: 335,358 ind m⁻³) was higher than that at the intake site (Mean: 84,634 ind m⁻³, SD: 101,409 ind m⁻³). A total of 121 mesozooplankton taxa were recorded from 2001 to 2005. Copepoda constituted the most common taxon and comprised more than 57% of the total zooplankton, followed by cladocers and gastropod larvae. A seasonal zooplankton cycle was observed only during 2004; in other years, the plankton varied only between years. Overall mesozooplankton abundance at the discharge site was similar to levels reported from the inner zone of this estuary in 1991–1993. Surface temperature was the important factor structuring the zooplankton community at the discharge site. No effect on the mesozooplankton by passage through the condensers could be discerned, and no permanent negative influence on the plankton populations could be detected. Guest editors: U. M. Azeiteiro, I. Jenkinson & M. J. Pereira Plankton Studies     

Relative growth and morphological sexual maturity of the caridean shrimp Nematopalaemon schmitti (Decapoda: Caridea: Palaemonidae) in an upwelling region in the Western Atlantic*

In crustaceans, successful reproductive processes, such as the transition from juvenile to adult, exhibit important morphological changes that can be detected by analyzing relative growth. This study describes the relative growth of body structures in Nematopalaemon schmitti and its secondary sexual characteristics, and also estimates the morphological sexual maturity of this species in a region influenced by upwelling. The carapace length (CL), second pleuron length (PlL), cheliped carpus length (CaL), cheliped propodus length (PrL) and the length of appendix masculina (AML) of the shrimp were measured. The relationships that best demonstrated the changes in allometric coefficient between demographic categories were AML vs. CL for males, and PlL vs. CL for females. The estimated CL for morphological sexual maturity in males was 8.51 mm and 9.30 in females. Our results showed the appendix masculine and the second pleuron were secondary sexual characteristics that play roles in reaching the morphological sexual maturity necessary for reproductive success and to assure the life cycle of this species.     

Three new species of Parhydraenida Balfour-Browne, 1975 (Coleoptera: Hydraenidae) from southeastern Brazil

ABSTRACT

Three new species of Parhydraenida Balfour-Browne, 1975 are described from southeastern Brazil: P. plesioformis, sp. n.; P. proboscis sp. n.; P. unicornis sp. n. The genus Parhydraenida is recorded from Minas Gerais for the first time, and P. hygropetrica Perkins, 1980 is recorded from Rio de Janeiro for the first time. We describe one secondary sexual character, which enables, for the first time, unambiguous distinction of males and females. Habitus photographs as well as line drawings of the male and female genitalia of all three new species, and of P. hygropetrica are provided.

urn:lsid:zoobank.org:pub:CCA56FB6-897A-4128-B8DD-3892464A8D1B     

Cycles of coral reef ‘turn‐on’, rapid growth and ‘turn‐off’ over the past 8500 years: a context for understanding modern ecological states and trajectories

Human activities threaten reef ecosystems globally, forcing ecological change at rates and scales regarded as unprecedented in the Holocene. These changes are so profound that a cessation of reef accretion (reef ‘turn‐off’) and net erosion of reef structures is argued by many as the ultimate and imminent trajectory. Here, we use a regional scale reef growth dataset, based on 76 core records (constrained by 211 radiometric dates) from 22 reefs along and across the inner‐shelf of the Great Barrier Reef, Australia, to examine the timing of different phases of reef initiation (‘turn‐on’), growth and ‘turn‐off’ during the Holocene. This dataset delineates two temporally discrete episodes of reef‐building over the last 8500 years: the first associated with the Holocene transgression‐early highstand period [～8.5–5.5 k calibrated years bp (cal ybp )]; the second since ～2.3 k cal ybp . During both periods, reefs accreted rapidly to sea level before entering late evolutionary states – states naturally characterized by reduced coral cover and low accretion potential – and a clear hiatus occurs between these reef‐building episodes for which no records of reef initiation exist. These transitions mimic those projected under current environmental disturbance regimes, but have been driven entirely by natural forcing factors. Our results demonstrate that, even through the late Holocene, reef health and growth has fluctuated through cycles independent of anthropogenic forcing. Consequently, degraded reef states cannot de facto be considered to automatically reflect increased anthropogenic stress. Indeed, in many cases degraded or nonaccreting reef communities may reflect past reef growth histories (as dictated by reef growth–sea level interactions) as much as contemporary environmental change. Recognizing when changes in reef condition reflect these natural ‘turn‐on’– growth –‘turn‐off’ cycles and how they interact with on‐going human disturbance is critical for effective coral reef management and for understanding future reef ecological trajectories.     

Two new species of <Emphasis Type="Italic">Calathea</Emphasis> (<Emphasis Type="Italic">Marantaceae</Emphasis>) from South-eastern Brazil

Summary   Calathea dryadica and Calathea reginae are described, circumscribed and illustrated. These new species are probably endemic to the Atlantic Forest of Rio de Janeiro State in Southeast Brazil and are considered critically endangered because of the restricted geographic area of occurrence, sometimes enclosed by densely urbanised areas.     

Towards a ‘Sea‐Level Sensitive’ dynamic model: impact of island ontogeny and glacio‐eustasy on global patterns of marine island biogeography

A synthetic model is presented to enlarge the evolutionary framework of the General Dynamic Model (GDM) and the Glacial Sensitive Model (GSM) of oceanic island biogeography from the terrestrial to the marine realm. The proposed ‘Sea‐Level Sensitive’ dynamic model (SLS) of marine island biogeography integrates historical and ecological biogeography with patterns of glacio‐eustasy, merging concepts from areas as diverse as taxonomy, biogeography, marine biology, volcanology, sedimentology, stratigraphy, palaeontology, geochronology and geomorphology. Fundamental to the SLS model is the dynamic variation of the littoral area of volcanic oceanic islands (defined as the area between the intertidal and the 50‐m isobath) in response to sea‐level oscillations driven by glacial–interglacial cycles. The following questions are considered by means of this revision: (i) what was the impact of (global) glacio‐eustatic sea‐level oscillations, particularly those of the Pleistocene glacial–interglacial episodes, on the littoral marine fauna and flora of volcanic oceanic islands? (ii) What are the main factors that explain the present littoral marine biodiversity on volcanic oceanic islands? (iii) How can differences in historical and ecological biogeography be reconciled, from a marine point of view? These questions are addressed by compiling the bathymetry of 11 Atlantic archipelagos/islands to obtain quantitative data regarding changes in the littoral area based on Pleistocene sea‐level oscillations, from 150 thousand years ago (ka) to the present. Within the framework of a model sensitive to changing sea levels, we discuss the principal factors affecting the geographical range of marine species; the relationships between modes of larval development, dispersal strategies and geographical range; the relationships between times of speciation, modes of larval development, ecological zonation and geographical range; the influence of sea‐surface temperatures and latitude on littoral marine species diversity; the effect of eustatic sea‐level changes and their impact on the littoral marine biota; island marine species–area relationships; and finally, the physical effects of island ontogeny and its associated submarine topography and marine substrate on littoral biota. Based on the SLS dynamic model, we offer a number of predictions for tropical, subtropical and temperate volcanic oceanic islands on how rates of immigration, colonization, in‐situ speciation, local disappearance, and extinction interact and affect the marine biodiversity around islands during glacials and interglacials, thus allowing future testing of the theory.     

The sandfly fauna (Diptera: Psychodidae: Phlebotominae) of the Parque Estadual da Serra da Tiririca,Rio de Janeiro, Brazil

Cutaneous leishmaniasis (CL) in the state of Rio de Janeiro is sporadic and canbe characterised as a peridomestic transmission that occurs in modified naturalenvironments. The aim of this work was to study the fauna and ecologicalcharacteristics of sandflies in an environmentally protected area (the StatePark of Serra da Tiririca) within the remnants of the Atlantic Forest in themunicipalities of Niterói and Maricá and their possible relationship withleishmaniasis. Captures were performed using light traps during the night once amonth for one year in both sylvatic environments and areas surrounding homesnear the park. A total of 1,037 sandflies were captured, belonging to ninegenera and 12 species: Evandromyia tupynambai (34.1%),Migonemyia migonei (20.6%), Brumptomyiacunhai (13.8%), Micropygomyia schreiberi (9.7%),Psathyromyia lanei (6.5%), Brumptomyianitzulescui (5.7%), Evandromyia edwardsi(5.4%), Nyssomyia intermedia (2.8%), Evandromyiacortelezzii (0.6%), Pintomyia bianchigalatiae(0.5%), Lutzomyia longipalpis (0.2%) and Sciopemyiamicrops (0.1%). Both Mg. migonei and Ny.intermedia may be acting as vectors of CL in this area.     

A Quantitative Photographic Survey of ‘Spoke-Burrow’ Type Lebensspuren on the Cape Verde Abyssal Plain

A photographic survey of the seafloor was conducted during RRS Discovery cruise 204 in September/October 1993 at the oligotrophic site of the French EUMELI programme at c. 21° N: 31° W (4650 m) on the Cape Verde Abyssal Plain. Two systems were used, a wide angle system (WASP) obtaining usable photographs covering 10 to 30m², and an epibenthic sledge mounted system obtaining photographs of about 2 m². A total of more than 3000 usable frames were obtained, covering about 64000 m² of seafloor. The photographs indicated an epibenthic invertebrate megafaunal density of 5.2 hectare⁻¹ and a demersal fish density of 1.8 hectare ^−1. In contrast, the photographs were dominated by lebensspuren, particularly of the ‘spoke-burrow’ type usually attributed to echiurans. Spoke-burrow density, where related burrows and other features such as sediment mounds are considered unitary, is estimated at 871 hectare ^−1. We describe and quantify the five principal types of spoke burrow encountered in this study and, on the basis of these observations and some morphometric studies of the lebensspuren, discuss the possible behaviour of their producers. We have no direct knowledge of the production rate of, or longevity of, lebensspuren in the study area. However, based on a number of assumptions and limited data from other areas we suggest that trace producer (mainly echiurans) density is on the order of tens of individuals per hectare. These trace producers probably greatly outnumber the rest of the invertebrate megabenthos, and we can only guess at the role this potential large population of hidden benthos might have in the overall structure and function of the deep-sea floor ecosystem.     

Evolutionary history of the Maltese wall lizard Podarcis filfolensis: insights on the ‘Expansion–Contraction’ model of Pleistocene biogeography

The expansion–contraction (EC) model predicts demographic and range contraction of temperate species during Pleistocene glaciations as a consequence of climate‐related habitat changes, and provides a paradigm for explaining the high intraspecific diversity found in refugia in terms of long‐term demographic stability. However, recent evidence has revealed a weak predictive power of this model for terrestrial species in insular and coastal settings. We investigated the Pleistocene EC dynamics and their evolutionary consequences on temperate species using the Maltese archipelago and its endemic lizard Podarcis filfolensis as a model system. The evolutionary and demographic history of P. filfolensis as inferred from mitochondrial and nuclear sequences data does not conform to the EC model predictions, supporting (i) demographic and spatial stability or expansion, rather than contraction, of the northern and southern lineages during the last glacial period; and (ii) a major role for allopatric differentiation primed by sea‐level dynamics, rather than prolonged demographic stability, in the formation of the observed genetic diversity. When combined with evidence from other Mediterranean refugia, this study shows how the incorporation of Pleistocene sea‐level variations in the EC model accounts for a reverse demographic and range response of insular and coastal temperate biotas relative to continental ones. Furthermore, this cross‐archipelago pattern in which allopatric diversity is formed and shaped by EC cycles resembles that seen between isolated populations within mainland refugia and suggests that the EC model, originally developed to explain population fluctuations into and out‐of refugia, may be appropriate for describing the demographic and evolutionary dynamics driving the high genetic diversity observed in these areas.     

Phylogeny of the suborder Psocomorpha: congruence and incongruence between morphology and molecular data (Insecta: Psocodea: ‘Psocoptera’)

The largest suborder of bark lice (Insecta: Psocodea: ‘Psocoptera’) is Psocomorpha, which includes over 3600 described species. We estimated the phylogeny of this major group with family‐level taxon sampling using multiple gene markers, including both nuclear and mitochondrial ribosomal RNA and protein‐coding genes. Monophyly of the suborder was strongly supported, and monophyly of three of four previously recognized infraorders (Caeciliusetae, Epipsocetae, and Psocetae) was also strongly supported. In contrast, monophyly of the infraorder Homilopsocidea was not supported. Based on the phylogeny, we divided Homilopsocidea into three independent infraorders: Archipsocetae, Philotarsetae, and Homilopsocidea. Except for a few cases, previously recognized families were recovered as monophyletic. To establish a classification more congruent with the phylogeny, we synonymized the families Bryopsocidae (with Zelandopsocinae of Pseudocaeciliidae), Calopsocidae (with Pseudocaeciliidae), and Neurostigmatidae (with Epipsocidae). Monophyly of Elipsocidae, Lachesillidae, and Mesopsocidae was not supported, but the monophyly of these families could not be rejected statistically, so they are tentatively maintained as valid families. The molecular tree was compared with a morphological phylogeny estimated previously. Sources of congruence and incongruence exist and the utility of the morphological data for phylogenetic estimation is evaluated. © 2014 The Linnean Society of London     

Phylogenetic analysis of genera of ‘Akanthophoreinae’ (Crustacea: Tanaidacea)

Phylogenetic relations among tanaidacean genera within ‘Akanthophoreinae’ are addressed using computer-assisted parsimony methods. The morphology-based analysis includes 10 well-defined and described genera: Araphura, Chauliopleona, Collettea, Paragathotanais, Metagathotanais, Paraleptognathia, Paranarthrura, Portaratrum, and Tanaella in Tanaidomorpha, and Glabroapseudes in Apseudomorpha as the outgroup. Chauliopleona and Paraleptognathia form a monophylum; Portaratrum cannot be placed in any known family. These three genera are considered as incertae sedis. The analysis does not support the monophyly of ‘Akanthophoreinae’, and further questions the monophyly of Tanaellidae.     

Diversification of a ‘great speciator’ in the Wallacea region: differing responses of closely related resident and migratory kingfisher species (Aves: Alcedinidae: Todiramphus)

The Collared Kingfisher species complex is the most widespread of the ‘great speciator’ lineages of the Indo‐Pacific. They have shown a remarkable ability to spread and diversify. As a result of this rapid diversification, Todiramphus species are often found in secondary sympatry. In Southeast Sulawesi, Indonesia, two Todiramphus species are present, the breeding resident Collared Kingfisher Todiramphus chloris and the overwintering migratory Sacred Kingfisher Todiramphus sanctus. We investigated the effect of isolation on these closely related species by comparing their populations on mainland Sulawesi and its larger continental islands, with populations on the small, oceanic Wakatobi Islands. Within our wider analysis we provide further support for the distinctiveness of the Sulawesi Collared Kingfisher population, perhaps isolated by the deep water barrier of Wallace's line. Within Sulawesi we found that populations of Collared Kingfisher on the Wakatobi Islands had diverged from those on mainland Sulawesi, differing both in morphology and in mitochondrial DNA. In contrast, there was no divergence between Sacred Kingfisher populations in either morphology or mitochondrial DNA. We propose that a difference in habitat occupied by Collared Kingfisher populations between the mainland and continental islands vs. oceanic islands has caused this divergence. Mainland Collared Kingfishers are predominately found inland, whereas Wakatobi Collared Kingfishers are also found in coastal habitats. The larger body size of Wakatobi Collared Kingfisher populations may be a result of increased competition with predominantly coastal Sacred Kingfisher populations. The uniform nature of Sacred Kingfisher populations in this region probably reflects their consistent habitat choice (coastal mangrove) and their migratory nature. The demands of their breeding range are likely to have an even stronger selective influence than their Sulawesi wintering range, limiting their scope for divergence. These results provide insight into the adaptability of the widespread Todiramphus lineage and are evidence of the need for further taxonomic revision of Collared Kingfisher populations.     

Theraphosidae phylogeny: relationships of the ‘Ischnocolinae’ genera (Araneae,Mygalomorphae)

The mygalomorph spider subfamily ‘Ischnocolinae’ was originally established as a group based on the presence of divided tarsal scopula. Later, the divided condition of the scopula was considered the plesiomorphic state, which could not support the monophyly of ‘Ischnocolinae’. In Raven 1985, the subfamily was considered paraphyletic, pending a phylogenetic analysis to reinvestigate monophyletic groups. This study comprises such a phylogenetic analysis, based on morphological data, that includes representatives of all genera currently included in ‘Ischnocolinae’ as well as representatives of all other nine Theraphosidae subfamilies (Thrigmopoeinae, Ornithoctoninae, Eumenophorinae, Stromatopelminae, Harpactirinae, Selenogyrinae, Theraphosinae, Aviculariinae and Selenocosmiinae). The family Theraphosidae is considered monophyletic and expanded to include three additional genera previously considered as possible Barychelidae, namely Brachionopus (as Harpactirinae), Trichopelma and Reichlingia (as Ischnocolinae sensu stricto) while ‘Ischnocolinae’ as previously defined does not appear as monophyletic. However, two monophyletic groups were defined as subfamilies to include some former ‘Ischnocolinae’ representatives. The first group includes Acanthopelma rufescens, Trichopelma nitidum, Reichlingia annae, Ischnocolus spp., Holothele rondoni, Holothele culebrae and Holothele aff culebrae and is hereby named as Ischnocolinae (sensu stricto). The other subgroup comprises Sickius longibulbi, Holothele incei, Holothele aff incei, Guyruita spp., Schismatothele lineata, Hemiercus modestus, Holothele colonica and Holothele sp., together established as Schismatothelinae subfam. nov. Several genera included in former ‘Ischnocolinae’ appear as monophyletic (Catumiri, Oligoxystre, Heterothele, Nesiergus, Chaetopelma, Ischnocolus, Guyruita and Plesiophrictus). However, the genera Holothele, Schismatothele and Hemiercus deserve more attention in order to evaluate their intrarelationships and inclusion of species.     

Phylogeny and revision of the ‘Cubitermes complex’ termites (Termitidae: Cubitermitinae)

The phylogeny of the genus Cubitermes Wasmann was reconstructed using two mitochondrial genes (COI and COII) and a fragment of the nuclear 28S rDNA, including samples of Apilitermes Holmgren, Crenetermes Silvestri, Megagnathotermes Silvestri and Thoracotermes Wasmann. Based on our analyses, we recovered these genera within a paraphyletic Cubitermes clade. Cubitermes species are distributed between five main clades, highly reflective of their enteric valve armatures (EVAs). Consequently, Cubitermes is here divided into five monophyletic genera based on phylogenetic analyses and EVAs: Cubitermes (sensu stricto), Isognathotermes Sjöstedt, Nitiditermes Emerson, Polyspathotermes Josens & Deligne, gen.n. , and Ternicubitermes Josens & Deligne, gen.n. Moreover, the two species of Megagnathotermes included in this study, exhibiting different EVAs, are phylogenetically distant: M. katangensis Sjöstedt belongs in the genus Polyspathotermes, while M. notandus Silvestri remains as a monotypic genus. During the evolution of the Cubitermes complex, sclerotized EVA spatulae appeared twice independently: three or six spatulae in Polyspathotermes (sulcifrons and oblectatus patterns) and two jaw‐like spatulae in Nitiditermes (sankurensis pattern), which is absent in some species (oculatus pattern). Zoobank registration: http://zoobank.org/urn:lsid:zoobank.org:pub:576AAFF8‐63C6‐4962‐A8F5‐75C9317AEA7B .     

The Feeding Behaviour of Starlings (Sturnus vulgaris) in the Presence of ‘Eyes’

Previous studies that have looked at the aversive properties of simple, predator-related, eye-like patterns presented in an artificial context, have yielded conflicting results. The aim of the present series of experiments was to investigate whether such stimuli had potential for use as bird scarers. The starling (Sturnus vulgaris) was the subject species. The experimental procedure was designed to provide a relevant and objective, interval scale of aversiveness based upon the ability of eye patterns to deter hungry birds from a feeding area. Each subject was deprived of food for 5 h and then tested in an apparatus which exposed an eye pattern over a food trough immediately after the bird alighted on that trough. 10 measures of the starlings' feeding behaviour were monitored over a 1-h trial. It was predicted that five of these variables would be positively and five negatively correlated with an increase in the fear evoking properties of the stimulus. The 10 measures were analysed using principal component analysis and the first component in every experiment had the signs of the latent vectors of the variables exactly in the predicted pattern and therefore appeared to be an ‘aversiveness index’. The scores from this first component were then used in a Latin square ANOVA to distinguish subject, test order and stimulus effects. The main findings from the eight experiments are as follows. Simple eye-like patterns can indeed deter hungry starlings from feeding in their vicinity. The presence of a pair of eyes painted on a white card caused an 88% reduction in time spent on the nearby food trough and a 65% reduction in actual feeding time. The presence of ‘pupils’ in the patterns is essential for simple circles alone are not significantly more aversive than the control card. Once a pattern has the ‘pupil/iris’ distinction then a circular outline becomes important although the shape of the ‘pupil’ appears relatively unimportant. Eyes with coloured irises are more effective than black and white patterns but the degree of contrast between the ‘pupil’ and ‘iris’ appears irrelevant. Increasing the number of simultaneously presented eyes from one to three is correlated with an increasing trend in aversiveness which is destroyed if the same stimuli are presented within the context of a simple head outline. Changing the orientation of a pair of eyes from horizontal to vertical only slightly reduces their aversiveness. The size of the patterns appears to be unimportant within the range tested. The aversive properties of eyespots and of broadcast starling distress calls are positively additive. The effectiveness of a stimulus combining all the important features mentioned is as aversive as is a realistic model of the head of an owl.