Northeastern Coast Of Brazil Research Articles

Apolar Compounds in Seaweeds from Fernando de Noronha Archipelago (Northeastern Coast of Brazil)

Hyphenated techniques of gas chromatography coupled to mass spectrometer were used to determine fatty acids in eleven species of seaweeds from Fernando de Noronha archipelago. The main compounds detected in all studied species were the alcohol phytol and the fatty acids 14 : 0; 15 : 0; 16 : 0; 18 : 0; 18 : 1 n9; 18 : 2 Δ9,12; 20 : 4; 20 : 5. These fatty acids are commonly found in seaweeds present in warm regions. Thus, we found no specificity in the presence of a particular set of fatty acids and the studied species indicating that they are not useful as taxonomic indicators. However, they could be used in a comparative study with algae found in polluted area because many of the studied seaweeds are widespread and Fernando de Noronha has low human influence.

The Genus Herposiphonia (Ceramiales, Rhodophyta) in the Coral Reefs Environmental Protection Area, Northeastern Brazil, with new records for Brazil and the Atlantic Ocean

This paper presents a taxonomic study of taxa of the red algae genus Herposiphonia (Ceramiales) occurring on Maracajau Reef in the Coral Reefs Environmental Protection Area (CREPA - Area de Protecao Ambiental dos Recifes de Corais) in Rio Grande do Norte State, along the northeastern coast of Brazil. The CREPA comprises coastline and continental shelf areas of the municipalities of Touros, Rio do Fogo, and Maxaranguape and includes sand dunes, coastal lagoons, and the adjacent shoreline and offshore reefs. Detailed morphological studies were made, considering recent taxonomic criteria for species delimitation of Herposiphonia, and five species were identified: H. delicatula, H. nuda, H. parca, H. secunda, and H. tenella, thus increasing the number of species in the genus from three to six. Herposiphonia delicatula and H. parca represent new occurrences for Brazil, and H. nuda is reported for the first time for the Atlantic Ocean.

Satellite-tracking reveals multiple foraging strategies and threats for olive ridley turtles in Brazil

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 443:237-247 (2011) - DOI: https://doi.org/10.3354/meps09427 Satellite-tracking reveals multiple foraging strategies and threats for olive ridley turtles in Brazil Augusto C. C. D. da Silva1,*, Erik A. P. dos Santos1, Fábio L. das C. Oliveira2, Marilda I. Weber2, Jamyle A. F. Batista2, Thiago Z. Serafini3, Jaqueline C. de Castilhos2 1Centro Nacional de Conservação, Pesquisa e Manejo das Tartarugas Marinhas, ReBio Santa Isabel, 49190-000, Pirambu, SE, Brazil 2Fundação Centro Brasileiro de Proteção e Pesquisa das Tartarugas Marinhas, 49035-485, Aracaju, SE, Brazil 3Programa de Pós-Graduação em Meio Ambiente e Desenvolvimento - MADE, Universidade Federal do Paraná—UFPR, 80035-050, Curitiba, PR, Brazil *Email: cesar@tamar.org.br ABSTRACT: The state of Sergipe in northeastern Brazil is the largest nesting area for olive ridley turtles along this nation’s coast, and constitutes a major rookery in the western Atlantic as well. Conservation efforts with a focus on nesting activities have been implemented there since 1982, but little is known about other aspects of the life cycle, specifically post-nesting movements of females and the locations of foraging grounds. To address this issue, satellite transmitters were deployed on 10 females that nested between February and April 2006. The turtles were monitored for an average of 113 d (range: 14 to 297 d), and an average movement of 1669 km (range: 407 to 4265 km) was recorded. Of the 10 turtles monitored, 6 moved along the Brazilian continental shelf to neritic foraging areas. Five of these turtles utilized foraging areas along the northern and northeastern coasts of Brazil, while one foraged along the southeastern coastline. Two females were tracked to equatorial oceanic waters, with one first moving to an inshore foraging site where she remained for 34 d before migrating to oceanic waters off the Brazilian coast. Signal transmission of 3 of the 10 turtles tracked ceased during their post-nesting migrations, preventing identification of their feeding areas. Olive ridley turtles nesting on the coast of Sergipe displayed a range of post-nesting movements including to coastal sites along the continental shelf as well as offshore oceanic areas. Inter-nesting habitats, migration routes and foraging grounds showed great overlap with a variety of coastal fisheries, as well as with longline fishing in oceanic waters, a key consideration for developing conservation strategies for this species in the western Atlantic. KEY WORDS: Satellite tracking · Inter-nesting · Migration · Foraging grounds · Lepidochelys olivacea · Conservation · Brazil Full text in pdf format PreviousNextCite this article as: da Silva ACCD, dos Santos EAP, Oliveira FLdC, Weber MI, Batista JAF, Serafini TZ, de Castilhos JC (2011) Satellite-tracking reveals multiple foraging strategies and threats for olive ridley turtles in Brazil. Mar Ecol Prog Ser 443:237-247. https://doi.org/10.3354/meps09427 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 443. Online publication date: December 20, 2011 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2011 Inter-Research.

Detection of paramyxoviruses in Magellanic penguins (Spheniscus magellanicus) on the Brazilian tropical coast

Aquatic migratory birds are a major vectors by which influenza viruses and paramyxoviruses are spread in nature. Magellanic penguins (Spheniscus magellanicus) are usually present on the southern shores of South America and can swim as far as the southern coast of Brazil in winter. In 2008, however, several Magellanic penguins were observed on the northeastern coast of Brazil. Paramyxoviruses were isolated from Magellanic penguins on the Espírito Santo state coast, approximately 4000km from their breeding colonies, although influenza viruses were not detected. Among the paramyxoviruses, five Avulavirus isolates belonging to serotype APMV-2 and the serotype APMV-10, which was proposed by Miller et al. (2010), were identified. These results highlight the risks associated with the spread of paramyxoviruses between natural to non-natural habitats by birds exhibiting unusual migration patterns, and they document for the first time the presence of the APMV-2 and APMV-10 serotypes on penguins in Brazil. The local avifauna may become infected with these viruses through close contact between migratory and resident birds. Continued surveillance of virus incidence in these migratory populations of penguins is necessary to detect and prevent the potential risks associated with these unusual migration patterns.

First record of Flabellina dana Millen and Hamann, 2006 (Mollusca: Nudibranchia) in the South Atlantic Ocean

The nudibranch Flabellina dana Millen and Hamann, 2006 is reported from two localities in the northeastern Brazilian coast. These are the first records of this species, previously recorded from localities in the Bahamas and the Caribbean Sea, in South Atlantic Ocean, extending its known geographic distribution more than 3500 km southward.

Migratory flows and foraging habitat selection by shorebirds along the northeastern coast of Brazil: The case of Baía de Todos os Santos

Large numbers of Nearctic shorebirds migrate and winter along the coast of northeastern Brazil, but there is little information on their migratory flows, foraging ecology, and on the structure of the species assemblages that they form with resident shorebirds. We studied these issues on intertidal flats of Baía de Todos os Santos (Bahia), the second largest bay in Brazil. During a full year cycle we carried out weekly bird counts in an intertidal area of 280 ha divided in sectors, where we also measured environmental parameters. The analyses of weekly counts resulted in a detailed phenology of use of the area by shorebirds. Five species were resident and ten were Nearctic migrants. Several of the latter had clear peaks in numbers in March and October, revealing the use of the bay as a stopover during both the north-bound and south-bound migration flows. A canonical correspondence analysis of the relationship between environmental parameters and bird numbers indicated that the foraging bird assemblage could be divided into five main groups, occupying distinct ecological gradients in the study area. The most important factors driving this structure were invertebrate prey abundance, percentage of fine sediments, area of mangrove cover and distance to channels. Our findings imply that maintenance of the diversity of intertidal habitats in this bay is crucial to satisfy the particular habitat requirements of resident and migrant shorebirds using the northeastern coastal regions of Brazil.

Tuna fishing, capture and post-capture practices in the northeast of Brazil and their effects on histamine and other bioactive amines

The objective of this study was to investigate existing fishing, capture and post-capture practices and their effects on the quality of tuna with respect to histamine and other amines. The study was undertaken in the northeastern coast of Brazil from December 2007 until December 2008. Fishing was performed at high-ocean by pelagic trawl using stainless steel boats and Spanish or American technology. Illex argentinus was used as baits. The fish were insensibilized, bled, eviscerated, rinsed with seawater and stored with flake ice. Histamine was not detected in 95% of the samples whereas histamine levels ranged from 4.92 to 6.90 mg/kg in 5% of the samples. Every lot analyzed complied with the European legislation, and the fish were classified as good quality. No significant difference was observed on histamine levels between fish that were captured dead and alive. The results indicate that the prevalent fishing, capture and post-capture practices used by boats from Rio Grande do Norte, Brazil, are ensuring good quality tuna.

<strong>Additions to the epiphytic macroalgae flora of Bahia and Brazil</strong>

The northeastern coast of Brazil is one of the richest and most diverse areas for the marine algae, but there are still many gaps in our knowledge about the flora, especially in terms of deep-water taxa and small epiphytic macroalgae. Studies of the epiphytic algae on Acetabularia crenulata from the coastal waters along the beaches of Barra Grande and Penha, Vera Cruz, Itaparica Island, Bahia State, Brazil, revealed the occurrence of the genus Chroodactylon and the species Chroodactylon ornatum in the South Atlantic Ocean. The red algae Chondria collinsiana and green algae Phaeophila dendroides expanded its geographical distribution to the northeastern coast of Brazil; the green algae Ulva clathrata and U. flexuosa subsp . paradoxa are new records for the coast of Bahia. The vegetative and reproductive structures are described in detail and a comparison with similar species is provided.

Guanidine Alkaloids from Monanchora arbuscula: Chemistry and Antitumor Potential

AbstractFive guanidine alkaloids, mirabilin B (1), 8bβ‐hydroxyptilocaulin (2), ptilocaulin (3), and a mixture of the 8β‐ and 8α‐epimers, 4 and 5, of 8‐hydroxymirabilin (1,8a;8b,3a‐didehydro‐8‐hydroxyptilocaulin), were isolated from Monanchora arbuscula colonies collected off the northeastern Brazilian coast. All structures were elucidated by spectroscopic analysis, including 1D (1H‐, 13C‐ (BB), and 13C‐DEPT) and 2D (COSY, HSQC, and HMBC) NMR experiments, and comparison with the literature data. The cytotoxicity of the isolated compounds were evaluated against four tumor cell lines, showing that mirabilin B (1) and the two epimers were inactive, while 8bβ‐hydroxyptilocaulin (2) and ptilocaulin (3) presented IC50 values in the range of 7.9 to 61.5 μM, and 5.8 to 40.0 μM, respectively. Further studies on the mechanism of action of ptilocaulin, using HL‐60 leukemia cells, demonstrated that this guanidine compound induced apoptosis of the treated cells.

A new species of Elasmopus (Amphipoda: Hadzioidea: Maeridae) from Suape Harbor, Northeastern Brazilian coast

A new amphipod species of the genus Elasmopus Costa, 1853 is described based on material collected from intertidal rocky shore, near the Suape Harbor, coast of the Brazilian state of Pernambuco. The new species may be recognized by the propodus of gnathopod 2 suboval, slightly tapering distally, palmar margin not defined by a stout seta, spine, or palmar corner, with a subdistal blunt tubercle, posterior margin covered by a dense fringe of plumose setae, and posterior margin of basis of pereopod 7 castelloserrate. This is the ninety-fifth species of the genus Elasmopus described worldwide, the most diverse genus in the family Maeridae Krapp-Schickel, 2008, and the eighth species recorded from Brazilian waters. An identification key to Brazilian species of Elasmopusis also provided.

The spatial dynamics of diversity in Centridini bees: the abundance of oil-producing flowers as a measure of habitat quality

It is assumed that oil bees in the tribe Centridini and oil flowers in the family Malpighiaceae have a conservative evolutionary association, and it is postulated that they also have a tight ecological relationship. Here, we test the hypothesis that variations in the availability of Malpighiaceae flowers affect the abundance and richness of centridine bees. We measured oil availability and sampled bees on oil flowers of Byrsonima sericea DC. (Malpighiaceae) in 12 localities (habitats), along the Northeastern Brazilian Atlantic coast. The availability of floral oil was strongly correlated with richness and abundance of centridine bees. In contrast, no significant correlation was observed between abundance and richness of centridine bees after excluding the effect of oil abundance. We proposed that the asymmetry of density dependence is determining the observed pattern of regional diversity for centridine bees.

Luminescence dated Late Pleistocene wave-built terraces in northeastern Brazil

This study presents field and chronological investigations along the coast of northeastern Brazil from ~4°S to 9°S latitude, which corresponds to ~700 km of coastline under a semi-diurnal mesotidal regime. We investigated wave-built terrace deposits and dated sediments using the optically stimulated luminescence and thermoluminescence methods on quartz grains. The wave-built terraces yielded two main age groups: 200-230 ka and 100-130 ka, which we interpreted as depositional ages. We correlated these age groups with oxygen-isotope stages 7c and 5e, respectively. These events correspond to the antepenultimate and penultimate transgressions along the Brazilian coast. The deposits occur mainly in patches on low-lying flat plateaus along the littoral zone and incision valleys that cut across coastal tablelands. The altitude of the base of the 200-230 ka terraces ranges from 10 m above mean sea level (asl) to -2m asl, whereas the base of the 100-130 ka varies from 12 m asl to -2 m asl. Both terraces were deposited in the foreshore and upper shoreface zones. We noted a coincidence between sea-level highstand chronologies in northeastern Brazil and those in the Bahamas and Bermuda.

Nutrient budgets (C, N and P) and trophic dynamicsof a Brazilian tropical estuary: Barra das Jangadas

This paper focuses on the nutrient dynamics of a tropical estuary on the northeastern Brazilian coast, studied using the LOICZ biogeochemical budgeting protocol. We describe the methodology and assumptions underlying this model. Input data (monthly for rainfall, evaporation, river discharge, and concentrations of salt, phosphorus and nitrogen) were obtained during field campaigns in the Barra das Jangadas Estuary (BJE) over a 5 years period (1999 to 2003). Mass balance results indicate large inputs of nutrients to the system. The model shows that the seasonal variation of the Net Ecosystem Metabolism (NEM) indicates that the system passes from a stage of organic matter liquid production and mineralization during the dry season (-0.5 mmoles C m(-2) d(-1)) to liquid mineralization during the rainy season (-19 mmoles C m(-2) d(-1)). We suggest that the system varies slightly between autotrophy and heterotrophy during the year due to the rainfall regime, human activities in the basin (density population and sugarcane plantations), and associated DIP riverine loads. High per capita loads of N and P indicate a high population density and high runoff. The application of flux balance modeling was useful to understand the nutrient dynamics of this typical small tropical estuary.

The response of meiofauna to human trampling on coral reefs

Coastal environments are trampled by humans worldwide; however, there are few studies that evaluate the effect of trampling on the meiofauna of hard substrates, and none on meiofauna of reef environments. We investigated the effects of trampling due to tourism on the meiofauna of reef formations on the northeastern coast of Brazil. Samples were taken from five paired stations located in two areas on the reef: an area protected since 2004, and an area open to tourist visits. Trampling caused important changes in the meiofaunal assemblage. The densities of the total meiofauna and of the commonest groups were negatively affected in the trampled area. Among the major groups, Polychaeta proved to be very sensitive to this disturbance. The meiofauna groups showed different response patterns to trampling depending on the species of algae trampled. Reductions in animal densities were partly attributed to the loss of turf biomass and associated sand caused by trampling, and partly to the direct effect of people stepping on the animals. Considering the importance of meiofauna in the food web as well as its biodiversity, these results highlight the possible negative effects of human trampling on the ecological and economic “services” that coral reefs provide.

Research Spotlight: The traveling rings of the North Brazil Current

The North Brazil Current (NBC) moves northward along the northeastern coast of Brazil. Drawing from the South Equatorial Current and the outflow of freshwater from the Amazon River, the NBC carries warm, nutrient‐rich water north of the equator. Up near the coast of French Guiana, part of the NBC makes a hard right, flowing east along the equator. Once in a while the turn is especially sharp and the current loops around, pinching off an independently traveling parcel of ocean water. This portion of current travels northwest with a clockwise rotation, moving through the ocean like a Frisbee™ travels through air. (Journal of Geophysical Research‐Oceans, doi:10.1029/2010JC006575, 2011)

Age and growth of the blacknose shark, Carcharhinus acronotus (Poey, 1860) off the northeastern Brazilian Coast

From August 1994 to January 1999 and July 2004 to December 2007, 369 blacknose sharks, Carcharhinus acronotus (Poey, 1860), (43.5–146 cm in total length, TL) were collected from gillnets and surveys (longline) carried out on the continental shelf off the state of Pernambuco, Brazil (07°15′45″S and 09°28′18″S). Vertebral sections from 131 males (43.5–124 cm TL) and 146 females (46–146 cm TL) were analyzed. The index of average percentage of error ( IAPE) estimated from two readings ranged from 1.61 to 4.83 (mean: 6.66%) in the vertebrae of specimens between 0 and 15 years of age. The monthly marginal increment ( MIR) ratio revealed a tendency toward band pair deposition between autumn and winter, whereas the monthly distribution of growth band pair classes revealed that the first band pair appears on the vertebrae of newborns (ages 0+) about 6 months after birth (by June). This deposition pattern was found in specimens with one, two and three band pairs on the vertebrae (1+ and 2+, respectively), thereby confirming annual band pair completion in winter. Growth parameters were derived using the von Bertalanffy model ( VBGM), VBGM b (modified by length at birth: L 0), the Richards, Gompertz and Schnute models. No significant differences in growth rates were detected between the two sampling periods (1994–1999 and 2004–2008). Models with multiple inflection points, particularly the Richards model, best represented growth for the species, which was evidenced by the lowest Akaike information criterion value. There were no differences in growth between sexes. Growth parameters estimated for both sexes were L ∞ = 129.34 cm (SE = 1.35), k = 0.24 (SE = 0.009) and m = 1.34 with the Richards model. The “average” model was determined by averaging L ∞ between the models with substantial statistical support that were considered the most plausible (Richards and Gompertz). Thus, a consensual L ∞ = 130.69 cm, estimated by multi-model inference, was employed to minimize the effects of sample size and selectivity. Males and females mature at 6 years of age and the largest specimen in the catches was 15 years of age (132 cm TL). Local fishing exploitation using gillnets caught only juveniles, whereas the longlines used for the surveys caught subadult and adult specimens. C. acronotus is a relatively fast-growing species inhabiting shallow coastal waters throughout its lifespan, which is a pattern to be taken into account in the years to come for the management of coastal exploitation targeting valuable teleosts.

Taxonomic review of the genus Opaliopsis (Gastropoda: Nystiellidae) from Brazil, with description of a new species

The genus Opaliopsis from Brazil is revised based on three deep-water species. Opaliopsis atlantis (Clench & Turner, 1952) is confirmed as occurring in north-eastern and south-eastern localities. Opaliopsis opalina (Dall, 1927) is reported for the first time in the south-western Atlantic. A new species, Opaliopsis cearense, is described from the north-eastern Brazilian coast, and is distinguished by its large number of fine spiral cords per teleoconch whorl.

Comparative analysis of rating curve and ADP estimates of instantaneous water discharge through estuaries in two contrasting Brazilian rivers

AbstractThis work quantifies, using ADP and rating curve techniques, the instantaneous outflows at estuarine interfaces: higher to middle estuary and middle to lower estuary, in two medium‐sized watersheds (72 000 and 66 000 km2 of area, respectively), the Jaguaribe and Contas Rivers located in the northeastern (semi‐arid) and eastern (tropical humid) Brazilian coasts, respectively. Results from ADP showed that the net water balances show the Contas River as a net water exporter, whereas the Jaguaribe River Estuary is a net water importer. At the Jaguaribe Estuary, water retention during flood tide contributes to 58% of the total volume transferred during the ebb tide from the middle to lower estuary. However, 42% of the total water volume (452 m3 s−1) that entered during flood tide is retained in the middle estuary. In the Contas River, 90% of the total water is retained during the flood tide contributing to the volume transported in the ebb tide from the middle to the lower estuary. Outflows obtained with the rating curve method for the Contas and Jaguaribe Rivers were uniform through time due to river flow normalization by dams in both basins. Estimated outflows with this method are about 65% (Contas) and 95% (Jaguaribe) lower compared to outflows obtained with ADP. This suggests that the outflows obtained with the rating curve method underestimate the net water balance in both systems, particularly in the Jaguaribe River under a semi‐arid climate. This underestimation is somewhat decreased due to wetter conditions in the Contas River basin. Copyright © 2011 John Wiley & Sons, Ltd.

Salinity-induced mixed and barrier layers in the southwestern tropical Atlantic Ocean off the northeast of Brazil

Abstract. High-resolution hydrographic observations of temperature and salinity are used to analyze the formation and distribution of isothermal depth (ZT), mixed depth (ZM) and barrier layer thickness (BLT) in a section of the southwestern Atlantic (0°30´ N–14°00´ S; 31°24´–41°48´ W), adjacent to the northeastern Brazilian coast. Analyzed data consists of 279 CTD casts acquired during two cruises under the Brazilian REVIZEE Program. One occurred in late austral winter (August–October 1995) and another in austral summer (January–April 1997). Oceanic observations are compared to numerical modeling results obtained from the French Mercator-Coriolis Program. Results indicate that the intrusion of subtropical Salinity Maximum Waters (SMW) is the major process contributing to the seasonal barrier layer formation. These waters are brought by the South Equatorial Current (SEC), from the subtropical region, into the western tropical Atlantic boundary. During late austral winter southeastern trade winds are more intense and ITCZ precipitations induce lower surface salinity values near the equator. During this period a 5–90 m thick BLT (median = 15 m) is observed and BLT > 30 m is restricted to latitudes higher than 8° S, where the intrusion of salty waters between 8°–12.3° S creates shallow mixed layers over deep (ZT ≥ 90 m) isothermal layers. During austral summer, shallow isothermal and mixed layers prevail, when northeasterly winds are predominant and evaporation overcomes precipitation, causing saltier waters at the surface/subsurface layers. During that period observed BLT varies from 5 to 70 m and presents thicker median value of 35 m, when comparing to the winter. Furthermore, BLT ≥ 30 m is observed not only in the southernmost part of the study area, as verified during late winter, but in the latitude range 2°–14° S, where near-surface salty waters are transported westward by the SEC flow. These results indicate that the inclusion of salinity dynamics and its variability are necessary for studying mixed and barrier layer behaviors in the tropical Atlantic, where ocean-atmosphere coupling is known to be stronger.

Relative Growth and Its Use to Determine the Morphological Sexual Maturity of Ucides Cordatus (Linnaeus, 1763) (Decapoda, Brachyura, Ucididae) from Two Mangrove Areas on the Northeastern Brazilian Coast

AbstractThe study of relative growth was used to determine the dimensions that best demonstrate the morphological sexual maturity of Ucides cordatus from two mangrove areas (Ariquindá and Mamucabas) of the southern coast of the state of Pernambuco, northeast Brazil. The crabs were collected manually, with a capture effort by one person, during low tide in three different areas (each 25 m2) during a period of one year from April 2008 to March 2009. Crabs of both sexes were measured for the following dimensions: carapace width (CW); carapace length (CL); major cheliped propodus length (CPL); major cheliped propodus height (CPH); abdomen width (AW) and gonopod length (GL). These variables were subjected to an analysis of K-means clustering followed by a bivariate discriminant analysis, which separated the data into two groups: juveniles and adults. After separation of the groups, each age category was grouped into sixteen size classes of CW and the proportion of adults in each size class was calculated, and fitted to a logistic equation. Subsequently, an interpolation was performed to determine the size at which 50% of the males and females were mature. The relationships that best demonstrated the size at maturity were CPL vs. CW for males (Positive allometry; Ariquindá — juveniles b = 1.27, adults b = 1.41; Mamucabas — juveniles b =1.27, adults b = 1.44) and AW vs. CW for females (Positive allometry; Ariquindá — juveniles b = 1.34, adults b = 1.28; Mamucabas — juveniles b = 1.32, adults b = 1.44). The present analysis indicates that, for these same relationships, 50% of males and females are morphologically mature, at, respectively, 38.0 and 35.4 mm CW in Ariquindá, and 37.3 and 32.9 mm CW in Mamucabas. Positive allometry shown in the female abdomen is related to egg incubation, and the excessive growth observed in the male cheliped must be related to reproductive behaviour, including courtship and intra- or inter-specific agonistic interactions with other males.