Population structure and breeding season of the hermit crab Diogenes brevirostris Stimpson, 1858 (Decapoda,Anomura, Diogenidae) from southern Mozambique

In this study the population dynamics of the diogenid hermit crab Diogenes brevirostris is evaluated focusing on size structure, sex ratio and breeding season. Crabs were randomly taken on a monthly basis from January to December 2003 at Costa do Sol, Maputo Bay, southern Mozambique. A total of 622 crabs was obtained of which 290 were males (46.6%), 170 were non‐ovigerous females (27.3%) and 162 were ovigerous females (26.1%). The overall size frequency distribution was unimodal for males, non‐ovigerous females and ovigerous females. The overall sex ratio (1:1.15) differed from the expected 1:1 ratio. Sexual dimorphism was evidenced by the larger size attained by males in relation to both ovigerous and non‐ovigerous females. Breeding took place year‐round with three peaks of spawning (March, August and December). Despite the high reproductive activity, young recruits were scarce in the population. The present results suggest that the study area is suitable for reproduction and growth whereas recruitment may take place in different areas from that occupied by the adults.     

Breeding season of the hermit crab Dardanus deformis H. Milne Edwards, 1836 (Anomura,Diogenidae) in Maputo Bay,southern Mozambique

The breeding season of the hermit crab Dardanus deformis is studied based on the occurrence of ovigerous females, in relation to temperature and rainfall. Monthly samples were obtained between January and December 2003 at Costa do Sol, Maputo Bay, southern Mozambique. A total of 331 female individuals was analysed of which 164 were non‐ovigerous females and 167 were ovigerous females. Dardanus deformis breeds continuously in the study area with peaks of spawning from August to October and a slight decrease from May to July. Both temperature and rainfall were positively correlated with percentages of ovigerous females. Multiple linear regressions suggest that rainfall is the main factor controlling the breeding activity of D. deformis in Maputo Bay. This species may have a rapid and high reproductive activity in the study area as observed by the higher number of ovigerous females relative to non‐ovigerous females which may contribute to a constant larval supply and recruitment of this species in the study area.     

Population dynamics and reproduction of the hermit crab Calcinus gaimardii (Anomura: Diogenidae) at Inhaca Island,southern Mozambique

The population dynamics and reproduction of Calcinus gaimardii were studied at Inhaca Island, southern Mozambique. Crabs were sampled monthly from January to December 2003 in order to assess abundance, size distribution, sex ratio, reproduction and juvenile recruitment. Calcinus gaimardii is sexually dimorphic in relation to size, with males reaching larger size than females. Sex ratios were female‐biased during most months of the study period. Females were more abundant in the intermediate size classes while males outnumbered females in the largest ones. Reproduction took place year‐round with peaks of breeding activity between August and October. Likewise, juvenile recruitment was continuous for both sexes with high intensity in winter (May and June). Egg number increased with female size.     

Life cycle and population structure of the terrestrial isopod Hemilepistus klugii (Brandt, 1833) (Isopoda: Oniscidea) in Iran

The life cycle and population structure of Hemilepistus klugii were studied in a population in Varamin, Iran. The population was sampled monthly (or fortnightly during the breeding season) from February 2008 to June 2009 and a total of 7015 individuals, comprising 1069 males, 1079 females and 4867 juveniles, were collected. As in other Hemilepistus species, five distinct phenophases, namely pair formation, gestation, hatching, growth and stationary, were recorded during the life cycle of H. klugii. The overall sex ratio was 1 : 1 but varied over time. Ovigerous females were observed only in April, indicating a seasonal and very short breeding period. With a short lifespan after breeding, females demonstrated true semelparity. The mean cephalothorax width for ovigerous/post-ovigerous females was higher in 2008 than in 2009. These females attained the largest size in the population throughout the year. The number of eggs per female ranged from 28 to 147 (mean ± SE, 78 ± 1.8). There was a positive correlation between female size and fecundity. Recruitment occurred in late April and resulted in the highest population density in this month, whereas the lowest densities were observed during November to January. Despite a high percentage of ovigerous females carrying undeveloped eggs (72.3%), intramarsupial mortality was low (3.5%).     

Population and reproductive biology of the mud crab,Macrophthalmus hirtipes (Jacquinot, 1853) (Ocypodidae), from marine and estuarine habitats

Summary

Monthly samples of the burrowing mud crab, Macrophthalmus hirtipes (Jacquinot 1853) were collected from February 1978 to April 1979 from a marine inlet, Governors Bay (43°38′S, 172°39′E) and from the Avon-Heathcote Estuary (43°33′S, 172°44′E), New Zealand. The estuarine population was generally unimodal and had an annual cycle of population change sustained by a concentrated summer recruitment. In contrast, the bay population was unimodal and bimodal at different times, and recruitment occurred thoughout the year with no clear peak. Both populations had equal sex ratios and were dominated by immature individuals (crabs?10·0 mm carapace width). Estuarine crabs were generally smaller than marine crabs.

At both sites, ovigerous females were found from late May to early March; the smallest egg-bearing female measured CW 10·0 mm at the estuary and CW 10·5 mm at the bay. Cycles of ovarian and embryological development were similar at the two localities. Three peaks of egg oviposition occurred within one breeding season and mature females were capable of producing successive broods. Fecundity was related exponentially to female carapace width and did not alter during the breeding season. Estuarine females carried more eggs than marine females of similar size. Egg volume was related to time of oviposition and eggs laid at the start of the breeding season were significantly larger than those deposited during the second half; however, estuarine eggs showed less reduction in size at this time than did marine eggs.

Estuarine sediments had higher levels of chlorophyll ‘a’, ATP, and microorganism activity and productivity than marine sediments. As M. hirtipes is a detritus feeder, the differences in food resources are believed to account for the elevated fecundity of estuarine crabs. At the bay, salinities were high with little tidal and seasonal variation, whereas in the estuary sea water was diluted significantly both daily and seasonally. The differences in salinity characteristics between the sites may be affecting larval distribution; recruitment in the estuary was concentrated to the summer and in the bay it was almost continuous.     

Biology of a tropical intertidal population of Cerithium atratum (Born, 1778) (Mollusca,Gastropoda)

The spatial distribution, population structure and secondary production of the gastropod Cerithium atratum were studied in the intertidal region of Engenho d'Água Beach in São Sebastião Channel on the northern coast of the state of São Paulo, Brazil, from May 1998 to April 1999. Twenty monthly samples were taken using a 0.25×0.25?m quadrat in each of two areas. Individuals showed a clustered dispersion pattern in both areas throughout the sampling period. Abundance and mean shell size were related to period of the year and to intertidal level, although no consistent pattern could be described when comparing the two sampling areas due to significant interaction between time and locality. Migratory events were evidenced by marked year-round variations in the frequency of individuals in the largest size classes. Thus, the size structure of this population could not be explained by recruitment and growth alone. Almost 90% of the intertidal population was represented by small individuals that recruited at the beginning of the sampling period (May to August 1998). The intertidal population was composed of annual cohorts formed by individuals recruited in different years. Each annual cohort could be divided into sub-cohorts with closely spaced recruitment periods in the same breeding season. The growth and annual P/B ¯ ratio (where P is the annual somatic production and B ¯ is the mean annual biomass) was estimated for the first year of life of recruits from 1998 occupying the intertidal region. Individuals may have an increment of up to 15?mm in shell length during the first year of life, but the growth was more pronounced in summer months. The high value of P/B ¯ (3.902) was a consequence of the dominance of small-sized individuals, which have a higher weight-specific increment than large-sized ones.     

Breeding biology of the Pale-bellied Tyrant-manakin Neopelma pallescens (Aves: Pipridae) in south-eastern Brazil

The Pale-bellied Tyrant-manakin (Neopelma pallescens) inhabits semi-deciduous and riparian forests in central-north South America. Contrary to most manakins, there is no evident sexual dichromatism in the species and little is known about its breeding biology. We studied the breeding biology of a colour-banded population of the species from August to December 2016 and from August to October 2017 in the Campus Florestal of the Universidade Federal de Viçosa, south-eastern Brazil. The breeding season extended from early September to late November. The species is promiscuous, with males exhibiting simple courtship displays (exploded leks) in individual arenas. The nest (n = 13) is a cup attached by its top lip between forked branches and is very simple, with a structural layer made with dry grass stems and heads, attached to the branch with spider silk. The outer and lining layers are absent. The mean clutch size was 1.8 eggs (n = 11), which are oval and pale coloured, covered with spots of different shades of brown, often concentrated in the larger pole. Mean egg length and width (± SD) were 21.0 ± 0.9 × 15.8 ± 0.7 mm (n = 14) and the mean weight was 2.8 ± 0.4 g (n = 10). The incubation period could not be estimated, but the nestling period was 15 days (n = 2). The simple percentage of successful nests was 15.4%, with 76.9% of the nests depredated and 7.7% abandoned. This is the first detailed study about the breeding biology of any Neopelma species, providing relevant data for the study of the evolution of life history strategies not only for the genus, but for the whole family Pipridae.     

Seasonal abundance and life cycle of Archaeomysis articulata (Crustacea: Mysidacea) on a sandy beach of western Hokkaido,Japan

Abundance and life cycle of the sandy beach mysid Archaeomysis articulata Hanamura were investigated based on seasonal collections over 2 years in the surf-swept intertidal zone of an open sandy beach in Ishikari Bay, western Hokkaido, Japan. The number of mysids in the intertidal zone increased from winter to spring, reaching its maximum in summer to autumn (average >2500 ind. m^-2), and decreased considerably towards the end of the year. The abundance was maintained at a minimal level (<200 ind. m^-2) throughout the winter. The breeding season was March to November, during which the liberation of young was most intensive from late April to mid-October. Three alternate generations, including overwintering, spring and summer generations, were suggested from field observations. Archaeomysis articulata is primarily iteroparous; only a restricted number of breeding females could produce more than two broods. The life span was estimated to be 7–9 months for the overwintering generation and 2.5–3.5 months for the spring–summer generations. The sex ratio fluctuated at samplings, with a tendency toward female dominance (overall mean=54%). The number of embryos carried by a single female ranged from 16 to 94, tending to increase with increasing size of females. The size of egg-like embryos varied seasonally from 0.45 to 0.54mm, being largest in spring and smallest in summer, with no correlation to the length of females; water temperature is considered an important factor in regulating the size of eggs. The embryonic development time was estimated to be 1.5 months for the spring generation (water temperature 5–10°C), and much shorter than this for the summer and overwintering generations. The mortality rate during embryonic development was estimated to be 7–18%.     

Reproductive pattern in the southernmost populations of South American redbelly toads

Anurans can be classified as explosive or prolonged breeders. In species with short breeding periods, intensive competition for females is expected. We analyse the temporal breeding pattern of Melanophryniscus aff. montevidensis, a bufonid inhabiting grasslands of Argentina, whose breeding is concentrated in events of short duration. Males are active during more than one breeding event per season, whereas most females take part in a single event per season. Active males outnumbered females by up to 14 times at one event, operational sex ratio (OSR) ranged from 0.07 to 0.75 and intensity of sexual selection (I _s) from 0.69 to 24. We found a negative association between I _s and OSR, but no correlation between number of active males per event and OSR or I _s. The OSR is the key that regulates the intensity of sexual selection, nevertheless, and in spite of being explosive breeders, females would potentially have chances to select their mate.     

Gastropod shell utilisation pattern by the hermit crab Clibanarius symmetricus (Anomura: Diogenidae) in an Equatorial Amazon estuary

Clibanarius symmetricus is a diogenid hermit crab that is highly abundant in rocky intertidal environments, including the rock outcrops of Amazon estuaries. This study characterises the C. symmetricus shell utilisation pattern in the Marapanim River estuary, Pará, Equatorial Amazon, based on the hypothesis that occupancy patterns would differ, in relation to biometry and diversity of occupied shells, from those of other regions in the range of this species due to the distinct and unique environmental characteristics of equatorial estuaries. Monthly samplings were carried out from August 2006 to July 2007, in the upper and lower areas of the mid-littoral during low tides. A total of seven gastropod species were found as shells occupied by hermit crabs: 93.33% were Thaisella trinitatensis shells, 2.00% Nassarius vibex, 2.00% Neritina virginea, 1.33% Natica marochiensis, 0.67% Parvanachis obesa, and, occasionally, Littorina flava and Phalium granulatum shells, each with only one occurrence (0.33%). Juveniles [cephalothoracic shield length (CL) of less than 3.6 mm] occupied a higher variety of shells, while adults occupied shells with larger meristic variation. Males occupied larger shells. The length of the animal was influenced by shell measurements (total width, aperture width and weight). Clibanarius symmetricus showed occupancy patterns generally similar to those of specimens of the same species previously studied in the Brazilian subtropical region, and this occupancy is explained by shell availability, shell size and weight, and hermit size. However, this study showed lower occupied shell species diversity, and the species with highest occupancy frequency (T. trinitatensis) was not reported in any previous studies on this hermit crab. In addition, the studied population differed in occupancy patterns, with differences between males and females, and between juveniles and adults.     

Intertidal distribution,symbiotic association and reproduction of Pinnotheres bidentatus (Brachyura: Pinnotheridae) from Taiwan

The intertidal distribution of the free-swimming stage of the pea crab Pinnotheres bidentatus Sakai, 1939 was examined on the Kaomei coastal flat of western central Taiwan. Most pea crabs were collected from tidal waters close to the shoreline regardless of the collecting treatments utilized. Results suggest that the pea crab population remains inshore during ebb tides. Many hard and post-hard stages of P. bidentatus were discovered, including ovigerous females. The host was found to be Laternula marilina (Valenciennes, 1860) (Mollusca: Bivalvia: Laternulidae). The infestation percentage of the bivalve was high (average 74.6%). However, this may be due to collections of the bivalve population in the present study being skewed towards large sizes. This biased size structure of the host population may be also responsible for lack of positive size correlation between number and size of male and female symbiotic crabs and the host. Multiple infestations of P. bidentatus appear to be uncommon for L. marilina. Single infestations might be a model for bivalves in tropical regions where the sea is relatively oligotrophic and food competition may be enhanced with multiple infestations. In comparison to other members of the genus Pinnotheres or related species (e.g. P. pisum (Linnaeus, 1767) and Zaops ostreum (Say, 1817)), the body size of sexually mature female P. bidentatus is relatively small, even though it produces similar size eggs (approximately 300 microns). The general rule for size correlation between fecundity and female size in brachyuran crabs holds for P. bidentatus.     

Population study of the landhopper Talitroides topitotum (Crustacea: Amphipoda: Talitridae) in central Mexico

A population of Talitroides topitotum discovered in Mexico City in 1992, was studied during a one year cycle. Abundance varied significantly during the study period, high population numbers occurred from June to October with a marked decrease from November to May. Maximum total length was 12.5mm and the size at first reproduction for females was 4.0 mm total length. Talitroides topitotum reproduced all year, with a peak in June. A female biased sex ratio was obtained in every sample with a maximum of 4:1 in May. Fecundity varied from one to nine eggs per female and mean fecundity varied significantly through time. The results show that T. topitotum exhibits little variation among widely separated populations and, 4 years after introduction, a life cycle adjusted to the local weather.     

Leucocytozoon toddi in British sparrowhawks Accipiter nisus: observations on the dynamics of infection

Leucocytozoon infection was studied in a breeding population of sparrowhawks Accipiter nisus in the British Midlands. Almost all transmission was apparently ‘vertical’ from adults to young, at the nest site. Parasites largely disappeared from the blood between October and April, reappearing in a ‘spring relapse’ during the nesting season. Infected birds remained infected for life; there was no evidence of reduced longevity in infected birds once they had dispersed from their natal nest site. There was no assortative mating between infected or uninfected males and females. It is postulated that the concentration of birds at the nest site allows transmission by a non-specific vector breeding in the same vicinity. At other times of year the birds are more regularly dispersed and transmission in unlikely to occur.     

Relative growth and sexual dimorphism of the South Atlantic hermit crab Loxopagurus loxochelis (Anomura,Diogenidae) from Ubatuba,Brazil

Some morphometric relationships and biometric aspects were studied in the hermit crab Loxopagurus loxochelis (Moreira, 1901) with the purpose of describing its relative growth and finding the size where sexual maturity is reached. Specimens were collected monthly from September 1995 to August 1997 in the Ubatuba region using double-rig trawl nets. A total of 1084 specimens (625 males and 459 females) was analysed. In this species, sexual dimorphism is particularly evident in chela dimensions. Differences between males and females were found in the left propodus length versus cephalothoracic shield length (CSL) and wet weight (WW) versus CSL relationships. Males showed a high positive allometry in chela growth, while females were close to isometry/negative allometry. The size at which a clear differentiation occurred in the growth of the propodus length and in the increase in weight of males and females was between 4.5 and 6.0mm CSL, suggesting that size at which sexual maturity is reached.     

Natural history of the Lined Seedeater Sporophila lineola (Aves: Thraupidae) in southeastern Brazil

The Lined Seedeater (Sporophila lineola) is a migratory species that inhabits a variety of open habitats in South America. We studied the breeding biology and territorial behaviour of a colour-banded population of the species in the Universidade Federal de Viçosa, Campus Florestal (1–19,8808ºS, –44,4136ºW), during two breeding seasons (2014/2015 and 2015/2016), which spans from December to April. We monitored 74 nests of this species. The nest is a low cup supported between a fork. Nests are mainly built with grass stems and rootlets, with spider-web used to hold the material together and to bind the nest to its supporting plant. Females are solely responsible for nest building and incubation, which is synchronic, and also for feeding nestlings to a large extent. Males are responsible for defending the territory, which corresponds to a small portion of the home range restricted to the nest environs, and also feed the nestlings. Mean clutch size is two eggs (78% of nests monitored), with clutches of three (20.3%) and four (1.7%) eggs also observed. Eggs are whitish, covered with dark brown blotches and spots to a variable extent. Mean incubation period, considered as the period between the onset of incubation and hatching of the first egg, is 11 days. Nestling period, considered as the period between hatching of the first egg and fledging of the last young, is 10 days. During the first breeding season, the simple percentage of successful nests was 34.1%, while the Mayfield success was 29.8%, with slightly higher values observed during the second breeding season, with 39.4% and 35.7%, respectively. We recorded, for the first time, three cases of polygamy in the species. We also recorded breeding site fidelity for the first time in the species, with males returning to the same territory owned in a previous breeding season.     

Comparative population biology of Uca rapax (Smith, 1870) (Brachyura,Ocypodidae) from two subtropical mangrove habitats on the Brazilian coast

The structure of two populations of the fiddler crab Uca rapax in two subtropical mangrove habitats near Ubatuba, State of São Paulo, Brazil were compared. The size–frequency distribution, sex ratio, and recruitment were evaluated. Sampling was performed monthly from April 2001 to March 2002 in the Itamambuca and Ubatumirim habitats. Crabs were caught manually for 15?min by two collectors during low tide. The carapace width of each crab was measured with a digital caliper, and the sex and ovigerous state were recorded. The median size of the carapace width of males was greater than that of females at both sites (P<0.05). The median size of the crabs from Itamambuca was larger than at Ubatumirim (P<0.05). Only 28 ovigerous females were obtained from both mangroves, which suggested that females might remain in their burrows during the incubation period. The highest recruitment pulse occurred in winter for both populations, probably as a consequence of high reproductive activity during summer. The sex ratio in the size classes showed an anomalous pattern, with a higher frequency of females in the intermediate size classes. This may be related to a greater energy requirement for reproduction in females, thus delaying growth. The variable environmental conditions to which Uca rapax populations are subject appear to act directly or indirectly on the population, causing variations in growth and reproductive processes in the different populations investigated here.     

Breeding biology of the Sayaca Tanager (Thraupis sayaca) in southeast Brazil

ABSTRACT

Thraupis is a genus of the American endemic Thraupidae (subfamily Thraupinae), comprising seven species that inhabit tropical forests to urban centres. The Sayaca Tanager (Thraupis sayaca) is a disturbance-tolerant species of high representativeness in plant-frugivore networks, but information on its breeding biology is scarce and often restricted to non-systematic surveys. We studied the breeding biology of the T. sayaca, following 39 active nests in a periurban area of southeast Brazil during two breeding seasons (2017/2018, 2018/2019). The breeding season ranged from early September to middle December, and the nests were placed in native and exotic plants and human buildings (nest height above ground: 3.35 ± 1.73 m, mean ± SD). Only females incubated and brooded, but both adults built the nests, fed the nestlings, and removed their faecal sacs. Clutch size was 2.86 ± 0.38 eggs and nest attentiveness was 71.2%. The incubation and nestling periods were, respectively, 13.4 and 17.4 days. Males and females did not differ on nestling provisioning and nest sanitation rates. Nestling provisioning (13.35 ± 6.25 trips/hour) increased with nestling age, while mean brooding time was 37.2% and decreased with nestling age. Apparent nest success was 38.7%, and nest survival according to the Mayfield method was 27.2%. Five nests (20.8%) were parasitised by the Shiny cowbird (Molothrus bonariensis), and we recorded five events of nest-site reuse. We concluded that the most remarkable breeding traits of T. sayaca in comparison with close-related tanagers are the use of anthropogenic nest sites, the higher clutch size and number of feeding trips, and the longer nestling period.     

Breeding biology of the threatened Campo Miner Geositta poeciloptera (Aves: Scleruridae), a Neotropical grassland specialist

The Campo Miner is a threatened grassland passerine endemic to the South American Cerrado, whose life history is almost unknown. In this paper, we studied during three breeding seasons (2014 to 2016) the breeding biology of a colour banded population of the species found in the Upper Rio Grande Grasslands, south-eastern Brazil. We found 98 nests, 81 of which became active and were monitored. The Campo Miner breeds in frequently burnt-and-grazed natural grasslands, successfully nesting in highly disturbed sites, such as dirt banks along roads and even in mine pits. The species is socially monogamous and both parents build the nest, which is a cavity/with-tunnel/simple/platform type. The nest chamber is lined with a platform made of grass fragments, charcoal, hairs, and mammal faeces. The most common clutch size is three eggs (n = 66), with some nests containing one (n = 1), two (n = 12) or four eggs (n = 2). The egg is white and pyriform and the incubation, performed by both parents, lasts 17.5 days. Mean nestling period is 15.5 days, with both parents feeding the young. Breeding season lasted for about 125 days (August to December) and multiple breeding attempts in a single season were common, with a maximum of three attempts recorded. All species of Scleruridae built their nests inside cavities dug in the soil with an access tunnel to it, where they lay a small clutch (usually 2–3 white eggs), but no other species in the family has been studied in detail to date. Further studies are required to understand why a species apparently tolerant to anthropogenic impacts such as G. poeciloptera can be so rare, patchily distributed and threatened throughout its range.     

Asynchronous reproduction in three species of crocodilians in south-eastern Amazonia

We studied reproduction of three species of crocodilians, Paleosuchus trigonatus, Caiman crocodilus and Melanosuchus niger, in the Xingu River, near the Belo Monte hydroelectric dam. The periods of laying and hatching of eggs were estimated for each nest before (2013–2014) and after (2016–2017) the river was dammed and the reservoir was formed. Nesting of the three species occurred between August and December, but was largely asynchronous; nest construction peaked in September for P. trigonatus, October for M. niger and November for C. crocodilus. Reservoir filling had little effect on the laying period of any of the species. Nests of P. trigonatus and M. niger were always within 0–12 m of the bank, whereas nests of C. crocodilus, which nests later in the season when flooding is more likely, were up to 100 m from the nearest water body. There was no relationship between distance from water and the number of eggs in nests, suggesting that larger and presumably more experienced females do not lay at different distances from the bank in any of the species.     

Spatial distribution of the ghost crab Ocypode quadrata in low‐energy tide‐dominated sandy beaches

The spatial distribution of the ghost crab Ocypode quadrata (Crustacea, Brachyura) was described in five low‐energy tide‐dominated sandy beaches in São Sebastião Channel (Grande, Zimbro, Pitangueiras, Cabelo Gordo, and Segredo). On each beach, the zonation study was performed on five random 1?m wide transects sampled from the water line to the vegetation. In general, individuals occurred above 1?m in height in relation to mean low water (MLW) and peaked between 1.5 and 2.0?m. The sandy beaches were then divided into three 1?m wide strips (tidal levels) parallel to the water line and equivalent to medium intertidal, upper intertidal, and subterrestrial fringe to test the effect of beach and tidal level on the abundance and size of O. quadrata. These strips were sampled as a whole using adjacent 1 m² squares. A previous evaluation showed a positive significant relationship between carapace length and burrow diameter, thus supporting crab size estimates from burrow openings. The smallest individuals occurred mainly in the medium intertidal, but were also recorded in the subterrestrial fringe, which was occupied mainly by large‐sized individuals. In general, the individuals were randomly dispersed within the strips and concentrated in the upper intertidal zone on all the sandy beaches. The density of O. quadrata varied among beaches, with a tendency to lower values on areas with very fine and poorly or moderately sorted sediments and on that most used for recreational activities (Grande). Burrow size also varied among beaches, with a tendency of smaller burrows in areas with coarser sand grains and higher tourism. These results indicate that the effect of human impact on density and size of O. quadrata may be confounded by the high environmental heterogeneity of the studied system, which may make it difficult to utilize this species as an indicator of the conservation status of low‐energy tide‐dominated sandy beaches.