Trypauchen vagina (Bloch & Schneider, 1801) a new established species in the Southwestern Atlantic

Trypauchen vagina (Bloch & Schneider, 1801) is a goby that lives burrowed into the substrata feeding on small invertebrates. It is native to the Indo-pacific region, ranging from Kuwait to China. Recently, this fish has been reported outside the original range of distribution, being found in the Mediterranean Sea, and more recently in the northeastern Brazilian coast. The Mediterranean reports are usually associated with Lessepsian migration, while the reports from Brazil are possibly related to ballast water transportation. In the present work, we provide eight new records from southeastern Brazilian coast, all made in São Paulo state, far from the first record. These additional records raised concern since the presence of an alien species could implicate in environmental and economic losses. Thus, we decide to model the environmental suitability for this goby in the Brazilian coast, specially focusing on major ports, usually places with high ballast water propagule pressure. In addition, an analysis of the suitability in the Red Sea was also made, to verify the hypothesis of Lessepsian migration. The results revealed that temperature and primary productivity are among the most important parameters for the presence of T. vagina, also indicating a high environmental suitability for this species in the Red Sea and Brazilian coast, especially in southeastern region, where the new records were made. Due to the number of collected individuals, it is hard to affirm that this taxon presents a self-sustaining population in Brazilian waters, but the several registers, in different locations and different life stages point to an establishment of the species in this new region. This population status allied with a high environmental suitability is alarming and should motivate new studies concerning T. vagina in Brazilian waters.     

Genetic detection of cryptic species in the frillfin goby Bathygobius soporator

We compared, through allozyme and cytochrome b sequence analyses, populations of Bathygobius soporator from four localities on the Brazilian coast with those from the Oceanic Islands of the Rocas Atoll (230 km off the Brazilian coast) and the Bahamas (5700 km northwest from Rocas). The population from the Rocas Atoll was genetically more similar to the geographically distant Bahamas (gene identity, I=0.92) than to the supposedly conspecific populations from the Brazilian coast (I<0.55). Five diagnostic allozyme loci and a high level of nucleotide divergence (Kimura 2-parameters=0.21) were found between the two groups, further confirming that they must belong to different biological species. Since the type locality of B. soporator is in the Caribbean, the binomial should be maintained for the Bahamas/Rocas Atoll populations, and the other Brazilian populations of Bathygobius analysed either represent the occurrence of a species of Bathygobius hitherto not cited for the Brazilian coast, or belong to a new species. The coastal populations of this species were also found to be highly structured (F_ST=0.18; p<0.05), indicating that, even along the coast, levels of gene flow of this species are limited. This could be related to their reproductive biology (adhesive benthic eggs with parental care and short length of larval life). The timings of the divergence between the Caribbean and the Southwest Atlantic species found here, estimated from allozymes and cytb sequences, are very similar (around 10 MY bp), and correlate with the start of the Amazon river outflow. The colonization of the Rocas Atoll by populations from the Caribbean seems to have taken place after the Caribbean and the Brazilian coast species had diverged.     

Metazoan parasites as biological indicators of population structure of Halobatrachus didactylus on the Portuguese coast

A total of 366 Halobatrachus didactylus (Bloch & Schneider 1801) individuals were obtained between September and November 2000 from commercial fishing vessels at eight localities along the Portuguese coast and analysed for metazoan parasite infections. Three species of parasites were found: Hysterothylacium aduncum Rudolphi 1802; Progrillotia dasyatidis Beveridge et al. 2004, which constitutes a new host record for the Lusitanian toadfish; and Nerocila orbignyi Guérin‐Méneville 1829. Prevalence and mean abundance were determined for the total fish population in each sample. Significant differences in the number of parasitized fish among the samples analysed indicate differences in the H. didactylus populations along the Portuguese coast. The cluster analysis performed to prevalence data pointed out two major clusters: one composed of mainly estuarine samples and one that grouped samples from coastal areas; this is in agreement with the results obtained by morphological and genetic analysis performed for the same samples.     

Observations on the biology of the trochid gastropod Austrocochlea constricta (Lamarck) (Prosobranchia). I. Factors affecting shell-banding pattern

The intertidal trochid gastropod Austrocochlea constricta (Lamarck) has a variable pattern of banding, the shell ranging from white with fine black stripes, to completely black. The patterns may be classified into six arbitrarily chosen types. Populations were studied on three shores chosen to represent the range of habitat of the species (from estuarine to open coast localities) to determine the relative importance of environmental factors on shell-banding. The densities and proportions of the population in different stripe-classes were estimated from quadrat samples on the three shores. Analysis of size-frequency distributions indicates recruitment to be continuous. In all months, juveniles were at greater densities at the upper level of the distribution of the species on open coast and estuary-mouth platforms; the estuarine population was uni modal in size-frequency distribution. Growth rates of caged animals were higher on the open coast than in the estuary, and changed in animals transferred between the two localities, but did not differ between animals of different banding pattern. The proportion of the population in each stripe-class was constant on each shore, suggesting large scale selection processes do not cause the observed differences in frequencies of each banding pattern.The concentrations of the major shell pigment, uroporphyrin I, varied between stripe-classes and, within classes, between shores, with a trend for more densely pigmented shells on the open coast, a trend also evident for the proportion of the population in the darker stripe-classes. Caged animals transferred from the open coast to the estuary showed changes in banding pattern which were consistent with the differences in pigmentation of the two populations. Animals transferred from the estuary to the open coast changed pattern at the same frequency as controls from the open coast; a result of severe storms which caused the open coast population to show changes of banding pattern.It is concluded that shell-banding patterns in A. constricta are primarily governed by environmental factors.     

Genetic and morphological variation in two littorinid gastropods: evidence for recent population expansions along the East African coast

In marine species, population diversity and differentiation is affected by the population history and by the complex interaction between oceanographic dynamics and ecological traits. In the present study, we examined two species of marine gastropods (the mangrove periwinkle Littoraria scabra and the rocky shore Littoraria glabrata) along the East African coast, using both genetic and geometric morphometric methods. We report a greater variation of shell shape in L. scabra compared to the slightly smaller variation in L. glabrata. This variation was probably associated with variation of environmental factors along the coast, such as temperature and hydrodynamics. Despite morphological variation, we found low mitochondrial genetic differentiation among samples from different localities for both species, which is probably a consequence of the ongoing gene flow during the free‐swimming larval stage of these gastropods. Additionally, high levels of haplotype diversity, low nucleotide diversity, and ‘star‐like’ genealogies were found in both species. These observations and the results from mismatch distributions, indicate a possible signature of recent population expansions in both species, which probably started during interglacial periods of the Pleistocene and led to the colonization of the Indian Ocean coast. © 2013 The Linnean Society of London     

Oxidative stress in Perna perna and other bivalves as indicators of environmental stress in the Brazilian marine environment: Antioxidants, lipid peroxidation and DNA damage

Oxidative stress can take place in marine bivalves under a series of environmental adverse conditions. The study of different systems related to oxidative stress in these organisms can give important information about their physiological status and also about environmental health. Bivalves have been proposed as good sentinel organisms in pollution monitoring studies through the analysis of biochemical biomarkers, and most of the biomarkers analyzed are those related to oxidative stress. However, it is very important to know how other environmental factors not associated to the presence of pollutants might affect these parameters. We have studied a series of mechanisms related to oxidative stress in mussels which inhabit the Brazilian coast, especially in Perna perna species, subjected to different stress conditions, such as the exposure to different contaminants in the laboratory and in the field, the exposure of mussels to air and re-submersion, simulating the tidal oscillations, and in mussels collected at different seasons. Both oxidative damage levels and antioxidant defense systems were strongly affected by the different environmental stress. This review summarizes the data obtained in some studies carried out in bivalves from the Brazilian coast.     

When an advantageous reproductive trait turns bad: Eggs of the threatened fish Genidens barbus as a natural bait in recreational fisheries

This study describes a recreational fishing method focused on marine catfish, Genidens barbus (Endangered-EN), through the adoption of its eggs as a natural bait during its reproductive period in southern Brazilian ecosystems. Male G. barbus perform parental care, collecting eggs released by females after the fertilization process. Owing to this male behavioural pattern during the reproductive period, these individuals are easily caught in recreational fisheries that use eggs of the same species as a natural bait. The current adoption of G. barbus eggs as bait may intensify the fishing pressure on its populations.     

Salmonella spp., Vibrio spp., Clostridium perfringens, and Plesiomonas shigelloides in Marine and Freshwater Invertebrates from Coastal California Ecosystems

The coastal ecosystems of California are highly utilized by humans and animals, but the ecology of fecal bacteria at the land–sea interface is not well understood. This study evaluated the distribution of potentially pathogenic bacteria in invertebrates from linked marine, estuarine, and freshwater ecosystems in central California. A variety of filter-feeding clams, mussels, worms, and crab tissues were selectively cultured for Salmonella spp., Campylobacter spp., Escherichia coli-O157, Clostridium perfringens, Plesiomonas shigelloides, and Vibrio spp. A longitudinal study assessed environmental risk factors for detecting these bacterial species in sentinel mussel batches. Putative risk factors included mussel collection near higher risk areas for livestock or human sewage exposure, adjacent human population density, season, recent precipitation, water temperature, water type, bivalve type, and freshwater outflow exposure. Bacteria detected in invertebrates included Salmonella spp., C. perfringens, P. shigelloides, Vibrio cholerae, Vibrio parahaemolyticus, and Vibrio alginolyticus. Overall, 80% of mussel batches were culture positive for at least one of the bacterial species, although the pathogens Campylobacter, E. coli-O157, and Salmonella were not detected. Many of the same bacterial species were also cultured from upstream estuarine and riverine invertebrates. Exposure to human sewage sources, recent precipitation, and water temperature were significant risk factors for bacterial detection in sentinel mussel batches. These findings are consistent with the hypothesis that filter-feeding invertebrates along the coast concentrate fecal bacteria flowing from land to sea and show that the relationships between anthropogenic effects on coastal ecosystems and the environmental niches of fecal bacteria are complex and dynamic.     

Geographic Variations in Reproductive Parameters of the Starspotted Dogfish, Mustelus manazo, from Five Localities in Japan and in Taiwan

Geographic variations in reproductive parameters of Mustelus manazo from five localities, four in Japan (Aomori, Tokyo Bay, Maizuru, and Shimonoseki) and one in Taiwan, were investigated from November 1994 to June 1996. Differences in age and length at sexual maturity from Aomori to Taiwan were approximately 3 years and 300mm in TL, respectively, for both sexes. The sharks appeared to mature at a later age and to a larger size in the most northern population, Aomori, and to be faster and smaller in the southern population, Taiwan. Mating, ovulation and fertilization periods were generally during May and August in the four localities of Tokyo Bay, Maizuru, Shimonoseki, and Taiwan. In Aomori, males showed no clear monthly variation in gonad index, and females in Aomori had a protracted reproductive period. Females became pregnant every year in all localities, except Aomori. The Aomori population apparently has a different reproductive cycle. The Tokyo Bay population is distinctive regarding growth and sex ratios of embryos. In all localities, the number of embryos per litter increased relative to total length of mother. There was no geographic variation in reproductive parameters between Maizuru and Shimonoseki. The coast between these last two sampling areas is a continuous steep slope with similar environmental conditions, and mixing of the two populations may occur.     

Spatial and temporal variability of early post-settlement survivorship and growth in the barnacle Balanus glandula along an estuarine gradient

The performance of a species can be significantly altered by subtle changes in the physical environmental. The intertidal barnacle Balanus glandula is predominantly an open coast species in the Northeast Pacific. However, B. glandula commonly inhabits estuaries where environmental conditions such as salinity and temperature drastically differ from the open coast. We used survivorship and growth rates as a measure of performance in recently metamorphosed laboratory reared juvenile B. glandula outplanted along an environmental gradient at the mouth, mid-estuarine, and riverine end of the South Slough Estuary, Oregon, USA. Juvenile performance was highly variable over spatial and temporal scales and dependent upon existing environmental conditions. Surprisingly, along this estuarine gradient, juveniles performed better at a mid-estuarine location than at the mouth of the estuary. Typically, the riverine end of the estuary was the least suitable habitat along the estuarine gradient due to high juvenile mortality and a low growth rate. Although seasonally variable, survivorship and growth decreased with height along a vertical intertidal gradient as well. In a reciprocal transplant experiment, populations from both ends of the estuarine gradient displayed similar survivorship and growth rates. Our results demonstrate that the interactions of environmental conditions that vary temporally and spatially along a gradient strongly affect the success of an individual surviving and prospering during the early juvenile period.     

Body temperatures of Tropidurus torquatus (Squamata,Tropiduridae) from coastal populations: Do body temperatures vary along their geographic range?

We studied the geographic variation in body temperatures of the lizard Tropidurus torquatus in 10 restinga populations along approximately 1500 km of Brazilian coast. The mean activity body temperatures (30.8–36.2 °C) seems to remain constant along the populations and the differences recorded among them result from the adjustment of each lizard population to the local thermal environment (i.e. may express in part the local microhabitat temperatures occurring in each of the localities in that particular moment). Forested and open habitat tropidurine species have different mean activity body temperatures, probably as a consequence of the different thermal environments in these habitats.     

Can otoliths of <Emphasis Type="Italic">Genidens genidens</Emphasis> (Cuvier 1829) (Siluriformes: Ariidae) reveal differences in life strategies of males and females?

Determination of the growth parameters of a species is an indispensable requirement for understanding its biology and consequently the management of its fishery. In fisheries science, calculations of longevity, mortality rates and stock-assessment models depend on the availability of growth data. Genidens genidens, one of the most common ariid species in Brazil, is a potential sentinel species for biomonitoring in Guanabara Bay, one of the country’s most degraded estuarine ecosystems. The present study investigated the growth parameters of G. genidens, providing supporting information for its management. Individuals were measured, sexed, and the number of rings in the lapillus otolith counted. The periodicity in the formation of the rings was determined by the monthly ratio of the edge type. Individuals from 0 to 11.5 years old were captured, including some that were older than previously reported. Our results showed that this species forms two growth rings per year, one in summer and the other in winter; the summer ring is formed one month later in males than in females, due to the period of oropharyngeal incubation of the offspring. Sexual dimorphism was also observed in the growth rate: males grow faster, and consequently reach smaller sizes than females. We can conclude that the parental care carried out by males and the high energy expenditure in forming the large oocytes by females mark the otoliths and reveal life-cycle particularities differently in each sex.     

Ecological distribution of Penaeus schmitti (Dendrobranchiata: Penaeidae) juveniles and adults on the southern coast of São Paulo state,Brazil

This study aimed to describe the ecological distribution and the effects of environmental factor oscillation on the life cycle of Penaeus schmitti along the coastal area of Cananéia, southern São Paulo state, Brazil. Shrimp and environmental factors were sampled monthly from July 2012 to May 2014 at seven sampling stations covering both estuarine and marine environments, using a shrimp fishing boat. A total of 273 juveniles and 829 adults were sampled, and there were significant differences in body size (carapace length – CL) throughout the seasons (ANOVA). Smaller animals were observed in the summers of 2013 and 2014 and at sites closer to the coast, which are under estuarine influence. Adults were observed mostly in autumn and in deeper regions. We detected a positive relationship between the abundance of juveniles and the bottom water temperature and between the abundance of adults and the water salinity (canonical correspondence analysis). The presence of smaller individuals in the estuarine environment demonstrates the importance of the estuaries in this species’ ontogeny, mainly concerning their protection, since this environment has physical (depth) and physiological (salinity) barriers to their predators. Furthermore, we could clearly observe the plasticity of juveniles in relation to environmental factors. In contrast, adults were less tolerant to the variation in environmental factors, mainly water salinity, which was a limiting factor of their distribution. Seasonal differences in body size could be directly associated to this species’ life cycle and characterized by trophic and reproductive migratory activities between the estuarine and oceanic regions.     

Factors affecting seasonal variations in demersal fish assemblages at an ecocline in a tropical–subtropical estuary

Seasonal changes of fish species composition in terms of biomass, density and number of species in three areas of the main channel of the Paranaguá Estuary (axis east–west) are described in relation to seasonal fluctuations in salinity, water temperature and dissolved oxygen in the main channel. Two hundred and thirty‐four samples were collected monthly, between July 2000 and June 2001, in the main channel. Seventy‐nine species of 29 families were captured with a total estimated mean density and biomass of 1513 individuals ha^?1 and 34 kg ha^?1, respectively. The number of species and total mean density differed significantly among areas and seasons, but the total mean biomass differed only significantly throughout the ecocline (areas) of the Paranaguá Estuary. For the most abundant species, the mean densities of Stellifer rastrifer, Aspistor luniscutis, Menticirrhus americanus, Sphoeroides testudineus, Cynoscion leiarchus and Symphurus tesselatus (with the exception of Cathorops spixii and Genidens genidens) differed significantly among seasons. The mean biomass of these species, with the exception of G. genidens, S. rastrifer, A. luniscutis and S. testudineus, also differed significantly for the factor seasons. Area was a significant factor for the eight most abundant species (density and biomass), except S. testudineus (density), G. genidens, C. leiarchus and S. tesselatus (biomass). The season v. area interaction term was significant for C. leiarchus (density). Most of these differences occurred during the rainy season when fishes concentrated principally in the middle of the estuary, where the salinity remained stable. It is suggested that the salinity stability in the middle of the estuary is the main reason why the most estuarine resident fish species move downstream and remain there, regardless of the increased freshwater runoff. Moreover, canonical correspondent analysis output detected that during the late rainy season, the variable dissolved oxygen (P < 0·01) was the most important environmental variable, responsible for structuring patterns of fishes assemblages in the west–east axis of Paranaguá Estuary. During the end of the dry season, both salinity (P < 0·01) and dissolved oxygen (P < 0·05) were responsible for this ecological feature in the estuary. Finally, it was possible to detect that juveniles and adults of some important species respond differently to seasonal fluctuations of the ecocline‐determining environmental factors. This behaviour is suggested as a strategy to avoid competition and predation during the rainy season in the middle estuary. The Paranaguá Estuary did not fit with the pre‐existing models described in the tropical and subtropical estuarine fish literature since its main channel fish assemblages remained within its bounds even during the rainy season.     

Genetic variation and population structuring in two brooding coral species (<Emphasis Type="Italic">Siderastrea stellata</Emphasis> and <Emphasis Type="Italic">Siderastrea radians</Emphasis>) from Brazil

Siderastrea stellata and S. radians are scleractinian coral species that present a remarkable overlap of diagnostic characteristics and sympatric distribution. Moreover, both are viviparous with similar reproductive strategies and with a gregarious larval behavior. Samples of both species from the Brazilian coast were analyzed using 18 isozymic loci to quantify their genetic variability and populational structure. Results confirmed species identity, high intrapopulational variability and revealed moderate genetic structuring among all samples (S. stellata: F (ST) = 0.070; S. radians: F (ST) = 0.092). Based on genotypic diversity analysis, there was evidence that local recruitment may have a minor role in the populations (mean, G ( o ):G ( e )= 1.00 +/- 0.0003 SD for S. stellata and 0.99 +/- 0.0023 SD for S. radians). Deviations towards heterozygote deficiencies found in both Siderastrea species could be explained by the Wahlund effect, since there was evidence that populations might be composed of colonies of different ages. In S. radians it is also likely that there is some inbreeding occurring in the studied populations. Despite the brooding pattern and the gregarious larval behavior, our data suggest the occurrence of gene flow along the Brazilian coast. This is the first study on population genetics of Brazilian reef corals.     

Predictions of establishment risk highlight biosurveillance priorities for invasive fish in New Zealand lakes

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Influences of the reproductive cycle on condition of marine catfishes (Siluriformes,Ariidae) in a coastal area at southeastern Brazil

Marine catfishes use estuaries and bays as part of their life cycle for spawning and feeding purposes, respectively. We compared three species of the family Ariidae in two different environmental conditionsa relatively stable bay site and an estuarine site, which differ in their magnitudes of environmental fluctuation to investigate effects of spawning on condition. We found differences in the parameter b (slope) and in they-intercept (a) among species and sexes. The parameter a varied inversely with b, and thus is not a good indicator to express fish condition. Condition factor did not differ between the two sites for species and sexes when we consider the pooled data. Overall condition was associated with reproductive process, and was high before the reproductive season, decreasing during the spawning period in the estuary, then increasing when fish move to the bay (April–August). Temporal segregation in the reproductive period and small-scale movements between the estuary and the bay are the probable mechanisms to optimize the use of the area by the three species.     

A phylogeographical analysis across three biogeographical provinces of the south-eastern Pacific: the case of the marine gastropod Concholepas concholepas

Aim The present‐day population structure of a species reflects the influence of population history as well as contemporary processes. Little is known about the mechanisms that have shaped the geographical distribution of genetic diversity in marine species present on the south‐eastern Pacific (SEP) coast. Here we provide the first comprehensive phylogeographical study of a species distributed along the SEP coast by analysing the endemic and emblematic muricid gastropod Concholepas concholepas. Location The study localities were distributed along the SEP coast ranging from Matarani (11° S) to Puerto Eden (49° S), crossing three major biogeographical provinces: Peruvian Province, Intermediate Area and Magellanic Province. Methods A total of 337 specimens of C. concholepas were collected from 14 localities in the three biogeographical provinces/areas. Mitochondrial cytochrome oxidase I (COI) gene partial sequences (658 bp) were obtained and analysed using coalescence‐based methods to infer molecular diversity and phylogeographical patterns. Results Across the 337 individuals, we found a large diversity, with a total of 179 haplotypes at the COI gene fragment. Although a slight decrease in gene diversity was observed from north to south, an analysis of molecular variance did not reveal any significant spatial population differentiation from Peru to the tip of Chile, not even across the recognized biogeographical boundaries at 30° S and 42° S. In addition, a star‐like haplotype network suggested the past occurrence of a rapid demographic and geographical expansion over the total range examined. Calculations of the onset of this expansion suggest that it might be due to climatic conditions during the period of the marine isotope stage 11 (MIS 11, 400,000 years ago), the longer and warmer interglacial episode during the Pleistocene epoch. Main conclusions Our phylogeographical analyses indicate that in the recent past C. concholepas mitochondrial DNA lineages underwent a sudden population expansion event. In addition, our data do not support the hypothesis of concordance between biogeographical barriers and phylogeographical breaks along the SEP coast. These two results are in accordance with the paradigm of high larval dispersal ability in marine species with an extended pelagic larval phase.     

Environmental interpretation using insoluble residues within reef coral skeletons: problems,pitfalls, and preliminary results

Insoluble residue concentrations have been measured within colonies of four massive reef corals from seven localities along the Caribbean coast of Panama to determine if detrital sediments, incorporated within the skeletal lattice during growth, record changes in sedimentation over the past twenty years. Amounts of resuspended sediment have increased to varying degrees at the seven localities over the past decades in response to increased deforestation in nearby terrestrial habitats. Preliminary results of correlation and regression analyses reveal few consistent temporal trends in the insoluble residue concentration. Analyses of variance suggest that amounts of insoluble residues, however, differ among environments within species, but that no consistent pattern of variation exists among species. D. strigosa and P. astreoides possess high concentrations at protected localities, S. siderea at localities with high amounts of resuspended sediment, and M. annularis at the least turbid localities. Little correlation exists between insoluble residue concentration and growth band width within species at each locality. Only in two more efficient suspension feeders (S. siderea and D. strigosa) do weak negative correlations with growth band width exist overall.These results indicate that insoluble residue concentrations cannot be used unequivocally in environmental interpretation, until more is known about tissue damage, polyp behavior, and their effects on the incorporation of insolubles in the skeleton during growth in different coral species. Insoluble residue data are highly variable; therefore, large sample sizes and strong contrasts between environments are required to reveal significant trends.     

Diversity of tiger moths in a Neotropical hotspot: determinants of species composition and identification of biogeographic units

The Brazilian Atlantic Forest is a Biodiversity Hotspot, yet many biological groups in this biome are poorly known. We compiled information on the diversity of Atlantic Forest tiger moths (Arctiidae) and assessed the resemblance among localities, whether arctiid assemblages are concordant with major vegetation types, and the importance of environmental factors in structuring the variation among assemblages. Additionally, we developed a procedure composed of subsampling and procrustean analysis to assess the robustness of the results from community composition ordinations when localities differ in species richness. To do this, we built a database from specimens deposited in the ten most important Brazilian entomological collections, and mapped species richness in one-degree latitude/longitude grid cells. We employed Principal Coordinates Analysis to assess similarities among the best-sampled localities. We obtained 8,667 records including 1,193 species, representing 60 and 20% of the estimated Brazilian and Neotropical faunas, respectively. Our subsampling procedure indicated that the ordination was not greatly affected by differences in species richness, and was congruent with major vegetation types. Lowland localities on the seacoast were quite distinct in species composition. A second group included localities in montane areas in the southeast part of the biome. The last group included localities in the southern region and those with Araucaria forests, and was associated with long distances from the ocean, even distribution of precipitation throughout the year, and large annual temperature ranges.