Inferred global connectivity of whale shark Rhincodon typus populations

Ten years have passed since the last synopsis of whale shark Rhincodon typus biogeography. While a recent review of the species' biology and ecology summarized the vast data collected since then, it is clear that information on population geographic connectivity, migration and demography of R. typus is still limited and scattered. Understanding R. typus migratory behaviour is central to its conservation management considering the genetic evidence suggesting local aggregations are connected at the generational scale over entire ocean basins. By collating available data on sightings, tracked movements and distribution information, this review provides evidence for the hypothesis of broad‐scale connectivity among populations, and generates a model describing how the world's R. typus are part of a single, global meta‐population. Rhincodon typus occurrence timings and distribution patterns make possible a connection between several aggregation sites in the Indian Ocean. The present conceptual model and validating data lend support to the hypothesis that R. typus are able to move among the three largest ocean basins with a minimum total travelling time of around 2–4 years. The model provides a worldwide perspective of possible R. typus migration routes, and suggests a modified focus for additional research to test its predictions. The framework can be used to trim the hypotheses for R. typus movements and aggregation timings, thereby isolating possible mating and breeding areas that are currently unknown. This will assist endeavours to predict the longer‐term response of the species to ocean warming and changing patterns of human‐induced mortality.     

Northernmost record of a whale shark Rhincodon typus from the Sea of Okhotsk

The whale shark Rhincodon typus is the world's largest fish and it occurs in tropical, subtropical and warm temperate waters. Here, the northernmost record of R. typus is reported, when it was found in the Sea of Okhotsk for the first time. This occurrence can be explained by the unusually high sea surface temperature during the summer of 2012.     

New insights into whale shark <Emphasis Type="Italic">Rhincodon typus</Emphasis> diet in Brazil: an observation of ram filter-feeding on crab larvae and analysis of stomach contents from the first stranding in Bahia state

We describe the first record of a whale shark, Rhincodon typus, feeding in Brazilian coastal waters, and the first stranding record in the state of Bahia, Northeast Brazil. In April 2008, an individual of R. typus was observed surface feeding on Dromiidae crab larvae in the continental shelf off the coast of Bahia, near a gas platform. Other fishes were observed foraging in association with the whale shark. We also document the first stranding of R. typus on the coast of Bahia in October 2013. Biometric data confirmed that the stranded whale shark was a juvenile. Stomach content analysis revealed the ingestion of Geryonidae crab larvae. Plastic debris were also found in the gastric lumen of the stranded juvenile whale shark, and we speculate that it could have been a contributing factor to the stranding, and subsequent death of the whale shark. Crab larvae were observed in both of our records and likely to consist as relevant prey items for R. typus in Brazilian continental shelf. Our study provided a contribution on the diet and feeding behaviour of whale sharks in tropical oligotrophic waters and highlights the risks of marine pollution for the species conservation.     

Do they stay or do they go? Acoustic monitoring of whale sharks at Ningaloo Marine Park,Western Australia

Whale sharks Rhincodon typus were monitored via acoustic transmitters at the northern end of Western Australia's Ningaloo Marine Park to establish the extent to which the species inhabits the region beyond the whale‐shark ecotourism industry season, which usually extends from March to August in each year. Despite the vast majority (c. 98%) of photographic submissions of R. typus from Ningaloo Reef being between March and August, acoustic detections from the tagged R. typus at Ningaloo were recorded in all months of the year, but do not preclude the occurrence of extended absences. It is concluded that as a species, R. typus occurs year round at Ningaloo, where it generally remains in close proximity to the reef edge, but that some individuals move outside of the detection range of the array for extended periods.     

A new record of whale shark Rhincodon typus in Brazilian waters: a report of association with Caranx crysos

In May 2011, a Rhincodon typus was sighted on the continental shelf of the central Brazilian coast, in the vicinity of a gas platform. During the video record, an interspecific following association was observed between a Caranx crysos school and the R. typus.     

Population structure and residency of whale sharks Rhincodon typus at Utila,Bay Islands,Honduras

There were 479 reported whale shark Rhincodon typus encounters between 1999 and 2011 at the island of Utila, which forms part of the Meso‐American Barrier Reef System (MBRS) in the western Caribbean Sea. The majority of R. typus were found to feed on small bait fish associated with various tuna species. Ninety‐five individual R. typus, ranging from 2 to 11 m total length (L_T), were identified through their unique spot patterns. A significant male bias (65%) was present. There was no significant difference between the mean ± s.d . L_T of female (6·66 ± 1·65 m) and male (6·25 ± 1·60 m) R. typus. Most R. typus were transient to Utila, with 78% sighted only within a single calendar year, although some individuals were sighted in up to 5 years. Mean residency time was modelled to be 11·76 days using maximum likelihood methods.     

Distribution and feeding habitat characterization of whale sharks Rhincodon typus in a protected area in the north Caribbean Sea

The relationship between the distribution of the whale shark Rhincodon typus and hydrobiological variables in the Caribbean Sea during 2005–2009 was analysed. Monthly trips were made to the R. typus aggregation area during the months when this species is present in the region (May to September) to record sightings and hydrological data and to collect samples to determine nutrients, chlorophyll a (Chl a) and zooplankton biomass. A total of 2104 R. typus were counted and three zones of high abundance were identified: Cabo‐Catoche, Contoy (both within the Whale Shark Biosphere Reserve, WSBR) and the zone knows as Afuera. The zones of greatest R. typus density within the WSBR were characterized by high Chl a concentrations (median: 1·1 mg m^?3, interpercentile range: 0·5–1·8 mg m^?3) and high nutrient concentrations, such as ammonium (median: 2·5 µmol l^?1, interpercentile range: 0·5–6·4 µmol l^?1), due to the influence of local upwelling. A generalized additive model (GAM) was used to explore the relationship between R. typus distribution and the environmental variables inside WSBR. Zooplankton biomass was the most influential environmental variable, supporting the close relationship between R. typus distribution and biological productivity. Copepods were the dominant zooplankton group within the WSBR. In the Afuera zone, there were large R. typus aggregations (>80 individuals) associated with zooplankton dominated by fish eggs and significantly higher mean ± s.d. biomass (3356·1 ± 1960·8 mg m^?3) compared with that recorded inside the WSBR (103·5 ± 57·2 mg m^?3). The differences among zones generated changes in R. typus distribution patterns and provided opportunities to develop local management strategies for this species.     

Age and growth parameters of shark‐like batoids

Estimates of life‐history parameters were made for shark‐like batoids of conservation concern Rhynchobatus spp. (Rhynchobatus australiae, Rhynchobatus laevis and Rhynchobatus palpebratus) and Glaucostegus typus using vertebral ageing. The sigmoid growth functions, Gompertz and logistic, best described the growth of Rhynchobatus spp. and G. typus, providing the best statistical fit and most biologically appropriate parameters. The two‐parameter logistic was the preferred model for Rhynchobatus spp. with growth parameter estimates (both sexes combined) L_∞ = 2045 mm stretch total length, L_ST and k = 0·41 year^?1. The same model was also preferred for G. typus with growth parameter estimates (both sexes combined) L_∞ = 2770 mm L_ST and k = 0·30 year^?1. Annual growth‐band deposition could not be excluded in Rhynchobatus spp. using mark‐recaptured individuals. Although morphologically similar G. typus and Rhynchobatus spp. have differing life histories, with G. typus longer lived, slower growing and attaining a larger maximum size.     

Comparison of the lateral line and ampullary systems of two species of shovelnose ray

The anatomical characteristics of the mechanoreceptive lateral line system and electrosensory ampullae of Lorenzini of Rhinobatos typus and Aptychotrema rostrata are compared. The spatial distribution of somatic pores of both sensory systems is quite similar, as lateral line canals are bordered by electrosensory pore fields. Lateral line canals form a sub-epidermal, bilaterally symmetrical net on the dorsal and ventral surfaces; canals contain a nearly continuous row of sensory neuromasts along their length and are either non-pored or pored. Pored canals are connected to the surface through a single terminal pore or additionally possess numerous tubules along their length. On the dorsal surface of R. typus, all canals of the lateral line occur in the same locations as those of A. rostrata. Tubules branching off the lateral line canals of R. typus are ramified, which contrasts with the straight tubules of A. rostrata. The ventral prenasal lateral line canals of R. typus are pored and possess branched tubules in contrast to the non-pored straight canals in A. rostrata. Pores of the ampullae of Lorenzini are restricted to the cephalic region of the disk, extending only slightly onto the pectoral fins in both species. Ampullary canals penetrate subdermally and are detached from the dermis. Ampullae occur clustered together, and can be surrounded by capsules of connective tissue. We divided the somatic pores of the ampullae of Lorenzini of R. typus into 12 pore fields (10 in A. rostrata), corresponding to innervation and cluster formation. The total number of ampullary pores found on the ventral skin surface of R. typus is approximately six times higher (four times higher in A. rostrata) than dorsally. Pores are concentrated around the mouth, in the abdominal area between the gills and along the rostral cartilage. The ampullae of both species of shovelnose ray are multi-alveolate macroampullae, sensu Andres and von Düring (1988). Both the pore patterns and the distribution of the ampullary clusters in R. typus differ from A. rostrata, although a basic pore distribution pattern is conserved.     

Whale shark Rhincodon typus strandings in the Gulf of California,Mexico

The present study analyses whale shark Rhincodon typus stranding in the Gulf of California, Mexico, reported by the public, scientists, authorities and artisanal fishermen. A total of 14 strandings were documented during the period 2001–2018. The total length of R. typus ranged from 350–1,102 cm, with a sex ratio of 3.5:1 (males: females). This study highlights potential stranding areas, the need for better stranding protocols to assist R. typus recovery and the importance of samples from dead animals for scientific research.     

Hoarding patterns of sigmodontine rodent species in the Central Monte Desert (Argentina)

Hoarding food is an important strategy of rodents in desert environments characterized by unpredictable and poor food resource availability. In the Monte Desert, Prosopis produces abundant food, unevenly in time and space, in the form of pods and seeds. Sigmodontine rodents (Graomys griseoflavus, Akodon molinae, Eligmodontia typus and Calomys musculinus) use Prosopis propagules extensively, and they could be predators or dispersers depending on how they handle and where they leave the propagules. The objectives of this study were: (1) to know what rodent species transported propagules; (2) to evaluate what hoarding pattern was used by species that transport propagules (larder and scatterhoarding); and (3) to analyse in which condition were propagules left by the rodent species, both at the food source and in caches. Our results showed that all four species transported propagules, with G. griseoflavus and E. typus being the species that carried more seeds. Our study supported the evidence that food caching is common among species and that many species both larderhoard and scatterhoard food. Graomys griseoflavus and A. molinae, the largest species, larderhoarded more than did the smaller E. typus. These results uphold the hypothesis that larger species will show greater propensity to larderhoard than smaller species. Considering the interaction between seed‐hoarding patterns and plants, E. typus was the species that could most improve germination because it scatterhoarded propagules and left seeds out of pods. In contrast, G. griseoflavus could have a negative impact on plant populations because this was the species that predated more seeds and larderhoarded a high percentage of them. The smallest C. musculinus was the species that transported propagules least, and left them as seeds inside pods or pod segments mainly at the food source, which makes seeds more vulnerable to predation.     

Seasonal trends in whale shark Rhincodon typus sightings in an established tourism site in the Gulf of California,Mexico

The number of individual whale shark Rhincodon typus sightings registered October 2015–March 2018 in Bahia de La Paz, Mexico, ranged from 73 to 129 animals per season (total 1662), with the majority of them identified as males. Density plot maps showed high correlations in sightings between sample seasons and revealed potential hotspot areas. Our study provides an essential baseline of information for the management of whale shark conservation and the associated touristic activities in the region.     

A new species of Dermopristis Kearn, Whittington & Evans-Gowing, 2010 (Monogenea: Microbothriidae), with observations on associations between the gut diverticula and reproductive system and on the presence of denticles in the nasal fossae of the host Glaucostegus typus (Bennett) (Elasmobranchii: Rhinobatidae)

Dermopristis cairae n. sp. (Monogenea: Microbothriidae) is described from the skin and possibly from the nasal fossae of the giant shovel-nosed ray Glaucostegus typus (Bennett). The new species is distinguished from D. paradoxus Kearn, Whittington & Evans-Gowing, 2010 by its larger size, body shape, lack of transverse ridges on the ventral surface and absence of a seminal receptacle. Extensive short gut branches lie dorsal to the testes and adjacent to the coiled region of the vas deferens and the o?type, possibly reflecting high metabolic demand in these areas. Denticles are present in the lining of the nasal fossae of G. typus, providing a firm substrate for the cement-based attachment of a microbothriid. However, confirmation that D. cairae inhabits the nasal fossae of G. typus is required.     

Acoustic telemetry reveals cryptic residency of whale sharks

Although whale sharks (Rhincodon typus) have been documented to move thousands of kilometres, they are most frequently observed at a few predictable seasonal aggregation sites. The absence of sharks at the surface during visual surveys has led to the assumption that sharks disperse to places unknown during the long ‘off-seasons’ at most of these locations. Here we compare 2 years of R. typus visual sighting records from Mafia Island in Tanzania to concurrent acoustic telemetry of tagged individuals. Sightings revealed a clear seasonal pattern with a peak between October and February and no sharks observed at other times. By contrast, acoustic telemetry demonstrated year-round residency of R. typus. The sharks use a different habitat in the off-season, swimming deeper and further away from shore, presumably in response to prey distributions. This behavioural change reduces the sharks'' visibility, giving the false impression that they have left the area. We demonstrate, for the first time to our knowledge, year-round residency of unprovisioned, individual R. typus at an aggregation site, and highlight the importance of using multiple techniques to study the movement ecology of marine megafauna.     

Population structure of a whale shark Rhincodon typus aggregation in the Red Sea

The presence of whale sharks Rhincodon typus were recorded around Shib Habil, a small, coastal reef off the Red Sea coast of Saudi Arabia, from 2010 to 2015. A total of 267 suitable photographs resulting in the identification of 136 individuals, were documented from 305 encounters. Sharks were divided evenly between the sexes with no evidence of temporal or spatial segregation. All individuals were immature based on size estimates and, for males, juvenile clasper morphology. Scars were reported for 57% of R. typus with 15% showing evidence of propeller trauma. Estimates of population size and patterns of residency were calculated by modelling the lagged identification rate. Multiple models were run simultaneously and compared using the Akaike information criterion. An open population model was found to best represent the data and estimates a daily abundance between 15 and 34 R. typus during the aggregation season, with local residence times ranging from 4 to 44 days. Residence times away from Shib Habil range from 15 to 156 days with a permanent emigration–death rate between 0·07 and 0·58 individuals year^?1. These results are broadly similar to those from other aggregations of R. typus, although the observed sexual parity and integration found at this site is unique for the species and needs further study.     

Association of the whale shark Rhincodon typus with the cannonball jellyfish Stomolophus meleagris

This study describes the first record of the whale shark Rhincodon typus association with the cannonball jellyfish Stomolophus meleagris in the Gulf of California, Mexico. Whale sharks were observed swimming and feeding among swarms of jellyfish, suggesting competition and predatory behaviour given the overlap in food preferences between both species. This finding is relevant because of the species‐wide distribution and the importance of these interactions, which should be considered in conservation strategies of R. typus and management of cannonball jellyfish fisheries.     

Taxonomic revision of the genus Cutleria proposing a new genus Mutimo to accommodate M. cylindricus (Cutleriaceae,Phaeophyceae)

Molecular phylogenetic analyses of representative Cutleria species using mitochondrial cox3, chloroplast psaA, psbA and rbcL gene sequences showed that C. cylindrica Okamura was not included in the clade composed of other Cutleria species including the generitype C. multifida (Turner) Greville and the related taxon Zanardinia typus (Nardo) P.C. Silva. Instead, C. cylindrica was sister to the clade composed of the two genera excluding C. cylindrica. Cutleria spp. have heteromophic life histories and their gametophytes are rather diverse in gross morphology, from compressed or cylindrical‐branched to fan‐shaped, whereas the sporophytes are rather similar. In contrast, the monotypic species Z. typus has an isomorphic life history and resembles fan‐shaped Cutleria in morphology. Morphological comparisons of these taxa revealed that C. cylindrica is morphologically distinct from other Cutleria spp. and Z. typus in having cylindrical gametophytes with multiseriate trichothallic filaments instead of uniseriate filaments (hairs) characteristic of Cutleriales (or Cutleriaceae, Tilopteridales), and in lacking rhizoidal filaments in the crustose sporophytes. Therefore, based on the molecular and morphological data, the establishment of a new genus Mutimo to accommodate C. cylindrica, and the new combination of M. cylindricus, is proposed.     

Locomotion and Escape Modes in Rodents of the Monte Desert (Argentina)

Modes of locomotion and escape tactics are attributes that affect the structure of animal communities, promoting exploitation of different microhabitats and the coexistence of different species. Bipedal locomotion is considered to be more effective than a quadrupedal gait in escaping attacks by predators because it allows for higher speed, a faster response to attack, sudden changes of direction and better detection of aerial raptors. The aim of this study was to determine the type of locomotion used at the moment of escape by three rodent species of the Monte desert –Eligmodontia typus, Akodon molinae and Graomys griseoflavus. The study was carried out in three plant communities of the Ñacuñán Reserve (Mendoza). All three species showed differences in both mode of escape and locomotory pattern. Graomys griseoflavus exhibited the highest proportion of escapes using quadrupedal saltation. The mode of locomotion employed by E. typus varied according to the type of plant communities it inhabited. Those occurring at open sites (Medanal community) exhibited a greater propensity to jump during escapes than those from more sheltered habitats (Algarrobal community). Akodon molinae relied primarily on a quadrupedal gait when fleeing from predators, which would explain its greater dependence on plant cover. Therefore, the morphological and behavioural characteristics of these species are related to their mode of locomotion and the strategies they employ to diminish the risk of predation.     

Sperm-egg interactions in the shrimp Rhynchocinetes typus

Sperm-egg interaction in Rhynchocinetes typus was studied with the phase-contrast and scanning electron microscopes. R typus spermatozoa present in the vas deferens have the shape of a round-headed nail. After contact with seawater it is possible to observe the unfolding of the rays or stellate arms, giving the spermatozoon the appearance of an inverted umbrella. From the center of the flat face of the umbrella emerges a spike with longitudinal striations. Ovarian eggs and spermatozoa were mixed in vitro by agitating them for two minutes in Millipore-filtered seawater. The first gamete contact was established by the spermatozoon through the tip of the spike, which exerted a lytic action on the egg envelopes. After the rigid spike was completely inside the egg, the rays became aligned parallel to each other and began to enter the eggs. Toward the final stages of ray entry, it was possible to observe fusion of the ray membranes with one another, and later the fusion process continued toward the tip of the radial spines. Concomitantly, the egg surface that surrounds the sperm swelled in a circular fashion and formed a fertilization cone. After the spermatozoon entry was complete, a scarlike mark appeared at the place on the egg surface through which penetration occurred. The whole penetration process was completed within 45-60 minutes.     

Protein polymorphism inEligmodontia typus. Genetic divergence with other phyllotine cricetids

By means of gel electrophoresis of tissue extracts we have studied protein polymorphism inEligmodontia typus. The analysis was performed on specimens from five population samples collected at different sites in Patagonia (Argentina). Mean heterozygosity (\-h) and proportion of polymorphic loci (P) were determined on the basis of 19 loci. Considering all individuals as one sample, \-h gave a value of 0.16 and P was 70%. Although these values are much higher than those reported for most rodent species, they are very similar to those obtained by us for four species of the genusCalomys and forGraomys griseoflavus. There is a striking genetic identity (I_N=0.99) among populations from regions with different environmental conditions, indicating that the species possesses a common genic pool. Genetic distance with other species of the Phyllotini was estimated. D_N was lower betweenE. typus andCalomys (mean D_N=0.88) than betweenE. typus andGraomys griseoflavus (D_N=1.01). The high morphological similarity between these last two species, especially regarding those characters related to desert life adaptation, could be assigned, at least in part, to convergent evolution.