Risks and advantages of using surface laser photogrammetry on free‐ranging marine organisms: a case study on white sharks Carcharodon carcharias

This study employed a non‐lethal measurement tool, which combined an existing photo‐identification technique with a surface, parallel laser photogrammetry technique, to accurately estimate the size of free‐ranging white sharks Carcharodon carcharias. Findings confirmed the hypothesis that surface laser photogrammetry is more accurate than crew‐based estimations that utilized a shark cage of known size as a reference tool. Furthermore, field implementation also revealed that the photographer's angle of reference and the shark's body curvature could greatly influence technique accuracy, exposing two limitations. The findings showed minor inconsistencies with previous studies that examined pre‐caudal to total length ratios of dead specimens. This study suggests that surface laser photogrammetry can successfully increase length estimation accuracy and illustrates the potential utility of this technique for growth and stock assessments on free‐ranging marine organisms, which will lead to an improvement of the adaptive management of the species.     

Comparison of fin ray sampling methods on white sturgeon Acipenser transmontanus growth and swimming performance

Effects of two fin‐ray sampling methods on swimming performance, growth and survival were evaluated for hatchery‐reared sub‐adult white sturgeon Acipenser transmontanus. Fish were subjected to either a notch removal treatment in which a small section was removed from an anterior marginal pectoral‐fin ray, or a full removal treatment in which an entire marginal pectoral‐fin ray was removed. Control fish did not have fin rays removed, but they were subjected to a sham operation. A modified 3230 l Brett‐type swim tunnel was used to evaluate 10 min critical station‐holding speeds (S_CSH) of A. transmontanus, immediately after the fin ray biopsies were obtained with each method. Survival and growth were evaluated over a 6 month period for a separate group of fish subjected to the same biopsy methods. Mean ± s.e . 10 min S_CSH were 108·0 ± 2·3, 110·0 ± 2·6 and 115·0 ± 3·5 cm s^?1 for the notch removal group, full removal group and control group, respectively, and were not significantly different among treatments. Behavioural characteristics including tail‐beat frequency and time spent hunkering were also not significantly different among treatment groups swimming at the same speeds. There were no mortalities and relative growth was similar among treatment groups. Average biopsy time for the notch removal method was lower and the wounds appeared to heal more quickly compared with the full removal method.     

Validation of daily microincrement deposition in otoliths of juvenile and adult Peruvian anchovy Engraulis ringens

Wild adult specimens of the Peruvian anchovy Engraulis ringens were captured and reared to validate the daily periodicity of otolith microincrement formation. The postcapture stress generated spontaneous spawning, making it possible to conduct a rearing trial on larvae first in an artificial nutrient‐enriched system (ANES) for 52 days followed by an artificial feeding regime in a culture tank until day 115 post‐hatch. Microincrements of the sagittal otoliths of sacrificed juveniles [mean ± s.d. total length (L_T) = 5·13 ± 0·37 cm, range 5–6 cm; c.v. = 7·5%] showed very distinct light and dark zones. The slope of the relationship between the total number of increments after the hatch check and days elapsed after hatching was not significantly different from 1. The transfer from ANES to the artificial feeding regime induced a mark in the sagittal otoliths. The number of microincrements after this induced mark coincided with the number of days elapsed after the transfer date. In parallel experiments, adult E. ringens (mean ± s.d. L_T = 14·92 ± 0·55 cm, range 13–16 cm) were exposed to one of two fluorescent marking immersion treatments with either alizarin red S (ARS; 25 mg l^?1 per 6 h) or oxytetracycline hydrochloride (OTC; 200 mg l^?1 per 10 h). The microincrements between fluorescent bands were distinct, ranging from 0·89 to 2·75 µm (mean ± s.d. =1·43 ± 0·28 µm; c.v. = 32%) and from 0·71 to 2·89 µm (1·53 ± 0·27 µm; c.v. = 35%) for ARS and OTC, respectively. The relationship between the number of microincrements between marks and the number of elapsed days for ARS and OCT treatments indicated that there was a significant correspondence between the number of increases observed and the number of days. Hence, daily microincrements of otoliths of E. ringens are likely to be formed in juveniles and adults under natural conditions.     

Discovery of a multispecies shark aggregation and parturition area in the Ba Estuary,Fiji Islands

Population declines in shark species have been reported on local and global scales, with overfishing, habitat destruction and climate change posing severe threats. The lack of species‐specific baseline data on ecology and distribution of many sharks, however, makes conservation measures challenging. Here, we present a fisheries‐independent shark survey from the Fiji Islands, where scientific knowledge on locally occurring elasmobranchs is largely still lacking despite the location's role as a shark hotspot in the Pacific. Juvenile shark abundance in the fishing grounds of the Ba Estuary (north‐western Viti Levu) was assessed with a gillnet‐ and longline‐based survey from December 2015 to April 2016. A total of 103 juvenile sharks identified as blacktip Carcharhinus limbatus (n = 57), scalloped hammerhead Sphyrna lewini (n = 35), and great hammerhead Sphyrna mokarran (n = 11) sharks were captured, tagged, and released. The condition of umbilical scars (68% open or semihealed), mean sizes of individuals (±SD) (C. limbatus: 66.5 ± 3.8 cm, S. lewini: 51.8 ± 4.8 cm, S. mokarran 77.4 ± 2.8 cm), and the presence of these species over recent years (based on fishermen interviews), suggest that the Ba Estuary area is a critical habitat for multiple species that are classified as “Near Threatened” or “Endangered.” Specifically, the area likely acts as a parturition ground over the studied period, and potentially as a subsequent nursery area. We identified subareas of high abundance and found that temperature, salinity and depth acted as small‐scale environmental drivers of shark abundance. The data suggests a tendency for species‐specific spatial use, both horizontally (i.e., between sampling areas) and vertically (i.e., across the water column). These results enhance the understanding of shark ecology in Fiji and provide a scientific basis for the implementation of local conservation strategies that contribute to the protection of these threatened species.     

The effect of spine postures on the hydrodynamic drag in Epinephelus ongus larvae

Laboratory behavioural observation and computational fluid dynamics (CFD) analysis were conducted to examine whether the movement of the elongated dorsal and pelvic spines changed the hydrodynamic drag in white‐streaked grouper Epinephelus ongus larvae. The behavioural observation in the tank revealed that the larvae extended the dorsal and pelvic spines during passive transport and retracted during swimming; the angles of the dorsal and pelvic spines in relation to the anteroposterior axis were larger during the passive transport (mean ± s.d . = 28·84 ± 14·27 and 20·35 ± 15·05°) than those during the swimming (mean ± s.d . = 2·59 ± 5·55 and 0·32 ± 6·49°). The CFD analysis indicated that the relative hydrodynamic drag acting on the larvae was approximately 1·25 times higher when the spines were extended (passive transport) than when the spines were retracted (swimming), suggesting that the E. ongus larvae have an ability to adjust their hydrodynamic drag depending on the behavioural context.     

Scarring patterns and relative mortality rates of Indian Ocean whale sharks

This study recorded the scarring rate and severity for whale sharks Rhincodon typus from three Indian Ocean aggregations (Australia, Seychelles and Mozambique), and examined whether scarring (mostly attributed to boat strikes and predator attacks) influences apparent survival rates using photo‐identification libraries. Identifications were based on spot‐and‐stripe patterns that are unique to individual whale sharks. Scarring was most prevalent in the Seychelles aggregation (67% of individuals). Predator bites were the most frequent source of scarring (aside from minor nicks and abrasions) and 27% of individuals had scars consistent with predator attacks. A similar proportion of whale sharks had blunt trauma, laceration and amputation scars, the majority of which appeared to be caused by ship collisions. Predator bites were more common (44% of individuals) and scars from ship collisions were less common at Ningaloo Reef than at the other two locations (probability of among‐site differences occurring randomly = 0·0007 based on a randomized multinomial contingency analysis). In all aggregations, scars occurred most often on the caudal fin, which may result from the fin being the body part closest to the surface when boats pass over, or they may provide a large target for predator attack. No evidence was found for an effect of scarring on apparent survival (φ; mean ±s .e .) for the Ningaloo (not scarred φ= 0·858 ± 0·033; scarred φ= 0·929 ± 0·033) or Seychelles populations (not scarred φ= 0·502 ± 0·060; scarred φ= 0·538 ± 0·070). The lower apparent survival of the Seychelles population may be attributed to a high number of transient whale sharks in this aggregation that might bias estimates. This study indicates that while scarring from natural predators and smaller vessels appears to be unrelated to whale shark survival, the effect of deaths related to ship strike need to be quantified to assist in future management of this species.     

Morpho‐histological and ultra architectural changes during early development of endangered golden mahseer Tor putitora

Ultrastructural and histological changes in the embryonic and larval surface during ontogenesis of the endangered golden mahseer Tor putitora is studied here for the first time. Embryonic development was completed 91–92 h after fertilization at an ambient temperature of 23° ± 1° C (mean ± s.d. ). The gastrula stage was characterized by presence of the Kupffer's vesicle, notochord, ectoderm and endoderm cells. Primordial germ cells were clearly identifiable from c. 55 h post‐fertilization at the organogenesis stage. Mean total length of newly hatched larvae was 7·0 ± 0·5 mm. Scanning electron microscopy of newly hatched larvae demonstrated vitelline arteries, microridged epithelial cells and mucous gland openings over much of the body surface. Eye, oral cavity, pharyngeal arches, heart, intestinal loop, prosencephalon, cephalic vesicle and nasal epithelium were clearly distinguished in 3 day old hatched individuals. In 6 day old individuals, caudal‐fin rays and internal organs were evident. The dorsal fin became prominent at this stage and larvae began swimming at the surface. The reserved yolk material was totally absorbed 8–11 days after hatching and larvae began feeding exogenously. Tor putitora exhibited a longer early developmental period than other cyprinids reared at similar temperatures.     

Population biology of an endangered species: the common guitarfish Rhinobatos rhinobatos in Lebanese marine waters of the eastern Mediterranean Sea

This study focuses on the population biology of the common guitarfish Rhinobatos rhinobatos, a cartilaginous fish listed as Endangered in the International Union for the Conservation of Nature (IUCN) Red List. Between December 2012 and January 2014, 67 individuals were collected by bottom longlining in coastal Lebanese marine waters at different ports at depths ranging from 10 to 110 m. The total length (L_T) of the specimens ranged from 50 to 143 cm, and the mean ± s.d . was 76·2 ± 19·7 cm. The most common L_T classes were between 60 and 70 cm. The total mass of the specimens ranged from 410 to 10 000 g, and the mean ± s.d . was 1841 ± 1987 g. A total of 34 males and 33 females were collected, and the sex ratio was not significantly different from 1:1. The mass and L_T relationship showed positive allometric growth (b = 3·096 and r² = 0·99), and the mean ± s.d . L_T at which 50% of the individuals were sexually mature was 84·73 ± 5·81 cm for females and 78·57 ± 4·88 cm for males. The gonado‐somatic and hepato‐somatic indices were determined along with a condition factor, and parturition appeared to occur in winter. The primary prey items found in the fish stomachs during the autumn and winter seasons were Penaeidae. The results of this study will help to parameterize models of the population dynamics for this exploited fish stock to ensure the long‐term sustainability of its fishery.     

Larval and juvenile development of dusky grouper Epinephelus marginatus reared in mesocosms

The larval development of the dusky grouper Epinephelus marginatus up to the benthic juvenile stage is described in detail to establish a reference for their larval identification. Development is described in terms of ontogenetic changes in morphology, growth, pigmentation, fin structure and skeletal structure. Larvae were reared in mesocosms at a mean temperature of 24·3° C, salinity of 36·5, dissolved oxygen of 6·4 mg l^?1 and pH of 8·2. Newly hatched larvae had an estimated total length (L_T) of 2·3 mm. On the second day post hatching the yolk was almost fully absorbed with traces of the oil globule still present, the eyes were already pigmented and mouth and gut functional. At this stage the cranial skeletal elements for feeding and breathing (mouth and gills) and the pectoral‐fin support were already present. About 50% of the observed larvae had food in their guts. Pigmentation was very characteristic, consisting of two large chromatophores visible on the edge of the primordial fin, close to the midpoint of the post‐anal region of the body and over the midgut and hindgut and post‐anal portion of the body. At 2·9 mm L_T the emergence of the second dorsal‐fin spine, characteristic of the Epinephilinae, was clearly visible. The pre‐flexion stage started in larva of 3·2 mm L_T. At 5·5 mm L_T the larvae possessed posterior preopercular angle spines, and the dorsal and pelvic spines presented serrated edges and were pigmented. The water surface‐tension‐related death of the yolk sac and pre‐flexion larvae described in the rearing of several other grouper species did not occur during E. marginatus culture. Notochord flexion, with initial ossification of the caudal‐fin supporting elements, started at 6·6 mm L_T. At this stage the major melanophores, preopercular, dorsal and pelvic spines and mandibular teeth were already present. Transformation of larvae into juveniles occurred when larvae averaged 13·8 mm L_T. Juveniles with a mean L_T of 20·1 mm started to settle and most of them were benthic with a mean L_T of 26·8 mm.     

Validation of photo‐identification as a mark–recapture method in the spotted eagle ray Aetobatus narinari

The spotted eagle ray Aetobatus narinari is characterized by pigmentation patterns that are retained for up to 3·5 years. These pigmentations can be used to identify individuals through photo‐identification. Only one study has validated this technique, but no study has estimated the percentage of correct identification of the rays using this technique. In order to carry out demographic research, a reliable photographic identification technique is needed. To achieve this validation for A. narinari, a double‐mark system was established over 11 months and photographs of the dorsal surface of 191 rays were taken. Three body parts with distinctive natural patterns were analysed (dorsal surface of the cephalic region, dorsal surface of the pectoral fins and dorsal surface of the pelvic fins) in order to determine the body part that could be used to give the highest percentage of correct identification. The dorsal surface of the pectoral fins of A. narinari provides the most accurate photo‐identification to distinguish individuals (88·2%).     

Evaluating the effectiveness of teeth and dorsal fin spines for non‐lethal age estimation of a tropical reef fish,coral trout Plectropomus leopardus

This study investigated whether teeth and dorsal fin spines could be used as non‐lethal methods of age estimation for a vulnerable and highly valued tropical fisheries species, coral trout Plectropomus leopardus. Age estimation of individuals from 2 to 9 years old revealed that dorsal spines represent an accurate ageing method (90% agreement with otoliths) that was more precise [average per cent error (APE) = 4·1, coefficient of variation (c.v .) = 5·8%] than otoliths (APE = 6·2, c.v . = 8·7%). Of the three methods for age estimation (otoliths, dorsal spines and teeth), spines were the most time and cost efficient. An aquarium‐based study also found that removing a dorsal spine or tooth did not affect survivorship or growth of P. leopardus. No annuli were visible in teeth despite taking transverse and longitudinal sections throughout the tooth and trialling several different laboratory methods. Although teeth may not be suitable for estimating age of P. leopardus, dorsal spines appear to be an acceptably accurate, precise and efficient method for non‐lethal ageing of individuals from 2 to 9 years old in this tropical species.     

Tracking the migratory success of stocked European eels Anguilla anguilla in the Baltic Sea

To investigate the extent to which European silver eels Anguilla anguilla, originating from stocking programmes in the Baltic Sea tributaries, effectively contribute to the spawning stock, two hundred and seventy‐four formerly stocked A. anguilla. emigrating from the Schwentine River near Kiel, Germany, were tagged with T‐Bar anchor tags. A total of 29 Anguilla spp. were recaptured (c. 11%) up to 14 months after release. Stocking history of recaptured A. anguilla. was confirmed by otolith microchemistry. Recapture locations were concentrated around the outlet of the Baltic Sea (Danish Belt Sea) with 62% of all recaptures reported here or in the Kattegat. Recaptured Anguilla spp. showed a reduction in both L_T and mass (mean ± s.d . = ?1·5 ± 0·9 cm and ?125·3 ± 50·1 g) while average total fat content remained in the order of values previously reported as high enough to provide energy resources to allow successful completion of the spawning migration (mean ± s.d . = 28·4 ± 4·4%). The documented mean rate of travel (0·8 km day^?1), however, indicated a delay in the target‐oriented migration that might be interpreted as a delayed initial migration phase of orientation towards the exit of the Baltic Sea.     

Summer stream habitat partitioning by sympatric Arctic charr, Atlantic salmon and brown trout in two sub-arctic rivers

Summer habitat use by sympatric Arctic charr Salvelinus alpinus, young Atlantic salmon Salmo salar and brown trout Salmo trutta was studied by two methods, direct underwater observation and electrofishing, across a range of habitats in two sub-arctic rivers. More Arctic charr and fewer Atlantic salmon parr were observed by electrofishing in comparison to direct underwater observation, perhaps suggesting a more cryptic behaviour by Arctic charr. The three species segregated in habitat use. Arctic charr, as found by direct underwater observation, most frequently used slow (mean ±s .d . water velocity 7·2 ± 16·6 cm s⁻¹) or often stillwater and deep habitats (mean ±s .d . depth 170·1 ± 72·1 cm). The most frequently used mesohabitat type was a pool. Young Atlantic salmon favoured the faster flowing areas (mean ±s .d . water velocity 44·0 ± 16·8 cm s⁻¹ and depth 57·1 ± 19·0 cm), while brown trout occupied intermediate habitats (mean ±s .d . water velocity 33·1 ± 18·6 cm s⁻¹ and depth 50·2 ± 18·0 cm). Niche overlap was considerable. The Arctic charr observed were on average larger (total length) than Atlantic salmon and brown trout (mean ±s .d . 21·9 ± 8·0, 10·2 ± 3·1 and 13·4 ± 4·5 cm). Similar habitat segregation between Atlantic salmon and brown trout was found by electrofishing, but more fishes were observed in shallower habitats. Electrofishing suggested that Arctic charr occupied habitats similar to brown trout. These results, however, are biased because electrofishing was inefficient in the slow-deep habitat favoured by Arctic charr. Habitat use changed between day and night in a similar way for all three species. At night, fishes held positions closer to the bottom than in the day and were more often observed in shallower stream areas mostly with lower water velocities and finer substrata. The observed habitat segregation is probably the result of interference competition, but the influence of innate selective differences needs more study.     

Morphological and morphometric aspects of early life stages of piabanha Brycon gouldingi (Characidae)

Adult specimens of piabanha Brycon gouldingi were collected from Rio das Mortes (Mato Grosso, Brazil), adapted to captivity and induced to spawn at Buriti Fisheries (Nova Mutum, MT, Brazil). The early developmental stages of B. gouldingi were then characterized. Samples were collected at pre‐determined times from oocyte extrusion to total yolk absorption. Oocyte diameter, total larval length (L_T) and yolk‐sac volume were measured. The mean ± s.d . duration of embryo developmental of B. gouldingi was 13·90 ± 0·06 h at 26·40 ± 1·13° C. The mean ± s.d . oocyte diameter was 1·13 ± 0·06 mm with 54% of oocytes ranging from 1·11 to 1·20 mm. Seven stages characterized the early developmental phase of this species: zygote, cleavage, morula, blastula, gastrula, histogenesis–organogenesis and hatching, with unique features related to each stage. At hatching, the larvae measured 3·40 ± 0·07 mm, presented an elongated shape with yolk‐sac volume of 0·46 ± 0·08 µl, non‐pigmented eyes and exhibited swimming ability. When the yolk was completely absorbed at 55 h post‐hatch, mean ± larval L_T was 6·68 ± 0·65 mm, the eyes were highly pigmented and the teeth were visible. These are the first reported findings on the initial developmental stages of B. gouldingi and could be used to improve captive breeding management and conservation practices.     

Reproductive biology of albacore Thunnus alalunga

Reproductive variables in albacore Thunnus alalunga were evaluated by gonad histology in samples of 132 males (58–118 cm fork length, L_F) and 112 females (59–101 cm L_F) that were collected from the western North Pacific Ocean from 2001 to 2006. In the sex ratio examination, males greatly outnumbered females in large adult fish (L_F > 100 cm). Thunnus alalunga exhibited a protracted spawning period from March to September in the waters off eastern Taiwan and the Philippines, and the peak spawning activity occurred in March and April. Minimum sizes associated with the classification of mature fish were 78 and 83 cm L_F for males and females, respectively. In addition, the largest L_F of immature fish were 93 cm for males and 94 cm for females. The spawning frequency estimate in April was 1·7 days. Batch‐fecundity estimates of 21 females (89–99 cm L_F) ranged between 0·17 and 1·66 million eggs (mean ±s.d . = 0·94 ± 0·43). The relative fecundity estimates of the 21 females ranged between 9·2 and 92·4 oocytes g^?1 body mass (mean ±s.d . = 50·5 ± 22·8). The results presented in this study provide increased information regarding this species' reproductive‐related characteristics than are currently available in stock status determinations.     

Fecundity regulation by atresia in turbot Scophthalmus maximus in the Baltic Sea

Down‐regulation of fecundity through oocyte resorption was assessed in Baltic Sea turbot Scophthalmus maximus at three locations in the period from late vitellogenesis in April to spawning during June to July. The mean ± s.d . total length of the sampled fish was 32·7 ± 3·1 cm and mean ± s.d . age was 6·2 ± 1·5 years. Measurements of atresia were performed using the ‘profile method’ with the intensity of atresia adjusted according to the ‘dissector method’ (10·6% adjustment; coefficient of determination was 0·675 between methods). Both prevalence (portion of fish with atresia) and intensity (calculated as the average proportion of atretic cells in fish displaying atresia) of atresia were low in prespawning fish, but high from onset of spawning throughout the spawning period. Atretic oocytes categorized as in early alpha and in late alpha state occurred irrespective of maturity stage from late prespawning individuals up to late spawning fish, showing that oocytes may become atretic throughout the spawning period. Observed prevalence of atresia throughout the spawning period was almost 40% with an intensity of c. 20%. This indicates extensive down‐regulation, i.e. considerably lower realized (number of eggs spawned) v. potential fecundity (number of developing oocytes), suggesting significant variability in reproductive potential. The extent of fecundity regulation in relation to fish condition (Fulton's condition factor) is discussed, suggesting an association between levels of atresia and fish condition.     

Life‐history traits of the long‐nosed skate Dipturus oxyrinchus

This work investigates life‐history traits of the long‐nosed skate Dipturus oxyrinchus, which is a common by‐catch in Sardinian waters. The reproductive variables were analysed from 979 specimens sampled during scientific and commercial hauls. Females (10·4–117·5 cm total length, L_T) attained larger sizes than males (14·5–99·5 cm L_T). To evaluate age and growth, a sub‐sample of 130 individuals (76 females and 54 males) were used. The age was estimated by annuli counts of sectioned vertebral centra. Four models were used for the length‐at‐age data: the von Bertalanffy, the exponential, the Gompertz and the logistic functions. According to the Akaike's information criterion, the Gompertz model seemed to provide the best fitting curve (L_∞ mean ± s.e. : 127·55 ± 4·90 cm, k: 0·14 ± 0·09, IP: 3·97 ± 0·90 years). The oldest female and male were aged 17 (115·5 cm L_T) and 15 years (96·0 cm L_T), respectively. Lengths at maturity were 103·5 cm for females and 91·0 cm for males, corresponding to 90% of the maximum observed length in both sexes. The monthly distribution of maturity stages highlighted an extended reproductive cycle, with spawning females and active males being present almost throughout the year, as confirmed by the gonado‐somatic index. Ovarian fecundity reached a maximum of 26 yolked follicles with a mean ± s.e. size of 19·7 ± 6·5 mm.     

Freshwater influences on embryos,hatching and larval survival of euryhaline Gulf killifish Fundulus grandis and potential constraints on habitat distribution

The influence of fresh water on potential habitat occupancy of early life‐history stages of euryhaline Gulf killifish Fundulus grandis was determined by evaluating fertilization of freshwater‐spawned eggs and subsequent survival of embryos and larvae in comparison with saline water (salinity 7). Overall per cent fertilization of eggs was low (mean ± s.e . = 20·21 ± 0·03%). Embryo survival was greater in saline water, but hatching rate (mean ± s.e . = 81·6 ± 0·1%) and post‐hatch survival of larvae in fresh water (mean ± s.e . = 74·5 ± 0·1%) was relatively high. Therefore, the relative limitation of fresh water on habitat distribution of F. grandis changes with development, stimulating further questions on factors that may constrain habitat distribution of euryhaline fishes.     

Balitora eddsi,a new species of hillstream loach (Ostariophysi: Balitoridae) from Nepal

A new species of hillstream loach Balitora eddsi is described from the Karnali River drainage in south‐western Nepal. The new species is distinguished from all its congeners by possessing the following combination of characters: six to seven unbranched pectoral‐fin rays, pelvic‐fin length 12–14% standard length (L_S), dorsal surface without circular or irregular shaped dark blotches, snout pointed, median lobe between anterior rostral barbels pointed posteriorly, dorsal‐fin origin posterior to pelvic‐fin origin, lateral line scales 66–67, caudal peduncle length 22–23·2% L_S, caudal peduncle depth 4·1–4·2 times its length.