Cynthia A. Awruch

Endocrine and morphological correlates of reproduction in the draughtboard shark Cephaloscyllium laticeps (Elasmobranchii: Scyliorhinidae)

This study examined the endocrine and reproductive correlates of reproduction in 636 female and 468 male draughtboard sharks (Cephaloscyllium laticeps) captured from southeastern Australia. Females were oviparous and displayed a single external-type ovary with a maximum follicle diameter of 35 mm. Vitellogenesis commenced at a follicle diameter of 10 mm. Females showed a constant overlap between follicular recruitment, ovarian growth, and egg laying. The male reproductive tract consisted of paired testes with spermatocysts undergoing diametric development. Plasma levels of the presumptive gonadal steroids, testosterone (T), 17beta-estradiol (E2), progesterone (P4), and 11-ketotestosterone (11-KT; males only) were correlated with morphological developmental stages of the gonads. In females, E2 increased as the follicle developed before declining as the follicle reached maturity. T remained low during the first stages of ovarian growth and increased as the follicle reached maturity. P4 showed a peak just before ovulation. In males, T was the only hormone that varied with maturity, increasing in adults; E2 and P4 were present at low plasma concentrations in males and did not change with stage of gonadal development. 11-KT was undetectable at all times. Endocrine changes in draughtboard sharks were consistent with hormonal correlates reported for other species and suggest roles for E2( in females) and T (in both sexes) in gametogenesis and P4 in maturational events in females.

An overview on the role of Hexanchiformes in marine ecosystems: biology, ecology and conservation status of a primitive order of modern sharks

The large size, high trophic level and wide distribution of Hexanchiformes (cow and frilled sharks) should position this order as important apex predators in coastal and deep-water ecosystems. This review synthesizes available information on Hexanchiformes, including information not yet published, with the purpose of evaluating their conservation status and assessing their ecological roles in the dynamics of marine ecosystems. Comprising six species, this group has a wide global distribution, with members occurring from shallow coastal areas to depths of c. 2500 m. The limited information available on their reproductive biology suggests that they could be vulnerable to overexploitation (e.g. small litter sizes for most species and suspected long gestation periods). Most of the fishing pressure exerted on Hexanchiformes is in the form of commercial by-catch or recreational fishing. Comprehensive stock and impact assessments are unavailable for most species in most regions due to limited information on life history and catch and abundance time series. When hexanchiform species have been commercially harvested, however, they have been unable to sustain targeted fisheries for long periods. The potentially high vulnerability to intense fishing pressure warrants a conservative exploitation of this order until thorough quantitative assessments are conducted. At least some species have been shown to be significant apex predators in the systems they inhabit. Should Hexanchiformes be removed from coastal and deep-water systems, the lack of sympatric shark species that share the same resources suggests no other species would be capable of fulfilling their apex predator role in the short term. This has potential ecosystem consequences such as meso-predator release or trophic cascades. This review proposes some hypotheses on the ecology of Hexanchiformes and their role in ecosystem dynamics, highlighting the areas where critical information is required to stimulate research directions.

Reproductive endocrinology in chondrichthyans: the present and the future.

The class Chondrichthyes, that includes Elasmobranchii and Holocephali, is a diverse group of fish occupying a key position at the base of vertebrate evolution. Their evolutionary success is greatly attributed to their wide range of reproductive strategies controlled by different endocrine mechanics. As in other vertebrates, hormonal control of reproduction in chondrichthyans is mediated by the neuropeptide gonadotropin-releasing hormone (GnRH) that regulates the brain control of gonadal activity via a hypothalamus-pituitary-gonadal (HPG) axis. Chondrichthyans lack of a direct vascular supply from the hypothalamus to the zone of the pituitary where the gonadotropic activity resides, thus transport between these two zones likely occurs via the general circulation. In the brain of elasmobranchs, two groups of GnRH, GnRH-I and GnRH-II were identified, and the presence of two immunoreactive gonadotropins similar to the luteinising (LH) and follicle stimulating (FSH) hormones was identified in the pituitary. In holocephalans, only GnRH-II has been confirmed, and while gonadotropin activity has been found in the buccal pituitary lobe, the presence of gonadotropin receptors in the gonads remains unknowns. The diversity of reproductive strategies display by chondrichthyans makes it difficult to generalize the control of gametogenesis and steroidogenesis; however, some general patterns emerge. In both sexes, androgens and estrogens are the main steroids during gonadal growth; while progestins have maturational activity. Androgens also form the precursors for estrogen steroid production. Estrogens stimulate the hepatic synthesis of yolk and stimulate the development of different part of the reproductive tract in females. The role of other gonadal steroids may play in chondrichthyan reproduction remains largely unknown. Future work should concentrate in filling the gaps into the current knowledge of the HPG axis regulation, and the use of reproductive endocrinology as a non-lethal technique for management of chondrichthyan populations.

Evaluation and use of a portable field kit for measuring whole-blood lactate in sharks

Measurement of lactate is becoming a common procedure in assessing the physiological effects of capture stress in sharks, although the necessity to measure the concentrations in the laboratory limits the ability for field assessments. Portable lactate analysers offer an alternative, but await validation against laboratory assays for sharks. The present study assessed the reliability of a portable Lactate Pro analyser for measuring whole-blood lactate in the school shark, Galeorhinus galeus, in the field. Laboratory determination of whole-blood and plasma lactate obtained by spectrophotometry was highly correlated with field determinations. Because shark lactate concentration can exceed the upper detection limit of the portable analysers, which were designed for mammalian use, a method for dealing with values greater than the maximum detection limit was evaluated. Whole-blood diluted by 50% with acidified saline solution, tap water and distilled water gave measured values of 55, 56 and 52%, respectively, of the original values, allowing accurate estimation of concentrations exceeding the upper detection limit of the analyser. These findings indicated that the Lactate Pro can be used to rapidly and reliably measure lactate for sharks in the field.

Endocrine Control of Embryonic Diapause in the Australian Sharpnose Shark Rhizoprionodon taylori

The reproductive cycle of the Australian sharpnose shark, Rhizoprionodon taylori, includes a temporary suspension of development at the commencement of embryogenesis termed embryonic diapause. This study investigated levels of 17β-estradiol (E2), testosterone (T) and progesterone (P4) in plasma samples of mature wild female R. taylori captured throughout the reproductive cycle and correlated them with internal morphological changes. Levels of T were elevated through most of the embryonic diapause period, suggesting a role of this hormone in the maintenance of this condition. Increasing plasma T concentrations from late diapause to early active development were associated with a possible role of androgens in the termination of embryonic diapause. As in other elasmobranchs, a concomitant increase of E2 with ovarian follicle size indicated a direct role of this hormone in regulating vitellogenesis, while a peak in P4 suggested this hormone is associated with preovulation and ovulation. Additionally, significant correlations between photoperiod or water temperature and maximum follicular diameter and hepatosomatic index suggest that these abiotic factors may also play a role triggering and regulating the synchrony and timing of reproductive events.

Non-lethal assessment of the reproductive status of broadnose sevengill sharks (Notorynchus cepedianus) to determine the significance of habitat use in coastal areas

Understanding the reproductive role that the habitat plays on animal behaviour is essential for conservation programs. Non-lethal endocrine methods were used to obtain reproductive information on the sevengill shark Notorynchus cepedianus habiting Tasmanian coastal areas to understand the reproductive significance that these systems might play on this large apex predator.

Reproductive seasonality and embryo development in the draughtboard shark Cephaloscyllium laticeps

The draughtboard shark Cephaloscyllium laticeps is a common oviparous catshark found in Australia. Studies on the reproductive seasonality of catsharks and the reproductive endocrinology of oviparous elasmobranchs are limited to a few species. The aim of the present study was to investigate the reproductive seasonality and embryo development of C. laticeps using morphological and endocrine parameters. Macroscopic examinations of the reproductive system and embryo stages were recorded. The steroid hormones 17β- oestradiol (E2) and progesterone (P4) were measured in females, and testosterone (T) in both sexes. Females laid eggs throughout the year, but with a peak in egg deposition between January and June. Female T levels were higher in the middle of the year. Elevated P4 values coincided with the period of egg deposition, whereas E2 was depressed in the middle of the egg deposition stage. Females laid eggs in the absence of males for 15 months, and eggs were laid in pairs at monthly intervals. Hatching occurred after 12 months. Variability in gonadosomatic index, percentage of sperm in seminal vesicles, and plasma T levels for males indicated that males were reproductively active throughout the year.

Movement patterns of the draughtboard shark Cephaloscyllium laticeps (Scyliorhinidae) determined by passive tracking and conventional tagging

A combination of passive tracking and conventional tagging was used to provide insight into the movement patterns of the draughtboard shark Cephaloscyllium laticeps, the most common catshark in coastal areas of southern Australia. A series of acoustic receivers deployed throughout south-eastern Tasmania as well as a receiver array along an isolated reef, Crayfish Point Reserve (CPR), passively tracked 25 C. laticeps from January to July 2003. Cephaloscyllium laticeps were present from 4 to 98 days. The majority of the C. laticeps stayed within the CPR where most individuals were active throughout the night. They were found actively moving (i.e. when a C. laticeps was consecutively detected by two or more non-overlapping receivers, suggesting the individual was moving) and spending periods of minor movements (i.e. when an individual was consecutively detected by only one receiver, suggesting it was at rest). The length of these minor movements periods, observed both day and night, ranged from 1 h to 5 days. In addition to passive tracking, 1552 conventionally tagged C. laticeps were released in the eastern and south-western coastal areas of Tasmania and within the CPR between January 2000 and April 2007. The CPR showed a higher recapture rate, 38%, than eastern and south-western areas where the recapture rates were 10 and 3%, respectively. Within the CPR, 36% of the sharks were recaptured on multiple occasions. The maximum time at liberty ranged from 1 month to 7 years. The majority of the C. laticeps were recaptured in the vicinity of where they were released (<10 km), although larger longer-term movements of up to 300 km were recorded. The large amount of multiple recaptures within the CPR, in addition to acoustic tagging results, indicated a high degree of site fidelity for C. laticeps. This isolated reef appears to be an important habitat for this species, and therefore, the current protection status of this area is probably beneficial for the conservation of C. laticeps.

A review on the morphology of ovarian follicles in elasmobranchs: A case study in Rhizoprionodon taylori

The identification of the elasmobranch secondary ovarian follicles and their function can be challenging and the inconsistent use of terminology derived from other taxa is a matter of ongoing debate. In this study, the available information on the histology of the elasmobranch secondary ovarian follicles derived from atresia (preovulatory follicles) or ovulation (postovulatory follicles) is reviewed highlighting their morphology and steroidogenic capacity. Based on this literature review, the ovarian follicles of the Australian sharpnose shark Rhizoprionodon taylori were classified according to their preovulatory or postovulatory origin. Two types of secondary follicles originating from atresia of developing follicles (atretic previtellogenic follicles) and ripe follicles (atretic vitellogenic follicles), and one type of postovulatory follicle were identified throughout the reproductive year of this species. Morphological similarities of the elasmobranch secondary ovarian follicles and their variations in different species denote the difficulty to classify them. Given the multiple origins of ovarian follicles, their poorly understood functions and capacity to supply steroids, visual identification of these structures and the use of terminology derived from mammalian and other vertebrate studies (with the exception of the corpora lutea as a temporary endocrine gland) is not advisable. J. Morphol. 278:486–499, 2017.

7 - Reproduction strategies

Elasmobranchs are an evolutionarily conserved group that has success- fully survived for over 400 million years. The permanence of elasmobranch populations has largely depended on the reproductive strategies of the population, as the primary requirement for successful propagation of any species and their individuals is the ability to reproduce. In vertebrates, reproductive strategies are regulated by the brain-gonadal axis, which controls the synthesis of reproductive hormones triggering all aspects related to reproduction. This chapter details the different reproductive strategies employed by elasmobranchs, from the wider range of reproductive modes including oviparity (egg-laying) and different forms of viviparity (live- bearing); followed by a description of the different reproductive cycles, from seasonal to continuous, displayed by both sexes. Finally, the role of reproductive hormones in both females and males regulating gametogenesis and the different reproductive cycles are discussed. The endocrine control of the elasmobranch reproductive strategies are preserved throughout vertebrate evolution, however they are distinct within this group.