Non-lethal dorsal fin sampling for stable isotope analysis in seahorses

Sampling collection for stable isotope analysis has traditionally involved the sacrifice of the animal. Seahorses (Hippocampus spp.) are listed as threatened by the Convention on International Trade in Endangered Species (http://www.cites.org) and consequently lethal sampling is undesirable. We evaluated the adequacy of dorsal fin tissue of adult seahorses Hippocampus guttulatus for stable isotope analysis as an alternative to lethal tissue sampling. Three seahorse tissues (dorsal fin, muscle, and liver) were analyzed for comparisons of δ¹⁵N and δ¹³C values. Similarities found between δ¹⁵N and δ¹³C values in dorsal fin and muscle tissue of H. guttulatus suggest that both tissues are adequate for stable isotope analysis to understand feeding ecology of seahorses. However, considering the threatened status of the species, dorsal fin tissue would be recommended in adult seahorses as a non-lethal sampling. The effect of lipid extraction on carbon and nitrogen stable isotope values was also evaluated in each seahorse tissues. Significant effects of lipids extraction did only occur for δ¹³C values in muscle and liver. It was found that lipid removal was not necessary to perform SIA in dorsal fin tissues. Due to the limited availability of fin tissue obtained from fin-clipping in seahorses, the relationship between the mass/surface of dorsal fin clip and stable isotope values was analyzed. δ¹⁵N and δ¹³C values in fin samples were found to be independent of the size of fin analyzed. According to our study, the use of fin-clipping sampling, with a minimum surface analyzed of 12.74?mm², was found to be an adequate method for SIA in seahorses.     

Serial monogamy in the European long-snouted seahorse Hippocampus guttulatus

Seahorses (Hippocampus spp.) are non-sex-role-reversed members of the Syngnathidae family that provide extensive brood care. Previous studies of seahorses have revealed monogamy within a single brood, but their longer term mating system had not been comprehensively evaluated. The parental contribution to 29 wild-born broods of Hippocampus guttulatus, sampled from six Portuguese populations with differing seahorse densities and sex ratios, was assessed using microsatellite DNA markers. To assess the longer term genetic mating system of this species parentage was determined in eleven broods sampled from a captive population over two breeding seasons. Genetic data suggest that this socially polygamous seahorse is serially monogamous across breeding seasons, i.e. monogamous within a season but may switch mates between seasons, and that differing population densities and sex ratios do not affect the mating system.     

Does a No-Take Marine Protected Area Benefit Seahorses?

Seahorses are iconic charismatic species that are often used to ‘champion’ marine conservation causes around the world. As they are threatened in many countries by over-exploitation and habitat loss, marine protected areas (MPAs) could help with their protection and recovery. MPAs may conserve seahorses through protecting essential habitats and removing fishing pressures. Populations of White''s seahorse, Hippocampus whitei, a species endemic to New South Wales, Australia, were monitored monthly from 2006 to 2009 using diver surveys at two sites within a no-take marine protected areas established in 1983, and at two control sites outside the no-take MPA sites. Predators of H. whitei were also identified and monitored. Hippocampus whitei were more abundant at the control sites. Seahorse predators (3 species of fish and 2 species of octopus) were more abundant within the no-take MPA sites. Seahorse and predator abundances were negatively correlated. Substantial variability in the seahorse population at one of the control sites reinforced the importance of long-term monitoring and use of multiple control sites to assess the outcomes of MPAs for seahorses. MPAs should be used cautiously to conserve seahorse populations as there is the risk of a negative impact through increased predator abundance.     

Conservation value of Codium tenue habitat for the endangered Knysna seahorse Hippocampus capensis

We investigated the density and size structure of the endangered Knysna seahorse Hippocampus capensis in the macroalga Codium tenue at three sites in a residential marina estate in the Knysna Estuary, South Africa, over an 18 month period from March 2017 to August 2018. Seahorses were present in C. tenue throughout the year. Average (± SE) densities of 0.33 (± 0.03) and 0.23 (± 0.03) seahorses per kg of C. tenue were found for 2017 and 2018 respectively. Density did not vary across sites but varied significantly across months with a decreasing trend from summer to spring. The average (± SE) adult seahorse height was 75.16 mm (± 0.63 mm) and 69.09 mm (± 0.64 mm) and mass was 2.05 g (± 0.04 g) and 1.65 g (± 0.04 g) for males and females, respectively, and an even sex ratio was observed throughout the sampling period. Height varied across months, with significantly smaller seahorses found in June and August of 2017. Tail length was highly correlated with height and therefore can serve as a potential proxy for determining the height of H. capensis. This study showed that C. tenue is consistently utilised by Knysna seahorse and should therefore be considered an important habitat to protect and facilitate the ongoing conservation of this endangered seahorse species.     

Conservation and management of seahorses and other Syngnathidae

This article analyses the pressures on seahorses and explores conservation responses. It focuses on seahorses (Hippocampus spp.) but also considers pipefishes and seadragons, especially where they can fill gaps in seahorse knowledge. The charisma of many syngnathids can make them good flagship species for threats and solutions in marine conservation. The article combines a synthesis of published literature with new data on the trade in seahorses for traditional medicine, aquarium display and curiosities. Most traded seahorses come from trawl by-catch, although seahorses are also targeted. The total extraction is large, tens of millions of animals annually, and unsustainable. A first review of the effect of habitat change on syngnathids raises many questions, while suggesting that some species may cope better than others. The combination of pressures means that many species of syngnathid are now included in the IUCN Red List of Threatened Species or national equivalents. In addition, seahorse exports from 175 countries are limited to sustainable levels under the Convention on International Trade in Endangered Species (CITES) of Wild Fauna and Flora. Possible conservation measures include marine protected areas, fisheries management, select aquaculture ventures, trade regulation, improved governance (particularly) and consumer engagement.     

Getting old: an endangered seahorse (Hippocampus whitei) lives for up to 7 years in the wild

A long-term monitoring programme from 2005 to 2021 has allowed the assessment of age and longevity in an endangered seahorse Hippocampus whitei in the wild. Seahorses were marked using visible implant fluorescent elastomer (VIFE) which allows for individual identification. The longest period from marking to last sighting was 6 years 8 months and 17 days for a female. Using a von Bertalanffy growth function model for the species, this individual would have been approximately 7 years 7 months old on last sighting. These observations suggest that seahorses in the wild can live for over 7 years and demonstrate the benefits of using VIFE in long-term movement, population abundance and life-history studies of seahorses.     

Fishers' knowledge and seahorse conservation in Brazil

From a conservationist perspective, seahorses are threatened fishes. Concomitantly, from a socioeconomic perspective, they represent a source of income to many fishing communities in developing countries. An integration between these two views requires, among other things, the recognition that seahorse fishers have knowledge and abilities that can assist the implementation of conservation strategies and of management plans for seahorses and their habitats. This paper documents the knowledge held by Brazilian fishers on the biology and ecology of the longsnout seahorse Hippocampus reidi. Its aims were to explore collaborative approaches to seahorse conservation and management in Brazil; to assess fishers' perception of seahorse biology and ecology, in the context evaluating potential management options; to increase fishers' involvement with seahorse conservation in Brazil. Data were obtained through questionnaires and interviews made during field surveys conducted in fishing villages located in the States of Piauí, Ceará, Paraíba, Maranhão, Pernambuco and Pará. We consider the following aspects as positive for the conservation of seahorses and their habitats in Brazil: fishers were willing to dialogue with researchers; although captures and/or trade of brooding seahorses occurred, most interviewees recognized the importance of reproduction to the maintenance of seahorses in the wild (and therefore of their source of income), and expressed concern over population declines; fishers associated the presence of a ventral pouch with reproduction in seahorses (regardless of them knowing which sex bears the pouch), and this may facilitate the construction of collaborative management options designed to eliminate captures of brooding specimens; fishers recognized microhabitats of importance to the maintenance of seahorse wild populations; fishers who kept seahorses in captivity tended to recognize the condtions as poor, and as being a cause of seahorse mortality.     

To eject or to abandon? Life history traits of hosts and parasites interact to influence the fitness payoffs of alternative anti‐parasite strategies

Hosts either tolerate avian brood parasitism or reject it by ejecting parasitic eggs, as seen in most rejecter hosts of common cuckoos, Cuculus canorus, or by abandoning parasitized clutches, as seen in most rejecter hosts of brown‐headed cowbirds, Molothrus ater. What explains consistent variation between alternative rejection behaviours of hosts within the same species and across species when exposed to different types of parasites? Life history theory predicts that when parasites decrease the fitness of host offspring, but not the future reproductive success of host adults, optimal clutch size should decrease. Consistent with this prediction, evolutionarily old cowbird hosts, but not cuckoo hosts, have lower clutch sizes than related rarely‐ or newly parasitized species. We constructed a mathematical model to calculate the fitness payoffs of egg ejector vs. nest abandoner hosts to determine if various aspects of host life history traits and brood parasites’ virulence on adult and young host fitness differentially influence the payoffs of alternative host defences. These calculations showed that in general egg ejection was a superior anti‐parasite strategy to nest abandonment. Yet, increasing parasitism rates and increasing fitness values of hosts’ eggs in both currently parasitized and future replacement nests led to switch points in fitness payoffs in favour of nest abandonment. Nonetheless, nest abandonment became selectively more favourable only at lower clutch sizes and only when hosts faced parasitism by a cowbird‐ rather than a cuckoo‐type brood parasite. We suggest that, in addition to evolutionary lag and gape‐size limitation, our estimated fitness differences based on life history trait variation provide new insights for the consistent differences observed in the anti‐parasite rejection strategies between many cuckoo‐ and cowbird‐hosts.     

Life history and ecology of the elusive European short‐snouted seahorse Hippocampus hippocampus

To improve the understanding of the life history and ecology of one of Europe's most elusive fishes, the short‐snouted seahorse Hippocampus hippocampus, data from wild populations in a shallow coastal lagoon in southern Portugal were analysed. The data were collected from 17 tagged seahorses on a focal‐study grid as well as from >350 seahorses encountered during underwater visual surveys and a fishery‐independent study using beach seines. These populations of settled juveniles and adults had a mean population density of 0·009 m^?2. During the study period (2000–2004), reproduction peaked in July and August. Juveniles recruited to the lagoon at c. 66 mm standard length (L_S) and 0·5 years of age and established small home ranges (0·8 to 18·2 m²). First reproduction was estimated at 100 mm and 1 year of age. Based on a fitted von Bertalanffy model, H. hippocampus grew quickly (growth coefficient K = 0·93) to a maximum theoretical size L_∞ = 150 mm and have a maximum lifespan of c. 3·2 years. Courtship behaviours were consistent with the maintenance of pair bonds and males brooded multiple batches of young per year. Estimated annual reproductive output averaged 871 young (±632). Together these analyses provide the first life‐history parameters for this species and indicate that H. hippocampus bears characteristics of opportunist and intermediate strategists. Such populations are predicted to exhibit large fluctuations in abundance, making them vulnerable to extended periods of poor recruitment.     

Seahorses in focus: local ecological knowledge of seahorse-watching operators in a tropical estuary

Background

Seahorses are endangered teleost fishes under increasing human pressures worldwide. In Brazil, marine conservationists and policy-makers are thus often skeptical about the viability of sustainable human-seahorse interactions. This study focuses on local ecological knowledge on seahorses and the implications of their non-lethal touristic use by a coastal community in northeastern Brazil. Community-based seahorse-watching activities have been carried out in Maracaípe village since 1999, but remained uninvestigated until the present study. Our goal is to provide ethnoecological understanding on this non-extractive use to support seahorse conservation and management.

Methods

We interviewed 32 informants through semi-structured questionnaires to assess their socioeconomic profile, their knowledge on seahorse natural history traits, human uses, threats and abundance trends.

Results

Seahorse-watching has high socioeconomic relevance, being the primary income source for all respondents. Interviewees elicited a body of knowledge on seahorse biology largely consistent with up-to-date research literature. Most informants (65.5 %) perceived no change in seahorse abundance. Their empirical knowledge often surpassed scientific reports, i.e. through remarks on trophic ecology; reproductive aspects, such as, behavior and breeding season; spatial and temporal distribution, suggesting seahorse migration related to environmental parameters.

Conclusions

Seahorse-watching operators were aware of seahorse biological and ecological aspects. Despite the gaps remaining on biological data about certain seahorse traits, the respondents provided reliable information on all questions, adding ethnoecological remarks not yet assessed by conventional scientific surveys. We provide novel ethnobiological insight on non-extractive modes of human-seahorse interaction, eliciting environmental policies to integrate seahorse conservation with local ecological knowledge and innovative ideas for seahorse sustainable use. Our study resonates with calls for more active engagement with communities and their local ecologies if marine conservation and development are to be reconciled.

     

Genetic monogamy despite social promiscuity in the pot-bellied seahorse (Hippocampus abdominalis)

Sexual selection theory predicts a positive correlation between relative parental investment and mate choice. In syngnathid fishes (seahorses and pipefish), males brood offspring in specialized brooding structures. While female-female mating competition has been demonstrated in some pipefishes, all seahorses (genus Hippocampus) studied to date have been found to have conventional sex roles with greater male-male competition for access to mates despite possessing the most complex brood structures in the family. Although multiple mating is common in pipefish, seahorses are again exceptional, exhibiting strict genetic monogamy. Both demographic and behavioural explanations have been offered to explain the lack of multiple mating in seahorse species, but these hypotheses have not yet been explicitly addressed. We investigated mating systems and brood parentage of the pot-bellied seahorse, Hippocampus abdominalis, a temperate-water species that is socially promiscuous with conventional sex roles in laboratory populations. We observed promiscuous courtship behaviour and sex-role reversal in high density, female-biased field populations of H. abdominalis. We hypothesize that sex roles are plastic in H. abdominalis, depending on local population density and sex ratio. Despite promiscuous courtship behaviour, all assayed male seahorses were genetically monogamous in both laboratory and wild populations. Physiological limitations associated with embryo incubation may explain the absence of multiple mating in seahorses and may have played an important role in the development of the unique reproductive behaviour typical in these species.     

Correlates of reproductive success in a wild population of Hippocampus whitei

An analysis of 27 broods born in situ to wild male seahorses Hippocampus whitei , with known partners, showed that female size was the key determinant of the number of young released by the male. There was also an apparent decline in both the number of young per brood, and the size of those young, over the breeding season. The size of the brood and of the young may be more proximate indicators of reproductive success in seahorses than in other fishes, because they can be measured at the time of release, here 20–22 days after fertilization, after which survival is presumed to be relatively higher than in other species. The best model, however, explained <40% of the brood size.     

Evolution of seahorses' upright posture was linked to Oligocene expansion of seagrass habitats

Seahorses (Syngnathidae: Hippocampus) are iconic marine teleosts that are readily identifiable by their upright posture. The fossil record is inadequate to shed light on the evolution of this trait because it lacks transitional forms. There are, however, extant syngnathid species (the pygmy pipehorses) that look like horizontally swimming seahorses and that might represent a surviving evolutionary link between the benthic seahorses and other, free-swimming members of the family Syngnathidae. Using sequence data from five nuclear loci, we confirm the sister taxon relationship between seahorses and pygmy pipehorses. Molecular dating indicates that the two taxa diverged during the Late Oligocene. During this time, tectonic events in the Indo-West Pacific resulted in the formation of vast amounts of new shallow-water areas and associated expansion of seagrass habitats that would have favoured the seahorses’ upright posture by improving their camouflage while not affecting their manoeuvrability negatively. The molecular techniques employed here provide new insights into the evolution of a taxon whose fossil record is incomplete, but whose evolutionary history is so recent that the major stages of morphological evolution are still represented in extant species.     

Comparative analysis of phylogenetic and fishing effects in life history patterns of teleost fishes

The effects of fishing on life history traits and life history strategies of teleost fishes are analysed by a new comparative method that splits traits into an allometric part (size effect), an autoregressive phylogenetic component, and an environmental component (fishing effect). Both intra- and inter-specific variation of age and size at maturity, fecundity, adult size and egg size are analysed by comparing 84 populations of 49 species submitted to various fishing pressures. Two axes of life history diversification are found among teleosts. One is the well-known slow-fast continuum separating short-lived and early maturing species (like Clupeiformes) from longer-lived species that mature late relative to their size and spawn larger eggs (like salmonids or Scorpaeniformes). An additional strategy involves the schedule of resource allocation to growth and reproduction. Indeterminate growth allows higher teleosts (e.g. Gadiformes) to reach a large size while maturing early and laying small eggs. Increasing fishing pressure decreases age at maturity and egg size, and increases fecundity at maturity, the slope of the fecundity-length relationship and relative size at maturity. These compensations for higher adult mortality differ among life history strategies. Indeterminate growth is associated with a greater flexibility in resource allocation to growth and reproduction that facilitates greater resilience to fishing mortality.     

Seahorse fingerprints: a new individual identification technique

Individual identification is particularly important for ethological studies and sampling design. Previous studies have developed various types of marking by tags and chemical marks, but these methods involve significant manipulation of the individuals. Other studies have reported natural marks as an efficient method for individual recognition. Our study aims to elucidate a new method for natural mark identification of seahorses, which we tested with the endangered Brazilian seahorse Hippocampus reidi. We avoid pseudoreplication by recognizing individuals. Seahorses have a hard bony structure on the top of their heads called the coronet, which has a different shape for each individual, corresponding to a fingerprint. We tagged seahorses in the field with collar tags and photographed their coronets. After two days, we identified seahorses by their photographs and verified their identification with the collar tags. We correctly identified all individuals by fingerprint identification. Although this method was only tested with adults, we suggest that it applies to seahorses in general, as all species possess the coronet structure.     

The catch and trade of seahorses in Vietnam

Catch monitoring and surveys were used to assess the seahorse trade in Vietnam. Despite low daily catch rates, potentially 6.5 t of dried seahorses (~2.2 million seahorses) were taken annually as bycatch by trawlers operating out of five coastal provinces of Vietnam. Individual seahorse catches were collated by a few local buyers, who supplied wholesalers in three major markets: Ho Chi Minh City, Hai Phong City and Da Nang. Domestic consumption was small and most seahorses were exported, generally through unofficial and unregulated channels across the northern border into Guangxi province of China. Overall, the seahorse trade was of low economic value to Vietnam, but may constitute an important source of income to upper level buyers and exporters. Most fishers and buyers surveyed reported that seahorse catch had declined over time. This paper should help in meeting the new CITES requirements – through implementation of an Appendix II listing in 2004 – that all international trade in seahorses must be monitored and managed for sustainability.     

Host stage selection of the mealybug parasitoid Anagyrus spec. nov near sinope

The mealybug parasitoid Anagyrus spec. nov near sinope (Hymenoptera: Encyrtidae) is an undescribed parasitoid of the Madeira mealybug, Phenacoccus madeirensis Green (Homoptera: Pseudococcidae). We investigated the preference of Anagyrus spec. nov near sinope for six developmental stadia (first‐ and second‐instar nymphs, third‐instar immature females, third‐ or fourth‐instar immature males, pre‐reproductive adult females, and ovipositing adult females) of P. madeirensis and the fitness consequences of the host stage selection behavior. In the no‐choice test, Anagyrus spec. nov near sinope parasitized and completed development in all host stadia except third‐instar immature males. When all host stadia were offered simultaneously, the parasitoids preferred third‐instar immature and pre‐reproductive adult females. Dissection of the stung mealybugs revealed that the clutch size (number of eggs per host) was approximately four and three in the third‐instar and pre‐reproductive females, respectively, and one egg per first‐instar nymph. Parasitoids emerged from P. madeirensis parasitized at third‐instar or pre‐reproductive adult female completed development in the shortest duration, achieved a higher progeny survival rate, larger brood and body size, and the lowest proportion of males. We showed that the continued development of mealybugs had significant influence on the fitness of the parasitoids. Although deposited as eggs in first‐ or second‐instar nymphs, parasitoids emerged from mummies that had attained third‐instar or adult development achieved similar progeny survival rate, brood size, body size, and sex ratio as those parasitoids deposited and developed in third‐instar or adult mealybugs. By delaying larval development in young mealybugs, Anagyrus spec. nov near sinope achieved higher fitness by allowing the parasitized mealybugs to grow and accumulate body size and resources. We suggest that the fitness consequence of host stage selection of a koinobiont parasitoid should be evaluated on both the time of parasitism and the time of mummification.     

Evolutionary ecology of pipefish brooding structures: embryo survival and growth do not improve with a pouch

For animals that reproduce in water, many adaptations in life‐history traits such as egg size, parental care, and behaviors that relate to embryo oxygenation are still poorly understood. In pipefishes, seahorses and seadragons, males care for the embryos either in some sort of brood pouch, or attached ventrally to the skin on their belly or tail. Typically, egg size is larger in the brood pouch group and it has been suggested that oxygen supplied via the pouch buffers the developing embryos against hypoxia and as such is an adaptation that has facilitated the evolution of larger eggs. Here, using four pipefish species, we tested whether the presence or absence of brood pouch relates to how male behavior, embryo size, and survival are affected by hypoxia, with normoxia as control. Two of our studied species Entelurus aequoreus and Nerophis ophidion (both having small eggs) have simple ventral attachment of eggs onto the male trunk, and the other two, Syngnathus typhle (large eggs) and S. rostellatus (small eggs), have fully enclosed brood pouches on the tail. Under hypoxia, all species showed lower embryo survival, while species with brood pouches suffered greater embryo mortality compared to pouchless species, irrespective of oxygen treatment. Behaviorally, species without pouches spent more time closer to the surface, possibly to improve oxygenation. Overall, we found no significant benefits of brood pouches in terms of embryo survival and size under hypoxia. Instead, our results suggest negative effects of large egg size, despite the protection of brood pouches.     

Reproductive biology of female shovelnose sturgeon in the upper Wabash River, Indiana

Shovelnose sturgeon Scaphirhynchus platorynchus are commercially important, supporting a viable roe fishery throughout much of the Mississippi River drainage. We examined the reproductive attributes of stage‐5 female shovelnose sturgeon captured from the upper Wabash River, Indiana, from March to June 2004. Shovelnose sturgeon were collected using boat electrofishing and experimental gill nets, measured for fork length (FL) and wet weight, and sexed externally if possible. Size‐ and age‐at‐maturity, absolute and relative fecundity, relative egg size, and gonadosomatic index (GSI) were determined for 49 female shovelnose sturgeon (range, 601‐ to 858‐mm FL). Female shovelnose sturgeon reached sexual maturity at approximately 600 mm and age‐at‐maturity ranged from ages 6 to 12 (median age = 9). Relative fecundity ranged from 11 220 to 23 956 eggs kg^?1 (mean = 18 156 eggs kg^?1). Absolute fecundity ranged from 14 294 to 65 490 eggs female^?1 (mean = 30 397 eggs female^?1) and was positively related to FL (r² = 0.76) and wet weight (r² = 0.82). The number of eggs g^?1 of ovary weight ranged from 72 to 170 (mean = 98 eggs g^?1) and was negatively correlated with GSI. GSI values ranged from 9.4 to 27.2 (mean = 19.3) and were positively correlated to FL (r² = 0.18). Our results increase our understanding of shovelnose sturgeon reproductive biology and recruitment dynamics and provide input for models to evaluate the effects of harvest on this species.     

Life history and population dynamics of an endangered seahorse (Hippocampus capensis) within an artificial habitat

Species-specific life-history information is critical for successful conservation, particularly in establishing an accurate baseline status. Obtaining such information is challenging for most species, but in particular for rare and threatened marine species. To facilitate future conservation of the endangered Knysna seahorse (Hippocampus capensis) this study aimed to determine important life-history information for this species. Visible implant fluorescent elastomer (VIFE) tags were used to mark 78 seahorses within a residential marina estate in the Knysna estuary, South Africa, in February 2018. Using a mark-resight approach, the size and movement patterns of the population and growth rate of seahorses were determined over a 14-month period. The closed population estimate for H. capensis, within Thesen Islands Marina, was estimated to be 134 (118–152 95% C.I. ) in February 2018 compared to only 72 (48–108 95% C.I. ) in February 2019. The species showed rapid initial growth with males and females having similar rates of growth based on the specialised von Bertalanffy growth function model. The importance of Reno mattresses as a habitat for H. capensis was confirmed based on the high abundance and site fidelity of the population, which emphasises the conservation potential of heavily modified environments for threatened seahorse species. The use of VIFE tags was deemed effective in studying this endangered seahorse and allowed the collection of important information for this species which can be used in future Red List assessments and conservation actions.