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.     

Monthly population density and structure patterns of an endangered seahorse Hippocampus capensis: a comparison between natural and artificial habitats

This study investigated aspects of the population ecology of the endangered Knysna seahorse Hippocampus capensis within different habitat types. High densities of H. capensis were found within artificial Reno mattress habitat, within the Knysna Estuary, South Africa. Monthly surveys at three sites were conducted from October 2015 to August 2016 to compare population densities of H. capensis in this artificial habitat with natural eel grass Zostera capensis habitat. Hippocampus capensis densities varied significantly across all sites and highest population densities were consistently observed within the Reno mattress habitat. Hippocampus capensis were also found to be significantly larger within the Reno mattress habitat and pooled data showed that males were significantly larger than females. The overall sex ratio for all three sites was female biased, although this varied across seasons at two sites. The findings suggest that artificial Reno mattresses provide novel habitat for this endangered species and consideration should be given to the usefulness of these structures in future conservation actions.     

First Field Studies of an Endangered South African seahorse, Hippocampus capensis

South Africa's endemic Knysna seahorse, Hippocampus capensis Boulenger 1900, is a rare example of a marine fish listed as Endangered by the IUCN because of its limited range and habitat vulnerability. It is restricted to four estuaries on the southern coast of South Africa. This study reports on its biology in the Knysna and Swartvlei estuaries, both of which are experiencing heavy coastal development. We found that H. capensis was distributed heterogeneously throughout the Knysna Estuary, with a mean density of 0.0089m^–2 and an estimated total population of 89000 seahorses (95% confidence interval: 30000–148000). H. capensis was found most frequently in low density vegetation stands (20% cover) and grasping Zostera capensis. Seahorse density was not otherwise correlated with habitat type or depth. The size of the area in which any particular seahorse was resighted did not differ between males and females. Adult sex ratios were skewed in most transects, with more males than females, but were even on a 10 by 10m focal study grid. Only three juveniles were sighted during the study. Both sexes were reproductively active but no greeting or courtship behaviours were observed. Males on the focal study grid were longer than females, and had shorter heads and longer tails, but were similar in colouration and skin filamentation. The level of threat to H. capensis and our limited knowledge of its biology mean that further scientific study is urgently needed to assist in developing sound management practices.     

An assessment of West African seahorses in fisheries catch and trade

This study provides the first assessment of a heavily traded West African seahorse species, Hippocampus algiricus, and the first information on short‐snouted seahorse Hippocampus hippocampus biology in Africa. A total of 219 seahorses were sampled from fisher catch in Senegal and The Gambia, with estimated height at reproductive activity for H. algiricus (161 mm) larger than mean ± s.d . catch height (150 ± 31 mm). Catch composition, height at reproductive activity and potential biases in fishery retention are discussed with regard to the current Convention on International Trade of Endangered Species (CITES) guidelines.     

An endangered seahorse selectively chooses an artificial structure

The development of a residential marina estate within the Knysna estuary, South Africa, introduced Reno mattresses (horizontal wire cages filled with rocks) as a novel habitat for the endangered Knysna seahorse Hippocampus capensis. Consistently high seahorse densities on these artificial structures, despite the availability of seagrass habitat, begged the question of whether this habitat was chosen by the seahorse in preference to natural vegetation. An in situ habitat choice experiment was conducted which focused on the choice made by adult H. capensis between natural vegetation (Zostera capensis) and artificial (Reno mattress) habitat within a choice chamber. Seahorses were significantly more likely to move away from Z. capensis onto a Reno mattress structure or remain on this structure. This study concludes that higher H. capensis densities on Reno mattresses within Thesen Islands Marina are owing to some positive feature of this habitat and the underlying processes responsible for the choice made by this species (additional food, holdfasts, protection) can now be investigated.     

Isolation of microsatellite markers for the endangered Knysna seahorse Hippocampus capensis and their use in the detection of a genetic bottleneck

We report the isolation and characterization of 15 (12 di‐, 1 tri‐ and 2 tetranucleotide) microsatellite markers from Hippocampus capensis, the Knysna seahorse. This marker set allows the detection of a genetic bottleneck as shown in a captive population. Furthermore, we test their genotyping potential in eight other seahorse taxa.     

Population genetics of the endangered Knysna seahorse,Hippocampus capensis

The evolutionary history of the endangered Knysna seahorse, Hippocampus capensis, and the extent of gene flow among its three known populations, were investigated using 138 mitochondrial DNA control region sequences. Similarly high levels of genetic diversity were found in two of the populations (Knysna and Keurbooms Estuaries), whereas diversity in the third population (Swartvlei Estuary) was lower. Although most haplotypes are shared between at least two populations, based on the haplotype frequency distributions the three assemblages constitute distinct management units. The extant population structure of H. capensis suggests that the Knysna seahorse originated in the large Knysna Estuary. The presence of seahorses in the two smaller estuaries is either the result of a vicariance event at the beginning of the present interglacial period, colonization of the estuaries via the sea, or a combination of the two.     

MASS MORTALITY OF MARINE AND ESTUARINE FISH IN THE SWARTVLEI AND WILDERNESS LAKE SYSTEMS,SOUTHERN CAPE.

Summary

Two incidences of mass mortality of fish were recorded in two saline waterbodies (Swartvlei estuary and Rondevlei) in the Wilderness National Park. Species affected were the Knysna seahorse (Hippocampus capensis) and longsnout pipefish (Syngnathus acus) in the Swartvlei estuary, and Cape stumpnose (Rhabdosargus holubi) and white steenbras (Lithognathus lithognathus) in Rondevlei. Comparison of water quality parameters during fish mortalities to past ranges and reported species tolerances indicated that the probable causes for the mortalities were high water temperature in the Swartvlei estuary, and low dissolved oxygen concentration in Rondevlei.     

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.     

Thanatosis in the Brazilian seahorse <Emphasis Type="Italic">Hippocampus reidi</Emphasis> Ginsburg, 1933 (Teleostei: Syngnathidae)

The present study reports thanatosis behavior in Hippocampus reidi seahorses. We observed two different types of tonic immobility during seahorse handling, both in the laboratory and in field studies. In the former, the seahorses assumed a stretched posture, and in the latter, pointed their heads towards their bellies and curled their tails in a spiral towards their abdomen. The displays lasted 124 ± 100 s (n = 3) in the laboratory and 155 ± 7.07 s (n = 5) in the field. Camouflage is the seahorses primary defense from predators and our observations suggest that thanatosis is a secondary strategy that could be more effective than retaliation or escaping predators, due to the fact that seahorses lack teeth and have limited swimming ability. The responses caused by fear occurred after manipulation, allowing for the conclusion that the seahorse H. reidi may use thanatosis as a secondary defense strategy under major stress events.     

Optimization of diet and culture environment for larvae and juvenile freshwater pearl mussels,Hyriopsis (Limnoscapha) myersiana Lea, 1856

Summary

Culture of the freshwater pearl mussel, Hyriopsis (Limnoscapha) myersiana, was carried out in three consecutive steps: (1) culture of glochidia larvae in artificial media, (2) rearing the early juveniles (0–120 days old) in a nursery, and (3) rearing the juveniles (120–360 days old) in an earthen pond. The percentage survival of glochidia in standard tissue culture medium (M199) supplemented with common carp plasma was 95±2.5. All surviving larvae (100%) transformed to juveniles, the duration of transformation being 8 days. The early juveniles (0–60 days old) were fed with a mixture of four selected phytoplankton species (Chlorella sp., Kirchneriella incurvata, Navicula sp. and Coccomyxa sp.). The survival rate of juveniles was 8±0.2%. The average length of these juveniles increased from 0.13±0.01 mm to 1.41±0.16 mm and the average height from 0.16±0.01 mm to 0.98±0.09 mm. Subsequently, 60–120-day juveniles were fed with one of the same four phytoplankton species or a combination of the four. Feeding the juveniles with K. incurvata resulted in the highest survival rate (65±8.32%), with an average length of 3.46±0.04 mm and an average height of 1.94±0.04 mm. Finally, the 120–360-day juveniles were cultured in an earthen pond. There were progressive changes in average weight (0.0037±0.002 g to 11.24±5.02 g), length (3.48±0.39 mm to 54.08±6.21 mm), height (1.97±0.24 mm to 25.09±2.48 mm) and width (0.98±0.06 mm to 12.28±3.21 mm) from 120 to 360 days. The average growth rates per day of these parameters were 0.0497±0.01 g, 0.2414±0.15 mm, 0.0975±0.08 mm and 0.0493±0.03 mm, respectively. H. (L.) myersiana juveniles developed the complete structural composition of the adult by 160 days, and at 360 days, gametogenesis was complete.     

Life history and ecology of seahorses: implications for conservation and management

We present the first synthesis of the life history and ecology of seahorses, compare relationships for seahorses with other marine teleosts and identify research needs. Seahorses occurred primarily amidst temperate seagrasses and tropical coral reefs. Population densities were generally low, ranging from 0 to 0·51 individuals m^?2, but reached 10 m^?2 in some patches. Inferred life spans ranged from 1 to 5 years. Seahorses consumed live prey and possibly changed diet as they grew. Growth rates are poorly investigated to date. Reproduction and mating systems are the best‐studied aspects of seahorse ecology. The relationship between size at first maturity and maximum size in seahorses conformed to that for other marine teleosts. All seahorse species were monogamous within a cycle, but some were polygamous across cycles. Direct transfer of clutches to the brood pouch of the male fish made it difficult to measure clutch size in live seahorses. After brooding, males released from c. 5 to 2000 young, depending on species and adult size. Newborn young measured from 2 to 20 mm in length, which was a narrower size range than the 17‐fold difference that occurred in adult size. Newborn body size had no relationship to adult size. Both eggs and young were larger than expected among marine teleosts, even when considering only those with parental care, but brood size at release was lower than expected, perhaps because the young were more developed. The size of adults, eggs and young increased with increasing latitude, although brood size did not. Considerable research is needed to advance seahorse conservation and management, including (a) fisheries‐dependent and fisheries‐independent abundance estimates, (b) age‐ or stage‐based natural and fishing mortalities, (c) growth rates and age at first maturity, and (d) intrinsic rates of increase and age‐ or size‐specific reproductive output. Current data confirm that seahorses are likely to be vulnerable to high levels of exploitation.     

Population dynamics of the long-snouted seahorse (Hippocampus guttulatus Cuvier, 1829) in the Mar Menor coastal lagoon

A 10-year monitoring program was developed to quantify the population dynamics of the long-snouted seahorse population in the Mar Menor coastal lagoon. Based on 985 underwater visual censuses, we estimated the long-snouted seahorse (Hippocampus guttulatus Cuvier, 1829) population size in the Mar Menor lagoon and its reduction in size in the last decades, as well as the effect of eutrophication crises in 2016 and 2019 on the species. The annual recruitment for the 2013–2020 period was estimated by comparing the relative abundance of early seahorse life stages in the ichthyoplankton. The density ranged from 0.0458 specimens/m³ at the beginning of the sampling period to 0.0004 at the end, showing a statistically significant difference between the three analyzed periods (H_gl=2 = 14.0, p = 0.001). The long-snouted seahorse population from the Mar Menor lagoon exemplifies the impact of fishing activities and human pressure, especially euxinic episodes and habitat destruction. As a result of this, the Mar Menor population has decreased from several million specimens to a few thousand, in only three decades. This species showed considerable resilience, the seahorse population began to recover once fishing activity stopped. In contrast, the long-snouted seahorse showed high vulnerability to habitat loss and an episodic flooding event. Adult seahorses showed preferences for highly complex habitats, especially Caulerpa prolifera–Cymodocea nodosa mixed meadows and habitats of high complexity and anthropogenic origin, such as harbors, jetties, or breakwaters. In contrast, juvenile seahorses preferred monotonous seabeds with low complexity, such as the sandy beds that are characteristic of the Mar Menor lagoon littoral.     

Novel life‐history data for threatened seahorses provide insight into fishery effects

Life‐history variables for three incidentally captured species of seahorse (Kellogg's seahorse Hippocampus kelloggi, the hedgehog seahorse Hippocampus spinosissimus and the three‐spot seahorse Hippocampus trimaculatus) were established using specimens obtained from 33 fisheries landing sites in Peninsular Malaysia. When samples were pooled by species across the peninsula, sex ratios were not significantly different from unity, and height and mass relationships were significant for all species. For two of these species, height at physical maturity (H_M) was smaller than the height at which reproductive activity (H_R) commenced: H. spinosissimus (H_M = 99·6 mm, H_R = 123·2 mm) and H. trimaculatus (H_M = 90·5 mm, H_R = 121·8 mm). For H. kelloggi, H_M could not be estimated as all individuals were physically mature, while H_R = 167·4 mm. It appears that all three Hippocampus spp. were, on average, caught before reproducing; height at 50% capture (H_C) was ≥H_M but ≤H_R. The results from this study probe the effectiveness of assessment techniques for data‐poor fisheries that rely heavily on estimates of length at maturity, especially if maturity is poorly defined. Findings also question the sustainability of H. trimaculatus catches in the south‐west region of Peninsular Malaysia, where landed specimens had a notably smaller mean height (86·2 mm) and markedly skewed sex ratio (6% males) compared with samples from the south‐east and north‐west of the peninsula.     

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.     

Two new species of seahorses (Syngnathidae, Hippocampus) from the Middle Miocene (Sarmatian) Coprolitic Horizon in Tunjice Hills, Slovenia: The oldest fossil record of seahorses

Two new and the only known extinct seahorse species Hippocampus sarmaticus and Hippocampus slovenicus are described from the Middle Miocene beds (Lower Sarmatian) in Slovenia, representing the oldest known fossil record of seahorses. Hippocampus sarmaticus was most similar to the extant seahorse species Hippocampus trimaculatus, while H. slovenicus can be most easily compared to the extant pygmy seahorses H. bargibanti, H. denise, and H. colemani. These Sarmatian seahorses lived among seagrasses and macroalgae in the temperate shallow costal waters of the western part of the Central Paratethys Sea.     

Species identification and genetic structure of threatened seahorses in Gran Canaria Island (Spain) using mitochondrial and microsatellite markers

A non-invasive DNA analysis of seahorse populations was carried out after extensive underwater surveys in Gran Canaria Island (Spain). In this geographical area, the presence of two species, Hippocampus hippocampus and H. guttulatus, has been previously reported. Sequencing of 16S ribosomal DNA (16S rDNA) was used for specific identification of live seahorses sampled in situ, as a previous step to evaluate genetic structure based on ten microsatellite markers. Phylogenetic analyses revealed the presence of a single species, H. hippocampus, in the seahorse communities found at Gran Canaria. No evidences of H. guttulatus or interspecific hybrids were found based on 16S rDNA and microsatellite data. The nuclear markers revealed low genetic diversity and lack of population structure across populations of Gran Canaria Island, with evidence of small population sizes. This study provides critical information to support conservation strategies of Gran Canaria seahorses.     

Is filament clipping an effective tool for tissue sampling in <Emphasis Type="Italic">Hippocampus guttulatus</Emphasis>?

Examining genetic diversity, population structure, and geographic distribution has become an important part in the conservation strategies of endangered populations. However, these studies rely on tissue samples collection for DNA analysis which may be problematic for species with a sensitive conservation status. Partial fin-clipping has been employed for tissue collection, due to the increased popularity of DNA-based analysis for ecological and fisheries studies, however, fin clipping can potentially affect behavior, swimming performance, predator avoidance and the ability to find and capture prey. This study aimed to test the effects of filament clipping using captive breed seahorses (Hippocampus guttulatus) as an alternative tool for tissue sample. Clipped filaments regrew on average 0.05?±?0.02 mm/day for 3 months and no mortality or disease was observed during the experiment. Filaments provided enough tissue for DNA analysis. This study provided valuable information regarding a new sampling technique that does not impair the seahorse locomotion. This methodology may be used in population’s genetic studies of other species that have skin filaments as a morphological trait.     

Interannual Variability in Survival Rates for Adult Female White-Tailed Deer

Adult female survival is an important component to population models and management programs for white-tailed deer (Odocoileus virginianus), but short-term survival studies (1–3 yrs) may not accurately reflect the variation in interannual survival, which could alter management decisions. We monitored annual survival and cause-specific mortality rates of adult female white-tailed deer (n = 158) for 6 years (2010–2012, 2016–2018) in southern Delaware, USA. Annual survival rate differed among years. Survival rates (±SE) and mortality causes were similar in 3 years (2011 = 0.72 ± 0.08, 2017 = 0.68 ± 0.08, 2018 = 0.74 ± 0.09) and comparable to previous research from mixed forest-agricultural landscapes. A relatively low survival rate in 2010 (0.48 ± 0.11) was influenced by hunter harvest and potentially compounded by abnormally severe winter conditions in the prior year. A peracute outbreak of hemorrhagic disease occurred during summer 2012, resulting in an annual survival rate of 0.38 ± 0.11, and to our knowledge is the first reported case of a hemorrhagic disease outbreak in a monitored wild population with known fates. In 2016, we did not observe any harvest mortality, resulting in high annual survival (0.96 ± 0.04). Our results demonstrate the degree of variability in annual survival and cause-specific mortality rates within a population. We caution against the use of short-term survival studies to inform management decisions, particularly when incorporating survival data into population models or when setting harvest objectives. © 2020 The Wildlife Society.     

Fisheries and trade of seahorses in Brazil: historical perspective, current trends, and future directions

Seahorses (Hippocampus spp.) are globally threatened by overexploitation and habitat destruction; they are also regarded as susceptible to heavy exploitation due to some of their life-history traits. From an economic perspective, they are fishes with high monetary value and marketability. Seahorses are now listed in Appendix II of CITES (Convention on International Trade in Endangered Species of Wild Fauna and Flora), to ensure that the international trade is not detrimental to the survival of wild populations; the effectiveness evaluation of these international controls needs comparable monitoring data, including evaluation of spatial and temporal trends. This study assesses the seahorse trade in Brazil, aiming to detect trends in catch levels, volumes and prices. Our main findings were: the dried trade was unregulated, without formal records, and primarily domestic, although records of unreported exports existed; it was primarily sustained by incidental captures in trawl nets. The live seahorse trade was mainly destined for exports, and regulated through national quotas. Between 2002 and 2009, mean prices for dried seahorses ranged from US$1.06 ± 0.46 (level 1 traders) to US$1.06 ± 0.46 (level 1 traders) to US2.78 ± 0.68 (end-sellers) each, while mean prices for each live seahorse traded on the domestic market (1997–2009) ranged from US$1.13 ± 0.02 (level 1 traders) to US$1.13 ± 0.02 (level 1 traders) to US10.08 ± 1.71 (retailers). Mean declared export price (2006–2008) was 15.57 ± 10.87. Enhanced implementation of the CITES listing in Brazil will require further research, and additional measures to address both direct and indirect fishing pressure on seahorse populations.