Amorphinopsis atlantica Carvalho, Hajdu, Mothes & van Soest, 2004 (Porifera: Halichondriidae): notes on its distribution in the Colombian CaribbeanAbstract
Amorphinopsis atlantica (Halichondriidae) is the only known species of its genus in the Atlantic Ocean. It is a brittle, massive sponge with variable projections and coloration ranging from yellow to dark green, characterized by oxeas and accessory styles. Originally described from Brazil’s southwestern coast, it has since been reported in several Caribbean sites, typically inhabiting submerged mangrove stilt-roots in coastal lagoons. Although shallow-water sponges in the Colombian Caribbean have been studied, this conspicuous species had not been previously documented in lagoonal systems of the region. This study reports its presence for the first time in the Mallorquín (Atlántico Department) and La Escollera (Magdalena Department) coastal lagoons. The report includes new information on its morphology, habitat, substrata, distribution, and ecology in the Colombian Caribbean. Two potential threats to the species were identified: in some areas of the Mallorquín coastal lagoon, a high degree of sedimentation was observed, causing individuals to become buried within the sediment, while in La Escollera, recent heavy rainfall and runoff events were followed by the disappearance of sponges from mangrove roots. These findings highlight the need to incorporate A. atlantica into local management plans to ensure its protection.
Key words
Coastal lagoon; Conservation; Management; Sediment
INTRODUCTION
The genus Amorphinopsis Carter, 1887 belongs to the family Halichondriidae Gray, 1867 and is represented by 18 valid species (De Voogd et al. 2025). To this genus belong sponges with encrusting to massive shape, occasionally with irregular branches that derive from a voluminous base; its skeleton is composed of smooth oxeas and styles, with their size being the main taxonomic characteristic for differentiating between species (Erpenbeck & Van Soest 2002, Alvarez & Hooper 2011). Amorphinopsis atlantica Carvalho, Hajdu, Mothes & Van Soest, 2004 is the only species within the genus with distribution in the Atlantic Ocean (Carvalho et al. 2004). This species has been recorded from Belize (Diaz & Rützler 2009), Costa Rica (Cortés et al. 2009) México (De La Cruz-Francisco et al. 2019, Castellanos-Pérez et al. 2020, Briceño-Vera et al. 2021), and Brazil (Carvalho et al. 2004, Santos et al. 2018, Veloso-Junior et al. 2023, Praxedes et al. 2024).
In Colombia, the presence of A. atlantica has been mentioned in various contexts, though often incidentally or within studies where the species was not the focus or were part of the grey literature. An early reference appears in Gómez & Rodríguez (2009), who noted its association with certain ophiuroid populations in Cispatá Bay (Córdoba Department). Montaño-Castañeda & Santafé-Patiño (2011) included the species in an analysis of the antioxidant activity of methanolic extracts from several sponges collected in the same bay. A more focused mention was made when A. atlantica was listed in a review of sponge diversity in the Corales del Rosario and San Bernardo National Natural Park, specifically for Tintípan Island mangrove channels (Zea & Díaz-Sánchez 2011). Additionally, Quirós-Rodríguez (2015) reaffirmed the association between A. atlantica and certain ophiuroid species, an interaction later cited by Veloso-Junior et al. (2023) as supporting evidence for the presence of the sponge in Colombia. However, a subsequent study by Quirós-Rodríguez et al. (2017) examined the sponges on the submerged roots of Rhizophora mangle in Cispatá Bay and did not record the presence of A. atlantica, raising questions about the status of the species in that area.
No records or information on the species were previously available from areas further north than the localities mentioned above, although the Magdalena Department is comparatively better known for its sponge diversity. The Atlántico and Magdalena Departments, located in the northern Colombian Caribbean, are key areas for monitoring marine and coastal ecosystems due to the strong influence of the Magdalena River. This river directly affects salinity, turbidity, and other environmental variables, shaping the biodiversity of the region’s aquatic systems. In the Atlántico Department, the Mallorquín coastal lagoon stands out as a strategic ecosystem that supports species of commercial importance, such as fish, mollusks, and crustaceans, on which nearby fishing communities rely. The lagoon also hosts a wide variety of bird species and has gained touristic significance in recent years, particularly with the development of the Mallorquín Eco Park and the restoration of Puerto Mocho Beach (CRA 2021). In the Magdalena Department, La Escollera is an artificial coastal lagoon excavated in the early 1960s on a former salt flat of the local coastal plain, its edges were planted with mangroves around 1985 (Zea & Espada Gómez-Lor 2020). The hard substrata within La Escollera support a diverse and dynamic epifauna with high colonization potential. This artificial lagoon holds economic relevance due to the presence of a commercial boat marina and a nightclub (Zea et al. 2017).
Continuous biodiversity monitoring is essential for understanding the species present in a given area, detecting environmental changes, identifying habitat characteristics, and developing management plans that support the protection and conservation of natural resources. In this context, the objective of this study was to formally document the presence and distribution of A. atlantica in Colombia, with a special focus on the Mallorquín and La Escollera coastal lagoons, two ecosystems of notable ecological and commercial significance.
MATERIALS AND METHODS
Study Area
In the Colombian Caribbean, the climate is influenced by the geographical position of the Intertropical Convergence Zone, allowing for the recognition of a dry season with strong winds (December to April), a transition season (May to July) and a rainy season (August to November) (Pujos et al. 1986, Amaya 2001).
The Atlántico Department is strongly influenced by discharges from the Magdalena River, which contributes 143.9 x 106 tons of sediments annually to the sea (Higgins et al. 2016). The Mallorquín coastal lagoon (Table I) is located in the northeastern margin of the Atlántico Department. It has an area of approximately 830 ha and an average depth of 1 m (CRA 2021) (Figure 1). The lagoon borders the Caribbean Sea to the north, where water enters naturally throughout the year or artificially when residents open one or more inlets in the sandbar. To the east, it is bordered by the Magdalena River, which remains connected to the lagoon through three box culverts that contribute to the inflow of freshwater. To the southeast, it borders the Las Flores neighborhood and to the southwest, the La Playa neighborhood (CRA 2021). Previously, Mallorquín had a permanent inflow of water from the mouth of the León creek in the northwest, but in 2018 this tributary was diverted to the Caribbean Sea. The temperature ranges between 27.6 °C and 31.9 °C (CRA 2021), while the average salinity is 16.4, with variations between 5.8 and 22 depending on the sector (CRA 2018), though over the years these values have fluctuated considerably as they are subject to the aforementioned hydrobiological variations of the coastal lagoon (CRA 2021). Other observation points of the species further south of the coastal lagoon were also included, such as Puerto Velero and Caño Dulce (Table I, Figure 1). Both sectors are on the open ocean but equally influenced by the Magdalena River, characterized by high turbidity and bay-like environmental conditions.
Geographic coordinates of the sampling sites and their respective stations in the Atlántico and Magdalena departments. MCL= Mallorquín coastal lagoon.
Occurrence of Amorphinopsis atlantica in Colombia. Previously documented records included Cispatá Bay and Tintípan Island (Zea & Díaz-Sánchez 2011, Quirós-Rodríguez 2015) — shown as green circles. New records from this study are indicated by red circles and include La Escollera coastal lagoon (LECL) in the Magdalena Department, and in the Atlántico Department: Mallorquín coastal lagoon (MCL), Puerto Velero, and Caño Dulce. In Mallorquín, the six surveyed stations are marked: P1 – Old León Creek outlet; P2 – La Barra; P3 – San Nicolás Lighthouse; P4 – Polvorín; P5 – Punta Félix; and P8 – old garbage dump.
La Escollera coastal lagoon (Table I) is a human-made, small (2.2 ha), shallow lagoon (mean depth of 1.15 m) located located in the northern extreme of the Ensenada de Gaira, in the touristic sector El Rodadero, in Santa Marta city (Figure 1). The lagoon connects to the sea through a straight channel about 400 m long and 9–13 m wide. It has two drains that collect the rain runoff from the neighborhood and the hills adjacent to the area. The area is surrounded by well-developed and preserved mangroves that reach 6 m in height and whose roots are covered by epifauna and algae. The lagoon bottom is muddy with patchy accumulations of shells and gravel. The waters of La Escollera Lagoon are, on average, warm (30.7 °C) and saline (34.3), but become colder during the dry, upwelling season (down to 24.6 °C) and less saline during the rainy season (< 30), especially after downpours (Zea et al. 2017, Zea & Espada Gómez-Lor 2020).
Methods
In the Mallorquín coastal lagoon, two sampling events were conducted. The first, an exploration survey, took place on October 26, 2024, primarily in the Punta Félix area near the Eco Park, to confirm the presence of A. atlantica in the lagoon. The second campaign was carried out on January 27, 2025, during which six sampling mangrove stations were established throughout the study area, selected based on key points of interest: P1 – former León creek outlet, P2 – La Barra, inlet at the sandbar (connection with the sea), P3 – San Nicolás Lighthouse (near the first box culvert), P4 – Polvorín (near the second box culvert and the Las Flores neighborhood), P5 – Punta Félix (adjacent to the Eco park) and P8 – old garbage dump (Figure 1). At each station, salinity was measured using an Atago salinometer, and the presence of A. atlantica was assessed. When present, sponge fragments were collected and fixed in 96 % ethanol. The material was collected mainly on submerged roots of R. mangle, with smaller amounts found detached from the substratum (floating) and on a glass fragment. Additional collections from both natural (rocks) and artificial substrata (wooden piles) were carried out during sporadic field trips to Puerto Velero and Caño Dulce, following the same collection protocols described above. In La Escollera coastal lagoon samples were collected on September 20, 2018, by snorkeling around submerged roots of R. mangle, which were subsequently fixed in 96 % ethanol.
All collected material was deposited in the collection of the Museo de Historia Natural Marina de Colombia “Makuriwa” in Santa Marta, Colombia (acronym INV POR).
For the study of the sponge skeleton, permanent mounts of spicules and thick tangential and perpendicular sections were made following standard methodologies (Zea 1987). Spicules were observed in the laboratory to confirm the taxonomic identification. The width and length of 20 spicules of each type were measured for each specimen; skeletal structures were also measured. Measurements were taken using a micrometer adapted to one of the oculars of a Zeiss Primo Star microscope. Measurements are reported as minimum–mean–maximum length / width.
RESULTS
The presence of A. atlantica in the Mallorquín coastal lagoon was confirmed at four of the six surveyed sites (P2, P3, P5 and P8). At site P2, a high degree of sediment accumulation was observed to the extent that the sponges were found still alive but almost completely embedded within the sediment. The sites within the lagoon where the sponge was not found coincide with areas that have been documented as experiencing the greatest environmental stress. One such area is the former outlet of León creek (P1), which received freshwater after passing through the nearby urban center of Barranquilla, Colombia’s fourth most populous city. Another is the area near the Las Flores neighborhood, which is subject to significant anthropogenic pressure associated with dense human occupation and coastal use. In contrast, at the sites where the sponge was present, it was observed colonizing the roots of R. mangle, a mangrove species undergoing recovery within the lagoon ecosystem (Figure 2a, c and d); the sponges were found at very shallow depths (between 0.1 and 0.3 m). It is important to note that the coastal lagoon is undergoing sedimentation processes.
Amorphinopsis atlantica in situ at different localities in the Atlántico and Magdalena Departments. a: Mallorquín coastal lagoon (MCL), on a mangrove root (P5, October 26, 2024), specimen associated with oysters; b: MCL, on a rock adjacent to mangroves (P5, October 26, 2024); c: MCL, on a mangrove root (P5, October 26, 2024), showing a more branched morphology; d: MCL, on a mangrove root (P2, January 27, 2025); the sponge is partially buried in sediment; e: Caño Dulce (June 13, 2024); specimen attached to rocks in the intertidal zone, exposed during low tide but located in a shaded area under trees; f: Puerto Velero (June 13, 2024); specimen on wooden pilings, associated with macroalgae and exposed during low tide, although shaded by the floor of a nearby cabin; g: La Escollera coastal lagoon (LECL) (September 26, 2018); specimen on a mangrove root; h: LECL (September 20, 2018); specimen on a mangrove root at 0.5 m depth; i: LECL (September 26, 2018); specimen on rubble.
In other places further south of Mallorquín in the open Caribbean Sea, the species was also observed, although less developed. In Caño Dulce, A. atlantica was found among rocks at a depth of less than 0.2 m (Figure 2e). In Puerto Velero, it was widely observed on the tourist wooden cabin pilings near the shore (Figure 2f). In La Escollera Coastal lagoon A. atlantica was also found colonizing mangrove roots (Figure 2g-i).
Systematic treatment
Phylum Porifera Grant, 1836
Class Demospongiae Sollas, 1885
Order Suberitida Chombard & Boury-Esnault, 1999
Family Halichondriidae Gray, 1867
Genus Amorphinopsis Carter, 1887
Amorphinopsis atlantica Carvalho, Hajdu, Mothes & van Soest, 2004 (Figure 2a-i)
Examined materials
INV POR2151(13/06/2024, Caño Dulce), INV POR2152 (26/10/2024, Mallorquín Coastal Lagoon), INV POR2153 (26/10/2024, Mallorquín Coastal Lagoon), INV POR2154 (26/10/2024, Mallorquín Coastal Lagoon), INV POR2156 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2157 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2158 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2159 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2160 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2161 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2162 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2163 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2164 POR-028 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2165 (27/01/2025, Mallorquín Coastal Lagoon), INV POR2166 (08/03/2025, Puerto Velero), INV POR1409 (20/09/2018, La Escollera Coastal Lagoon), INV POR1410 (20/09/2018, La Escollera Coastal Lagoon).
Taxonomic description
External morphology
The color in vivo is usually bright yellow —the specimens found in Caño Dulce and Puerto Velero were greenish—; after fixation in ethanol, the sponges turn beige or completely white. Encrusting to massive shape, surface rugose, up to 10–15 cm in diameter. Some specimens develop more narrow-to-wide ramifications or fistules, or lobulate protrusions, some crowned with oscules. The ramifications may be sparse or dominate the form, making it somewhat bristle.
Skeleton
Ectosome easily detachable, as a tangential carpet of scattered and confusingly intersecting spicules, sometimes supported by subsurface tangential multi-spicule tracts. In the fistules, there is a kind of cortex of spicules perpendicular to the surface. Choanosome of ascending multi-spiculated tracts 27–58 μm in diameter, irregularly intersected by thinner ones, 26–40 μm in diameter (Figure 3a-d).
Skeleton and spicules of A. atlantica. a: Choanosome; b: Perpendicular section of the surface of a fistule; c-d: Tangential section of the ectosome; e: Styles; f: Oxeas; g: Spicules with malformations; h: Details of spicules malformations.
Spicules
Two types of spicules, both smooth and straight or slightly curved: oxeas (150–475–810 / 2.1–13–30 μm) and styles (88–205–710 / 3–7–25 μm), oxeas on average much larger than styles (Table II, Figure 3e-f).
Comparison of styles and oxeas dimensions of Amorphinopsis atlantica from Brazil, Mexico, and the present records from Colombia. Values are expressed in μm as minimum – mean – maximum length/ width (n = 20). MCL= Mallorquín coastal lagoon; LECL= La Escollera coastal lagoon.
Ecology
Specimens were found attached to R. mangle roots, rocks, wooden structures and debris, rubble, and glass fragments in very shallow waters, at depths of no more than 0.5 m. In some cases, parts of the sponge were exposed to air during low tide. Polychaete annelids were found inside the sponge, while small crabs and amphipods were observed on its surface. In several instances, the sponge was partially covered with sediment. In the Mallorquín coastal lagoon, salinity measurements at the time of collection ranged from 25.6 to 29.3. In Caño Dulce and Puerto Velero, salinities were 27.3 and 27.8, respectively. In La Escollera, salinity was 34.8.
Remarks
Some individuals found in the Mallorquín coastal lagoon exhibited a specific type of spicule malformation, regardless of spicule type. This variation involved a localized thickening in one or two areas of the spicule, forming a structure that resembled an incomplete sphere, but not a true tyle (Figure 3g-h). Due to their low frequency, considered rare, and their absence in many of the observed specimens, these altered spicules are interpreted as malformations rather than as a distinct spicule type for the species.
The presence of Amorphinopsis maza (de Laubenfels, 1954) in La Guajira, in northern Colombia, was reported by Díaz & Zea (2008). According to unpublished data from the undergraduate thesis from which this record originally derived (Díaz 2007), the specimen (INV POR1105) exhibits morphological characteristics as: preserved in alcohol displays a dark violet coloration, possesses an extremely fragile consistency with a smooth surface, incorporates gravel and sediment within the skeletal structure, presents three distinct spicule types: oxeas, strongyloxeas, and styles. These features are inconsistent with those of A. atlantica and given that A. maza is native to the Pacific Ocean, its occurrence in the Caribbean is unlikely. Therefore, it is plausible that the Colombian specimen represents a distinct, potentially undescribed species. A thorough re-examination of specimen INV POR1105 is needed to reassess its taxonomic identity and to enhance the current understanding of the genus Amorphinopsis in the Atlantic Ocean.
DISCUSSION
In the Atlántico Department there are no specific studies focused on the taxonomic diversity of sponges. The only available reports are from CRA (2016), which recorded the species Haliclona (Reniera) implexiformis (Hechtel, 1965) (formerly Adocia implexiformis), Xestospongia muta (Schmidt, 1870), Hyrtios proteus Duchassaing & Michelotti, 1864, and Cliothosa delitrix (Pang, 1973) (formerly Cliona delitrix). However, there are no reference specimens for these records. Notably, A. atlantica, a conspicuous species due to its size, was not previously observed, suggesting that its presence in the Department is relatively recent. Therefore, this study constitutes the first official record of A. atlantica in the Atlántico Department. In contrast, the Magdalena Department has been the focus of more extensive research on sponge diversity (Wintermann-Kilian & Kilian 1983, 1984, Zea 1987). Nevertheless, A. atlantica had not been previously reported there either, which again suggests a recent colonization. Thus, this study also contributes a new record for the sponge inventory of the Magdalena Department.
The spicule dimensions of some specimens were, on average, larger than those reported in the original description by Carvalho et al. (2004) (Table II). This observation is consistent with the findings of Santos et al. (2018) and supports the idea of intraspecific variation in spicule size. Additionally, our study documents variability in certain spicules, including potential malformations. Although the precise cause of these abnormalities remains unknown, they may be associated with environmental stressors, such as fluctuations in salinity, increased sedimentation, or exposure to pollutants, that could interfere with normal spiculogenesis.
Amorphinopsis atlantica has been primarily documented as an epibiont on the prop roots of R. mangle (Santos et al. 2018, Castellanos-Pérez et al. 2020, Veloso-Junior et al. 2023), which is consistent with observations in the Atlántico and Magdalena Departments. In these areas, the species was found mainly on mangrove roots, but also on glass fragments, wooden pilings, and rocks, all located in very shallow waters, less than half a meter deep, in environments characterized by high turbidity and fluctuating salinity. Similar environmental conditions were reported by De La Cruz-Francisco et al. (2019) in a lagoon system of the Gulf of Mexico, where the species was found partially covered by sediment at depths below 1 m, in muddy substrates with highly turbid waters, meso- to polyhaline salinity (ranging from 20 to 40), and surface temperatures between 25 and 30 °C. In Brazil, the species was observed during spring low tides, frequently exposed to sunlight and desiccation (Veloso-Junior et al. 2023). Notably, and in agreement with our findings, some specimens in Caño Dulce and Puerto Velero also experienced periodic exposure to solar radiation and desiccation.
It remains unclear whether the species appeared in the Mallorquín coastal lagoon only in recent years, gradually colonizing mangrove roots, or whether it has been present all along but was previously overlooked. The latter seems unlikely given the massive morphology of the species and its prevalence on mangrove roots. However, according to local fishermen the species has been observed in the area for at least the past five years. These accounts align with the observations of Veloso-Junior et al. (2023), who suggest A. atlantica as exhibiting invasive behavior in Iguape Bay, thriving in estuarine environments with fluctuating salinity. The same authors, along with other studies from Brazil and Gulf of Mexico (Santos et al. 2018, De La Cruz-Francisco et al. 2019), conclude that populations of this species act as opportunists, colonizing any available hard substrata, including ports, shipyards, and fishing traps, which facilitates their spread. In Mallorquín its apparent recent establishment and conspicuous development may be associated with environmental changes, such as fluctuations in salinity, increased sedimentation, and altered hydrodynamic conditions within the lagoon. A similar situation is reported for the mussel Mytella strigata (Hanley, 1843), which had no historical records in the area but has been observed over the past decade colonizing mangrove roots in the coastal lagoon and even forming banks on the sediment (personal observations). As with A. atlantica, it is not yet possible to determine whether its presence is truly recent or whether it had gone unnoticed until now.
In addition to the above, the high degree of sedimentation that has affected the coastal lagoon in recent decades (CRA 2021) poses a significant threat to its biodiversity. This is evident at P2, where A. atlantica specimens are almost completely buried in sediment. Similarly, recent complementary observations at La Escollera revealed the loss of A. atlantica on mangrove roots following heavy rainfall events during the rainy season of 2019. These findings underscore the importance of conducting faunal inventories, particularly in hydrologically dynamic ecosystems such as coastal lagoons. The potential loss of undescribed or unrecorded species represents a serious risk to the full recognition and conservation of the country’s biodiversity.
Given these threats, it is recommended to implement long-term monitoring programs that include both biological and physico-chemical variables to track ecosystem changes. Moreover, adaptive management strategies should be developed to mitigate sedimentation and hydrological alterations, prioritizing habitat protection for key and potentially vulnerable species such as A. atlantica. Such efforts will enhance our understanding of species distribution, resilience, and the impacts of environmental disturbances, ultimately contributing to more effective conservation planning.
CONCLUSIONS
The present study constitutes the first confirmed record of A. atlantica in the Atlántico and Magdalena Departments, expanding the known distribution of this species in the Colombian Caribbean. Its conspicuous presence on mangrove roots, as well as on artificial and natural substrata in highly turbid, shallow environments, suggests a recent colonization event potentially driven by environmental changes. Observations from local fishermen and environmental parallels with other regions support the hypothesis that A. atlantica behaves as an opportunistic species, capable of rapidly establishing in disturbed or fluctuating ecosystems.
The variability observed in spicule size and the presence of malformations reinforce the notion of intraspecific plasticity, possibly influenced by environmental stressors such as salinity fluctuations, sedimentation, and pollution. These findings not only highlight the ecological adaptability of A. atlantica but also emphasize the vulnerability of coastal lagoon ecosystems to anthropogenic and natural disturbances.
The lack of historical data and baseline inventories in these regions represents a significant barrier to understanding biodiversity dynamics and responding effectively to environmental change. The case of A. atlantica, along with similar observations for the bivalve M. strigata, demonstrates the urgency of establishing consistent monitoring programs and integrating local ecological knowledge to improve detection of species introductions or shifts in community structure.
Acknowledgements
We express our gratitude to the fishermen of the Mallorquín coastal lagoon for their assistance during fieldwork and for sharing their valuable knowledge. Sven Zea thanks to the Instituto de Estudios en Ciencias del Mar (CECIMAR) of Universidad Nacional de Colombia, Caribbean Campus, the José Benito Vives de Andréis Institute for Marine and Coastal Research (INVEMAR) and Roberto Lemaitre, tenant of La Escollera nightclub, who granted unrestricted access to the lagoon. The Atlántico Department material was collected under the biological specimen collection permit granted to Universidad del Atlántico by ANLA, based on Resolution 1214 of September 29, 2017, and Ethics Resolution 00594 of April 26, 2018, as part of the project “El litoral del Departamento del Atlántico (Colombia): LDA”. Sampling at La Escollera was conducted under Decree 309-2003 of the Ministry of Environment and Sustainable Development, as part of the ongoing project “Sponges of the Colombian Caribbean” of INVEMAR’s Makuriwa – Museo de Historia Natural Marina (MHNMC) de Colombia. We thank the MHNMC for accessioning the material, with special thanks to Erika Montoya for her support. This work is a contribution of the research group “Geology, Geophysics, and Marine - Coastal Processes” at Universidad del Atlántico (Colombia), and “Colombian Marine Fauna, Biodiversity and Uses” of Universidad Nacional de Colombia. Contribution 596 of CECIMAR and 1413 of INVEMAR.
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Edited by
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Handling editor
Claudio Riccomini
The data for this study are available from the corresponding author by reasonable request.
