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3 November 2025

Influence of Lunar Periodicity on Medusae (Cnidaria) Composition in a Western Caribbean Reef: Community Structure Before Sargassum Blooms

,
,
and
1
Centro de Investigación de los Sistemas Costeros y Continentales, Universidad Autónoma de Chiapas, Av. Las Escolleras s/n, Puerto Madero 38000, Chiapas, Mexico
2
Departamento de Sistemática y Ecología Acuática, El Colegio de la Frontera Sur (ECOSUR), Chetumal 77014, Quintana Roo, Mexico
3
Instituto de Ciencias del Mar y Limnología, Unidad Académica Puerto Morelos, Universidad Nacional Autónoma de México, P.O. Box 1152, Cancún 77500, Quintana Roo, Mexico
*
Author to whom correspondence should be addressed.
Diversity2025, 17(11), 769;https://doi.org/10.3390/d17110769
This article belongs to the Topic Taxonomy and Ecology of Zooplankton
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Abstract

The medusae of the Mahahual reef, in the Mexican Caribbean Sea, were studied to document changes in species composition and abundance over a lunar cycle in 2001–2002. Plankton was sampled during two months of the dry and rainy seasons, in the fore reef, channel, and reef lagoon. Fifty-two species were collected. The highest abundance and species richness occurred during the rainy season. Their composition and abundance were similar in the fore reef and channel, but different in the reef lagoon. Abundance and biomass changed among seasons, reef zones and lunar phases; the highest abundance and the lowest biomass were recorded during the full moon. The fore reef and channel were dominated by Liriope tetraphylla and Aglaura hemistoma, the reef lagoon by Cubaia aphrodite and Slabberia halterata. Pennaria disticha and Bougainvillia frondosa were exclusive to the new moon and Pelagia noctiluca and Aequorea macrodactyla to the full moon. The results suggest that the medusae assemblage do not change species composition during the lunar cycle of either season, and abundance increases during full moon. The oceanic influence and tide currents explain the presence of oceanic species and the similarities between localities, but they do not explain the increase in abundance during the full moon. This study was conducted prior to the arrival of Sargassum influxes in this region and can serve as a reference point for assessing its effects in recent years.

1. Introduction

Lunar periodicity has been reported to induce intrinsic behavioural rhythms in copepods [,,], annelids [,], and overall zooplankton biomass []. Although predation is the most widely supported hypothesis explaining lunar rhythms [], little attention has been given to fluctuations in the populations of primary zooplankton predators, such as gelatinous zooplankters. Medusae, for instance, are important predators of zooplankton [,] and can alter the entire planktonic food web [,].
Medusa reproduction has been shown to be triggered by both biotic and abiotic factors, including food availability, temperature, salinity, and light intensity [,,]. However, studies addressing the influence of lunar periodicity on medusae are scarce, despite evidence that medusae production is linked to the third quarter of the lunar cycle [].
Several studies conducted along the Mexican Caribbean coast have described the species composition, distribution, and spatiotemporal fluctuations of medusae [,,,,,,,,,]. Nevertheless, the potential role of lunar periodicity, which strongly influences other zooplankton groups, remains unexplored in this region of coral reefs. Despite the fact that these ecosystems have been shown to harbor a diverse population of medusae [,] and that variations in their abundance and diversity have been associated with factors such as circulation patterns, wind regimes, vertical migration, upwelling events, and food availability, among others [,].
Considering the recent changes in the Mesoamerican reef [], the aim of the present study was to document the variations in abundance and species composition of the reef-associated medusae across two seasons, three reef habitats, and three lunar phases, to identify potential lunar periodicity in the medusa assemblage dynamics during the years 2001–2002. This study was made in Mahahual reef, prior to the massive arrivals of Sargassum in the Mexican Caribbean [], so it can be a reference point before this periodic event.

2. Materials and Methods

The study area is located in the western Caribbean Sea, near the southeastern Yucatan Peninsula, Mexico. This coast is characterized by a narrow shelf [] and receives Caribbean waters that flow through the Yucatan Channel into the Gulf of Mexico. The main current flows northward; however, a near-shore counter-current flows southward. These flows interact to generate elongated eddies along the coast []. The second-largest barrier reef system in the world extends along the Caribbean coast from Isla Contoy in the north to the Belizean coast in the south [].
Mahahual is located at 18°42′26″ N and 87°42′87″ W, where the reef forms a narrow and shallow lagoon. The reef crest is interrupted by two channels that connect the reef lagoon with the Caribbean Sea (Figure 1). The reef lagoon is shallow, with a mean depth of 1.5 m, and is 30–180 m wide. Benthic communities consist mainly of Thalassia testudinum (Koening) in the lagoon, and coral cover is minimal in the shallowest part, but increases towards the reef []. The barrier and its diversity have been described as a high-priority system for conservation [].
Figure 1. Map of the Mahahual reef showing the position of sampling sites and the shape of the reef. The sampling sites were located in the three main reef zones (abbreviations as explained in Abbreviations Section) approximately 1 km from the coast, the continental shelf ends, and the ocean sinks to 200 m.
Sampling was carried during the night (after 21:00 h) in two regional seasons: rainy season (October–November 2001) and dry season (April–May 2002), each encompassing two lunar cycles, including new moon, full moon, and first quarter moon phases. Samples were taken from three typical reef zones: fore reef (Stations 1, 3), channel (Stations 2, 4), and reef lagoon (Stations 5, 6) (Figure 1).
Two replicates were taken at the surface using a circular-mouth net, with a diameter of 0.45 m and a mesh size of 330 µm. Water flow through the net was measured with an electronic flowmeter, E-Flow (Hydro-Bios), attached to the mouth. All sampling stations were less than 10 m deep. Samples were fixed in a 10% buffered formalin solution. Since no tidal data were available for that time in the Mahahual reef, we used data from a nearby locality, Isla de Cozumel, located 270 km north. According to Kerfve [], the whole Mexican Caribbean has a similar tidal range. Salinity was measured with a refractometer, and superficial temperature (°C) was recorded using a Taylor thermometer. Wet-weight zooplankton biomass was measured after removing medusae accordingly with the methods proposed by Smith and Richardson [].
Medusae were isolated from other zooplanktonic organisms, identified based on taxonomic descriptions and compilations available in the literature [,,,], and counted.
The theoretical maximum number of medusa species was estimated according to the stochastic theory of species accumulation [] using the exponential model. The model is appropriate when the taxon is poorly known, and thus the probability of finding a new species never reaches zero. This model calculates the expected number of species as a function of sampling effort []:
S ( t ) = 1 z l n ( 1 + z a t )
where
  • S(t) being the expected value of species;
  • z is curvature or saturation parameter;
  • a is the initial species discovery rate parameter;
  • t is the observed accumulated species number.
This information was used to estimate the effectiveness of the sampling effort, and the theoretical expected value was compared with the observed values. Data analysis was performed using STATISTICA v4.3 for Windows.
Biomass and abundance were standardized (g100 m−3 and org100 m−3, respectively) and log-transformed for statistical analysis. A three-way ANOVA was performed on abundance data grouped according to three factors: season, reef zones, and moon phases. A Tukey post hoc test [] was applied when significant differences were found. These analyses were performed using Statgraphics software version 7.
To describe the spatiotemporal structure of the medusa assemblage during each season, all taxa, including Obelia, were considered. The Importance Value Index (IVI) and Dominance Index were calculated for each reef zone. Diversity metrics, including Shannon–Wiener’s (H’), evenness (J’) [], and species richness were calculated in the same way.
Multivariate classification analysis was conducted to identify groupings associated with the measured factors. Clusters were determined using the Bray–Curtis similarity index and single-linkage clustering []. Multivariate analyses were performed using ANACOM software V3.0 [].

3. Results

3.1. Environmental Data

During rainy season (RS) and dry season (DS), the tide reached its maximum amplitude during two moon phases, new moon (NM) and full moon (FM), with an amplitude ranging from 0.18 to 0.19 m (above the minimum low tide level). The tidal amplitude during first quarter moon phase (FQ) was slightly lower, ranging from 0.14 to 0.16 m.
Surface water temperature ranged from 26.5 to 29.1 °C during RS and from 28.5 to 29.5 °C during DS; the mean difference between seasons was 1 °C. In the reef lagoon (RL), the temperature was lower, but in all cases the difference was less than 1 °C, compared to other reef zones. There were no significant temperature differences among the three moon phases (Table 1).
Table 1. Surface mean temperature (°C), salinity and zooplankton biomass (g100 m−3; without medusae) in reef zones (CH, FR, and RL), moon phases (FM, FQ, and NM) and seasons (RS, DS) in the Mahahual reef.
Salinity values were more uniform during RS (36 to 39) than DS (32 to 38) (Table 1). The average difference between seasons was 1.

3.2. Composition and Assemblage Structure

The taxa recorded included, two cubomedusae, five scyphomedusae, and 45 hydromedusae, 11 species are new records for the Mexican Caribbean. In total, 45 medusa species were recorded, of which 38 corresponded to RS and 35 to DS (Table 2).
Table 2. Total abundance (100 m−3), relative abundance in each season (%), and relative abundance in all sampling periods (% total) of medusae in the Mahahual reef, Mexico, during the rainy season of 2001 (RS) and dry season of 2002 (DS). * New records in the Mexican Caribbean.
According to the exponential equation, the expected number of taxa was 53 (R2 = 0.98 with 97.65% of the variance explained). The 52 taxa registered represent 98% of the theoretical maximum (Figure 2).
Figure 2. Species accumulation curve. Fitted line is exponential predicted species model. Sampling effort, take 10 min per trawl.
In both seasons, only two species were recorded exclusively during the new moon: Pennaria disticha restricted to RL in RS, and Bougainvillia frondosa, which occurred throughout the entire study area. Two species were exclusive to FM, Pelagia noctiluca and Aequorea macrodactyla, both found in FR and CH. No species were found exclusively during the FQ moon.
Important species after their dominance in the RS (IVI > 10) were Aglaura hemistoma, Cubaia aphrodite, Liriope tetraphylla, and Slabberia halterata. In the DS, they were A. hemistoma, Corymorpha forbessi, and L. tetraphylla. During RS, the FR, and CH zones were dominated by A. hemistoma, and L. tetraphylla, and S. halterata (DI > 10%) across all three moon phases; C. aphrodite and S. halterata were dominant (DI > 10%) in the RL across all moon phases. In DS, FR, and CH were dominated by A. hemistoma, C. forbessi, and L. tetraphylla (DI > 10%), whereas C. aphrodite and Obelia spp. were the dominant species in the RL across all moon phases (Figure 3).
Figure 3. Dominant species (dominance index ≥ 10%) in each reef zone at three different moon phases in both rainy season (A) and dry season (B). Reef zones (CH, FR, RL).
Diversity Index values were highest in the CH (3.3 bits ind−1) and FR zones (2.8 bits ind−1) during NM of RS. In the same season, the lowest diversity value was recorded at the FR (1.3 bits ind−1) during FQ moon. During DS, the highest diversity values were found in the FR during NM (2.7 bits ind−1) and the lowest diversity values in the RL (1.1 bits ind−1) in FQ moon. The evenness values were higher in NM than FQ or FM of both seasons. Richness index values were higher during FM in both RS and DS.
Based on the abundance and occurrence of taxa across reef zones and moon phases, four groups were identified in RS, and three in DS were found (Figure 4). RS group A (Figure 4A) included 15 taxa, mainly found in all reef zones, with no specific lunar preference, except two subgroups: A’, consisting of species found only in FR and CH during FM, and A’’ characterised by taxa recorded only in RL during NM. Group B included mainly neritic species collected in CH and LA during FM and NM. Group C was composed of taxa found only in FR and CH, with no lunar preference. The fourth group, D, included species restricted to CH during NM.
Figure 4. Similarity dendrograms according to the Bray–Curtis index. Groupings by species and abundance are shown for the rainy season (A) and the dry season (B). A’ species found only in FR and CH during FM. A” taxa recorded only in RL during NM.
In the DS (Figure 4B), group A included both oceanic and neritic species with no specific distribution or lunar preference. Group B consisted mainly of neritic species collected only in CH and RL during NM and FQ. The third cluster included oceanic and mixed (neritic and oceanic) species found in FR and CH during FM and FQ.

3.3. Biomass

Mean biomass values ranged from 7.7 to 15.3 g100 m−3 during RS and from 3.6 to 17.1 g100 m−3 during DS (Table 1). The differences in biomass values were statistically significant between seasons (p < 0.05), among reef zones (p < 0.05), and among moon phases (p < 0.05). No significant interactions among factors were found (p > 0.05). High biomass values were found during NM and slightly decreased towards FM in both seasons (Table 1).

3.4. Abundance

A total of 18,657 medusae were collected. The highest mean abundance was recorded in the FR, with 380.3 and 383.4 org100 m−3 during FM in RS and DS, respectively. The lowest mean abundance was recorded in the FR, with 45.0 and 71.8 org100 m−3 during NM of RS and DS, respectively.
Abundance values were significantly different between seasons (p < 0.05), moon phases (p < 0.05), and reef zones (p < 0.05). Abundance was higher in RS than in DS (Figure 5A). In both seasons, abundance was the highest during FM (Figure 5B,E) and the lowest in NM (Figure 5E). No statistical differences were found between FR and CH zones across seasons and moon phases; RL was distinct, with high data variability (Figure 5C). The interaction between factors was significant only between zones and moon phases (p < 0.05); no differences were found among moon phases (Figure 5E; p > 0.05) or reef zones during either season (Figure 5D; p > 0.05).
Figure 5. ANOVA analysis for each factor and their interactions: (A) Seasons; (B) moon phase; (C) reef zones; (D) between seasons and reef zones; (E) between seasons and moon phases. Abbreviations as explained in Abbreviations Section.
These results suggest that the medusae assemblage exhibits a similar temporal pattern across seasons, but with seasonal varying abundances (Figure 5D,E).

4. Discussion

Some new records for the Mexican Caribbean are notable considering their biogeographic patterns. For example, Pandeopsis ikarii, mainly reported in the Indo-Pacific [], was documented in the Atlantic Ocean, Florida, for the first time in 2021 []. Euphysilla pyramidata has a circumglobal distribution in tropical seas, including the Gulf of Mexico [,]. Laodicea marama, Turritopsis nutricula, and the scyphomedusa Nausithoe maculata, whose distributions correspond to the eastern Caribbean Sea [,,], highlight the limited taxonomic studies on these organisms and, consequently, the scarcity of records from Mexican waters, despite being previously recorded in neighbouring areas, as in Cuban waters [].
The number of species found in this study was high compared to previous research in the reef and adjacent areas. For example, in the reefs of Puerto Morelos and Mahahual reef [,], as well as in large adjacent estuarine embayments, such as Chetumal Bay [], Ascensión Bay [], Bojórquez lagoon near Cancún [], and the nearby Banco Chinchorro [], the number of recorded species was low (<20) in each case.
In contrast, in the reef of Carrie Bow Cay, Belize [], in the Yucatan Shelf and northern Caribbean [], and in the PROIBE oceanographic campaigns carried out on the Campeche Bank and the pelagic zone of the Caribbean, more than 40 species were found per sampling effort. These surveys were conducted with trawls on different environments, and the probability of finding more species in marine or reef habitats than in estuarine embayment’s is high. However, it must be assumed that the number of species recorded depends on sampling effort. Additional sampling would likely reveal the presence of rare species []. The higher number of species found in our study is attributed to a greater sampling effort, and it was made during the night. The results of the exponential model confirmed that our sampling effort was representative of the medusae composition.
The species found in the Mahahual reef represent 47.7% of the total medusa species reported for the Mexican Caribbean until 2003 []. It is important to note that 82.4% of the species reported by Suárez-Morales et al. [] were also found in this survey, and 36.6% of the species in Mahahual are shared with Carrie Bow Cay, in Belize [], located 208 km to the south. In total, we found 12 of the 16 species reported in Bojorquez lagoon []. Although Gasca et al. [] sampled 19 sites both inside and outside Banco Chinchorro reef, the lower number of species they recorded was likely because all their sampling was conducted during the day and restricted to a single region in each sampling effort. Diversity values were slightly higher in NM than FM, and evenness values were also higher in NM. This indicates that the number of species was similar between moon phases, but during FM in both seasons, the community was notably dominated by a few species. This pattern was also supported by the Importance Value Index (IVI) and Dominance Index (DI) values. In RS, A. hemistoma, C. aphrodite, L. tetraphylla, and S. halterata, were the most important species. In DS, no substantial differences in species composition were observed compared to RS; A. hemistoma, L. tetraphylla, and C. forbesii remained the dominant species. The dominance of holoplanktonic A. hemistoma and L. tetraphylla has also been reported in waters of the Campeche Bank, Yucatan shelf, and the Caribbean [,,], although both species are considered mainly of the pelagic zone [].
The temperature and salinity values were within the recorded range for the Mexican Caribbean [], and were similar across seasons, moon phases, and reef zones. Apparently, these factors did not influence the medusa composition. However, there are other factors that can influence the composition and abundance of medusae, for example, light affects many behavioral activities in medusae, including diel vertical migration and reproduction, whether or not distinct ocelli [,].
The Caribbean tide is considered microtidal (0.20 m); however, horizontal and vertical currents have been recognized as major factors in coastal regions, strongly influencing the distribution of organisms []. During the FQ, the outflow promotes vertical eddies in the FR, leading to increased neritic water influence across all reef zones; during NM and FM, the inflow promotes similar eddies in the RL zone, resulting in greater oceanic influence, particularly in the FR []. It is evident that tidal movements and the strong influence of the main oceanic current [,] may explain the persistence of pelagic species such as A. hemistoma and L. tetraphylla on the reef, and consequently the overlap of coastal and pelagic species in a relatively small area.
Nevertheless, tidal currents alone cannot fully explain the increased abundance observed during FM; in all reef zones, most species found in relatively low abundance during NM, were the same as those recorded in higher abundance during FM, despite similar tidal magnitudes.
Gili et al. [] and Suárez-Morales et al. [] suggested that the arrival of oceanic species causes only a local enrichment in species composition but does not significantly increase total abundance. In Mahahual, this was not the case. During FQ, when the tide was low and presumably the oceanic influence was lower, the diversity, evenness, and richness indices were lower than during the other moon phases. Conversely, during NM and FM, when the tides and oceanic influence were higher, these indices increased, particularly during NM, while abundance increased only during FM.
Cluster analyses revealed that few species were exclusive to NM (P. disticha restricted to RL, and B. frondosa found across all zones) or FM (A. macrodactyla and P. noctiluca found outside the reef). No species were found exclusively during FQ. These findings, supported by ANOVA statistical analyses and Tukey’s post hoc test, together with the results on richness and diversity, indicate that the specific composition of medusae on the reef remains relatively constant throughout the lunar cycle. However, significant variations in biomass and total abundance were detected, with a significant increase in the abundance of certain species during FM.
Moonlight appears to be more important for some zooplankton groups than others [,]. Zooplankton in North Atlantic waters, particularly copepods, appear to follow a moon-related cycle, with increased abundance around the FM []. Similar patterns have been observed in freshwater copepods [], Caribbean reef copepods [], total zooplankton biomass [], and cubozoan aggregations []. However, experimental studies exposing medusae to artificial light–dark cycles concluded that their vertical migration behaviour was erratic when photoperiods were altered []. These and recent results suggest that some medusae respond to daylight and lack intrinsic rhythms []. Thus, moonlight per se is probably not a direct stimulus for medusae. However, many zooplankton groups are affected by moonlight, and some of them are potential prey for medusae. An example of this is the greater abundance of copepods and fish larvae recorded during the full moon phase in tropical waters [].
Medusae appear to be particularly adapted to take advantage of favorable food conditions to increase their abundance, or, for example, the release of Aurelia aurita ephyrae coincides with the yolk-sac larval stage of herring (Clupea harengus) in the Kiel fjord []. The decline of this larval population is accompanied by a sharp increase in this scyphomedusa []. Hydroids of Obelia spp. released more medusae during the third quarter of the moon []. In addition, a marked decline in zooplankton biomass has been observed in the Canary Islands after peaking during the full moon, suggesting that predation is the main cause of this decline [].
In our study, the increase in medusa abundance during FM could be a response to increased prey availability. It has been proposed that predators like medusae are not strongly affected by light or visibility in the water column, as they are mainly tactile predators and do not stalk or wait cautiously to catch their prey as some fish []. Therefore, if their prey increases around the FM, medusae may respond by increasing their abundance [,,].

5. Conclusions

The medusae in the Mahahual reef comprises a mixture of coastal and pelagic species, with a composition that mostly remains consistent throughout the lunar cycle in both seasons. This study provides evidence that medusae fluctuations in abundance are associated with the lunar cycle, with numbers peaking during FM. However, the underlying causes of this pattern remain uncertain.
Our study can serve as a reference point, since this region in particular has experienced periodic massive inflows of pelagic Sargassum since 2011 []. Because these inflows could modify the specific composition of medusae, given that the medusozoans reported as epibionts of Sargassum and with a medusa phase [] were not recorded in this study, with the exception of medusae of the genus Obelia. This suggests a possible increase in species richness. However, the massive inflows of Sargassum can also have negative effects, since it deteriorates water quality and the local flora and fauna due to the rapid decomposition of the algae along the shoreline [], damaging key substrates such as seagrasses [], for medusa species with a polyp phase (e.g., Clytia spp., Obelia spp.).

Author Contributions

Conceptualization, E.T.-J., L.S.-P. and M.E.-G.; methodology, E.T.-J., M.E.-G. and L.S.-P.; validation, E.T.-J., M.E.-G. and M.A.M.-B.; formal analysis, E.T.-J.; investigation, E.T.-J., M.E.-G., L.S.-P. and M.A.M.-B.; resources, M.E.-G.; data curation, M.A.M.-B.; writing—original draft preparation, E.T.-J., M.E.-G. and M.A.M.-B.; writing—review and editing, E.T.-J., M.E.-G. and M.A.M.-B.; visualization, M.E.-G. All authors have read and agreed to the published version of the manuscript.

Funding

No external funding was received. Field trips and support were provided by personal funds of ME-G.

Data Availability Statement

All data are available with the correspondence author. All requests should be directed to him.

Acknowledgments

We are grateful to Tania Garfias for her valuable collaboration in the field work and her comments to the manuscript; Lourdes Vásquez-Yeomans for her support in statistical procedures and her comments; Rebeca Gasca and Eduardo Suárez-Morales for their comments and support during the development of the study; and Laura Celis for her assistance in medusa identification. We give special thanks to Rosa M. Hernández and Iván Castellanos for laboratory assistance. We also thank Jacobo Schmitter-Soto, Gerald Islebe, and Dorothy Pless for their comments on the manuscript. In memoriam of Lourdes Segura-Puertas, for her support of this study and her enduring legacy in medusae research in Mexican marine environments. We appreciate the editorial processing of our work by the Associate Editor and to the anonymous reviewers for their helpful comments.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this manuscript:
CHChannel
DSDry season
FMFull moon
FQFirst quarter moon
FRFore reef
NMNew moon
RLReef lagoon
RSRainy season

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