# A review of manta and devil ray courtship and mating

M. Carpenter, A. Klimley, A. Boggio-Pasqua, M. Palacios, A. Armstrong, S. Hilbourne, M. Guerrero, M. Lapinski, K. Fuentes, G. Stevens

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M. Carpenter, A. Klimley, A. Boggio-Pasqua, M. Palacios, A. Armstrong, et al. A review of manta and devil ray courtship and mating. Environmental Biology of Fishes, 2025, 109 (1), pp.25. 10.1007/s10641-025-01779-4 . hal-05480165

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# A review of manta and devil ray courtship and mating

M. Carpenter · A. P. Klimley · A. Boggio-Pasqua · M. D. Palacios ·
A. O. Armstrong · S. Hilbourne · M. Guerrero · M. Lapinski · K. Fuentes ·
G. M. W. Stevens

Received: 30 December 2024 / Accepted: 6 October 2025
© The Author(s) 2025

**Abstract** The family Mobulidae (genus *Mobula*), comprising manta and devil rays, represents a wide-ranging, threatened group of species. The courtship and mating behaviours of most mobulid species remain poorly understood, with the majority of the existing literature focusing on the reef manta ray (*M. alfredi*). For this study, new observations were compiled globally via Internet and YouTube searches as well as through engagement with the scientific

community, with particular focus on the lesser-studied species. Each video or photographic observation was analysed and categorised based on known courtship stages. Here, we present 87 new records of courtship/mating behaviour from eight of the ten described mobulid species across 18 countries/territories. All seven stages were recorded in the study: Initiation (n=17), Endurance (n=71), Evasion (n=52), Pre-copulation positioning (n=21), Copulation (n=9), Post-copulation holding (n=2), and Separation (n=6). The videos and photographs presented for the Atlantic pygmy devil ray (*M. hypostoma*) represent a first for this species, while the absence of any

**Supplementary Information** The online version contains supplementary material available at https://doi.org/10.1007/s10641-025-01779-4.

M. Carpenter · A. Boggio-Pasqua · M. D. Palacios · S. Hilbourne · M. Guerrero · K. Fuentes · G. M. W. Stevens
The Manta Trust, Catemwood House, Norwood Lane, Dorset DT2 0NT, UK
e-mail: mmcarpen@eckerd.edu

M. Carpenter (✉)
Department of Biological Sciences, University of Cape Town, Rondebosch, Cape Town 7700, South Africa

A. P. Klimley
Biotelemetry Laboratory, Department of Wildlife, & Conservation Biology, University of California Davis, Fish, CA 95616, USA

A. Boggio-Pasqua
Aix-Marseille Univ, Université de Toulon, CNRS, MIO UM 110, Marseille, IRD, France

M. D. Palacios
Mobula Conservation, 23075 La Paz, Mexico

A. O. Armstrong
Project Manta, School of Science, Technology and Engineering, University of the Sunshine Coast, Sippy Downs, Qld, Australia

M. Guerrero
Fundación Megafauna Marina del Ecuador (Proyecto Mantas Ecuador), Calle F y Calle A, Lote 19 Urb. El Edén, Collaquí, Tumbaco, Quito 170902, Ecuador

M. Guerrero
Galápagos Science Center, Universidad San Francisco de Quito and University of North Carolina at Chapel Hill, San Cristóbal, 200101 Galápagos, Ecuador

M. Lapinski
AILERONS, University of Montpellier, Montpellier, France

K. Fuentes
Manta México Caribe, 77400 Isla Mujeres, Mexico
courtship or mating observations for the longhorned pygmy devil ray (*M. eregoodoo*) and the lack of new observations for the bentfin devil ray (*M. thurstoni*) demonstrate that further investigation is needed for the lesser-studied species. New records were compared to the existing literature and novel behaviours, such as male-to-male pseudo-courtship (documented in four species), were described. Our results substantially expand the body of knowledge on mobulid courtship and mating behaviours. These observations align with the previously described sequential framework, with additional refinements for individual species and novel delineations of species-specific behaviours within the defined stages.

**Keywords** Mobulids · *Mobula* · Copulation · Fish behaviour · Elasmobranch

## Introduction

Elasmobranch reproductive activity varies across species, but shares commonalities in the acts of courtship and mating/copulation (Klimley 1987; Carrier et al. 1994; Pratt and Carrier 2001; McCallister et al. 2020). Reproductive behaviour in rays (Batoidea) typically involves multiple males following a single female (rare cases of two) before biting the female by the male/s to insert their external reproductive organ, the clasper, into the cloaca of the female, with minimal interaction after copulation (Chapman et al. 2003). Courtship behaviours are similar across ray species, including the family Mobulidae (manta and devil rays) (Coles 1910; Sobral 2013). Mobulidae consists of ten extant zooplanktivorous species (Marshall et al. 2009; White et al. 2017; Notarbartolo di Sciara et al. 2019; Bucair et al. 2025). One characteristic of the behaviour of these species is that they exhibit further complexities and intricacies in their courtship.

All mobulids form aggregations, with courtship and mating one of the five primary functions of these gatherings (Palacios et al. 2023). Courtship and mating behaviours of mobulids can occur at or near cleaning stations, in areas of shallow reef structure where symbiotic cleaner fish remove dead tissue and parasites off hosts (O’Shea et al. 2010; Carpenter et al. 2024), or at feeding aggregations (Stevens et al. 2018a; Mendonça et al. 2020) and other unidentified

aggregation sites (McCallister et al. 2020). Stevens et al. (2018a, b) suggested cleaning stations act as leks for mobulids, with 90% of observed courtship behaviour for the reef manta ray [*Mobula alfredi* (Krefft, 1868)] in the Maldives occurring at these sites. Select areas for elasmobranch mating often remain unchanged over decades, as seen in the 30-year persistence of mating areas for both nurse sharks [**Gingelymostoma cirratum* (Bonnaterre, 1788)] at Dry Tortugas, USA (Pratt et al. 2022) and *M. alfredi* at Lady Elliot Island, Australia (Couturier et al. 2018).

Mobulid reproductive periodicity varies between species and locations but occurs on average per adult female once every other year, up to once every seven years (Notarbartolo di Sciara 1988; Marshall and Bennett 2010; Deakos et al. 2011; Kashiwagi 2014; Stevens 2016; Setyawan et al. 2020; Kiyatake et al. 2024). The gestation periods have been identified for three species, with ranges that have been estimated from 10 months for longhorned pygmy devil ray [*Mobula eregoodoo* (Cantor, 1849)] (Broadhurst et al. 2018), to 12–13 months for *M. alfredi* (Marshall and Bennett 2010; Deakos et al. 2011; Stevens 2016), and over 15 months for the spinetail devil ray [*Mobula mobular* (Bonnaterre, 1788)] (Kiyatake et al. 2024). However, gestation periods are mostly unknown in mobulid species. Combined with a K-selected life history strategy, which includes late maturation (ranging from 6 to 15 years for species/sex) and low fecundity, mobulids are highly vulnerable to overexploitation (Dulvy et al. 2014; Lawson et al. 2017; Stevens et al. 2018a: Stewart et al. 2018; Kiyatake et al. 2024). The primary anthropogenic threat to mobulids is fisheries exploitation, including targeted fishing and bycatch (Croll et al. 2016; Pacoureau et al. 2021; Palacios et al. 2025). Mobulid aggregations at certain habitats often overlap with commercially more valuable species targeted by various fishing gears and, as a result, increase the impact of bycatch and contribute to global population declines (Rohner et al. 2013; Croll et al. 2016; Lezama-Ochoa et al. 2019; Palacios et al. 2024). Consequently, all mobulid species are currently listed as either Critically Endangered, Endangered, or Vulnerable on the International Union for the Conservation of Nature’s Red List of Threatened Species (Marshall et al. 2022a, b, c, d, e, f, g; Rigby et al. 2022a, b).

Mobulid mating/copulation was initially documented in the Atlantic pygmy devil ray [*Mobula*
hypostoma (Bancroft, 1831)] (Coles 1910), while underwater courtship and mating behaviours were first described through video observations of the oceanic manta ray [Mobula birostris (Walbaum, 1792)], revealing a five-step sequence: (1) Following, where males trail a lead female; (2) Biting, with males grasping the female's pectoral fin for leverage; (3) Copulation; (4) Post-copulation, where the male holds onto the female's pectoral fin after retracting the clasper; and (5) Separation (Yano et al. 1999; Table 1). Observations of reproductive activity, including courtship and mating, were originally focused on the manta rays, especially *M. alfredi*, a more coastal species with greater human interactions that increase the likelihood of observing and documenting these rare behaviours (Clark 2010; Marshall and Bennett 2010; Stevens et al. 2018a). More detailed observations expanded the understanding of mobulid courtship, leading Stevens et al. (2018a, b) to describe a seven-stage courtship sequence (Table 1), supported by studies on various species: e.g., the Atlantic manta ray [Mobula yarae (Bucair, 2025)] (Pate 2024); the shorthorned pygmy devil ray

[Mobula kuhlii (Müller & Henle, 1841)] (Carpenter and Griffiths 2023); *M. mobular* (Duffy and Tindale 2018); Munk's pygmy devil ray [Mobula munkiana (Notarbartolo-di-Sciara, 1987)] (Palacios et al. 2024); the sicklefin devil ray [Mobula tarapacana (Philippi, 1892)] (Sobral 2013; Mendonça et al. 2020); and the bentfin devil ray [Mobula thurstoni (Lloyd, 1908)] (McCallister et al. 2020).

Although previous studies have reported on one to three species of mobulid courtship and mating (Yano et al. 1999; Marshall and Bennett 2010; Deakos et al. 2011; Duffy and Tindale 2018; Stevens et al. 2018a; McCallister et al. 2020; Mendonça et al. 2020; Carpenter and Griffiths 2023; Palacios et al. 2024; Setyawan et al. 2024), to date no comprehensive synthesis of existing data across the family has been conducted. Indeed, so little is known about mobulid reproduction that natural birth has never been observed in the wild for any species and has only been documented for *M. alfredi* (Anon 2007; Nozu et al. 2017; Murakumo et al. 2020) and *M. mobular* (Kiyatake et al. 2024) in aquaria. Here, we add substantial observations of mobulid courtship behaviour as well

<table><caption>**Table 1** Definitions of courtship and mating stages or steps in mobulid rays, including source references, species observed, and the behaviour descriptions as used in this study. Existing definitions have not been updated since Stevens et al. (2018a, b)</caption><thead><tr><th>Reference</th><th>Species analysed</th><th colspan="6">Courtship and mating steps or stages</th></tr><tr><th>Yano (et al. 1999)</th><th>Mobula birostris</th><th>Step 1. Chasing</th><th>Step 2. Biting</th><th>Step 3. Copulating</th><th>Step 4. Post-copulating</th><th>Step 5. Separating</th></tr></thead><tbody><tr><td>Marshall and Bennett (2010)</td><td>Mobula alfredi</td><td>(1) Close pursuits of females. (2) Faster than average swimming speeds. (3) Multiple male participants performing movements in a line one after another 'in cannon'</td><td>(4) Veering and swooping behaviour. (5) Somersaulting or looping behaviour</td><td>Positioning over the female's dorsal surface</td><td>Copulation</td><td>Separation</td></tr><tr><td>Stevens et al. 2018a, b</td><td>Mobula alfredi<br>Mobula birostris</td><td>Stage 1. Initiation</td><td>Stage 2. Endurance</td><td>Stage 3. Evaporation</td><td>Stage 4. Pre-copulation positioning</td><td>Stage 5. Copulation</td><td>Stage 6. Post-copulation holding</td><td>Stage 7. Separation</td></tr></tbody></table>
as review the existing body of literature on their mat-
ing behaviours. We discuss the underlying reasons for
the observed behaviours and identify knowledge gaps
and areas for future research priority. Finally, we reas-
sess the mobulid courtship definitions which have not
been updated since 2018 (Stevens et al. 2018a, b).

Materials and methods

Scientific literature, observations by authors and citi-
zen scientists, social media, and anecdotal reports of
mobulid courtship and mating behaviours were con-
solidated and examined. The behaviours were analysed
and described following the seven courtship stages
outlined by Stevens et al. (2018a). This includes the
‘Initiation’ stage, recognised when a male(s) aligns his
swimming to shadow the movements of a female, trail-
ing closely behind, while maintaining a typical cruis-
ing speed. The male may attempt to position his body
on top of the female’s, facing the same direction. The
Endurance’ stage was identified by above cruising
swim speeds of individuals, initiating a swift pursuit
as males trail closely behind the female in a single-file
formation (termed a courtship train), with their cephalic
fins frequently extended (in manta rays) and typically
mouths shut. In “Endurance”, the female is usually
at the lead of the formation in which the speed of the
female is often matched by the males following. The
“Evasion” stage involves the female engaged in unpre-
dictable, high-speed swimming characterized by abrupt
twists and turns, strategically navigating around obsta-
cles. These manoeuvres could include backward som-
ersaults, forward flips, and breaching (detailed more
in the results), with the males mimicking the female’s
movements (Marshall and Bennett 2010; Pate 2024).
The “Pre-copulation positioning” stage was recog-
nised when the female reduced her swimming to below
cruising speed, frequently ascending near the water’s
surface. The “Copulation” stage was categorised by
the male maintaining an active rhythm of pectoral fin
movements while grasping hold of one of the female’s
left pectoral fins (primarily the left) in his mouth, while
the female typically remains still and the male executes
rapid pelvic thrusts, inserting the clasper into the clo-
aca, with both individuals gradually descending in the
water column, often in a spiralling motion. The “Post-
copulation holding” stage was identified when the male
withdrew his clasper from the female’s cloacal opening,

in which the male may briefly hold onto her pectoral.
However, this phase is usually omitted entirely, as the
subsequent stage, “Separation”, can occur immedi-
ately upon the conclusion of copulation. “Separation”
was recognised when the pair disengaged and swam in
opposite directions.

Each resultant video or imagery viewed was then
assessed and, if relevant, was categorised into one or
more of the stages. Because courtship processions
can endure for extended periods, occasionally transi-
tioning to subsequent phases of courtship or return-
ing to earlier ones, each video or set of images was
watched entirely and carefully described.

To address knowledge gaps for species such as *M. eregoodoo*, focused searches on YouTube, Instagram, and Facebook were conducted. The keywords “mobula”, “Raja Ampat”, “Indonesia”, “devil ray”, “pygmy manta”, and “diving” were used to search for videos of undocumented courtship behaviour of *M. eregoodoo*, which included 200 videos with only 16 videos correctly identifying *M. eregoodoo*. We conducted 35 searches on YouTube for other mobulid species using different combinations of the keywords “mobula”, “manta”, “devil ray”, “mating”, “diving”, “ray”, and “courtship” to search for additional videos of undocumented courtship and mating events. Each resultant video viewed (300) was then assessed and, if relevant, was also categorised.

Results

In this study, we add 87 records from eight of the ten
described mobulid species across 18 countries/terri-
tories contributed by 67 observers, including seven
observations for *M. alfredi*, 21 for *M. birostris*, 12 for
*M. yarae*, four for *M. hypostoma*, ten for *M. kuhlii*,
28 for *M. mobular*, one for *M. munkiana*, and four
for *M. tarapacana* (Supplementary Table I). Overall,
the scientific literature is weak on mobulid courtship,
particularly observations for *M. eregoodoo*, *M. hypos-
toma*, *M. tarapacana*, and *M. thurstoni*. The videos
and photographs provided for *M. hypostoma* represent
the first visual documentation of reproductive behav-
iour for this species. These 87 records (Supplemen-
tary Table I) are consolidated below with existing lit-
erature and grouped into the seven different stages of
mating behaviour (Table 2; Supplementary Table II).
## Initiation

A total of 20 Initiation (Supplementary video 1, MVI_9789, *M. birostris*, event no. 10, Supplementary Table I) events were added: *M. birostris* (n=3), *M. yarae* (n=1), *M. kuhlii* (n=4), *M. hypostoma* (n=1), *M. mobular* (n=7), and *M. tarapacana* (n=1) (Supplementary Table I; Fig. 1a). Initiation was similar across species, including the behaviours undertaken and where it occurred, such as near the surface of the water column (n=16) or at aggregation sites, such as cleaning stations (n=3) and seamounts (n=1) (Fig. 1b–d, f). “Initiation” was observed in all mobulid species except *M. eregoodoo* (Table 2). In devil rays, this stage often arises when two or more individuals separate from a larger schooling group (events no. 45, 69, 80). These larger aggregations (>100 individuals in *M. munkiana* and *M. kuhlii*, and >50 individuals in *M. tarapacana*) are often seasonal, contain many near-term pregnant females (Fig. 1c, f; Supplementary video 2, “Manta Mobula Encounter”, *Mobula kuhlii*, event 45, Supplementary Table I) and may form spiralling vortex formations (Fig. 1e; Palacios et al. 2023).

## Endurance

Endurance (Fig. 2a; Supplementary video 3, “Matting Trains”, *M. alfredi*, event no. 5, Supplementary Table I) was documented across all mobulid species except *M. eregoodoo* (Table 2). Here, we add 71 observations of Endurance: *M. alfredi* (n=4), *M. birostris* (n=15), *M. yarae* (n=10), *M. hypostoma* (n=4), *M. kuhlii* (n=11), *M. mobular* (n=24), and *M. tarapacana* (n=3) (Supplementary Table I). If observed, Endurance constituted the longest stage in duration within each event for all species (e.g., event nos. 46, 60, 76, and 79). All species exhibited courtship trains, sometimes forming circling rings as the lead female joins the trailing male (Supplementary video 3). Courtship trains typically consisted of one female and between one and three males (Supplementary Table I; event nos. 14, 21, 22, 23, 25, 26, 32, 33, 34, 39, 40, 41, 42, 43, 45, 46, 51, 52, 53, 54, 55, 61, 70, 76, and 79) sometimes up to 6–8 males (Supplementary Table I; event nos. 3, 5, 49, 56, 60, 63, 65, 69, 75, and 76); however, more than two dozen males have been recorded (Clark 2010). Females usually led the courtship trains, but sometimes they followed

one or more males, as seen in *M. mobular* (Fig. 2i; Supplementary video 15; event no. 82). The female was often observed to keep her cephalic fins furled (Fig. 2c–i). Endurance tended to occur on coral reefs in *M. alfredi* (Supplementary Video 3) and *M. kuhlii* (Supplementary Video 2; Supplementary Video 4, event no. 49), in coastal neritic waters for *M. yarae* (Supplementary Video 5, event no. 36, Supplementary Table I), *M. hypostoma* (Supplementary Video 6, 1258rp, event no. 37), and *M. munkiana* (Supplementary Video 7, event no. 73, Supplementary Table I), and in oceanic deep water at or near the surface for *M. birostris* (Supplementary Video 8, MVI_2830, event no. 7), *M. mobular* (Supplementary Video 9, A065C798, event no. 70, Supplementary Table I), and *M. tarapacana* (Supplementary Video 10, event no. 77, GH011015, Supplementary Table I) (Fig. 2).

## Evasion

Evasion (Fig. 3; Supplementary video 11, *M. yarae*, event no. 31, Supplementary Table I) has been documented across all mobulid species except *M. eregoodoo* (Table 2). Here, we add 57 observations of Evasion: *M. alfredi* (n=5), *M. birostris* (n=12), *M. yarae* (n=7), *M. hypostoma* (n=4), *M. kuhlii* (n=6), *M. mobular* (n=16), and *M. tarapacana* (n=2) (Supplementary Table I). While these stages are exhibited across the mobulid species, there are notably more somersault and flipping behaviours during the Evasion stage of manta ray species than by the devil rays (Supplementary video 3; Supplementary video 12, *M. birostris*, event no. 18, MV-MB-0716, Supplementary Table I) (Fig. 3a–c). In *M. yarae*, the motion of males somersaulting onto their backs seems to initiate the transition from Endurance to Evasion (Fig. 3c; Supplementary video 11). The swimming speeds of *M. kuhlii* and *M. hypostoma* during Evasion were particularly fast, especially when only one male and one female were engaged (Supplementary video 13, *M. kuhlii*, event no. 46; Supplementary video 14, *M. hypostoma*, event no. 37, Supplementary Table I) (Fig. 3d, e). In *M. mobular* (Supplementary videos 15 and 16, events no. 54, 62, Supplementary Table I), *M. hypostoma* (Supplementary video 14, Supplementary Table I), and *M. munkiana*, the evasive females consistently remained close to the surface (Fig. 3d, f–i), sometimes holding their pectoral fins above the water or slapping the male to prevent the male(s)
<table><caption>**Table 2** The number of observations of courtship and mating events for *Mobula* species published in the scientific literature, plus those added in this study for each species.</caption><thead><tr><th>Species</th><th>Published observations (N)</th><th>Added observations (N)</th><th>References</th><th>Near-term pregnant females</th><th>Left pectoral fin preference</th><th>Pseudo-courtship</th><th>Preferred habitat</th><th>Mass aggregations (&gt; 50 ind.)</th></tr></thead><tbody><tr><td><i>M. alfredi</i></td><td>295</td><td>7</td><td>Marshall &amp; Bennett 2010 (n=5), Stevens et al. 2018a (n=235), Germanov et al. 2019 (n=55)</td><td>✓</td><td>✓</td><td>✓</td><td>Reef, coastal neritic</td><td>✗</td></tr><tr><td><i>M. birostris</i></td><td>8</td><td>21</td><td>Yano et al. 1999 (n=4), Stevens et al. 2018a (n=4)</td><td>✓</td><td>✓</td><td>✓</td><td>Reef, oceanic deep-water</td><td>✗</td></tr><tr><td><i>M. eregoodoo</i></td><td>0</td><td>0</td><td>N/A</td><td>N/A</td><td>N/A</td><td>N/A</td><td>N/A</td><td>N/A</td></tr><tr><td><i>M. hypostoma</i></td><td>1</td><td>4</td><td>Coles 1910 (n=1)</td><td>✗</td><td>✓</td><td>✓</td><td>Coastal neritic</td><td>✗</td></tr><tr><td><i>M. kuhlii</i></td><td>3</td><td>10</td><td>Carpenter &amp; Griffiths 2023 (n=3)</td><td>✓</td><td>✓</td><td>✗</td><td>Reef, coastal neritic</td><td>✓</td></tr><tr><td><i>M. mobular</i></td><td>13</td><td>28</td><td>Duffy &amp; Tindale 2018 (n=1), Palacios et al. 2024 (n=10), Kiya-take et al. 2024 (n=1), Diamant et al. 2025 (n=1)</td><td>✓</td><td>✓</td><td>✗</td><td>Oceanic deep-water</td><td>✗</td></tr><tr><td><i>M. munkiana</i></td><td>208</td><td>1</td><td>Palacios et al. 2024 (n=208)</td><td>✓</td><td>✓</td><td>✗</td><td>Coastal neritic</td><td>✓</td></tr><tr><td><i>M. tarapacana</i></td><td>15</td><td>4</td><td>Sobral 2013 (n=1), Mendonça et al. 2020 (n=14)</td><td>✓</td><td>✗</td><td>✗</td><td>Oceanic deep-water</td><td>✗</td></tr></tbody></table>

<figure><img src="image_2.png" /></figure>
<table><caption>**Table 2 (continued)**</caption><thead><tr><th>Species</th><th>Published observations (N)</th><th>Added observations (N)</th><th>References</th><th>Near-term pregnant females</th><th>Left pectoral fin preference</th><th>Pseudo-courtship</th><th>Preferred habitat</th><th>Mass aggregations (&gt; 50 ind.)</th></tr></thead><tbody><tr><td><i>M. thurstoni</i></td><td>4</td><td>0</td><td>McCallister et al. 2020 (n=1), Palacios et al. 2024 (n=3)</td><td>x</td><td>✓</td><td>x</td><td>Coastal neritic</td><td>x</td></tr><tr><td><i>M. yarae</i></td><td>18</td><td>12</td><td>Pate 2024 (n=18)</td><td>x</td><td>✓</td><td>✓</td><td>Coastal neritic</td><td>x</td></tr></tbody></table>

from grasping hold of these fins (Fig. 3d). Near-term pregnant females were observed engaged in Endurance and Evasion stages in four species in this study, totalling 14 observations (*M. birostris*, *M. kuhlii*, *M. mobular*, and *M. tarapacana*) (event nos. 17, 20, 21, 23, 43, 42, 48, 49, 53, 59, 60, 67, 74, 77, and 82; Supplementary Table I).

### Pseudo-courtship

Male to male courtship behaviour in mobulids has been described for the first time in this study, observed across four species (Supplementary Material I). This involves between two and four males exhibiting Initiation, Endurance, and Evasion behaviours (Fig. 4). Male to male courtship has been observed in *M. alfredi* at Nusa Penida, Indonesia (Fig. 4a, Event 1) and twice at Lady Elliot Island, Australia (Fig. 4b–d, Supplementary videos 17–18, event nos. 4 and 6), in *M. birostris* at the Revillagigedo Archipelago, Mexico (Fig. 4e, event no. 13), twice in *M. yarae* at Quintana Roo, Mexico (Fig. 4f, Supplementary Video 19, event no. 29, Supplementary Table I) and West Palm Beach, Florida, USA (juvenile males—Supplementary video 20, event no. 35, Supplementary Table I), and in captive *M. hypostoma* at the Frost Museum of Science, Florida, USA (event no. 40, Supplementary Table I).

### Pre-copulation positioning

A total of 28 observations of pre-copulation positioning (Fig. 5a–d, I; Supplementary video 7, Supplementary video 21, *M. hypostoma*, event no. 40, Supplementary Table I) were made for *M. alfredi* (n=2), *M. birostris* (n=2), *M. yarae* (n=3), *M. hypostoma* (n=2), *M. mobular* (n=18), and *M. tarapacana* (n=1). There was consistency of this stage across species. However, in *M. hypostoma* (Supplementary video 22, event no. 41), *M. munkiana* (Supplementary video 7), and *M. yarae* (Supplementary video 23, event no. 33, Supplementary Table I), the males undertook accelerated ‘leaps/lunges’ to position themselves onto the dorsal surface of the females, sometimes jumping out of the water to do so (Fig. 5d, e, h). In *M. mobular*, instead of the males positioning themselves dorsally on the female, the males used their heads and bodies to elevate the slowly moving female further towards the surface, sometimes while
<figure><img src="image_3.png" /><figcaption>**Fig. 1** Initiation stage and associated aggregation behaviours between: **a** *Mobula alfredi* at a cleaning station in the Maldives (Stevens et al. 2018a); **b** *M. birostris* at a cleaning station at Isla de La Plata, Ecuador, Event 11; **c** a large schooling aggregation of *M. kuhlii* containing near-term pregnant females in Ari Atoll, Maldives, Event 46; **d** an oceanic atoll aggregation of *M. mobular*, North Malé Atoll, Maldives, Event 80; **e** a</figcaption></figure>

cyclone formation aggregation of *M. munkiana* in Baja California, Mexico (Palacios et al. 2024); and **f** a seamount aggregation of *M. tarapacana* in the Azores, Portugal, containing near-term pregnant females (Stevens et al. 2024). © **a** Guy Stevens, **b** Michel Guerrero, **c** Dominik Ruhl, **d** Carlo Gargioni, **e** Sidharta Velázquez-Hernández, and **f** João Vieira Rodrigues

<figure><img src="image_4.png" /><figcaption>**Fig. 2** Endurance stage behaviour between: **a** *Mobula alfredi* in the Maldives (Stevens et al. 2018a); **b–c** *M. birostris*, Galápagos Islands, Ecuador and Revillagigedo Archipelago, Mexico, Events 7 and 14, respectively; **d–e** *M. hypostoma*, Quintana Roo, Mexico, and Florida, USA, Events 43 and 44,</figcaption></figure>

respectively; **f** *M. kuhlii*, Lhaviyani Atoll, Maldives, Event 51; and **g–i** *M. mobular* at Malapascua Island, Philippines and Toulon Canyon, France, Events 55 and 56, respectively. © **a**, **c**, **d** Guy Stevens, **b** Fabricio Morocho, **e** Jacob Rosner, **f** Lisa Bauer, **g** Joost van Uffelen, and **h**, **i** Frédéric Bassemayousse

upside-down, whereupon the male(s) would attempt to grasp the female's left pectoral fin and orientate himself in a position to enable copulation (Fig. 5f–g;

Supplementary video 24, event no. 58, Supplementary Table I). While there are variations in pre-copulation positioning strategies among species, and within
<figure><img src="image_5.png" /><figcaption>**Fig. 3** Evasion stage behaviour between: **a** *Mobula alfredi* in the Maldives (Stevens et al. 2018a); **b** *M. birostris* in the Maldives, Event 19; **c** *M. yarae* in Quintana Roo, Mexico, Event 35; **d–e** *M. hypostoma*, Georgia, USA and Quintana Roo, Mexico, Events 41 and 42, respectively; **f–h** *M. mobular* at</figcaption></figure>

Palawan, Philippines, Similan Islands, Thailand, and Mahé, Seychelles, Events 58, 60, and 63, respectively; and i *M. tara-pacana* in the Azores, Portugal, Event 82. © a, e Guy Stevens. b Najihu Mohamed, c José Ancona, d Robin Perrtree, f Duncan Murrell, g Richard Carey, h Darren Henn, and iAna Sobral

<figure><img src="image_6.png" /><figcaption>**Fig. 4** Pseudo-courtship (Evasion and Endurance stages) between **a** two male *Mobula alfredi* at Nusa Penida, Indonesia, Event 1; **b–c** two and **d** four male *M. alfredi* at Lady Elliot Island, Australia, Events 4 and 6, respectively; **e** two male *M.*</figcaption></figure>

*birostris* in the Revillagigedo Islands, Mexico, Event 14; and **f** two male *M. yarae*, Quintana Roo, Mexico, Event 33. © **a** Ana Sobral, **b–c** Olivia Cornale, **d** Conor Mulvenna, **e** Guy Stevens, and **f** Karen Fuentes

individual courtship events, the ultimate objective of the male(s) remains consistent across all species: to enable the alignment of his body with that of the female so that he can grasp hold of her pectoral fin (primarily the left) and then insert one of his claspers into the female’s cloacal opening and copulate.

## Copulation

Nine copulation events (Fig. 6) were observed in *M. alfredi* (n=3), *M. birostris* (n=3), *M. yarae* (n=1), *M. hypostoma* (n=1), and *M. munkiana* (n=1) (Table 2). All copulation observations were under 2
<figure><img src="image_7.png" /><figcaption>**Fig. 5** Pre-copulation positioning stage behaviour between *a–b Mobula alfredi* in the Maldives (Stevens et al. 2018a); *c M. birostris* at Isla de La Plata, Ecuador, Event 13; *d M. yarae* in Quintana Roo, Mexico, Event 37; *e M. hypostoma*, Georgia, USA, Event 42; *f–g M. mobular*, Corsica and Sanary-sur-Mer, France, Events 66 and 67, respectively; *h M. munkiana*, Baja California, Mexico (Palacios et al. 2024); and *i M. tarapacana* in the Azores, Portugal, Event 85. © a–b Guy Stevens, c Michel Guerrero, d Jahzeel Elizarraraz, d Robin Perrtree, f Kevin Sempé, g Remi Demarthon, h Marta Palacios, and iFlávia Corrêa</figcaption></figure>

min (i.e., 43 sec *M. munkiana* event no. 73; 50 sec *M. alfredi* event no. 2, Supplementary Table I), and the male had a consistent preference for the left pectoral fin during actual copulation (Supplementary video 7, Supplementary video 25, *M. alfredi* event no. 2, Supplementary Table I). In all mobulid species, except *M. tarapacana* (Fig. 7g), the males prefer to select the left pectoral fin. In all instances,

<figure><img src="image_8.png" /><figcaption>**Fig. 6** Copulation and Separation stages: **a–c** *Mobula alfredi* copulating in the Maldives (Stevens et al. 2018a); **d–e M. alfredi** copulation and separation at Lady Elliot Island, Australia, Event 2; **f–g** *M. birostris* copulating at the Galápagos Islands, Ecuador, Events 8 and 9, respectively; **h M. birostris** copulating in Suva, Fiji, Event 28; **i M. munkiana** copulating in Baja California, Mexico, Event 83. © **a–c** Guy Stevens, **d–e** Remy Suffredini, **f–g** Fabricio Morocho, **h** Luke Gordon, and **i**Koe Mucha Vida</figcaption></figure>
the male rotated his body beneath the female until aligned abdomen to abdomen for clasper insertion (Fig. 6a–d, f–i) (Supplementary Table I).

## Post-copulation holding

Post-copulation holding is often omitted entirely (Fig. 6). Of the nine copulation events documented in our results, only two exhibited any post-copulation holding [M. yarae (n=1) event 40 and M. hypostoma (n=1) (event 44, Supplementary Table I), and in each of these instances, the holding lasted less than two seconds (Table 2).

## Separation

Separation was observed six times: in M. alfredi (n=2), M. birostris (n=1), M. yarae (n=1), M. hypostoma (n=1), and M. munkiana (n=1) (Supplementary Table I). In Separation, the female usually initiated a rapid departure from the male, possibly to escape secondary copulation or the attentions of other pursuing males that may attempt to start a new courtship and mating encounter with the female (Fig. 6e, event nos. 2, 39, 43, and 76; Supplementary videos 5, 21, 24, 25, Supplementary Table I). This urgency by the female to “escape” could further be triggered by the trauma of the courtship and copulation, which

<figure><img src="image_9.png" /><figcaption>**Fig. 7** Courtship and mating stage behaviour frequency across 10 mobulid ray species. Stages include Initiation, Endurance, Evasion, Pre-copulation positioning, Copulation, Post-copulation Holding, and Separation. The corresponding data and literature can be found in Supplementary Table II</figcaption></figure>
sometimes leaves the females with significant wounds (see following section). As obligate ram ventilators, when multiple males are attempting to mate with a female, the physical restraint and harassment by the males may also prevent the females from respiring sufficiently. These factors, when combined, may occasionally result in mortality and could be the case for the female *M. munkiana* observed copulating in Event 82 in this study (Supplementary video 7, Supplementary Table I).

## Overall comparison between mobulid species

As the mating sequence progresses past Endurance, observations become increasingly rare (Fig. 7, Supplementary Table II) with each successive stage observed with decreasing frequency, although Initiation may be underreported. The frequency of observations varies greatly depending on both species behaviour and research effort. *Mobula alfredi* and *M. munkiana*, with higher levels of diver interaction and extended in-water observation time, yielded large numbers of documented Endurance and Evasion (Fig. 7). In contrast, rarer, more elusive, data-deficient, and critically endangered species contributed far fewer records due to their limited accessibility and cryptic nature (Fig. 7, Supplementary Table II).

Male following was consistently observed in all studied mobulids (*M. alfredi*, *M. birostris*, *M. yarae*, *M. hypostoma*, *M. kuhlii*, and *M. mobular*), aligning with previous literature (Table 2). *Mobula munkiana* demonstrated spiral vortex formation during initiation in literature, suggesting potential species-specific courtship displays. Both literature and this study confirm that *M. alfredi*, *M. birostris*, *M. yarae*, *M. hypostoma*, *M. kuhlii*, *M. mobular*, *M. munkiana*, and *M. tarapacana* consistently engage in mating trains. While largely similar across species, *M. mobular* exhibited a slower variant of the mating train, a behaviour observed only in this study, which may indicate species-specific adjustments in endurance strategies.

In Evasion, common behaviours across species include flips, breaches, pectoral displays, circular swimming, and high-speed maneuvers. *Mobula alfredi*, *M. birostris*, and *M. yarae* displayed flips and breaches, consistent with literature, whereas this study documented new evasive behaviours, such as

circle swimming in *M. hypostoma* and *M. mobular*, and high-speed bursts in *M. hypostoma* and *M. kuhlii*. *Mobula hypostoma* exhibited breach, circle, pectoral display, and high-speed swimming. In contrast, *M. thurstoni* and *M. tarapacana* show evasion patterns largely consistent with prior literature, breaches and pectoral displays, although observations were limited. Male-to-male endurance and evasion were observed across *M. alfredi*, *M. birostris*, *M. yarae*, and *M. hypostoma*, with mature male-male courtship in *M. alfredi* and *M. birostris*, and juvenile male courtship and male-to-male initiation were novel observations in *M. yarae*.

In Pre-copulation positioning, left pectoral fin positioning is ubiquitous across nearly all species, observed in literature and confirmed in this study for *M. alfredi*, *M. birostris*, *M. yarae*, *M. hypostoma*, *M. mobular*, *M. munkiana*, *M. tarapacana*, and *M. thurstoni*. Novel behaviours included leap positioning in *M. yarae*, *M. hypostoma*, and *M. munkiana*, observed for the first time in this study. *Mobula mobular* displayed a combination of female stall and male push behaviours, reflecting a more interactive pre-copulatory positioning than in other species.

Copulation, consistent with the literature and this study, was a brief mating event (<120 s) and was observed consistently for *M. alfredi*, *M. birostris*, *M. yarae*, *M. hypostoma*, and *M. munkiana*. Post-copulation holding, observed primarily as quick holding, was noted across *M. alfredi*, *M. birostris*, *M. yarae*, *M. hypostoma*, and *M. mobular*. Both the literature and this study confirm its presence, although it is often skipped. Separation was immediate, following copulation in nearly all species. This was consistently observed across the literature and in this study for *M. alfredi*, *M. birostris*, *M. yarae*, *M. hypostoma*, *M. munkiana*, and *M. mobular*. A novel observation in *M. munkiana* was female mortality following copulation, a behaviour not previously documented in any mobulid species.

Mating habitats varied across species: reef-associated *M. alfredi* and *M. kuhlii*, schooling aggregations for *M. kuhlii* and *M. munkiana*, surface or deep-water oceanic locations for *M. mobular*, *M. birostris* and *M. tarapacana*, and coastal neritic zones for *M. yarae* and *M. hypostoma*. These habitat preferences were largely consistent with the literature but were confirmed and expanded by this study.
Comparing reproductive wounds and scarring in mobulids to other elasmobranchs

Courtship and mating wounds and scars provide evidence that reproductive activity has transpired. Our observations of mobulid behaviours revealed a diverse range of courtship injuries, copulation wounds, and mating scars, depending on species. Being planktivores, mobulids possess teeth solely for mating (Motta and Wilga 2001). There is a variety in the appearance of the wounds and scars due to differences in toothband and mouth morphology (Notarbartolo di Sciara 1987; Marshall et al. 2009; White et al. 2017; Notarbartolo di Sciara et al. 2019). However, the overall parts of the body where they occur are similar.

In all the mobulid species for which data exist, the ventral and dorsal distal ends of the pectoral fins of females are often injured by the male's coarse denticles and mouth structure as he bites onto the fin during copulation (Fig. 7, Table 3) (Yano et al. 1999; Marshall and Bennett 2010; Duffy and Tindale 2018; Mendonça et al. 2020; Palacios et al. 2024). In the manta ray species, the males only have toothbands in the lower jaw (Marshall et al. 2009). In *M. alfredi*, cartilaginous ridges in the upper buccal cavity result in distinct black and/or white dorsal mating scars for this species (Stevens et al. 2018b) (Fig. 7k, l). Male devil rays have toothbands in both the upper and lower jaws (Notarbartolo 1989; Stevens et al. 2018b) (Fig. 7a). These rows of tiny teeth leave abrasions on the female's fin at an angle offset

by the degree to which the male's body had been positioned to enable copulation. The angle varies by species, but usually forms an abrasion line at an angle of 50–110 degrees from the leading edge of the female's pectoral fin (Fig. 7b–j). These abrasions are often visible as red wounds for several weeks after copulation (Fig. 7j). Similarly, in other elasmobranchs, differences in dentition were found between male and female in red stingrays [*Hemitrygon akajei* (Bürger, 1841)], with adult males having sharper teeth than females (Taniuchi and Shimizu 1993). Kajiura et al. (2000) also observed similar patterns in Atlantic stingrays [*Hypanus sabini* (Lesueur, 1824)] and noted a periodic shift in male dental morphology correlated with the mating season. Although male biting is the primary source of wounds and mating scars, mature females in some elasmobranch species also engage in biting behaviour during mating (Gilmore et al. 1983). These observations suggest the existence of sexual hierarchies that are maintained through aggressive interactions (Gordon 1993). These hierarchies may play a role in synchronizing the mating sequences of certain elasmobranchs.

In mobulids, the courtship and mating process can result in contact with the seafloor, especially when multiple males are engaged by the female. The female (and sometimes the males) often has cuts and scratches on her body, especially the dorsal surface, which likely resulted from collisions with the reef during copulation (Fig. 8b, c). Mobulid rays, like all elasmobranchs, have rough sandpaper-like skin. Therefore, the physical

<table><caption>**Table 3** Types of courtship and mating injuries documented for each *Mobula* species published in the scientific literature, plus those observed in this study: *X* = observed behaviour; = no available information</caption><thead><tr><th rowspan="2">Species</th><th colspan="4">Type of injury</th><th rowspan="2">Reference/corresponding figure</th></tr><tr><th>Pectoral fin scars/ wounds</th><th>Body abrasions/ wounds</th><th>Clasper abrasions/swelling</th><th>Cephalic fin/ mouth abrasions</th></tr></thead><tbody><tr><td><i>M. alfredi</i></td><td>X</td><td>X</td><td>X</td><td>X</td><td>Marshall & Bennett 2010; Stevens et al. 2018a; Germanov et al. 201; Figs. 7 and 8</td></tr><tr><td><i>M. birostris</i></td><td>X</td><td>-</td><td>-</td><td>-</td><td>Yano et al. 1999; Stevens et al. 2018a; Fig. 7</td></tr><tr><td><i>M. eregoodoo</i></td><td>-</td><td>-</td><td>-</td><td>-</td><td>N/A</td></tr><tr><td><i>M. hypostoma</i></td><td>X</td><td>-</td><td>-</td><td>X</td><td>Coles 1910; Figs. 7 and 8</td></tr><tr><td><i>M. kuhlii</i></td><td>X</td><td>-</td><td>-</td><td>-</td><td>Carpenter & Griffiths 2023; Fig. 7</td></tr><tr><td><i>M. mobular</i></td><td>X</td><td>-</td><td>-</td><td>-</td><td>Duffy & Tindale 2018; Palacios et al. 2024; Fig. 7</td></tr><tr><td><i>M. munkiana</i></td><td>X</td><td>X</td><td>-</td><td>-</td><td>Palacios et al. 2024; Fig. 8</td></tr><tr><td><i>M. tarapacana</i></td><td>X</td><td>-</td><td>X</td><td>-</td><td>Sobral 2013; Mendonça et al. 2020; Fig. 7</td></tr><tr><td><i>M. thurstoni</i></td><td>X</td><td>-</td><td>-</td><td>-</td><td>McCallister et al. 2020; Palacios et al. 2024</td></tr><tr><td><i>M. yarae</i></td><td>X</td><td>-</td><td>-</td><td>-</td><td>Figure 8</td></tr></tbody></table>
<figure><img src="image_10.png" /><figcaption>**Fig. 8** Mating scars and wounds: **a** *Mobula birostris* toothband, **b–d** dorsal, and **e–i** ventral mating scars or **j** fresh wounds on **b**, **d**, **h** *M. mobular*, **c** *M. kuhlii*, **e** *M. birostris*, **f** *M. yarae*, **g** *M. tarapacana*, **i** *M. hypostoma*, and **j** *M. alfredi* caused by the male's teeth during copulation; **k** dorsal mating scars on *M. alfredi* caused by **l** cartilaginous ridges in the male's mouth during copulation. © **a** Doug Perrine, **b** Darren Henn, **c** Andrea Marshall, **d** Pierre Leo Paul, **g** Danny Cope-land, **h** Duncan Murrell, and **e, f, i, j, k, l** Guy Stevens</figcaption></figure>

contact, primarily on the dorsal surface, by multiple males during lengthy courtship behaviours also results in the removal of skin pigmentation and often causes abrasions to the female's body (Fig. 8d).

During courtship and copulation, the inside of the male's cephalic fins may also be wounded where they have rubbed against the female's pectoral fin or dorsal surface (Fig. 8a). Male mobulids may also exhibit markings, such as clasper abrasions or swelling, due to mating friction, as observed in *M. tarapacana* and *M. alfredi* (Mendonça et al. 2020; Marshall and Bennett 2010).

Fresh reproductive wounds are distinguished by being red in colour, while older mating scars are off-white to grey in colour (Kajiura et al. 2000) (Fig. 9). These wound and scarring patterns can be species-specific, influenced by male biting, grasping, and mating behaviours, as well as the presence and utilization of dentition, horns, spines, and denticles (Pratt and Carrier 2005). Within a month, the wounds heal completely, either leaving little or no visible trace, or permanent white scar lines or patches (Stevens et al. 2018a; Kiya-take et al. 2024). Therefore, the distinct fresh reproductive wounds and mating scars can be used to aid identification of reproductive activity and assess the maturity status of individuals in the field (Table 3).

Comparing courtship behaviour and mating in mobulids to other elasmobranchs

The structure of mobulid courtship appears to share common elements with all batoids and even chondrichthyans as a whole, including courtship trains, biting during copulation, and resultant scarring. Our

<figure><img src="image_11.png" /><figcaption>**Fig. 9** Reproductive injuries: **a** abrasions to the inside of the cephalic fins of a male *Mobula hypostoma* resulting from copulation, **b–c** dorsal abrasions and scratches in *M. alfredi* resulting from contact with the reef during courtship and copulation, and **d** dorsal abrasions and the removal of skin pigmentation on a female *M. munkiana* caused by courting and copulating males. © **a** Jacob Rosner, **b** Julie Hartup, **c** Beth Taylor, and **d** Koe Mucha Vida</figcaption></figure>
results aligned with the general sequence described by Stevens et al. (2018a, b) with some updates for each species and new descriptions of species-specific behaviours, within the stages described. Building upon initial observations of mobulid species, this study can serve as a comprehensive resource for understanding courtship behaviour within the Mobulidae family.

Elasmobranchs, a taxonomically diverse group, share reproductive systems such as claspers, siphon sacs, ovaries with large follicles, and oviducts with specialized tissues, as well as fundamental courtship reproductive behaviours (Pratt and Carrier 2005). Chapman et al. (2003) notably provided the initial comprehensive account of mating in the free-living southern stingray [*Hypanus americanus* (Hildebrand & Schroeder, 1928)], which consists of five sequential stages: (1) close-following, (2) pre-copulatory biting, (3) insertion/copulation, (4) resting, and (5) separation. This consistent mating sequence is observed across various elasmobranchs, encompassing batoids such as the marbled ray [*Taeniura meyeni*; (Müller & Henle, 1841)] (Arnés-Urgellés et al. 2018), the whitespotted eagle ray [*Aetobatus narinari*; (Euphrasen, 1790)] (Tricas 1980), the yellow stingray [*Urobatis jamaicensis* (Cuvier, 1816)] (Young 1993), the round stingray [*Urobatis halleri* (Cooper, 1863)] (Nordell 1994), and sharks, including the scalloped hammerhead shark [*Sphyrna lewini*; (Griffith & Smith, 1834)] (Klimley 1980), oceanic whitetip shark [*Carcharhinus longimanus* (Poey, 1861)] (Talwar et al. 2023), basking shark [*Cetorhinus maximus* (Gunnerus, 1765)] (Sims et al. 2000), whitetip reef shark [*Triaenodon obesus* (Rüppell, 1837)] (Whitney et al. 2004), blacktip reef shark [*Carcharhinus melanopterus* (Quoy & Gaimard, 1824)] (McCauley et al. 2010), lemon shark [*Negaprion brevirostris* (Poey, 1868)] (Clapis-Garla et al. 2022), and silky shark [*Carcharhinus falciformis* (Bibron, 1839)] (Clarke et al. 2013).

## Behavioural sequence

The beginning of courtship, referred to as “Initiation” in mobulids (Stevens et al. 2018a), which leads to the “following” behaviour in male elasmobranchs, is widely suggested to be olfaction-mediated sexual attraction (Klimley 1980; McCauley et al. 2010; Stevens 2016). In *M. alfredi*, this courtship stage has

been suggested to be influenced by male social groups (Stevens et al. 2018a). Breaching behaviours (by both males and females) are associated with reproductive activity and likely serve as a means of communication to attract potential mates, such as a precursor for mating or a prerequisite to the Initiation stage (Marshall and Bennett 2010; Stevens et al. 2018b; Klimley et al. 2024; Stevens et al. 2024). The number of breaches in a single observation can be up to 286 for *M. munkiana* (Klimley et al. 2024), or in lesser-studied species such as *M. yarae*, up to 26 times (Pate 2024). Other behaviours such as vortex swimming may play an important role in facilitating Initiation in the case of *M. munkiana* (Palacios et al. 2024). Furthermore, in interspecies courtship between *M. alfredi* and *M. birostris*, “Initiation” was facilitated by a second male *M. birostris* who blocked the female *M. alfredi* from swimming away from the other *M. birostris* male (Setyawan et al. 2024).

In mobulids, “Initiation” progresses to “Endurance” and “Evasion” stages, also known as courtship trains, while in other elasmobranchs, behaviours like “nose-to-tail following”, “flank approach”, and “parallel swimming” are observed in *C. maximus* (Sims et al. 2000); “arch”, “avoid”, “following”, “group behaviour”, “pectoral grasp”, “positioning and alignment” in *T. obesus* (Whitney et al. 2004) and “echelon or parallel swimming”, “following”, “shielding”, “circle-back”, “nosing”, and “rotating” behaviours in *C. longimanus* (Talwar et al. 2023). Breaching may also occur during these stages, whereby males may be asserting dominance over one another (Klimley et al. 2024). The extended duration of the Endurance and Evasion stages is likely attributed to mate selection and competition, as females aim to have as many males following them as possible to diversify their gene pool (Stevens 2016).

Mobulid courtship trains are also likely facilitated by olfaction, as in other elasmobranchs (Johnson and Nelson 1978; Pratt and Carrier 2005). An increased number of males likely results in increased courtship train duration due to competition between male individuals (Marshall and Bennett 2010). The number of males following the female ranges from a minimum of one (e.g., Stevens et al. 2018a; Pate 2024; Diamant et al. 2025; current study) up to 40 in *M. alfredi* (Clark 2010). The average number of males engaged in Endurance/Evasion in this study was three, which supports past studies on manta rays (Stevens et al.
2018a, b). Endurance and Evasion have been most widely observed in *M. alfredi* (Marshall and Bennett 2010; Stevens et al. 2018a; Germanov et al. 2019) and have been recorded to last for up to at least 48 h (Clark 2010); in *M. yarae* at least 73 min (Pate 2024); and in *M. mobular*, at least 147 min (Duffy and Tindale 2018). Following behaviours differ between ben-thic and pelagic species, with skates and some rays mating on the sea floor, while pelagic species like *A. narinari* (Tricas 1980), *C. maximus* (Sims et al. 2000), and mobulids utilize the water column and surface (Pratt and Carrier 2005). Evidently, the use of a larger space for courtship results in more acro-batic and exaggerated movements in such species, as seen in mobulids. However, it appears that mobulid courtship can exist in a small, confined space, such as aquaria, where all stages have been observed for *M. alfredi* (Anon 2007), *M. hypostoma* (Jacob Rosner, pers. comm.), and *M. mobular* (Kiyatake et al. 2024).

After “following” behaviours, the stage known as “Pre-copulation positioning” in mobulids is also observed in other elasmobranchs, such as *C. longimanus* and *C. leucas*, whereby the female signals her readiness to mate by exhibiting a “stall” (Talwar et al. 2023). Similarly, in mobulids, the female slows down while the males employ various tactics to assist in lifting or grasping her pectoral fin. Breaching may be used as a tactic for slowing down the female, as seen in this study and others (Klimley et al. 2024). However, breaching has never been observed after copulation or immediately before copulation, which indicates its specific involvement in courtship when related to reproductive behaviours (Klimley et al. 2024). During copulation, elasmobranch mating always involves male “biting” and “holding” behav-iour, supported by our findings. This behaviour has been documented in numerous elasmobranchs, including roughtail stingrays [*Bathytoshia centro-ura* (Mitchill, 1815)] (Reed and Gilmore 1981), *U. jamaicensis* (Dugger 1987), and *A. narinari* (Tricas 1980). In *T. obesus*, these stages are described as “pectoral grasp”, “positioning and alignment”, “clasper flexion”, “insertion and copulation”, and “separating” (Whitney et al. 2004).

The distinct preference for biting the left pectoral fin is apparent in all mobulids where observations have been documented except for *M. eregoodoo* and *M. tarapacana* (Marshall and Bennett 2010; Stevens

2016; current study). Other elasmobranchs, such as *T. meyeni*, also employ the right side (Arnés-Urgellés et al. 2018). For instance, during nurse shark (*G. cir-ratum*) mating, if a female is approached from her left side, her left pectoral fin is grasped, and the male’s left clasper crosses his cloaca and is inserted (Klim-ley 1980; Carrier et al. 1994). The male then thrusts and turns his tail, lasting from seconds to minutes in larger shark species, and even extending to hours in certain skates (Pratt and Carrier 2004). Mobulid copulation occurs for approximately 1 to 2 min (i.e., 60–90 s *M. birostris*; Yano et al. 1999), which our findings support. To ensure fertilization and optimise fecundity, repeated copulations (sometimes with dif-ferent males) occur in some species, a phenomenon also observed in mobulids (Yano et al. 1999; Stevens 2016). Some species copulate on the seafloor, like *T. obesus* (Whitney et al. 2004) and *G. cirratum* (Klim-ley 1980; Carrier et al. 1994), in mid-water, as seen in *A. narinari* (Tricas 1980), or at the sea surface, similar to *N. brevirostris* (Johnson and Nelson 1978; Clapis-Garla et al. 2022), and *C. maximus* (Sims et al. 2000). Many mobulid copulations also primarily occur at the water’s surface, potentially due to the ani-mals sinking during copulation due to their negative buoyancy (Stevens et al. 2018a).

Following copulation, the subsequent stage may be “Post-copulation Holding” or immediate “Separation” (Stevens et al. 2018a), with the occurrence of these stages being variable and dependent on individual events. Similarly, in *G. cirratum*, after insemination, thrusting ceases and both parties become motionless; then the male slowly withdraws his clasper (Klimley 1980; Carrier et al. 1994). A viscous mucus mass, likely containing some sperm, extends between the clasper and female cloaca before breaking and retract-ing while the female remains still, cradling her pel-vic fins (Carrier et al. 1994). The pair subsequently becomes active again and either rapidly separates and swims away, or the male releases his pectoral grasp and remains motionless on the seafloor, seemingly in a state of recovery (Carrier et al. 1994). This study’s comprehensive observations show that courtship behaviour is largely similar across elasmobranchs, with the mobulids exhibiting complexity and more species-specific variation than previously thought, likely due to sociality and intelligence, traits also found in many elasmobranchs.
Importance of sociality in mobulid reproduction compared to other elasmobranchs

The substantial energetic investment required for increased brain size in mobulids (Ari 2011; Arostegui 2024) indicates the potential importance of social behaviour in their reproductive success, as supported by the existing evidence. In that regard, at cleaning stations there is an increased amount of sociality among *M. alfredi* (Perryman et al. 2021, 2022), with these interactions possibly relating to reproduction (Marshall and Bennett 2010; Stevens et al. 2018a; Germanov et al. 2019; Perryman et al. 2019). For instance, specific *M. alfredi* individuals are regularly seen together on certain cleaning stations (Perryman et al. 2019). Other observations indicate that males follow females for days to weeks, feeding and cleaning (Stevens 2016). In elasmobranchs, social hierarchies and interactions can occur both before and after active mating, extending into non-reproductive periods, particularly in species that remain together or repeatedly use specific mating sites over several annual cycles, as seen in *C. taurus* (Gilmore et al. 1983) and *G. cirratum* (Klimley 1980; Carrier et al. 1994).

The significance of sociality in mating behaviours may offer insights into the complex behaviours observed in *M. alfredi* (Perryman et al. 2022) and *M. munkiana* (Palacios et al. 2024), and with more research, likely other species too. Social mating behaviours included male-to-male courtship with one another, a behaviour described in several species for the first time in this study (see above). Fish such as the Mozambique tilapia, *Oreochromis mossambicus*, practice male-male courtship that presents the full courtship repertoire (tilting, signaling the nest, circling and quivering) (Oliviera and Almada 1998); as well as birds, more than 130 species (MacFarlane et al. 2010). One such species is the white-ruffed manakin [*Corapipo altera*, (Corapipo altera Hellmayr, 1906)], whereby young males often practice displays, sometimes in groups of 4–5, typically on tree logs that are not used by adults in that season (Rosselli and Vasquez 2002). These social behaviours occur at leks, in which cleaning stations have been proposed to function as (Stevens et al. 2018a, b). In mobulids, the only female-female courtship display observed is when two females briefly become engaged in the same courtship train with males (Stevens 2016), this may be because female mate choice is the primary driver for copulation. The

novel male-male observations therefore designate additional sub-stages (pseudo-initiation, pseudo-evasion and pseudo-endurance) for mobulids to those described by Stevens et al. (2018a, b).

## Mobulid reproductive plasticity/flexibility

The similarity of social and courtship behaviour, and the microsympatric use of certain habitats, like cleaning stations, may have resulted in interspecies courtship (Setyawan et al. 2024) and hybrids (Walter et al. 2013) between *M. alfredi* and *M. birostris*. The existence of the latter might indicate additional cases in other mobulid species (e.g., *M. birostris* and *M. yarae*, or *M. birostris* and *M. tarapacana* in multiple locations; Stevens et al. 2018a, b). Furthermore, with over half of mobulid species where near-term pregnant females engaged in courtship behaviour were observed, combined with field observations, suggested copulation often occurs within hours or days of parturition (Stevens 2016; Germanov et al. 2019; Carpenter and Griffiths 2023; Palacios et al. 2024; current study). Given the low fecundity of these species, the multiple copulation events recorded for the same female within a few days/weeks with different males (Yano et al. 1999; Stevens 2016), and the average interbirth intervals reported (> 1 year), this raises the likelihood of female sperm storage and/or embryonic diapause in these wide-ranging species (Stevens 2016; Jordan et al. 2021).

## Conclusions

The findings presented here offer the most comprehensive insight into mobulid courtship and mating behaviour to date, revealing patterns that appear consistent across the family and more broadly all elasmobranchs. We provide insights for identifying and contrasting the reproductive patterns and behaviours among mobulids. This included the first description of courtship and full mating observations for *M. hypostoma*, as well as complete cycle data for several species (e.g., *M. yarae*) and novel behaviours such as male–male courtship. We identify gaps in the literature, such as the reproductive behaviour of *M. eregoodoo* and a lack of complete observations of mating for *M. kuhlii*, *M. mobular*, and *M. tarapacana*, including very minimal observations for *M. thurstoni*. Reproduction is essential for the recovery of threatened species, such as these, and we hope this study will allow researchers better identification of both behaviour and indicators of reproduction in the field for continued monitoring.
**Acknowledgements** The authors thank all who contributed data to this study, of which this would not be possible without.

**Funding** The authors did not receive specific funding for this work. This work was facilitated by large-scale citizen science photography/videography.

**Data availability** All the data used in this study are presented in its text and figures. Additional videos are available upon request.

## Declarations

**Ethics approval** No approval of research ethics committees was required as this study only analyses opportunistic encounters in the field.

**Conflict of interest** The authors declare no competing interests.

**Open Access** This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/.

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