# New acute chromatic components during prey attack in juvenile cuttlefish (Sepia officinalis, Linnaeus 1758): the “Leopard spots”

Manon Peyrafort, Anne-Sophie Darmaillacq, Chuan-Chin Chiao, Ludovic

Dickel

► To cite this version:

Manon Peyrafort, Anne-Sophie Darmaillacq, Chuan-Chin Chiao, Ludovic Dickel. New acute chromatic components during prey attack in juvenile cuttlefish (Sepia officinalis, Linnaeus 1758): the “Leopard spots”. Behavioural Processes, 2025, 226 (2), pp.105164. 10.1016/j.beproc.2025.105164. hal-04939842

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# New acute chromatic components during prey attack in juvenile cuttlefish (*Sepia officinalis*, Linnaeus 1758): The “Leopard spots”

Manon Peyrafort<sup>a</sup>, Anne-Sophie Darmaillacq<sup>a</sup>, Chuan-Chin Chiao<sup>b</sup>, Ludovic Dickel<sup>a,*</sup>

## ARTICLE INFO

**Keywords:**
Mollusc
Cephalopod
Cuttlefish
Predation
Body pattern

## ABSTRACT

Cephalopod molluscs exhibit sophisticated colour changes that allow them to camouflage themselves in a dynamic environment, and to communicate with conspecifics, prey or predators. The present study reports the description of previously undescribed three chromatic body components that were observed by coincidence during prey attack by young cuttlefish. These three components are formed by several units in the form of dark spots located inside the white square on the mantle (L1), between the eyes along the white head bar (L2) and on the arms (L3). They were called the “Leopard spots.” They appeared very transiently during the attack of a shrimp, alone or in group, and their expression was variable within and between individuals. We hypothesise that the Leopard spots play a role in reducing the risk of cuttlefish predation during the shrimp attack, but it is not known whether this is a form of camouflage or a warning signal to predators.

## 1. Introduction

Coleoid cephalopods (octopus, cuttlefish and squid), and in particular, common cuttlefish (*Sepia officinalis*), are common models for camouflage studies. These molluscs have developed a complex system of dermal units (chromatophores, leucophores, iridophores) to produce a variety of body patterns that can change in less than 1 s (Chiao and Hanlon, 2019). To describe this system, Packard and Sanders proposed a hierarchical organisation for the octopus *Octopus vulgaris* (Packard and Sanders, 1971), which was adopted and adapted for the cuttlefish *Sepia officinalis* by Hanlon and Messenger (1988). This classification orders the system into elements (chromatophores, leucophores, iridophores, papillae), units (a particular morphological arrangement of elements in the skin), components (a group of units that allow the display of a shape or a texture modification at a given location) and patterns (the total appearance of the animal at a given moment, made up of several components) (Hanlon and Messenger, 1988). In *S. officinalis*, 13 basic body patterns have been observed, based on as many as 54 components, including 34 chromatic components with varying intensities of expression. Six patterns described in this species are “chronic” and seven are “acute” (Hanlon and Messenger, 1988). Chronic patterns may last for minutes to hours and are used for cryptic purposes (primary defence;

Hanlon and Messenger, 1988). Acute body patterns last only a few seconds or minutes and are used during communication episodes, either in response to a conspecific, to predator threat (secondary defences; Hanlon and Messenger, 1988; How et al., 2017; Palmer et al., 2006; Shook et al., 2024) or to deceive prey.

Paradoxically, studies on cuttlefish camouflage have mainly been carried out under stable laboratory conditions, whereas cuttlefish inhabit a permanently dynamic environment. Movement can be a disadvantage for cuttlefish as it can make them more visible to predators. Few studies investigated the camouflage of *S. officinalis* in motion (Josef et al., 2015, 2017; Zylinski et al., 2009) or changes in acute body patterns when they attack prey. Three of these acute patterns are: the “passing cloud” and the “flashing” patterns (dynamic patterns lasting a fraction of a second to a few seconds), displayed on waving arms during attention and positioning (Holmes, 1940; Hanlon and Messenger, 1988; How et al., 2017) and the “tentacle shot pattern” during tentacle striking (Kim, et al., 2022). In addition, two studies mentioned transient colour changes during prey seizure. These were described as “dark uniform” (Messenger, 1968) or “dark or light mottle” (Adamo et al., 2006).

The present study reports the discovery and the description of dark spots appearing in clusters in three locations on the body of juvenile cuttlefish (*Sepia officinalis*), that were fortuitously observed when they

<footnote><sup>a</sup> Université de Caen Normandie (UNICAEN), Université de Rennes 1, CNRS, EthoS (Ethologie Animale et Humaine), UMR 6552, Luc-sur-Mer 14530, France</footnote>

<footnote><sup>b</sup> Laboratory of Visual Neuroscience, Institute of Systems Neuroscience, National Tsing Hua University, Hsinchu 30013, Taiwan</footnote>

<footnote>* Corresponding author.</footnote>

<footnote>E-mail addresses: peyrafortmanon@yahoo.fr (M. Peyrafort), anne-sophie.darmaillacq@unicaen.fr (A.-S. Darmaillacq), ccchiao@life.nthu.edu.tw (C.-C. Chiao), ludovic.dickel@unicaen.fr (L. Dickel).</footnote>
attack a shrimp. Using a frame-by-frame analysis, we focused on the variations in chromatic expression that occurred on the cuttlefish body from the introduction of a shrimp to its seizure. We propose to describe the dark spots as new chromatic components according to the original Packard and Sanders (1971) hierarchical classification of body patterns and named them the "Leopard spots."

## 2. Method

### 2.1. Subjects and housing

Cuttlefish eggs (*Sepia officinalis*) from several egg clusters were collected in the English Channel in the vicinity of Luc-sur-Mer (Normandy, France). Newly hatched cuttlefish were reared in groups (n = 30 up to two months then n = 15 up to three months), in a tank measuring 60 cm × 40 cm × 20 cm, until three months of age at the CREC Marine Station (Luc-sur-Mer, France). Containers were supplied with natural sea water (open circuit, temperature range from 16 to 22 °C) under a natural photoperiod. Cuttlefish were fed daily with live shrimps and crabs of suitable size.

### 2.2. General procedure

At three months, nine cuttlefish (cuttlefish #1 to #9; Dorsal Mantle Length mean ± Standard Deviation = 3.65 ± 0.24 cm) were selected at random from cuttlefish reared at the marine station from wild eggs and isolated in circular uniform grey tanks (10 cm water depth x 24 cm diameter) for one week. Each tank was enriched with an artificial seaweed and a natural beige rock. Before behavioural recording, the seaweed was removed and a LED band (200 lm, 4000 K) installed around the tank to provide homogeneous lighting. A Panasonic Lumix DMC G80 camera with Panasonic 12–60 mm / F2.8–4.0 lens was suspended above the circular band. Cuttlefish were then left 15 min to acclimatise. Recording started and after 1 min 20 the experimenter introduced a single live shrimp (*Crangon crangon*) into the tank. Recording of the attack of the shrimp by the cuttlefish (thereafter referred as shrimp attack) stopped at the end of the shrimp consumption, and the seaweed was returned to the tank. This procedure was repeated (once a day) over ten days.

### 2.3. Data collection and analysis

The 4 K videos (30 frames per second) obtained were viewed with VLC media player 3.0.8 Vetinari® software supplemented by Time v3.2® plug-in. Firstly, videos were observed in their entirety, from the introduction of the shrimp to the end of its consumption. Subsequently, six phases of the attack (Messenger, 1968) were identified: resting (phase 0), positioning (phase 1), tentacle strike (phase 2), tentacle seizure (phase 3), arm seizure (phase 4) and prey consumption (phase 5). Using a frame-by-frame method (8 million pixels per image), an image was captured for each phase.

The dark spots appeared in groups that occurred in three locations on the mantle, L1, L2 and L3. For each location, we scored the expression of the spots from 0 to 3 (0, not expressed, 1 weakly, 2 moderately and 3 strongly expressed; Mathger et al., 2006).

We assessed inter-rater reliability using Cohen's kappa on half of the videos. One of the observers was familiar with cephalopods and the other naïve. The observed kappa was 0.91 (95 % CI: 0.88–0.94), indicating substantial agreement between observers.

The variables recorded were analysed using nonparametric statistical tests due to small sample size. Statistical analysis and graphical representations were carried out with R studio v4.0.4® software (R Core Team, 2021). Results were interpreted at a 95 % confidence limit.

### 2.4. Ethical approval

Experiments were conducted in accordance with directive 2010/63/EU concerning the protection of animals used for scientific purposes. Procedures were approved by the ethical committee of Normandy for animal experimentation (CENOMEXA) and have received authorization from the French ministry of research and innovation (MESRI, #22095).

## 3. Results

### 3.1. Spot display and location

The new display observed was composed of small dark spots concentrated in single areas, located inside the white square on the back (location L1), between the eyes along the white head bar (location L2) and on the arms (location L3). L1 comprised from 1 to about 20 spots of different shapes, L2 from 2 to 7 oval spots, and L3 from 1 to about 10 spots of round shape (Fig. 1).

### 3.2. Frequency of spot display

Observations ranged from the time cuttlefish rested to the end of shrimp consumption. In all video recordings L1, L2 and L3 were observed only during shrimp attack: positioning (phase 1), tentacle striking (phase 2), seizure by tentacles (phase 3) and subsequent seizure in arms (phase 4) (Duval et al., 1984; Messenger, 1968; Sanders and Young, 1940; Wilson, 1946). Spots on L1 appeared over 58.62 % of the observations (observed 51 times out of 87 videos), on L2 over 5.63 % (observed 31 times out of 87 videos) and on L3 over 10.34 % (observed 9 times out of 87 videos) during shrimp attacks (Fig. 2). The simultaneous display of L1, L2 and L3 was observed four times, the combination of L1 and L3 was observed six times, the combination of L1 and L2 was observed 12 times and the combination of L2 and L3 was observed five times. For all the spot appearances, flashing represents 29.41 % of L1 observations (15 appearances out of 51) and 12.90 % of L2 observations (4 appearances out of 31). Due to the speed of the attack, some pictures were blurred, making spots difficult to observe and consequently some may have been missed by the observer. Furthermore, spots on L2 were observed twice at the end of prey consumption.

Three cuttlefish failed to catch the shrimp once out of the 10 capture sequences and none leopard spot was observed during the first phases of attack.

### 3.3. Duration of spot display

Spot display was extremely transient: for L1 428 ± 321 ms, for L2 1452 ± 3312 ms (extreme values that disrupt the mean), and for L3 445 ± 319 ms. In addition, flashing of L1 and L2 was sometimes observed at an average interval time between two successive appearances of 499 ± 671 ms for L1 and 877 ± 590 ms for L2.

### 3.4. Inter-individual differences of spot display

Out of the nine cuttlefish, only one (cuttlefish #1) never expressed spots and only four (cuttlefish #4, #5, #6 and #8) expressed spots at three locations. Out of these four individuals, cuttlefish #5 most often expressed spots on L1 (25.49 % of the time), cuttlefish #6 most often expressed spots on L2 (25.81 % of the time) and cuttlefish #4 most often expressed spots on L3 (44.44 % of the time) (Fig. 3).

### 3.5. Dynamic of spot display during shrimp attack

Spots on L1 and L2 were observed during all phases. Spots on L3 were observed only during phase 2 (tentacle striking) and phase 3 (seizure by tentacles) (Fig. 2). Spot appearance changed during prey attack (Fisher's exact test: p < 0.001; Fig. 3): during phase 1 (positioning), spots
<figure><img src="image_4.png" /><figcaption>**Fig. 1.** Description of the Leopard spots. The Leopard spots consists of spot clusters in three locations, each composed of several units (dark spots): from two to seven spots on the white square (L1, some spots are shown with arrows on the top image), from one to about 20 spots on the white head bar (L2, some spots are shown with arrows on the middle image) and from one to about ten units on the arms (L3, some spots are shown with arrows on the bottom image).</figcaption></figure>

<figure><img src="image_5.png" /><figcaption>**Fig. 2.** Frequency of appearance of Leopard spots. After the analysis of 87 video recordings of nine cuttlefish, three spot locations (L1: inside the white square, L2: along the white head bar and L3: on the arms) were observed over four phases of attack behaviour (phase 1: positioning, phase 2: tentacle striking, phase 3: seizure in tentacles, phase 4: seizure in arms).</figcaption></figure>

appeared on L2 (38.71 % of the time), and subsequently during phase 2 (tentacle striking) on L1 (43.14 % of the time), L2 (41.94 % of the time) and L3 (88.89 % of the time). During phase 4 (seizure in arms), spots on L1 appeared (41.18 % of the time). Only one cuttlefish (cuttlefish #5) showed this in a dynamic way, four showed it discreetly (cuttlefish #2,

#6, #7 and #8) and three hardly showed it (cuttlefish #3, #4 and #9) (Fig. 3).

## 4. Discussion

We observed several dark spots with different sizes and shapes that appeared together or independently but repeatedly during attacks on shrimp. We propose to consider each spot as a “unit” according to Packard and Sanders classification (1971). These units are concentrated in three distinct locations: on the mantle inside the white square (L1), between the eyes along the white head bar (L2) and on the arms (L3). The appearance of the dark spots on the three locations can be simultaneous or independent, they can be associated with various stage of prey catching phase, included in the body patterns associated with prey capture in cuttlefish. Based on the hierarchical system of cephalopod body patterns (Packard and Sanders, 1971 adapted for Sepia officinalis by Hanlon and Messenger, 1988), we can hypothesise that the spots are chromatic components occurring in three locations (L1, L2 and L3). They may be part of an acute body pattern associated with prey capture in cuttlefish (Adamo et al., 2006; Kim et al., 2022). The Leopard spots did not appear when the cuttlefish failed to catch the shrimp. However, it is premature to conclude that they never appear in the event of failure, as the 3 cuttlefish to which this happened (and it happened only once per cuttlefish) were the cuttlefish that expressed these components the least. The discrete and highly transient nature of the Leopard spots explains why it has not previously been observed by the human eye, without training, and thus described. The use of high-resolution video equipment with frame-by-frame analysis was necessary to bring out its presence.

A similar display, the “spotted display” (Warren et al., 1974), was described in *Octopus rubescens* during crab attack. It appears occasionally during prey detection and also after the attack (Barbato et al., 2007; after Fig. 1 from Warren et al., 1974). Analogies between octopus and cuttlefish need to be further investigated (Barbato et al., 2007). However, Warren and colleagues (1974) suggested that colour changes during attack were related to muscular constraint during locomotion. Cuttlefish observed in the present study showed strong inter-individual differences in Leopard spots display (in frequency and duration) although their prey-attack behaviour is largely stereotyped (Messenger,
<figure><img src="image_6.png" /><figcaption>**Fig. 3.** Dynamics of Leopard spots display during shrimp attack. The three components L1, L2 and L3 were observed over four attack phases (phase 1: positioning; phase 2: tentacle striking; phase 3: seizure in tentacles; phase 4: seizure in arms) for nine cuttlefish. The frequency of spot displays (expressed as a percentage of the total number of observations per component) was measured for each cuttlefish, each location and each phase. Four classes were established (0 %, 0 % - 5 %], 5 % - 10 %] and > 10 %). In brown, the phases when the spots appeared the most often.</figcaption></figure>

1968; Zoratto et al., 2018). Consequently, it is unlikely that the appearance of the Leopard spots is linked to global muscular constraints during the pre-attack phase (Warren et al., 1974). However, a precise study of body postures and muscular activity during each Leopard display would be needed to test this hypothesis.

We can only speculate about the function of these new components. Dynamic skin patterns are considered to fulfil the function of either deceiving or communicating with a target observer which can be a prey, a congener or a predator (How et al., 2017). In our study, all component locations are visible in dorsal views, except L3 component which is potentially seen from the front by a prey. However, the L3 component is poorly expressed and occurs mainly during tentacle striking. Therefore, it is unlikely that the Leopard spots are used to deceive prey. The present study did not observe any change in the behaviour of shrimps during Leopard spots display, but this remains to be verified. Other prey with a better visual acuity than shrimps, such as fish, could also be tested to assess the effect of the display on prey behaviour.

Movements or transient colour changes can be used as a behavioural indicator of intent to attract attention and convey information (Tan and Elgar, 2021). During the search for food and in the presence of conge-ners, a very cannibalistic species of squid *Dosidicus gigas* (Markaida et al., 2008), emits a flashing and flickering display over its entire body (Rosen et al., 2015). This display could be seen as a warning signal to other squids (How et al., 2017) or as a threat pattern directed towards con-specifics in competitive situations (e.g., see agonistic behaviour of *Sepia officinalis*: Adamo and Hanlon, 1996; Palmer et al., 2006). However, cuttlefish were alone in their tank and this hypothesis would therefore seem unlikely.

Cuttlefish are more conspicuous to predators during a prey attack because of their movement. Kim et al. (2022) showed that adult cuttlefish displayed a highly contrasted, and hence conspicuous, body pattern during the capture of a piece of frozen food and suggested that it would act as a signal to halt predators for a few seconds. We have not

observed such a pattern in our study. This is probably because we used younger cuttlefish. During development the number of chromatophores increases (Reiter et al., 2018; Yacob et al., 2011) and body patterns change (some appear while others disappear; Hanlon and Messenger, 1988). We also used live shrimp which may be more motivating and ecologically relevant for cuttlefish. However, the appearance of the Leopard spots can be viewed as a signal to deter predators from attacking.

Last, the variability observed between individuals cannot be explained by individual personality (Carere et al., 2015; Zoratto et al., 2018) as it can also vary from one day to another within the same individual placed in standardised conditions. The observed difference may instead be due to the chosen camouflage strategy (Hanlon and Messenger, 1988; Zylinski et al., 2009). The Leopard spots were observed during prey attack in cuttlefish reared in the laboratory, so they were not subjected to any previous predation pressure. In their natural environment, an incorrect choice can lead to predator success and thus to strong selection in favour of movement patterns that optimise signal effectiveness (e.g., revealing defence) or dissimulation of the colour pattern (Tan and Elgar, 2021). Smaller differences between individuals in colour pattern display may be observed in wild-caught animals but further studies are needed to test this hypothesis. Additional observations on wild juvenile and adult cuttlefish subjected to strong selection pressure are also necessary.

## 5. Conclusion

The Leopard spots may be a form of signal to keep other animals at distance during prey capture (see also Shashar et al., 1996 about disappearance of polarisation pattern in cuttlefish during hunting). Even if individuals were tested alone, these signals may have developed before three months of age, while cuttlefish were kept in groups. Leopard spots are likely to play a role in reducing the risk of cuttlefish
predation but it is not known whether this is a form of camouflage or deception. The question is whether the pattern displayed by cuttlefish in motion can reduce their detection? For example, the flicker-fusion effect could be tested on predators with different visual systems, such as fish, to verify the function we have attributed to it. However, could this pattern be involved in a completely different function? Shook and colleagues asked the following question: “Are the dynamic skin patterns we observe on the skin a direct readout of changes in internal state?” (Shook et al., 2024, p.6). Therefore, more behavioural and physiological experiments are needed to address the function and development of the Leopard pattern, under other predation conditions and in other contexts.

## Funding

This work was supported by ANR (Agence Nationale de le Recherche): ETHiCs project [grant numbers 18-CE02-0022].

## CRediT authorship contribution statement

**Peyrafort Manon:** Writing – review & editing, Visualization, Methodology, Investigation, Formal analysis, Conceptualization. **Darmaillaqc Anne-Sophie:** Writing – review & editing, Supervision. **Chiao Chuan-Chin:** Writing – original draft, Conceptualization. **Dickel Ludovic:** Writing – review & editing, Writing – original draft, Supervision, Conceptualization.

## Acknowledgements

Thanks to the French National Agency for research (ANR) for funding this work., to Zoé Poupinet for validation of our observations, to Maëlle Peyrafort for illustrations of the components, to Jane Martin for proof-reading and to Manche 20–21 for equipment support.

## Data availability

Data will be made available on request.

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