Individual recognition in a jumping spider (Phidippus regius)

Recognising familiar individuals helps animals navigate their social lives, including finding mates, avoiding rivals, and caring for their young. This skill, called individual recognition, is often linked to social species with large brains, and it supports higher cognitive functions such as empathy, reputation tracking, and attributing mental states.

Recognising an individual requires memory and flexible matching across viewpoints and lighting, and larger brains have more neurons and brain areas that respond strongly to faces. Jumping spiders are tiny hunters with excellent vision, but they are mostly solitary and rarely encounter the same spider twice. Previous studies showed that these animals can distinguish species and sex using colours and patterns. However, it was less known whether a jumping spider can also recognise a specific individual.

To find out more, Dahl and Cheng set up face-to-face meetings between regal jumping spiders (Phidippus regius) while keeping them safely separated by transparent panels. Overhead video tracked how close the pair chose to be. Mutual distance was treated as a simple readout of interest, while moving closer suggested stronger attention to the other spider. Keeping distance indicated reduced interest or growing familiarity.

When a spider first saw another individual, it often approached. If the same pair met again minutes later, they tended to keep a bit more distance, consistent with becoming familiar with that specific spider. But when the next encounter featured a different individual, the spiders moved closer, showing increased interest. Across many controlled pairings, this clear behavioural switch between familiar and new repeated reliably.

The effect weakened as the same individuals were shown repeatedly across sessions, even hours later, potentially indicating that the spiders were building up familiarity. Introducing a completely new individual at the end of the experiment – after hours of testing – produced the strongest renewed interest. This late rebound is inconsistent with general fatigue or waning motivation. If spiders were merely tired, an approach would be expected to drop for all stimuli. Instead, this points to sensitivity to individual novelty. In other words, the spiders retain information about who they have seen and retrieve this information after delays – a pattern consistent with encoding and storing individual-specific visual features and comparing new views against that record.

These findings invite a shift in perspective. If a tiny, mostly solitary spider can recognise individuals, how much brain is really needed for flexible social memory? We may underrate animals with compact nervous systems because we treat brain size as a stand-in for cognition. What do such abilities imply for debates about animal consciousness? Behaviour cannot prove subjective experience, but it narrows what kinds of minds are plausible. Future work can map individual recognition across animal groups using the same test. It can then test whether the pattern tracks lifestyle and sensory systems more than brain size, and whether the trait evolved many times independently or from an ancient common origin.

Recognising individuals is a complex cognitive process requiring flexible learning and recognition memory. Arthropod species possessing the social ability of individual recognition would, thus, stand in stark contrast to the commonly accepted notion that animals with smaller brains are cognitively less advanced due to reduced computational power of nervous systems with smaller and fewer neurons (Niven and Farris, 2012). And yet, there is evidence for an arthropod species displaying face learning (Sheehan and Tibbetts, 2011) and long-term social memory (Sheehan and Tibbetts, 2008). That is, a social wasp species (Polistes fuscatus) showed mammal-like face learning (Sheehan and Tibbetts, 2011; Tibbetts, 2002), arguably providing social benefits by reducing aggression and stabilizing social interactions. As one of the few reported instances of individual recognition in arthropods (see also Dreier et al., 2007), this has contributed to the prevailing view that non-social arthropod species are unlikely to evolve such complex cognitive processes. The underlying reasoning is that individual recognition entails high energetic costs, longer processing times, and consequently an increased risk of predation – costs that would not be outweighed by the limited number of social encounters or the marginal survival benefits they provide (Gherardi et al., 2012; Dale et al., 2001). The general consensus, thus, is that a certain degree of sociality sensu Wilson (Wilson, 2000) is required for the emergence of individual recognition (Dale et al., 2001). Here, we challenge this consensus by testing for individual recognition in Phidippus regius, a non-social, miniature-brained jumping spider. In a controlled experimental procedure, we confronted subjects with live conspecifics, intensifying the social encounter and ensuring ecological relevance in the stimulus presentation. While jumping spiders have been shown to use visual and chemical cues for species, sex, or rival recognition (Tedore and Johnsen, 2013; Cross et al., 2020), there is no direct evidence for memory-based recognition of individual identity, particularly in taxa such as jumping spiders, where repeated encounters with the same conspecific are infrequent.

As a first step, we assessed the ability of P. regius to individually recognise other members of its species, commonly referred to as individual recognition (Tibbetts and Dale, 2007) or individuation of conspecifics (Bonatti et al., 2002). To test this, we employed a habituation-dishabituation paradigm, in which one individual habituates to the extended presence of another individual in its close proximity. Following a brief phase of visual separation, a different individual is introduced. If the focal individual can discriminate the current individual from the former, it is expected to exhibit dishabituation (Humphrey, 1974; Dahl et al., 2007). In other words, in this habituation-dishabituation paradigm, we expect the rebound in ’interest’ to be greater when the identity of the spider changes than when the same individual is presented again. To experimentally control the animal pairs, we placed each individual in a separate container with one transparent side and a transparent top panel. We then pairwise confronted the individuals by placing the containers such that the transparent sides faced each other, allowing the spiders to visually explore and approach one another at close range while preventing any direct physical interaction.

Each trial followed the same procedure: Two spiders, say individuals A and B, were exposed to one another for 7 min, eliciting an initial ’interest’ in each other. They were then visually separated for 3 min using an opaque slider. Following this separation, the same pair could be re-exposed to one another (A vs B, habituation trial) or either individual could be paired with a new individual (A vs C, or B vs D, dishabituation trial), again for 7 min, followed by another 3-min separation period. Relative interest was quantified by approximating spatial distances between the spider pairs in the xy-plane from video recordings captured through the transparent top panel, where high interest is reflected in smaller values (i.e. spiders move closer) and low interest in larger values (spiders maintain distance). Under the assumption that spiders are capable of individuating each other, we predict that in the habituation condition, where the same individuals are re-encountered, relative interest decreases, resulting in an increase in distance (Figure 1a–c; ‘Habituation’, hypothetical example 1: spiders seek maximal distance to each other [dashed line]; example 2: spiders seek medium distance to each other [solid line]). Conversely, in the dishabituation condition, where a new individual is introduced, relative interest increases and spiders approach each other, leading to a decrease in distance (Figure 1a–c; ‘Dishabituation’). It is important to clarify the use of the term ‘Baseline’ as illustrated in Figure 1a. The baseline trial shown there represents only the first trial in each session, before any habituation or dishabituation has occurred. For all subsequent comparisons, relative changes in proximity were computed by comparing each trial to the immediately preceding one. Specifically, each dishabituation trial was analysed relative to the preceding habituation trial (Figure 1c: ‘Dishabituation - Habituation’), while each habituation trial (except for the first in each session) was analysed relative to the preceding dishabituation trial (Figure 1c: ’Habituation - Baseline’, equivalent to ‘Habituation - Dishabituation’). Thus, habituation and dishabituation are not absolute responses, but are defined by how much the behaviour shifts relative to a reference point. As a result, the difference scores reflect relative changes in the frequency distribution of distances. Some values in the hypothetical distributions (Figure 1c) are negative, indicating that the number of times a spider was at a given distance was lower compared to the preceding trial. A decrease in interest can be inferred when this underrepresentation (i.e. negative values) occurs in the more proximal distance bins, accompanied by an overrepresentation (i.e. positive values) in the medium or distal distance bins. Conversely, an increase in interest can be inferred when there is an overrepresentation of occurrences (i.e. positive values) in the proximal distance bins, coupled with an underrepresentation (i.e. negative values) in the medium or distal distance bins. These relative changes in proximity describe the behavioural signature of habituation and dishabituation (Figure 1c) and constitute the basis for our statistical analyses.

Figure 1

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In the first experiment, we divided a total of 20 individuals into five groups of four individuals each. Each individual of each group was exposed to the three group members in both habituation and dishabituation trials, resulting in six trials per session, equivalent to one hour of recording. We repeated this procedure twice, resulting in 18 trials across three sessions and an exact repetition of a given trial (and pairing of individuals) in 1 hr intervals (for a detailed description of the procedure see Materials and methods and Table 1 and Table 2).

We found that spiders adjusted their proximity depending on whether they encountered a familiar or a new individual. In habituation trials, where the same individual was presented again, spiders tended to maintain greater distances. In contrast, in dishabituation trials, where a new individual was introduced, spiders were more likely to stay at close distances. This pattern was statistically robust: habituation and dishabituation trials (i.e. predictor variable condition) differed significantly as a function of inter-individual distances (i.e. predictor variable distance), leading to a significant improvement of model fitting the interaction of the predictors distance and condition (LRT: ${\chi }_{\mathrm{\Delta }3}^2$ = 63.66, p< 0.001; Figure 2a, exemplar trials: Figure 1d–f, Video 1, Figure 2—videos 1; 2; for model parameter estimation: Appendix 1—figure 1, Appendix 1—table 1): dishabituation trials (blue discs) showed a greater proportion of close-distance values than habituation trials (red discs), whereas habituation trials showed a greater proportion of far-distance values.

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Furthermore, the strength of this effect varied across sessions. The interaction between distance and condition was significantly modulated by session, indicating that the dissociative effect of condition changed over the course of testing periods. This effect was strongest in Session 1 and weakened progressively, with the weakest modulation observed in Session 3 (LRT: ${\chi }_{\mathrm{\Delta }6}^2$ = 34.14, p< 0.001; Figure 2a, exemplar trials: Figure 1d–f, Video 1, Figure 2—videos 1; 2; for model parameter estimation: Appendix 1—figure 1, Appendix 1—table 1, Dahl and Cheng, 2024).

The systematic dissociation of distance values between habituation and dishabituation trials suggests that P. regius is capable of individual recognition. If spiders did not differentiate between conspecifics, we would not expect such a consistent divergence in distance patterns between conditions (habituation vs dishabituation). In addition, this dissociation diminished across sessions, indicating a progressive habituation effect towards the already encountered individuals. That is, in Session 1, the initial exposure to each individual elicited a pronounced dishabituation response. By Session 2, this dishabituation response was still present but attenuated. By Session 3, the dishabituation response had largely disappeared, indicating that spiders no longer differentiated between familiar and novel individuals (Video 1, Figure 2—videos 1; 2). While this progressive reduction is consistent with the formation and retrieval of individual-specific memories, alternative explanations, such as general fatigue due to extended testing periods, cannot be ruled out at this stage.

As a second step, we therefore examined whether P. regius’s decreasing interest across repeated sessions is driven by a general fatigue effect due to the prolonged testing procedure, or whether P. regius recognises the current individual after having encountered it at least once (by Session 2) or twice (by Session 3), and as a result, no longer dishabituates. Such recognition capability would support the presence of long-term memory representations in the individuation of conspecifics, given the extended retention interval spanning from minutes to hours.

To this end, we assessed whether the presentation of a completely novel individual - unseen during any of the three experimental sessions - would elicit a rebound in interest when introduced at the end of Session 3. We refer to this condition as dishabituation [long-term] trials, in contrast to the dishabituation trials conducted during Sessions 1–3, henceforth referred to as dishabituation [short-term] trials (see Table 3). Importantly, the labels ‘short-term’ and ‘long-term’ do not imply specific memory systems, but rather reflect the retention interval at which the respective dishabituation trials were administered.

If such rebound occurs, it would indicate that the observed habituation across sessions results from the recognition of repeatedly presented individuals, rather than from physical fatigue due to the prolonged testing procedure. In other words, such rebound would suggest a form of ‘cognitive’ fatigue – a diminished response elicited by the repeated re-encounter of familiar individuals – subserved by long-term memory formation, rather than a ‘physical’ fatigue effect. To test this, we re-ran the experiment in an additional 16 spiders, arranged into four groups and added a memory dishabituation [long-term] trial at the end of Session 3. The memory dishabituation [long-term] trials were generated by cross-combining individuals from two groups (group 1: A, B, C, D; group 2: E, F, G, H; Table 3) that had been run in parallel: At the end of Session 3, each spider was paired with a novel individual from the other group (e.g. A - E, B - G, C - F, D - H), resulting in previously unseen pairings.

First, we observed that spiders responded with renewed interest toward these unfamiliar individuals (dishabituation [short-term] trials), approaching them more closely than they had approached previously encountered ones (habituation trials). Thus, we replicated our earlier findings and found a dissociation between the factors distance and condition (LRT: ${\chi }_{\mathrm{\Delta }3}^2$ = 29.52, p < 0.001; Figure 2b, Figure 2—video 3; for model parameter estimation: Appendix 1—figure 1, Appendix 1—table 2). Specifically, dishabituation [short-term] trials (blue discs) showed a greater proportion of close-distance values, whereas habituation trials (red discs) showed a greater proportion of far-distance values. Most critically, we found that the dishabituation [long-term] trials at the end of Session 3 elicited a rebound in interest that clearly exceeded the rebound in the dishabituation [short-term] trials of the same session. This was reflected in a significant interaction between condition (i.e. dishabituation [short-term] vs dishabituation [long-term]) and distance (F(3,127) = 3.91, sum sq.=0.92, mean sq.=0.31; p< 0.01, Figure 2b: right subfigure, white diamonds; exemplar trials: Figure 1g–i and j–l, Videos 2 and 3, Figure 2—video 4, Dahl and Cheng, 2024). This finding suggests that the reduction in effect size across sessions was not due to general fatigue, but rather reflects a long-term memory for previously encountered individuals: That is, when presented with a truly novel conspecific, the spiders’ interest rebounded to an unprecedented level.

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Our findings show, first, that P. regius can recognise individuals to which it was exposed to for a short period of 7 min and that reoccurred after a visual separation period of 3 min. Second, P. regius exhibited long-term habituation, that is a sustained reduction in interest when encountering the same individuals again 1 or 2 hr later. This result pattern is only possible if spiders remember the encountered individuals across sessions, suggesting the formation and retention of individual-specific memory representations. Third, despite this long-term habituation, P. regius showed an unprecedented rebound in interest when confronted with an entirely novel individual, ruling out a physical fatigue effect in favour of a ‘cognitive’ fatigue based on long-term memory. For these reasons, our results are the first evidence that P. regius, a non-social arthropod species, possesses long-term memory that allows it to individuate conspecifics and recognise novel individuals.

Before addressing the evolutionary implications of these findings, it is important to consider a key methodological question: Do the observed behavioural changes reflect true recognition memory, or could they result from simpler mechanisms such as transient familiarity or physical fatigue? To distinguish these possibilities, we employed a stringent habituation–dishabituation paradigm (Fantz, 1961), widely used in animal (Dahl et al., 2007) and infant cognition (Kavsek and Bornstein, 2010) to isolate memory-based recognition from short-term novelty effects. Unlike paradigms with static images or single exposures, our approach required subjects to recognise a dynamic, interacting conspecific across repeated encounters, making low-level novelty detection or general desensitisation unlikely explanations. In our design, evidence for individual recognition comes from both within-session and across-session comparisons: Within sessions, dishabituation [short-term] trials consistently elicited closer proximity-seeking behaviour than habituation trials, indicating discrimination between familiar and novel individuals. Across sessions, this effect showed a structured decline: strong in Session 1, attenuated in Session 2, and absent in Session 3. This temporal pattern is inconsistent with a general desensitisation, which would reduce proximity-seeking behaviour across all trial types regardless of stimulus identity. Instead, our results support the formation of identity-specific memory representations. Further support comes from the long-term dishabituation trials at the end of Session 3, where introducing a novel individual elicited the strongest rebound in interest. If prior responses reflected transient familiarity or physical fatigue, such a pronounced effect would not be expected. This indicates that spiders encoded previous individuals as distinct representations and recognised novelty at the level of individual identity.

Our results provide clear evidence of memory-based individual recognition in P. regius. If a spider does not dishabituate when re-encountering an individual, this indicates that a memory representation for that individual has been formed and retrieved. Conversely, if a marked dishabituation response occurs upon introduction of a novel individual, it implies that the current experience does not match any existing memory representation, prompting renewed interest. The clear dissociation between habituation and dishabituation trials within sessions, the progressive reduction in dishabituation responses across sessions, and the pronounced rebound upon introduction of a truly novel individual together indicate the formation and retrieval of individual-specific memories, rather than transient familiarity or physical fatigue.

Recognising members of one’s own species is a crucial cognitive ability that underpins various adaptive behaviours. Individual recognition allows animals to distinguish between friend and foe, to identify a mating partner, its offspring, or a kin member. Individual recognition is achieved via the production of individually distinct features (e.g. visual) or signals (e.g. acoustic) by the sender, and the ability of the receiver to extract and process these features and signals (Yorzinski, 2017). In social species, individual recognition bears particular significance in contexts such as territoriality, aggressive competition, and parental care (Tibbetts and Dale, 2007). It is precisely because jumping spiders, including P. regius, are generally considered non-social, solitary, and aggressive towards conspecifics that the presence of individual recognition is unexpected. This raises the question about the biological relevance of individual recognition in P. regius: One of the few social interactions in the life of a jumping spider occurs during mating, encompassing a highly structured visual courtship display composed of coordinated body movements and distinct morphological features. It is believed that the colouration of the appendages (the chelicerae), along with the facial hair patterns, serves as important visual features for species and sex identification in jumping spiders and may also signal individual quality during mate assessment (Zhou et al., 2021; Hill et al., 2006). Hence, the colouration of body parts (sender) and the ability to discriminate colours (receiver) appear sufficient for sexual selection, such as species and sex identification or mate assessment, without necessarily supporting individual recognition (Lim et al., 2007; Zhou et al., 2021). Similarly, in aggressive interactions, such as territorial disputes, fighting ability is largely associated with the size and colouration of the chelicerae (Lim et al., 2007; Tedore and Johnsen, 2012). In line with this, research has shown that the demands of mating and aggression in jumping spiders are met by cue recognition - by definition a basic-level classification - allowing classification of species, sex, or general rival status. For example, jumping spiders have been shown to use a combination of visual and chemical (pheromonal) cues to differentiate both species and sex, enabling them to identify potential mates or competitors (Tedore and Johnsen, 2013). Some species recognise rivals by relying on distinctive colour signals, such as a red facial patch, which serves as a key visual cue in aggressive interactions (Cross et al., 2020). In each case, it is basic-level category recognition rather than individual identity that guides behaviour, and thus subordinate-level discrimination appears unnecessary for these core ecological functions. As such, territoriality and aggressive behaviour appear insufficient as ultimate explanations (Tinbergen, 1963) for the evolution of individual recognition in P. regius. Instead, decisions in these contexts can be based on general physical features rather than identity-specific information.

In addition, most jumping spiders exhibited limited parental care (Gardner, 1965), protecting the nest through the spiderlings’ first molt. A notable exception is Toxeus magnus, which provides a nutrient-rich, milk-like substance to its offpsring, a behaviour functionally and behaviourally analogous to lactation in mammals (Chen et al., 2018). Despite these examples, there is little evidence that memory-based recognition of individual offspring is required to support parental care in salticids. In more specific contexts, however, some jumping spiders have been shown to discriminate their own eggsacs from those of conspecifics, presumably to avoid filial cannibalism (Clark and Jackson, 1994a), or to distinguish their own silk draglines from those of other conspecifics, representing a form of self-recognition (Clark and Jackson, 1994b). Both forms of recognition, however, are context-dependent and do not extend to memory-based identification of conspecifics across repeated encounters. Hence, although jumping spiders demonstrate a range of basic-level recognition abilities adapted to specific ecological challenges, these abilities, however, do not extend to the subordinate-level individual recognition demonstrated in the present study. In other words, even in this context, there is no clear evidence that memory-based individual recognition solves a critical survival problem.

Nevertheless, it is worth considering that individual recognition could provide adaptive benefits in certain social contexts, even in taxa where repeated encounters are typically rare. For instance, the so-called ‘dear enemy effect’, a phenomenon observed in territorial animals, involves reduced aggression towards familiar neighbours while maintaining heightened vigilance against unfamiliar individuals (Temeles, 1994; Bee and Gerhardt, 2002). This form of recognition can minimise the costs of unnecessary conflict and enhance social stability. While the natural history of P. regius suggests rather infrequent repeated encounters with the same conspecific, the current findings raise the intriguing possibility that cognitive capacities for identity recognition facilitate the emergence of ‘dear enemy’ relationships if territorial dynamics arise. Future experiments could directly test whether the ’dear enemy effect’ can be experimentally elicited in jumping spiders.

Taken together, the generally solitary nature of P. regius offers little ecological pressure to favour individual recognition. According to the ’minimum needs hypothesis’ (Wiley, 2013; Gherardi et al., 2012), selection will only favour specific recognition abilities when basic-level classification (Rosch et al., 1976; e.g. species, sex, or quality assessment via colouration or size cues) no longer suffices to solve ecologically relevant problems. In the case of P. regius, basic discrimination, such as chelicerae size or colouration, may be sufficient in most known contexts of social interaction. Moreover, the neural implementation of subordinate-level classification (Rosch et al., 1976), such as identity recognition, requires more detailed and exhaustive processing than basic-level categorisation. This type of fine-grained discrimination is typically associated with specialised neural structures and increased cognitive and neural resource requirements (Weiner and Grill-Spector, 2015). Given these costs and the limited ecological need for identity recognition in P. regius, it is more plausible to assume that this ability arises not through direct selection, but through pleiotropy, as a byproduct of general cognitive architecture evolved for other purposes - a view consistent with the ’general learning hypothesis’ (Wiley, 2013). Jumping spiders are known for complex foraging and navigation strategies (Jakob et al., 2007; Jakob et al., 2011; Skow and Jakob, 2006), requiring flexible learning and high behavioural adaptability. These domain-general cognitive capabilities may enable recognition of conspecifics with a degree of detail and specificity that exceeds their minimum recognition needs (Yorzinski, 2017).

In contrast to social animal species, including eusocial arthropod species (Tibbetts, 2002; Sheehan and Tibbetts, 2011; Dreier et al., 2007), where individual recognition is often maintained by direct selective advantage, we cannot conclusively identify a survival benefit for individual recognition in P. regius. Instead, we put forward the idea that individual recognition in P. regius emerges as a pleiotropic consequence, namely a byproduct of an already fairly sophisticated, broader learning capability. Critically, individual recognition relies on recognition memory, a form of long-term memory, in which a previously encountered event or entity, here an individual, is stored as a representation and reactivated upon re-encounter (Norman and O’Reilly, 2003). This memory mechanism may also underlie complex behaviours observed in other salticids, such as when Portia fimbriata navigates towards prey after an initial visual scan of the environment - even without visual input during execution (Tarsitano and Jackson, 1997) - or when Menemerus semilimbatus recognises biological motion, demonstrating sophisticated temporal integration and dynamic cue recognition (De Agrò et al., 2021).

Thus, our study challenges the notion of spiders being stimulus-response driven automata, by not only contributing to an increasing body of evidence that spiders and salticids in particular produce a wide spectrum of intelligent behaviour (Jackson and Cross, 2011), but by pinpointing the presence of two fundamentally important mechanisms for any higher cognitive processing: flexible learning and recognition memory. The key building blocks of these mechanisms are representations, mental images of external entities, that are not present to the sense organs, allowing more elaborate information processing, such as in complex decision making and goal-directed behaviour. The existence of which in arthropods in general and spiders in particular triggers rethinking of miniature brain cognition (Jackson and Cross, 2011).

Supplementary information is available for this paper. Codes and materials are available (https://osf.io/gpnct/).

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Author details

1. Christoph D Dahl

   1. Institute of Biology, University of Neuchâtel, Neuchâtel, Switzerland
   2. Graduate Institute of Mind, Brain and Consciousness, Taipei Medical University, Taipei, Taiwan

   Contribution

   Conceptualization, Resources, Data curation, Software, Formal analysis, Supervision, Funding acquisition, Validation, Investigation, Visualization, Methodology, Writing – original draft, Project administration, Writing – review and editing

   For correspondence

   christoph.d.dahl@gmail.com

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-4296-1526

2. Yaling Cheng

   Graduate Institute of Mind, Brain and Consciousness, Taipei Medical University, Taipei, Taiwan

   Contribution

   Conceptualization, Resources, Investigation, Methodology, Project administration

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0009-0002-5198-5797

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