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Prey seldom rely on a single type of antipredator defence, often using multiple defences to avoid predation. In many cases, selection in different contexts may favour the evolution of multiple defences in a prey. However, a prey may use multiple defences to protect itself during a single predator encounter. Such “defence portfolios” that defend prey against a single instance of predation are distributed across and within successive stages of the predation sequence (encounter, detection, identification, approach (attack), subjugation and consumption). We contend that at present, our understanding of defence portfolio evolution is incomplete, and seen from the fragmentary perspective of specific sensory systems (e.g., visual) or specific types of defences (especially aposematism). In this review, we aim to build a comprehensive framework for conceptualizing the evolution of multiple prey defences, beginning with hypotheses for the evolution of multiple defences in general, and defence portfolios in particular. We then examine idealized models of resource trade-offs and functional interactions between traits, along with evidence supporting them. We find that defence portfolios are constrained by resource allocation to other aspects of life history, as well as functional incompatibilities between different defences. We also find that selection is likely to favour combinations of defences that have synergistic effects on predator behaviour and prey survival. Next, we examine specific aspects of prey ecology, genetics and development, and predator cognition that modify the predictions of current hypotheses or introduce competing hypotheses. We outline schema for gathering data on the distribution of prey defences across species and geography, determining how multiple defences are produced, and testing the proximate mechanisms by which multiple prey defences impact predator behaviour. Adopting these approaches will strengthen our understanding of multiple defensive strategies. peerReviewed

Genus Corcovetella Galiano, 1975 The genus Corcovetella is characterised by an elevated cephalic part of the cephalothorax, chelicera with many teeth, forelegs with 2.2.2 spines, and dense plumose setae on the palps of both sexes (Galiano 1975)., Published as part of Pekár, Stano, 2022, Two new ant-mimicking spiders (Araneae: Salticidae) from Costa Rica, pp. 31-42 in European Journal of Taxonomy 852 (1) on page 37, DOI: 10.5852/ejt.2022.852.2005, http://zenodo.org/record/7448120, {"references":["Galiano M. E. 1975. Salticidae (Araneae) formiciformes. XV. Descripcion de Corcovetella aemulatrix, genero y especie nuevos. Physis, Revista de la Sociedad Argentina de Ciencias Naturales (C) 34: 33 - 39."]}

Genus Myrmapana Prószyński, 2016 Males of Myrmapana are characterised by long, flattened and broad chelicerae, with a row of teeth on ventral side, a circular coiled bulbus, and a short RTA, whereas females are characterised by spermatheca with a broad anterior dilation (Prószyński 2016)., Published as part of Pekár, Stano, 2022, Two new ant-mimicking spiders (Araneae: Salticidae) from Costa Rica, pp. 31-42 in European Journal of Taxonomy 852 (1) on page 33, DOI: 10.5852/ejt.2022.852.2005, http://zenodo.org/record/7448120, {"references":["Proszynski J. 2016. Delimitation and description of 19 new genera, a subgenus and a species of Salticidae (Araneae) of the world. Ecologica Montenegrina 7: 4 - 32. https: // doi. org / 10.37828 / em. 2016.7.1"]}

Corcovetella galianoae sp. nov. urn:lsid:zoobank.org:act: 150EC988-F9A0-439D-987B-8D13D5C1A1A0 Figs 1F–G, 4A–E, 5A–F Diagnosis Males of C. galianoae sp. nov. can be distinguished from C. aemulatrix by enlarged chelicera and a triangular and bent RTA. Females can be distinguished by an atrium wider than long and spermathecae aside of the atrium. Etymology The name is a matronym in honour of Maria Galiano, who has advanced salticid taxonomy in South America. Type material Holotype COSTA RICA – Heredia Province • ♂; Puerto Viejo de Sarapiquí, La Selva Biological Station and Reserve; 10°25.87145′ N, 84°0.45123′ W; 16 Apr. 2015; G. Corcobado leg.; litter; NMP (P6A 7399). Paratypes COSTA RICA – Heredia Province • 1 ♀; same collection data as for holotype; 2 May 2015; NMP (P6A 7400) • 1 ♂, 1 ♀; same collection data as for holotype; MUB • 1 ♀; same collection data as for preceding; 2 May 2015; MUB. Other material examined COSTA RICA – Heredia Province • 2 juvs; same collection data as for holotype; MUB • 1 juv.; same collection data as for holotype; 7 May 2015; MUB. Description Male MEASUREMENTS (N = 2). Total body 3.56 (4.6) mm long; carapace 1.60 (1.87) mm long, 0.92 (1.19) mm wide; cheliceral basal segment 0.68 (1.02) mm long, 0.35 mm wide; cheliceral fang 0.64 mm long; abdomen 1.50 (1.60) mm long, 0.89 mm wide. Diameter of eyes: AME = 0.28 mm, ALE = 0.11 mm, PME = 0.04 mm, PLE = 0.14 mm; sternum 0.69 mm long, 0.35 mm wide. Leg segments: palp (0.35, 0.18, 0.35, 0.35), I (0.94, 0.45, 0.99, 0.64, 0.40), II (0.69, 0.39, 0.69, 0.49, 0.30), III (0.74, 0.30, 0.59, 0.69, 0.30), IV (1.14, 0.35, 0.94, 0.94, 0.35). COLOUR. Chelicerae brown covered with white setae (Fig. 1F), carapace brown with cephalic part dorsally black, white setae around eyes, endites and palps brown, labium and sternum dark-brown, most leg segments with longitudinal black stripes on sides, all Cx pale, all Fe mostly black, Pt and Ti I+II pale, other black, Mt I-III pale, Mt IV black, Ta I black, other Ta pale; abdomen brown to black, dorsally and laterally with a transverse wide white stripe in the middle (Fig. 1F). MORPHOLOGY. Basal segment of chelicera enlarged, protruding anteriorly with 3 teeth on prolateral margin, 1 tooth on retrolateral margin (Fig. 5A); fangs straight, distally bent; cephalic part of carapace dorsally flat, higher than thoracic part, without constriction (Fig. 4A). Leg spines: Ti I ventrally with 2.2.2, Mt I, II, III ventrally 2.2, Ti II ventrally with 2.1. Abdomen elipsoid, with a dorsal scutum, without constriction. Palpal Pt, Ti and Ta with dense plumose setae on the ventral side. RTA triangular with bent tip (Figs 4E, 5D), cymbium with elongated apophysis (Figs 4C, 5C), bulbus oval split in the middle, embolus coiled (Figs 4D, 5B). Female MEASUREMENTS (N = 2). Total body 4.86 (4.32) mm, carapace 1.84 (1.73) mm long, 1.12 (0.95) mm wide; cheliceral basal segment 0.48 (0.54) mm long, 0.35 mm wide, cheliceral fang 0.32 mm long; sternum 0.89 mm long, 0.45 mm wide, abdomen 2.24 (2.38) mm long, 1.21 mm wide. Diameter of eyes: AME = 0.35, ALE = 0.14, PME = 0.04, PLE = 0.14. Leg segments: palp (0.39, 0.35, 0.52, 0.60), I (0.89, 0.45, 0.74, 0.59, 0.39), II (missing), III (0.99, 0.45, 0.74, 0.79, 0.34), IV (0.94, 0.50, 1.24, 1.24, 0.44). COLOUR. As in male (Fig. 1G). MORPHOLOGY. As in male but chelicerae smaller, with 5 teeth on prolateral and 1 tooth on retrolateral margin. Palpal Ti and Ta bulbous with dense plumose setae on the ventral side. Epigyne with a single oval atrium, wider than long (Figs 4B, 5E); spermathecae composed of two chambers with ducts twisted in the middle and with fertilisation ducts in front (Fig. 5F). Natural history The species was collected from litter. Both sexes appear to resemble Camponotus planatus Roger, 1863 ants (Fig. 1H). Barcode COI of this species is deposited in GenBank, accession no. OM321372. Distribution Known only from type locality in Costa Rica., Published as part of Pekár, Stano, 2022, Two new ant-mimicking spiders (Araneae: Salticidae) from Costa Rica, pp. 31-42 in European Journal of Taxonomy 852 (1) on pages 37-40, DOI: 10.5852/ejt.2022.852.2005, http://zenodo.org/record/7448120

Myrmapana costaricaensis sp. nov. urn:lsid:zoobank.org:act: 4FBB9658-87A4-4519-B9D7-CB83DDAA37AA Figs 1A–B, D, 2–3 Diagnosis Males of M. costaricaensis sp. nov. are most similar in the shape of sexual organs to M. parallela. Males of M. costaricaensis differ from this species by a protuberance at the anterior margin of the cheliceral fang, the shape of the first strong tooth, and a sinusiodal RTA (Galiano 1969: figs 33, 43). Females can be recognised by spermathecae without prominent insemination ducts (“pipes”, see Prószyński 2016). Etymology The name is derived from the country of origin. Type material Holotype COSTA RICA – Heredia Province • ♂; Puerto Viejo de Sarapiquí, La Selva Biological Station and Reserve; 10°25.87145′ N, 84°0.45123′ W; 25 Apr. 2015; G. Corcobado leg.; leaves of tropical plants; NMP (P6A 7397). Paratypes COSTA RICA – Heredia Province • 2 ♀♀; same collection data as for holotype; 15 Apr. 2015; NMP (P6A 7398) • 1 ♂; same collection data as for preceding; MUB • 3 ♂♂, 5 ♀♀; same collection data as for preceding; 16 Apr. 2015; MUB • 3 ♀♀; same collection data as for preceding; 25 Apr. 2015; MUB • 3 ♀♀; same collection data as for preceding; 7 May 2015; MUB • 2 ♂♂; same collection data as for preceding; 8 May 2015; MUB. – Puntarenas Province • 1 ♂; La Gamba; Feb. 2012; E. Líznarová leg.; MUB. Other material examined COSTA RICA – Heredia Province • 2 juvs; same collection data as for holotype; 16 Apr. 2015; MUB • 3 juvs; same collection data as for preceding; 25 Apr. 2015; MUB • 1 juv.; same collection data as for preceding; 7 May 2015; MUB • 1 juv.; same collection data as for preceding; 8 May 2015; MUB. – Puntarenas Province • 3 juvs; La Gamba; Feb. 2012; E. Líznarová leg.; MUB. Description Male MEASUREMENTS (N = 9). Total body length 9.18 (6.70–10.70) mm; carapace 2.55 (1.49–2.97) mm long, 1.52 (1.16–1.67) mm wide; basal segment of chelicera 3.43 (2.27–4.05) mm long, 0.48 mm wide, cheliceral fang 3.35 mm long; abdomen 2.71 (1.53–3.67) mm long, 1.20 mm wide; labium 0.88 mm long; sternum 1.44 mm long, 0.40 mm wide. Diameter of eyes: AME = 0.42 mm, ALE = 0.21 mm, PME = 0.07 mm, PLE = 0.25 mm. Leg segments (mm): palp (0.77, 0.39, 0.49, 0.63), I (1.99, 1.04, 2.15, 0.79, 0.40), II (1.44, 0.72, 1.11, 0.72, 0.48), III (1.36, 0.56, 0.96, 0.72, 0.48), IV (1.84, 0.64, 1.60, 1.60, 0.56). COLOUR. Chelicerae basal segment dark-brown with orange-brown distal tip, reticulated dorsally (Fig. 1A), covered with white sparse setae, fangs orange-brown, carapace dark-brown mostly covered by white setae, with brown spot in front of fovea, eyes surrounded by black patches; wedge of white setae on lateral sides behind head; sternum, endites light-brown; palpal Fe and Pt light-brown, palpal Ti and Ta dark-brown, all Cx pale, all Fe light-brown to dark-brown, all Pt pale, Ti I and II pale, Ti III and IV brown, Mt I dark-brown, all other Mt pale, all Ta pale, labium and gnathocoxae brown; abdomen dark-brown with white setae. MORPHOLOGY. Chelicerae protruding, longer than cephalothorax, with 14 teeth on prolateral margin, the first teeth strong, triangular; fangs straight, at base with dorsal protuberance, tip curved (Fig. 3A); cephalothorax elongated (Fig. 1A), cephalic part dorsally flat and slightly higher than thoracic part which is hump-shaped; sternum elongated. Leg spines: Ti I ventrally 2.2.2.2.2.2.2, Mt I ventrally 2.2, Ti II ventrally 2.1, Mt II ventrally 2.2, Mt III and IV ventrally 2. Abdomen elongated with a constriction about one third of its length and two dorsal scuta (Fig. 1A). Palpal cymbium oval, with one apical spine, tegulum large, oval as wide as cymbium, embolus coiled 2 times, tip of embolus straight, pointing laterally (Figs 2B, 3B); RTA elongated, sinusoid apically pointed (Figs 2D, 3C). Female MEASUREMENTS (N = 11). Total body 7.18 (5.40–7.34) mm, carapace 2.55 (2.48–2.96) mm long, 1.27 (1.08–1.39) mm wide, chelicera 0.63 (0.58–0.71) mm long, abdomen 2.87 (2.11–3.78) mm long, 1.20 mm wide; labium 0.49 mm long; sternum 1.27 mm long, 0.40 mm wide. Diameters of eyes: AME = 0.46 mm, ALE = 0.25 mm, PME = 0.07 mm, PLE = 0.25 mm. Leg segments (mm): palp (0.60, 0.31, 0.42, 0.60), I (1.52, 0.79, 1.60, 0.64, 0.40), II (1.12, 0.56, 1.04, 0.56, 0.48), III (1.28, 0.56, 0.96, 1.12, 0.48), IV (1.68, 0.56, 1.60, 1.52, 0.48). COLOUR. As in males but abdomen ventrally pale (Fig. 1B). MORPHOLOGY. As in males but chelicerae small, lateral margin of chelicerae with 6 teeth, retrolateral margin with 5 teeth; palpal Ta and Ti flattened (Fig. 1B, D); thoracic part of cephalothorax only slightly lower than cephalic part (Fig. 2A); abdomen with a single scutum. Epigyne with two large atria separated by a septum, atria approx. 1.5 times as wide as septum, median pocket rather triangular, situated below septum (Figs 2C, 3D); spermathecae large, oval, situated in the upper part of septum (Fig. 3E), without prominent insemination ducts. Variation The dark form is represented by both sexes (Fig. 1A–B), whereas the light form is known only in females (Fig. 1D). The two forms differ only by colouration. In the light form, palpal segments and carapace are orange-brown with dark spots around eyes, Fe I pale, frontal half of abdomen yellow, hind part dark-brown. Natural history This species was collected from tree foliage. The dark form appears to resemble Neoponera unidentata Mayr, 1862 ants (Fig. 1C), while the light form resembles Pseudomyrmex sp. ants (Fig. 1E). Barcode COI of this species is deposited in Genbank, accession no. OM321384. Distribution Known only from type locality in Costa Rica., Published as part of Pekár, Stano, 2022, Two new ant-mimicking spiders (Araneae: Salticidae) from Costa Rica, pp. 31-42 in European Journal of Taxonomy 852 (1) on pages 33-37, DOI: 10.5852/ejt.2022.852.2005, http://zenodo.org/record/7448120, {"references":["Galiano M. E. 1969. Salticidae (Araneae) formiciformes. VII. El genero Myrmarachne Mac Leay, 1839, en America. Revista del Museo Argentino de Ciencias Naturales Bernardino Rivadavia (Entomologia) 3: 107 - 148.","Proszynski J. 2016. Delimitation and description of 19 new genera, a subgenus and a species of Salticidae (Araneae) of the world. Ecologica Montenegrina 7: 4 - 32. https: // doi. org / 10.37828 / em. 2016.7.1"]}