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Developmental plasticity can alter expression of sexual signals in novel environments and is therefore thought to play an important role in promoting divergence. Sexual signals, however, are often multimodal and mate choice multivariate. Hence, to understand how developmental plasticity can facilitate divergence, we must assess plasticity across signal components and its cumulative impact on signalling. Here, we examine how developmental plasticity influences different components of cabbage white butterfly Pieris rapae multimodal signals, its effects on their signalling phenotypes and its implications for divergence. To do this, we reared P. rapae caterpillars under two different light environments (low-light and high-light) to simulate conditions experienced by P. rapae colonizing a novel light habitat. We then examined plasticity in both visual (wing coloration) and olfactory (pheromone abundance) components of male sexual signals. We found light environments influenced expression of both visual and olfactory components and resulted in a trade-off between signal modalities. The ‘low-light’ phenotype had duller wing colours but higher abundance of pheromone-indole, whereas the ‘high-light’ phenotype had comparatively brighter wings but lower abundance of indole. These results show that by simultaneously altering expression of different signal components, developmental plasticity can produce multiple signalling phenotypes, which may catalyse divergence.

Interspecific competition can occur when species are unable to distinguish between conspecific and heterospecific mates or competitors when they occur in sympatry. Selection in response to interspecific competition can lead to shifts in signalling traits—a process termed as agonistic character displacement. In two fan-throated lizard species—Sitana laticeps and Sarada darwini—females are morphologically indistinguishable and male agonistic signalling behaviour is similar. Consequently, in areas where these species overlap, males engage in interspecific aggressive interactions. To test whether interspecific male aggression between Si. laticeps and Sa. darwini results in agonistic character displacement, we quantified species recognition and signalling behaviour using staged encounter assays with both conspecifics and heterospecifics across sympatric and allopatric populations of both species. We found an asymmetric pattern, wherein males of Si. laticeps but not Sa. darwini showed differences in competitor recognition and agonistic signalling traits (morphology and behaviour) in sympatry compared to allopatry. This asymmetric shift in traits is likely due to differences in competitive abilities between species and can minimize competitive interactions in zones of sympatry. Overall, our results support agonistic character displacement, and highlight the role of asymmetric interspecific competition in driving shifts in social signals.

Chiromachetes sahyadriensis Mirza, Sanap et Zambre, sp. nov. (Figs. 1–8, Table 1) http://zoobank.org/urn:lsid:zoobank.org:act:D872BC FA-90E6-4F28-85DA-6AA162ACD8D4 Holotype: male NCBS AG-873, INDIA, Maharashtra State: Pune District, Tamhini Ghat, 18.764720°N, 73. 367294°E, 558 m a.s.l., under boulders along a dry stream bed, 8 November 2013, coll. Rajesh Sanap, Varun Vaze, Devavrat Joshi and Zeeshan Mirza. Paratypes: 2 males (NCBS AG-876–877) and 3 females (NCBS AG-872, 874, 875), same label as holotype. Diagnosis: Medium-sized scorpions, total length reaching 62.9 mm, carapace flat, three pairs of lateral eyes, manus length/width in males 4.6–4.8 and 3.4 in females; leg tarsi with three very small spinoid setae at the base (Fig. 3D); the ventral tarsal spines in males are stout, unlike those in other related genera. Dentate mar- gin of pedipalp chela finger with two rows of granules fused at the base (Fig. 3C). Pectinal teeth number 8–9 in males, 6–8 in females. Lamellar hook positioned distinctly in basal half of hemispermatophore in proximity to the lamellae. Etymology: The specific epithet refers to the Sahyadri hills synonymous to the Western Ghats where the type locality is situated. Description of holotype male NCBS AG873 (Figs. 1A–B, Table 1): Coloration (when alive) (Figs. 6 & 7): Carapace, mesosoma, and metasoma deep brown to black with a beady gloss. Edges of each segment darker than the central portions. Legs, vesicle, pectines, genital operculum yellow to tan. Carapace: Brown throughout with dark blackish brown reticulations throughout; anterior border, region near lateral eyes, and median eyes almost black (Fig. 2A). Smooth with a beady gloss with the exception of a few small sparse granules overall. Anterior margin with a moderately deep U-shaped indentation. Carapace horizontal, median eyes not on elevated tubercle. Lateral ocular tubercle with three eyes, first and the second lateral eyes are placed at a distance from the third lateral eye (Fig. 3B). Carapace lacks carinae. Lateral margins of carapace parallel posteriorly for more than half of its length. Cheliceral fixed finger with median and a basal bifid denticle; cheliceral movable finger dorsal edge with four denticles: distal denticle, one subdistal, one median, and one basal (Fig. 3A). Mesosoma: Tergites tan-brown overall with dark reticulations; sternites paler, almost tan except for the last sternite, which is darker. Sternum, genital operculum and pectines pale yellow; legs tan with a brownish tinge. All segments smooth, lacking carinae. Each segment medially with a longitudinal elevation. Sternum pentagonal with a median longitudinal furrow. Genital operculum sub-pentagonal; basal piece with a median furrow (Fig. 2B). Pectines well developed, their teeth 8/ 9 in number, with fulcra; lamellae indistinct. Legs: Two apical spinoid setae and one median spinoid seta present on tarsi of all legs; the ventral tarsal spines are stout, unlike those in other related genera; tarsi with three very small spinules at the base (Fig. 3D). Pedipalp: Manus with five carinae, patella and femur each with four carinae. Patella and femur smooth throughout, with beady gloss, manus lacking beady smooth gloss but smooth throughout. Prolateral aspect of patella with a large spine-like protrusion along with a few blunt spines along the anterior ridge. Trichobothrial pattern of type C (Figs. 5). Trichobothrium Db present on the external aspect of manus and not on dorsal aspect; dt positioned on the internal aspect of the fixed finger; patellar trichobothrium est located on ventral side of the external surface. Metasoma: smooth throughout, lacking granulation and carinae on segments I–IV. Ventrolateral carinae on segments II–IV present merely as depressed ridges. Segment V with ventrolateral carinae present, composed of rounded and spine-like granules. Anal arch with sparsely placed spine-like tubercles. Each segment dorsally with a shallow median furrow. Ventrally with sensory pits as in the genus Liocheles. All segments laterally compressed. Telson bulbous, hirsute ventrally. Aculeus short, nearly one-fourth the length of vesicle. Hemispermatophore: lamelliform with a complex capsule; distal lamella slender and long. Lamina length twice that of the capsule region and trunk. A single lamellar hook arising from above the transverse ridge; hook short, curved and pointing upwards. Lamella arising from the posterior lobe is clearly apart from the lamellar hook, vertical in orientation. Lamellar hook positioned distinctly in basal half of hemispermatophore in proximity to the lamellae. Variation. The species exhibits a high degree of sexual dimorphism, especially in the length and robustness of the pedipalp manus. Manus length/width in males 4.6– 4.8 and 3.4 in females. Females are darker in coloration as opposed to the males, and the leg coloration in males is much lighter. Tarsal spines in males are thick and long whereas they are short and thin in females. For other details see Table 1. Ecological Notes The new species is presently known only from the type locality where it occurs under boulders along forest streams (Fig. 8). The type locality lies close to the Tamhini Wildlife Sanctuary, which supports similar habitats and it is likely that the new species is also distributed in the adjoining protected areas. The type specimens were collected within an hour at around 2200 hours with the aid of a flashlight. Most individuals were found sitting at the entrance of small slit-like burrows under boulders with their pedipalp chelae exposed. Several of these burrows had remains of conspecific pedipalps around the entrance indicating that the species might be highly cannibalistic. This is further attested by the fact that specimens maintained in captivity preferentially fed on smaller individuals of the same species leaving pieces of pedipalps (Devavrat Joshi, pers. comm.).

Orthochirus bastawadei Zambre, Mirza, Sanap, Upadhye et Javed, sp. nov. (Figs. 3���17) Type material: Holotype female, INDIA: near Maharashtra Industrial Development Corporation (MIDC) area, Jalna District, Maharashtra, India, 23 November 2009, coll. Durgesh Pangarkar (BNHS SC- 51). Paratypes: 2 males (BNHS SC- 52 and BNHS SC- 53), same data as above. Etymology. Patronym in honor of Dr. Deshbhushan Bastawade, India for his immense contribution to the field of Indian scorpiology. Diagnosis. A species of moderate size in relation to other species of the genus; total length 36.54 mm in the female and 31.72 mm in male. General coloration dark coffee-brown to black; fingers and legs clear yellow with black pigmentation (Figures 3���6). Pedipalps with 9 rows of denticles on the fixed and movable fingers; external accessory denticles moderate. Trichobothriotaxy A-�� (beta), orthobothriotaxic. Distance between trichobothrium d 1 and d 3 less than distance between d 3 and d 4 ; trichobothrium e 1 in line with d 3 (Fig. 17). Complete lack of bristlecombs on legs (Figure 16). Telson almost glabrous. Metasomal segments IV and V without median carinae. Spaces among punctae on ventral surface of metasomal segments IV and V smooth. Dorsal surface of metasomal segment IV and V mesially densely granulated. Pectines six times longer than wide, with 18���19 teeth in females, 20��� 20 in males (Figures 9, 11 & 13). Orthochirus bastawadei sp. nov. can be distinguished from the mainland Indian species of this genus on the basis of the following character states: mesosoma and metasoma entirely blackish (metasomal segments are yellowish-brown in O. flavescens and O. pallidus; metasomal segments I���II are yellow and the rest black in O. bicolor); dorsal surface of metasomal segments mesially with granulation arranged in a distinct stripe (nearly smooth or with sparse granulation not arranged in a distinct stripe in O. pallidus); bristlecombs on legs I���III absent (bristlecombs on legs I���III present in O. fuscipes , O. bicolor, O. pallidus, O. krishnai). Pectinal teeth 18���19 (female) and 20-20 (male) in number [23-23 (male) and 22-22 (female) in O. krishnai; 16-16 (female) in O. pallidus]. For more details, see Table 2. Description based on female holotype BNHS SC - 51: Measurements in Table 1. Coloration (in alcohol). Carapace, mesosoma and metasoma black; telson reddish black with black aculeus; pedipalps dark brown to black, fingers yellow. Legs are pale yellowish, with both femur and patella mostly blackish. Sternites dark brown while presternal region brownish yellow. Coloration in life (Fig. 3���5). Carapace and mesosomal entirely black; metasoma blackish brown with a beady gloss; pedipalp femur, patella and manus coffee-brown, manus with dark reticulated markings; movable and immovable fingers pale. Vesicle reddish brown. Prosoma (Fig. 8, 10 & 12). Carapace densely and evenly granulated; interocular region granulated; median ocular tubercle very weakly granulated, without prominent carinae. Anterior margin straight and granulated with few short red setae. Lateral margin composed of minutely crenulate granules. Posterior margin granulated, composed of unevenly sized granules. Lat-eral ocular tubercle with five eyes of which four (three large and one small) are contiguous. Mesosoma (Fig. 6 & 14). Tergites I���VII with a median T-shaped carina; segment III���VII with poorly developed lateral carinae. Tergite VII pentacarinate, both pairs of lateral carinae moderate; median carinae present on the anterior half, weakly marked. Intercarinal spaces granulated. Sternites III���VI without carinae; sternite VII granulated on posterior and lateral margins, with two pairs of moderate carinae. Pectinal tooth count 18���19. Metasoma (Fig. 6, 7 & 15). The segment I with 8 granulated carinae, lateral carinae smooth. Segment II bears six carinae while segment III bears five carinae. Segment IV with a pair of weak dorsal carinae. Segment V with weak dorsal carinae and two incomplete dorsolateral carinae only in the posterior half, composed of large denticles. Dorsal surface of metasomal segments I���III with mesial granulation arranged in a distinct stripe. All segments punctate, intercarinal regions on segments II���III granulated. Punctation well developed on all segments except segment I, which is comparatively weakly punctate. Spaces among punctations smooth. Dorsal surface of all segments is mesially distinctly granulated. The entire metasoma and telson almost glabrous expect for a few red setae on the telson and dorsal ridge of the segments III���V. The telson is punctate and lacks granules. Chelicerae. Basal piece yellowish brown with black ornamentation, anterior margin dark black. Dentition as characterised in the family and genus. A few short silky hairs present. Pedipalp (Fig. 17). Trichobothrial pattern orthobothriotaxic, Type A (Vachon 1974). Dorsal trichobothria on femur in �� (beta) configuration (Vachon, 1975). Femur pentacarinate, all carinae crenulate. Patella with 7 smooth carinae. Dentate margins on fixed and movable fingers with 9 imbricate rows of granules; external accessory granule moderate. Legs (Fig. 16). The femur conspicuously granular. Tarsi and basitarsi I and II furnished with setae. Bristlecombs absent on all legs. The inner sides of legs I���III bear rows of spines. Variation. Two male (Fig. 10) paratypes are similar to female holotype but differ in the following set of characters: (a) a slightly higher pectinal tooth count of 20���20 (Figure 11); (b) mesosoma with rudimentary lateral carinae on segments II��� VI; (c) carinae on metasomal segments I��� III on metasoma weakly developed as compared to female (Figure 10); (d) a comparatively smaller size, 25.22���31.72 mm. Distribution and natural history (Fig. 1 & 2). The type locality, MIDC area in the Jalna District of Maharashtra, is characterized with open area with a few trees sparsely dotting the landscape. The new species was collected under boulders. The new species is common throughout the district in similar ecological conditions with other sympatric species, namely Heterometrus xanthopus (Scorpionidae) and Hottentotta tamulus (Buthidae). The species may yet be found in adjoining districts which share similar habitats. Specimens observed in the wild and in captivity would remain stationary in a position with their tails curled and pressed down on their mesosoma as observed in O. scrobiculosus by E. Fet et al. (2003) as well as other Orthochirus species in India (Zambre, pers. obs.) and Buthoscorpio rayalensis (see Javed et al., 2010). While collecting the types, one of us (RU) got stung by one of the specimens. Pain was agonizing and lasted for at least five hours and was much more painful compared to the sting of Hottentotta tamulus. Additionally, numbness on the stung finger was much more developed than in the sting of H. tamulus, and lack of sensation persisted for a considerable period of time. Taking into account the severity of the envenomation, it would be imperative to consider this species to be medically important., Published as part of Zambre, Amod M., Mirza, Zeeshan A., Sanap, Rajesh V., Upadhye, Raman & Maqsood Javed, S. M., 2011, A new species of Orthochirus Karsch, 1892 (Scorpiones: Buthidae) from Maharashtra, India, pp. 1-12 in Euscorpius 107 (107) on pages 2-6, DOI: 10.18590/euscorpius.2011.vol2011.iss107.1, http://zenodo.org/record/4649109, {"references":["VACHON, M. 1974. Etude des caracteres utilises pour classer les familles et les genres de Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et types de trichobothriotaxie chez les Scorpions. Bulletin du Museum National d'Histoire Naturelle, Paris, 140: 857 - 958.","VACHON, M. 1975. Sur l'utilisation de la trichobothriotaxie du bras des pedipalps des Scorpions (Arachnides) dans le classement des genres de la famille des Buthidae Simon. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, Paris, ser. D, 281: 1597 - 1599.","FET, E. V., D. NEFF, M. R. GRAHAM & V. FET. 2003. Metasoma of Orthochirus (Scorpiones: Buthidae): are scorpions evolving a new sensory organ? Revista Iberica de Aracnologia, 8: 69 - 72.","JAVED, S. M. M., K. THULSI RAO, Z. A. MIRZA, R. V. SANAP & F. TAMPAL. 2010. A new species of scorpion of the genus Buthoscorpio Werner, 1936 (Scorpiones: Buthidae) from Andhra Pradesh, India. Euscorpius, 98: 1 - 11."]}

Orthochirus Karsch, 1892 Orthochirus: Vachon, 1974: 910, 936; Tikader & Bastawade, 1983: 113; Sissom, 1990: 102; Kovař��k, 1996: 177; Fet & Lowe, 2000: 193; Fet E. et al., 2003: 69; Kovař��k, 2004: 4. Type species: Orthodactylus olivaceus Karsch, 1881 = Orthochirus scrobiculosus (Grube, 1873). Diagnosis: Dorsal trichobothria of femur arranged in beta-configuration. Tibial spurs present on legs III and IV. Movable fingers of pedipalps with 7���10 rows of granules and 2���5 distal granules. Carapace, in lateral view, distinctly inclined downward from median eyes to anterior margin. Metasomal segments I to III with carinae. Metasomal segments IV and V ventrally punctate., Published as part of Zambre, Amod M., Mirza, Zeeshan A., Sanap, Rajesh V., Upadhye, Raman & Maqsood Javed, S. M., 2011, A new species of Orthochirus Karsch, 1892 (Scorpiones: Buthidae) from Maharashtra, India, pp. 1-12 in Euscorpius 107 (107) on pages 1-2, DOI: 10.18590/euscorpius.2011.vol2011.iss107.1, http://zenodo.org/record/4649109, {"references":["VACHON, M. 1974. Etude des caracteres utilises pour classer les familles et les genres de Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et types de trichobothriotaxie chez les Scorpions. Bulletin du Museum National d'Histoire Naturelle, Paris, 140: 857 - 958.","SISSOM, W. D. 1990. Systematics, biogeography and paleontology. Pp. 64 - 160 in Polis, G. A. (ed.): The Biology of Scorpions. Stanford, California: Stanford University Press.","KOVARIK, F. 1996. Baloorthochirus becvari gen. et sp. n. from Pakistan, and taxonomic position of Orthochirus luteipes (Scorpiones: Buthidae). Acta Societatis Zoologicae Bohemicae, 60: 177 - 181.","KOVARIK, F. 2004. Revision and taxonomic position of genera Afghanorthochirus Lourenco & Vachon, Baloorthochirus Kovarik, Butheolus Simon, Nanobuthus Pocock, Orthochiroides Kovarik, Pakistanorthochirus Lourenco, and Asian Orthochirus Karsch, with descriptions of twelve new species (Scorpiones, Buthidae). Euscorpius, 16: 1 - 33."]}