Revue Suisse de Zoologie. vol. hors serie: 515-522; aout 1996 Precopulatory male ethograms ofthree species ofLycosa Latreille 1804 (Araneae: Lycosidae) ofthe Iberian peninsula Juan Manuel ORTA-OCANA. Jordi MOYA-LARANO & Jose A. BARRIENTOS Unitat de Zoologia. Departament de BiologiaAnimal, Biologia Vegetal Ecologia. i UniversitatAutonoma de Barcelona. E-08193 Bellaterra (Barcelona), Spain. Precopulatory male ethograms of three species of Lycosa Latreille, 1804 (Araneae: Lycosidae) of the Iberian peninsula. - In order to revise the representatives of the genus Lycosa from the Iberian Peninsula, the courtship behaviour of the males of the three morphotypes was analysed from a total of46 male-female interactions. The three ethograms presented here are subdivided into four parts: locomotory behaviours, foreleg move- ments and postures, palpal movements and postures and otherbehaviours. From the qualitative analysis of the repertoires, several differences in movements and postures can be seen, so we can conclude that the three morphotypes can be considered as three different species. While L. radiata shows extensions with vibration, L. tarentula fasciiventris shows static extensions and arches. There also are differences in the palpal movements. Lycosa sp. shows exclusive behaviours such as flexions, foreleg balance and sudden advances, as well as behaviours also seen in L. radiata and L. tarentulafasciiventris. Key-words: Lycosa tarentulafasciiventris - Lycosa radiata - Lycosa sp. - ethogram- courtship - isolation mechanisms - biospecies. INTRODUCTION This study is part ofa project aiming to revise taxonomically the genus Lycosa in the Iberian Peninsula using behavioural and ecological criteria, as well as the classical morphological ones. The representatives of the genus Lycosa in the Iberian Peninsula are Lycosa radiata Latreille. 1817 and L. tarentulafasciiventris Dufour. 1835. Nevertheless, new morphotypes that do not fit into either of these two species have been found. The genus Lycosa. which names the family Lycosidae, comprises 1 16 species distributed over the hot and temperate regions of the World. Nearly 20 nominal species have been recorded from the Iberian Peninsula, although their biological Manuscriptaccepted 08.12.1995. Proceedings ofthe Xlllth International CongressofArachnology, Geneva, 3-8.IX.1995. 516 JUAN MANUELORTA-OCANA, JORDI MOYA-LARANO & JOSEA. BARRIENTOS entity is questionable. Following Barrientos (1981), we can conclude that only two ofthem can be considered as biological species: Lycosa radiata Latreille, 1817 and L. tarentulafasciiventris Dufour, 1835. Nevertheless, new morphotypes (Lycosa sp.) have been found that do not fit in any of the accepted forms. So, a revision of the genus is necessary, bearing in mind the species concept in a wider point of view than the one given by taxonomy on its own. The biological species concept requires the existence of isolation mechanisms to guarantee the existence of separated species. The ethological barriers may be regarded as the most important isolating mechanisms (Mayr 1970). Lycosidae is a morphologically homogeneous group and due to the fact that the species we are studying are syntopic, the ethological mechanisms might explain the speciation pro- cesses in the group. Ethological barriers are due to behavioural incompatibilities. The male of one species performs some courtship patterns that are recognised just by the adequate female. These mechanisms have been observed in a vast group of insects and spiders. The validity of these ethological patterns as isolation mechanisms has already been proved in other Lycosa species (Costa & Caposacale 1984; Costa & Francescoli 1991), as well as in many other lycosid species (Uetz & Stratton (1982), Schizo- cosa spp.) and otherfamilies Vlijm 1986). ( MATERIAL AND METHODS The spiders were collected asjuveniles from different populations in the Bar- celona area and kept in isolated cages. After becoming adults, male-female pairs were chosen at random and the female put into the observation cage 24h prior the intro- duction of the corresponding male. Observation cages were made of transparent plastic and consist in two adjacent areas of 15 x 15 cm. separated by a removable opaque barrier. This barrier was removed at the beginning of the experiment. The interactions were recorded on video tape for further analyses. After 22.5 hours of observation, we constructed three ethograms of the dif- ferent movements and postures of the three forms. Some of the terms used in the ethograms were taken from the literature (Nossek & Rovner 1984; Aspey 1977). Within those 22.5 hours, 26 male-female interactions were recorded for L. radiata (409 min), 13 interactions for L. tarentula fasciiventris (502 min) and 17 for Lycosa sp. (440 min). RESULTS Due to the fact that there are a lot of details that can be recorded in one interaction, we have divided the ethogram in different subrepertoires. to allow the independence of the categories in each subrepertoire. The categories can overlap between subrepertoires, but not in the same subrepertoire. So, different observers could record different subrepertoires or the same observer could record different PRECOPULATORY MALE ETHOGRAMS OFTHREE SPECIES OFLYCOSA LATREILLE 5]7 subrepertoires in different video reviewing sessions. At the end of the sessions, it would be possible to obtain a multiple behaviour chart. We have only elaborated the male's premating ethogram, although we are aware of the relevant importance of the female's behaviour, so ethograms should be elaborated forthe latter. LOCOMOTORY BEHAVIOURS 1. Common behaviours 1.1. Stationary: the male remains motionless, with the four pairs of legs in contact with the substrate. 1.2. Approach: the male advances so it reduces the distance between itself and the female. 1.3. Retreat: the male moves away so it increases the distance between itself and the other one. It is assumed that the other spider is stationary. It is possible that the female goes after the male, so there is no actual increase in the distance between them. 1.4. Orientation: the male turns around an imaginary vertical axe so as to end up facing the female. 1.5. Side movement: the male moves towards its right or left side, without dis- placement forward or backwards. 1.6. Step-Wave: hyperextension of one or typically both legs I and II during for- ward motion. At the top ofthe raise, the femora are held at an angle of40-60° relative to the substrate. Forward motion continues as the forelegs are lowered to the substrate and again hyperextended and raised. 1.7. Random activity: running without an specific orientation. L. tarentulafasciiventris exclusive behaviours 1.8. Side movement around the burrow: similar to the previous category ( 1.5). but with the burrow as the centre ofthe side movement. Lycosa sp. exclusive behaviours 1.9. Fast approach: the spider advances very quickly reducing the distance between itselfand the female.. 2. Foreleg movements and postures Common behaviours 2.1. Stationary: the leg is in contact with the substrate. 2.2. Horizontal extend (left/right/both): the extended leg is raised and held parallel to the substrate, (fig. 1. a) (not in L. tarentulafasciiventris). 518 JUAN MANUELORTA-OCANA, JORDI MOYA-LARANO& JOSE A. BARRIENTOS 2.3. Oblique extend (left/right/both): the leg is extended straight, the femur is held at an angle of45-60° relative to the substrate, (fig. L. b) 2.4. Vertical extend (left/right/both): the leg is extended straight and the femur is held at an angle of60-90° relative to the substrate, (fig. 1. c) L. radiata exclusive behaviours In categories 2.2 to 2.4, it is common to observe some vibration of the legs raised. This vibration is not quantified in the ethogram. 2.5. Tapping (left/right/both): the leg is raised straight about 30-40° relative to the substrate and subsequently lowered to contact the substrate, as if it were probing the substrate. It is similar to 2.2 but here the tip ofthe leg touches the substrate or silk threads (laid by the female), (fig. 1. d) Fig. 1 Foreleg movements and postures: Extends, a.- Horizontal extend (2.2); b.- Oblique extend (2.3); c.- Vertical extend (2.4); d.- Tapping (L. radiata exclusive behaviour) (2.5). Exclusive foreleg movements and postures, e.- Vertical arch (2.6); f.- Obtuse arch (2.7); g.- Arch balancing (2.8). Lycosa sp. exclusive foreleg movements and postures, h.- Oblique flex (2.9);i.- Vertical flex (2.10);j.- Obtuse tlex (2.1 1); k.- Foreleg balance (2.12); I.- Metatarsal oscillation (2.14). Numbers in brackets refertothecategory numberintext. PRECOPULATORY MALEETHOGRAMS OFTHREE SPECIES OFLYCOSA LATREILLE 519 ||| 1 | Lycosa Lycosa Lycosa Lycosa Lycosa sp. form sp. form nuliata / nuliata / tarentula behaviour behaviour fasciiventris - number number j IJ 3.1 Stationary Stationary 1.1 Palpal 3.2 Palpal drumming 1.2 Approach 3.3 Silk touch 1.3 Retreat Locomotory ^.4^- P1'UlllLi'vcilll (1I1n1Mim11. on11 ihi1 1rtuilnl movements 1.4 Orientation I.J OIt-lC lUUVClllGUL 2.1 Stationary 1.6 Step-wave behaviours 2.2 Horizontal extend 1.7 Randomactivity 2.3 Obliqueextend 1.8 Side mov. burrow 2.4 Vertical extend 1.9 Fast approach Foreleg 2.5 Tapping 2.6 Vertical arch 2.7 Obtuse arch movements 2.8 Arch balancing and 2.9 Oblique flex 2.10 Vertical (lex 4.1 Contact postures Other 2.1 1 Obtuse Ilex 4.2 Mount 2.12 Foreleg balancing 4.3 Retreat 2.13 Sudden advance 4.4 Mating position behaviours 2.14 Metatarsal oscillation 4.5 Female'sburrow entry ) ) 520 JUAN MANUELORTA-OCANA, JORDI MOYA-LARANO & JOSE A. BARRIENTOS L. tarentulafasciiventris exclusive behaviours This species doesn't show horizontal extends (category 2.2) Opposite to L. radiata, vibration is not observed in categories 2.3 and 2.4. 2.6. Vertical arch (left/right/both): the leg I femur is raised at 60-90° relative to the substrate, and the femoro-patellarjoint is flexed, as well as the tibio-metatarsal joint, although this one is less flexed, (fig. I. e) 2.7. Obtuse arch (left/right/both): the leg I femur is pointing posteriorly at 95-140°. The femoro-patellar joint is flexed, as well as the tibio-metatarsal joint. This category is usually seen when the male is at the female's burrow, (fig. 1. f) 2.8. Arch balancing (left/right/both): the leg is raised in one of the arch categories while is slowly moving up and downwaixis at the tibio-metatarsal joint. The movement is slow, it is not a vibration movement, (fig. 1. g) In categories 2.3, 2.4 and 2.6 to 2.8, the most usual is that the spider moves both forelegs. Lycosa sp. exclusive behaviours In categories 2.2 to 2.4 is not usual to observe vibration. Ifit occurs, it is short, on the contrary as it happens in L. radiata. Illustrations are the same as forL. radiata. 2.9. Oblique flex (left/right/both): the leg I femur is raised and held at 45-60° rela- tive to the substrate. The femoro-patellarjoint is flexed, as well as the previous joints, while the rest ofthe leg segments remains parallel to the substrate, (fig. l.h) 2.10. Vertical flex (left/right/both): the leg I femur is raised and held at 60-90° relative to the substrate. The femoro-patellar joint is flexed, as well as the previous joints, while the rest of the leg segments remain parallel to the substrate, (fig. 1. i 2.11. Obtuse flex (left/right/both): the leg I femur is raised and held pointing pos- teriorly at 95-140° relative to the substrate. The femoro-patellarjoint is flexed, as well as the previous joints, while the rest of the leg segments remains parallel or in an acute angle to the substrate, (fig. 1.j 2.12. Foreleg balancing: While one leg is being flexed or extended at the femoro- patellarjoint, the other may l'emain motionless or moving in an opposite way (e.g.: one leg is being extended and the otherflexed), (fig. 1. k) 2.13. Sudden advance: while the forelegs are extended or flexed, the spider suddenly jumps forward, while legs II (right/left/both) are raised briefly. A brief vibration oflegs is often observed prior to thejump. 2.14. Metatarsal oscillation (right/left/both): a slight oscillation at the tibia-metatarsal joint is performed. It rarely occurs, (fig. 1.1) PRECOPULATORY MALE ETHOGRAMS OFTHREESPECIES OFLYCOSA LATRE1LLE 521 3. Palpal movements and postures Common behaviours 3.1. Stationary: the palps remain motionless, more or less perpendicular to the substrate. 3.2. Palpal drumming: palps alternately lifted and lowered in rapid succession, usually contacting the substrate. L. radiata exclusive behaviours 3.3. Silk touch (left/right/both): the palps touch a silk thread. L. tarentulafasciiventris exclusive behaviours 3.4. Palpal drumming on the rim: the palpal drumming is performed on the rim surrounding the female's burrow. 4. Other behaviours Common behaviours 4.1. Contact: one spider's foreleg (usually the male'.-,) contacts some part of the body ofthe other spider. 4.2. Mount: the male climbs over the female's body but does not adopt the mating position (4.4.). 4.3. Retreat: the male moves slightly backwards. 4.4. Mating position: the male places itself in the mating position (type 3, Foelix 1982), although actual copulation may not take place. L. tarentulafasciiventris exclusive behaviours 4.5. Female's burrow entry: the male gets into the female's burrow. DISCUSSION In this qualitative study ofcourtship, we can see several differences among the species. L. radiata and L. tarentulafasciiventris show clearly different categories in the subrepertoire of leg movements and postures. While L. radiata shows extensions with vibration, L. tarentulafasciiventris shows static extensions and arches. There are also differences in the palpal movements repertoire, probably associated with other communication channels as acoustic (Uetz & Stratton 1982; Stratton & Uetz 1983) or chemical communication (Tietjen & Rovner 1980, 1982). On the other . . 522 JUAN MANUELORTA-OCANA, JORDI MOYA-LARANO & JOSE A. BARRIENTOS hand. Lycosa sp. shows behaviours seen in L. radiata and L. tarentula fasciiventris (extensions and arch balancing), but also exclusive behaviours as flexions, foreleg balancing and sudden advance. The absence/presence ofbehaviours is summarised in table I. We can conclude, from a qualitative point of view, that we are dealing with a different species. A comprehensive study of the behaviour's variation is needed, as well as carrying out reproductive isolation experiments. Since sequential analyses have not been performed yet, it would be too speculative to state which of the behaviours observed may be the important in the species recognition process, this is, which one transports the signals to the female. Moreover, part of the courtship takes place via non-visual channels that haven't been analysed in the present work. Acknowledgements We want to thank to Prof. P.J.B. Slater the review ofthe ethograms. REFERENCES Aspey, W.P. 1977. Wolf Spider Sociobiology: I. Agonistic Display and Dominance subordi- nancerelations in adult male Schizocosacrassipes. Behaviour62:103-141 Barrientos, J.A. 1981. Discussion preliminaire du genre Lycosa Latr., 1804 dans la Peninsule Iberique. MemoriedellaSocieta toscanadiscienzenaturali, ser. 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