Abstract
Here we describe the hitherto unknown details of the highly unusual mating behaviour of Saltella sphondylii—a widely cited model for male longevity costs caused by multiple copulations. When compared to the known mating behaviour of 28 sepsid species, we find five unique behavioural elements based on frame-by-frame analyses of video-recordings. These new behaviours are documented with video clips. We suggest that the male longevity costs could be due to copulation bouts that involve multiple insertions of a comparatively membranous phallus into the female. We compare the phallus of the Saltella sphondylii to those from three other species (Themira putris, Parapaleosepsis plebeia, Sepsis punctum).
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Acknowledgements
We would like to thank all members of the Evolutionary Biology Lab, especially Martin Chew for his assistance in generating the molecular phylogeny, and Nalini Puniamoorthy for her helpful comments for the manuscript. This study was financially supported by grant R154-000-476-112 from the Ministry of Education in Singapore.
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Appendix
Appendix
Character descriptions modified from Puniamoorthy et al. (2009)
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(1)
‘Circling’—0: Absent (Male approach without gliding motion); 1: Present (Male circling female in a gliding motion, head and thorax leading the change of direction with abdomen bent at an angle)
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(2)
Initial mount—0: Male jumps or climbs onto female; 1: Male bends abdomen anteriad to establish direct genital contact
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(3)
Effect of struggle on in-copula position of pair—0: Females do not flip over; 1: Pair flipping over resulting in male on his back and female with her legs in air)
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(4)
Male proboscis–female interaction—0: No contact between male proboscis and female; 1: Male extends proboscis to ‘kiss’ female ocelli; 2: Male extends proboscis to tap dorsal part of female thorax
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(5)
Male grasp of female wingbase—0: Forelegs resting on female thorax; 1: Forelegs release wingbase only at separation; 2: Forelegs release female wings well before separation
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(6)
Male foreleg after release—0: Resting against female thorax (no movement); 1: Male foreleg interacting with female thorax and, or foreleg
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(7)
Midleg position—0: Male midlegs always in contact with female; 1: Male mid legs are stretched out away from female body i.e. ‘balancing’ (for extended period); 2: Male midlegs are not stretched out and are not in contact with female
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(8)
Motion restricted to mid tarsus—0: No independent tarsal movements; 1: Curling (movement of the tarsi 2–4 against the barsitarsus); 2: Quiver (vibration of entire tarsus without movement of tibia or femur)
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(9)
Non-contact midleg movement—0: Simultaneous usage of both midlegs; 1: Simultaneous and alternate; 2: Alternate usage of both midlegs
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(10)
Direction of midleg movement—0: Male midleg stretched out and stationary; 1: Male midleg towards female eye; 2: Male midlegs move posteriad
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(11)
Male midleg movement away from female head—0: Smooth return without any interruptions; 1: Return interrupted by midleg waves
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(12)
Midleg rotation during tarsal curl—0: Midleg curling in a horizontal plane; 1: Curling direction shifting from horizontal to vertical plane through leg rotation; 2: Curling in a vertical plane
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(13)
Number of tarsal curls per midleg movement—0: Single curl per midleg movement; 1: Multiple curls
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(14)
Midleg interaction with female head—0: No with contact female head; 1: Male uses his mid tarsi to rub head; 2: Male uses his mid tarsi to tap head (singular movements)
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(15)
Midleg interaction with female abdomen—0: No with contact female abdomen; 1: Male uses his midlegs to tap female abdomen
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(16)
Midleg interaction with female thorax—0: Absent; 1: Midlegs tap lateral surface (singular movements); 2: Midlegs ‘stroke’ lateral surface (extended rubbing); 3: Midlegs ‘stroke’ dorsal surface (extended rubbing)
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(17)
Female wings bent down by male midlegs—0: Male midlegs rest on female wings; 1: Male midlegs used to forcibly bend down female wings
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(18)
Midleg to midleg grasp—0: Male midlegs not holding female mid legs; 1: Male uses midlegs to hold onto female midlegs
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(19)
Contact of male hindleg with substrate—0: Hindleg not in contact with substrate when female moves; 1: Mounted male’s hindleg is dragged along substrate when female moves; 2: Mounted male walks in tandem with female instead of being dragged
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(20)
Non-contact movement of male hindlegs—0: Absent; 1: Hindlegs curl backwards 360° like a backward ‘butterfly stroke’; 2: ‘Cycling’ ( i.e. like a peddling motion); 3: Curling (Hindlegs stretched out and vibration of the 2–4 tarsal segments against the barsitarsus)
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(21)
Usage of hindlegs—0: No direct contact with female; 1: Tap ventral side of female abdomen; 2: Rub repeatedly on dorsal side of female wing; 3: Rub repeatedly wing margin
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(22)
Midleg–hindleg rub—0: No contact between male mid and hindlegs; 1: Males rub their hindlegs with their midlegs
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(23)
Part of female body male rubs after rubbing his hindlegs—0: Wing; 1: Thorax; 2: Head; 3: Abdomen; 4: Midlegs
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(24)
Movement of male abdomen—0: Abdomen maintained horizontally without any movement; 1: Abdomen lifted >90° over the thorax when mounted on female; 2: Male shakes abdomen vigorously from side to side (prior to mounting)
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(25)
Surstylus stimulation prior to genital contact—0: Males only lower abdomen to establish genital contact; 1: Male repeatedly lowers abdomen to stimulate female using the surstylis, either by vibration or by tapping on dorsal surface of female abdomen; 2: Male repeatedly lowers surstylus to stimulate female close to her genital opening; 3: Male lowers surstylus to stimulate on ventral surface of female abdomen
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(26)
Male tapping female with modified fourth sternites—0: Modified 4th sternites of males used to tap or stroke dorsal part of female abdomen; 1: Ventral part of female abdomen
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(27)
Separation after copulation—0: Quick (one or two 180° turns by the male); 1: Long (involving a prolonged struggle between male and female in trying to break genital contact); 2: Quick but not involving turns by the male
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(28)
Female Shake—0: Absent (no violent side to side movement); 1: Present
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(29)
Type of female shake—0: Horizontally; 1: Vertically
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(30)
Female foreleg movements—0: No significant movements of forelegs; 1: Female intermittently lifts forelegs off the substrate; 2: Female repeatedly lifts forelegs above head
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(31)
Female hindleg movements—0: Female hindlegs not used to interact with male; 1: Female hindleg used to ‘kick’ male; 2: Female hindleg ‘rubbing’ male hindlegs
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(32)
Female ejection of ovipositor when male is mounted—0: Absent; 1: Female ejects ovipositor when after genital contact; 2: Female ejects ovipositor prior to genital contact
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(33)
Use of male midleg—0: No forceful contact with female head; 1: Male uses midleg to ‘beat’ female head
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Tan, D.S.H., Ng, S.R. & Meier, R. New information on the evolution of mating behaviour in Sepsidae (Diptera) and the cost of male copulations in Saltella sphondylii . Org Divers Evol 11, 253–261 (2011). https://doi.org/10.1007/s13127-011-0054-2
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DOI: https://doi.org/10.1007/s13127-011-0054-2