Research Article |
Corresponding author: Christian Kronmüller ( ck@scolopendromorpha.com ) Academic editor: Ivan H. Tuf
© 2015 Christian Kronmüller, John G.J. Lewis.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kronmüller C, Lewis JGE (2015) On the function of the ultimate legs of some Scolopendridae (Chilopoda, Scolopendromorpha). In: Tuf IH, Tajovský K (Eds) Proceedings of the 16th International Congress of Myriapodology, Olomouc, Czech Republic. ZooKeys 510: 269-278. https://doi.org/10.3897/zookeys.510.8674
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The function of the variously shaped ultimate legs of Scolopendridae is briefly reviewed. Their function in Scolopendra heros Girard, 1853, Scolopendra subspinipes Leach, 1815, Scolopendra morsitans (Linnaeus, 1758), Scolopendra galapagoensis Bollman, 1889, Scolopendra hainanum Kronmüller, 2012, Scolopendra spinosissima Kraepelin, 1903 Cormocephalus aurantiipes (Newport, 1844) and Ethmostigmus trigonopodus (Leach, 1817), in which they are least specialised has been investigated. Specimens were tapped with forceps on different parts of the trunk to simulate the attack of a predator. When tapped on the first third of the trunk (near the head), the centipedes attacked the forceps with their forcipules. When tapped on the last third or the ultimate legs, they adopted a warning position, raising the ultimate legs to display the ventral and medial prefemoral spines as well as the spined coxopleural processes. In some cases the centipedes attacked the forceps with the claws of the ultimate legs by chopping down on them after lifting the legs high into the warning position. When tapped in the mid part of the trunk, the centipedes curled sideways to reach the forceps with their forcipules and ultimate legs simultaneously. Scolopendra galapagoensis not only lifted the ultimate legs into the warning position but also the last 3-4 pairs of locomotory legs, presenting their distodorsal prefemoral spines. This resembles the warning posture of some spiders. In addition to their function in warning behaviour, defensive stabbing, ritualised meeting reactions and during courtship behaviour, the ultimate legs may in addition act as hooks and perhaps be involved in species recognition. No evidence was found that the ultimate legs are used to catch prey, nor of prey or predators being held between the prefemora.
Chilopoda , Scolopendromorpha , ultimate legs, prey capture, defence reaction, courtship behaviour, mating
The ultimate legs of scolopendrids exhibit a variety of shapes, the majority being what
In this paper we review the functions of ultimate legs in those species that have the least modified ultimate legs and report the results of laboratory experiments on species of Scolopendra, Cormocephalus and Ethmostigmus.
Field observations have shown that the ultimate legs of scolopendrids are used as hooks from which the animal can hang. Thus
Kronmüller (2013, unpublished data) reported and photographed a Scolopendra dehaani that escaped the plastic container in the laboratory and was later found hanging on the camera tripod just using the ultimate legs (Figure
Warning posture
Lewis (unpublished data) noted in Nigeria that a small Scolopendra morsitans L., 1758, raised and splayed its ultimate legs when approached from behind by a large Mecistocephalus (Geophilomorpha).
Autotomy and sound production
As noted above, in case of danger the members of the genus Alipes swing the ultimate legs from side to side and stridulate perhaps to distract or frighten potential predators.. They also use autotomy to detract potential predators like birds, lizards or mammals. In case of an attack, the centipede starts to stridulate which directs the attention of the predator to the ultimate legs which can be autotomised to make the escape easier. When detached with forceps the leaf-like ultimate legs of an Alipes grandidieri from Tanzania continued to stridulate for more than half a minute.
Video clips of Alipes sp. showing the typical defence behaviour including the sound of the stridulation as well as the bending movements of the detached ultimate legs of Rhysida sp. can be seen on the website http://www.scolopendromorpha.com.
Meeting reactions
Ritualised meeting reactions have been described for Scolopendra cingulata Latreille, 1829 (
A Warning posture of Scolopendra spinosissima B Warning posture with lifted ultimate legs as well as the last pairs of locomotory legs in Scolopendra galapagoensis C Interlocked ultimate legs in a pair of Scolopendra galapagoensis just before mating D antennae tapping under the lifted ultimate legs in Scolopendra dehaani in courtship behaviour.
In head to head contact both rapidly swing round the posterior part of body and attempts to grip the other with the ultimate, followed by the more posterior legs. Brunhuber’s plate 1A shows that the legs are parted and hooked over the body of the other animal. In this reaction the median spines on the ultimate legs seem to aid the grip. Kronmüller (unpublished data) observed that when two Scolopendra dehaani meet and they grasp each other in the previous described way, they hold this position for several minutes and up to more than half an hour. This ritual may be to avoid any aggressive behaviour.
A short video clip of this meeting reaction can also be seen on the website http://www.scolopendromorpha.com.
Courtship behaviour
During courtship behaviour in Cormocephalus anceps anceps the “defence posture” is adopted both partners use their antennae to tap the posterior part of the body especially the ultimate legs of the opposite sex (Figure
Species recognition
Laboratory experiments were carried out to further investigate behaviour in prey capture as well as possible defence reactions in case of a predator attack. To simulate a the attack of mammal or bird predator, a piece of foam held with 25cm forceps was used to tap the centipedes from above.
The centipedes that were used for the experiments were either adult wild caught specimens (Scolopendra heros, Scolopendra subspinipes, Ethmostigmus trigonopodus, Scolopendra morsitans, Scolopendra hainanum, Cormocephalus aurantiipes) or, in one case, an adult captive bred specimen (Scolopendra galapagoensis).
The specimens were kept for at least 12 weeks before the experiments started in plastic containers of different sizes depending on the size of the specimen, with an air temperature of 25 degrees Celsius in day-time (around 10 hours illumination with a neon lamp) and 20–22 degrees Celsius at night-time. The average air humidity was around 70%. All plastic containers had a layer of humus and a piece of bark as a hiding-place. Once a week, the humus was sprayed with water to keep it moist and the specimen was fed one adult cricket (Gryllus assimilis).
For the experiments, the centipedes were moved to a 20 × 20 cm plastic container with a layer of about 2cm humus and left for at least one hour without disturbance before the experiments started. After each experiment, the centipedes were left for a further hour to recover before the next experiment.
The first experiment involved tapping the centipedes at different parts along the length of the body (anterior third, middle third and posterior third) to observe possible differences in defense reactions and to compare the responses of the investigated species.
In a second experiment a piece of bark was placed in the container as a refuge and the layer of humus was increased to 10cm. A centipede was placed in the container and as soon as the first third of the trunk was hidden either by the centipede crawling under the bark or by it digging into the humus, the tapping experiments described above were repeated.
Photos and short video clips are made with a Canon EOS 60D and a Canon EOS 6D, connected either to a Canon standard lens (18-55mm) or a Canon macro lens (100mm), three Canon Speedlight EX550/EX430 flashes (photos only), a tripod and a remote control (video clips).
Initial experiments were carried out on an adult Ethmostigmus trigonopodus (Leach, 1817) from Tanzania, length 13 cm. When tapped on the first third of the trunk (near the head), the centipede attacked the forceps with the forcipules. When tapped on the posterior third, or even at the ultimate legs, it adopted the warning position, raising the ultimate legs to display the ventral and medial prefemoral spines as well as the coxopleural processes which are also equipped with spines. In the most cases the centipede directed the warning position towards the forceps by turning the posterior segments. In some cases, depending on the strength and speed of the simulated attack, the centipede counter-attacked the forceps with the claws of the ultimate legs by chopping down on them after lifting them high into the warning position.
When the E. trigonopodus was tapped in the mid part of the trunk, the centipede curled sideways to reach the forceps with the forcipules and the ultimate legs simultaneously. These observations were repeated in experiments with Scolopendra heros var. arizonensis Kraepelin, 1903 (USA), Scolopendra galapagoensis (Peru), Scolopendra morsitans (Pakistan), Scolopendra cingulata (Spain), Scolopendra subspinipes (Indonesia), Scolopendra hainanum Kronmüller, 2012 (China), Scolopendra spinosissima Kraepelin, 1903 (Philippines), and Cormocephalus aurantiipes (Newport, 1844) (Australia). All showed the same warning posture raising the ultimate legs (Figure
In addition Scolopendra galapagoensis lifted not only the ultimate legs into the warning position but also the last 3-4 pairs of locomotory legs (Figure
In the experiments with bark and a thicker layer of humus, no difference was observed in the behaviour depending on whether the specimen was tapped on the mid part of the trunk or on the last third. The result was the same when the centipede was fed (Schistocerca sp.) and the forcipules as well as the anterior pairs of locomotory legs were involved in feeding. In this situation the centipede could not use the forcipules to carry out a warning bite and the only defence behaviour was hitting with the ultimate legs and stabbing with the ultimate leg pretarsi (claws). The experiment was repeated with Ethmostigmus trigonopodus, Scolopendra galapagoensis, Cormocephalus aurantiipes and Scolopendra subspinipes, which all showed the same results.
The scolopendrid species investigated here have unmodified and relatively robust ultimate legs. Species with very long thin ultimate legs such as some Otostigmus (Otostigmus) and Rhysida species, have not been investigated.
We conclude that in the case of the species investigated the use of the ultimate legs is primarily for defence, however, should potential prey make contact with the centipede, the defensive response and defensive biting will lead to feeding whereas in the case of a predator, the centipede will try to escape after one or more warning bites.
Rowland Shelley (email dated 03 January 2011) notes “that there are frequently spines on the medial surface of the scolopendrid (ultimate leg) prefemora and I’ve always presumed that these must function to hold something steady I had always thought this would be prey... ” We were unable to confirm that scolopendrid centipedes hold prey or predators between the ultimate leg prefemoral, although in the ritualised meeting reaction in some species, the median spines on the ultimate legs seem to aid the grip.
The four main functions in species in which the ultimate legs are little specialised are:
Acting as hooks to suspend the animal for example from cave ceilings.
Warning position, when the legs are raised and splayed to reveal the prefemoral and coxopleural spines.
Stabbing as a defence, especially when the front part of the trunk and the head are unable to carry out any defensive action.
Grasping, the legs being flexed as when two animals assume the defence posture or as part of the mating ritual. In this case posteriorly directed spines would aid the grip. No evidence was found of objects being held between the prefemora.
The investigated centipedes (S. heros, S. subspinipes, S. morsitans, S. hainanum, S. galapagoensis, S. spinosissima, C. aurantiipes, E. trigonopodus) are from different genera, different continents and different types of habitat. Nevertheless, their behaviour in prey capture and defense reaction do not differ. Also the meeting reaction seems to be very similar the same in all observed species.
In addition to the behavioural functions of the ultimate legs, two visual signals should be considered:
Some scolopendrid centipedes, especially the giant Scolopendra sp. have striking colour patterns. In several species, the colouration of the last trunk segments and the ultimate legs is the same as the colouration of the head and antennae (Figure
The warning posture, especially that of Scolopendra galapagoensis, where the last three or four pairs of locomotory legs are raised in addition to the ultimate pair, resembles that of spiders such as Phoneutria sp. or tarantulas and may be a case of possible signal co-evolution.