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Short Communication Matriphagy in The 2005. The Journal of Arachnology 33:873±877 SHORT COMMUNICATION MATRIPHAGY IN THE NEOTROPICAL PSEUDOSCORPION PARATEMNOIDES NIDIFICATOR (BALZAN 1888) (ATEMNIDAE) Everton Tizo-Pedroso and Kleber Del-Claro: LaboratoÂrio de Ecologia Comportamental e de InteracËoÄes, Instituto de Biologia, Universidade Federal de UberlaÃndia. C.P.593, Cep 38400-902, UberlaÃndia, MG, Brasil. Telfax: 55(34)32182243. E-mail: [email protected]. ABSTRACT. We studied the natural history and social behavior of Paratemnoides nidi®cator (Balzan 1888) in a tropical savanna system. Females were responsible for all nymphal care. We observed, for the ®rst time in pseudoscorpions, the occurrence of matriphagy behavior by the offspring. During conditions of food deprivation, the mother went out of the nest and passively awaited the protonymphs' attack, not reacting to the capture nor to the nymphs feeding on her body. We suggest that this extreme form of parental care, matriphagy, can reduce cannibalism among protonymphs and facilitate the evolution of social behavior in pseudoscorpions. RESUMO. NoÂs estudamos a histoÂria natural e o comportamento social de Paratemnoides nidi®cator (Balzan 1888) na regiaÄo dos cerrados. As feÃmeas foram responsaÂveis por todo o cuidado aÁs ninfas. NoÂs observamos, pela primeira vez em pseudoescorpioÄes, a ocorreÃncia de matrifagia pela prole. Em condicËoÄes de fome, a maÄe deixa o ninho e passivamente espera que as protoninfas a ataquem, naÄo reagindo nem aÁ captura, nem aÁ alimentacËaÄo das ninfas sobre seu corpo. NoÂs sugerimos que esta forma extrema de cuidado parental, matrifagia, possa reduzir o canibalismo entre as protoninfas e assim facilitar a evolucËaÄo de comportamento social em pseudoescorpioÄes. Keywords: Social behavior, maternal care, Arachnida, cannibalism, tropical savanna The order Pseudoscorpiones is highly diversi®ed may occur in the same rock ®ssure (Gabbutt 1962, with more than 3,239 described species in 425 gen- 1966). Females of Neobisium muscorum (Leach era and 24 families, representing around 3.3% of all 1817) can have chambers placed side by side (Wey- arachnids (Harvey 1991, 2002). In general, pseu- goldt 1969). In Paratemnoides elongatus (Banks doscorpions are small (2±8 mm) and are non-social 1895) and P. minor (Balzan 1892), both species that animals that behave aggressively in intraspeci®c occur beneath tree bark, nymphs can build molt contacts (Weygoldt 1969; Zeh 1987). Zeh (1987) chambers cooperatively and adult females bearing reported ®ghts between males of a Chernetidae spe- a brood sac can use molt chambers for brood care cies during contests for food or females, resulting as well (Brach 1978; Hahn & Matthiesen 1993b). in cannibalism. Some Atemnidae species, however, This cooperative building of nests saves time and show a high level of sociality, living in groups, silk, maintains appropriate humidity conditions and sharing food and hunting cooperatively (Brach protects the brood from predators (Brach 1978). In 1978; Zeh & Zeh 1990). both those Paratemnoides species females in the All pseudoscorpion species present some level of nest remove the brood sac from the genital opening parental care. Indeed, females of all species take after secreting the nutritive ¯uid to the embryos care of embryos that are maintained inside a brood (Brach 1978; Hahn & Matthiesen 1993a). Pro- sac attached to her genital opening (Levi 1953; Ga- tonymphs of Pselaphochernes scorpioides (Herman butt 1970b; Weygoldt 1969). Females can also build 1804) remain 2±3 days inside the nest receiving silk chambers in which they rest with the brood sac care until they disperse (Weygoldt 1969). until the emergence of the protonymphs (Brach Pseudoscorpions are widespread. For instance, in 1978; Gabbutt 1962, 1966, 1970a; Levi 1948, 1953; Central Amazon, Adis & Mahnert (1985) recorded Harvey 1986; Zeh & Zeh 2001). In Neobisium ma- 60 species belonging to 25 genera in 10 families. ritimum (Leach 1812) the silk chamber is built and The Brazilian cerrado savanna originally covered occupied by one individual, and several chambers approximately 25% of the country and is currently 873 874 THE JOURNAL OF ARACHNOLOGY Table 1.ÐComposition of all colonies (by sex and age classes) of Paratemnoides nidi®cator (Atemni- dae) studied. Colony Males Females Tritonymphs Deutonymphs Protonymphs Total 1348ÐÐ15 2451614342 3 8 12 15 5 Ð 40 4 7 5 4 12 21 49 5 9 10 4 Ð Ð 23 6 4 7 1 28 30 70 7121173Ð33 X 6 SD 6.71 6 3.25 7.71 6 3.25 7.85 6 5.7 8.86 6 10.08 7.71 6 12.47 38.86 6 17.98 one of the most endangered tropical ecosystems until the nymphs were adults or were forced from (Oliveira & Marquis 2002). To our knowledge there the nest. The female also guarded the nest entrance is no study about pseudoscopions in the cerrado. against enemies. In some cases, the female pro- Here, we studied the biology and natural history of duced an additional brood and then forced nymphs Paratemnoides nidi®cator (Balzan 1888) an atem- from the ®rst brood out of the chamber by touching nid species that occurs under the bark of living trees them with her pedipalps. After chasing the original of Caesalpinia pelthophoroides (Caesalpiniaceae), brood, the female sealed the exit with new silk and a tree found throughout the cerrado domain. produced the new brood sac. The ``displaced'' Observations for this study were conducted from nymphs then cooperatively built another chamber October 2001 to December 2003 in UberlaÃndia, in which they molted. The reproductive and the Brazil (188 539S, 488 159W; 863 m el.), in the south- molting chambers were built side by side sharing eastern limit of the cerrado distribution. Seven col- the vertical walls. This behavior has also been ob- onies of pseudoscorpions (Table 1) were collected served in P. elongatus (Banks 1895) and P. minor from the ®eld and maintained in captivity during all (Balzan 1892) (Brach 1978; Hahn & Matthiesen the study. Each colony was kept in a glass bottom 1993a). In the ®eld, we found colonies of P. nidi- culture dish (12 cm of diameter) having the original ®cator with nests composed of 3±20 chambers. piece of tree bark and was fed twice a week with We identi®ed 95 distinct behavioral acts, of live termites (Armithermes sp.) and beetles (Acant- which 16 were related to reproductive behavior, hocelides obtectus, Bruchidae). Moisture was pro- mainly parental care (Table 2). Parental behaviors vided by a small piece of water-soaked cotton. Be- comprised 10.13% of the behaviors seen in this spe- havioral observations were made using the ``all cies. However, the acts in this category are per- occurrence samples'' method (Altmann 1974). Us- formed only by adult females, so more than 75% ing this method, everything that a group or individ- of adult female behaviors are related to taking care ual does during an observation session is recorded of embryos and young. Other important behavioral ad libitum. This method is particularly useful to be- acts of females were self-grooming and feeding. gin a study or to observe rare or fortuitous behav- Males directly cooperated in parental care by catch- iors (see also Martin & Bateson 1993; Del-Claro ing and offering prey to all members of the colony 2004). Individual observation sessions lasted 30± (94 records during 50 observation sessions). How- 40 min and were made during the day (mainly be- ever, nymphs in general were fed by the mother. tween 09:00h and 15:00h) using natural light. As Our data revealed that females left the nest to hunt colonies are built under the bark of trees, during the prior to feeding the brood (56%, or in 42 out of 75 observation sessions the petri dishes with the col- times that we observed females leaving the nest onies were put on a wire stand with a mirror below during the observations). The mother ate the rest of that enable the observations without be disturbing the carrion left by the nymphs. Tritonymphs hunted the animals. In the present paper we describe the cooperatively with the mother (3 in 42 observations social and reproductive behavior of P. nidi®cator of the mother hunting), or without the mother (n 5 based on 50 observation sessions (34 hours total 14 records during 50 observation sessions). observations). Voucher specimens have been At the end of our observations we recorded col- lodged with the Museu de Zoologia de SaÄo Paulo ony behavior under food deprivation by depriving (MZUSP). the colony food for one week. On the seventh day We observed that females wove the reproductive we observed matriphagy behavior in three of the nest alone. Inside the chambers, the female provid- seven colonies. The mother exited the nest, raised ed parental care continuously to embryos and her pedipalps and passively waited for her nymphs nymphs by feeding and grooming them. This lasted to attack. Nymphs (9 6 3 protonymphs, X 6 SD, TIZO-PEDROSO & DEL-CLAROÐMATRIPHAGY IN A PSEUDOSCORPION 875 Table 2.ÐBehavioral acts of Paratemnoides nidi®cator (Atemnidae) associated with ``Parental Care''. Data recorded from a clutch of 90 individuals (33 adults and 57 nymphs) reared in captivity (n 5 34 hours of observations). Percent frequency of Number of the behavioral act Behavioral act observations (total 453 acts) 1-Female weaving chamber. 179 30.4 2-Female occupying a woven chamber previously built by anoth- 5 0.85 er individual. 3-Female excluding conspeci®cs from chamber. 4 0.68 4-Female resting in the nest. 76 12.9 5-Female moving in the nest. 19 3.23 6-Female touching the nest wall with pedipalps. 10 1.7 7-Female touching the embryos with pedipalps. 84 14.3 8-Female touching the protonymphs with pedipalps. 29 4.92 9-Female transporting wood pieces inside the nest. 4 0.68 10-Female inserting fragments of wood in the nest walls.
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