©Staatl. Mus. f. Naturkde Karlsruhe & Naturwiss. Ver. Karlsruhe e.V.; download unter www.zobodat.at andrias, 13: 17-22, 2 Figs; Karlsruhe, 30. 9. 1994 17 C h r is t o p h e r M a r t iu s , H u b e r t H o f e r , M a n f r e d V e r h a a g h , J o a c h im A d is & V o l k e r M a h n e r t Terrestrial arthropods colonizing an abandoned termite nest in a floodplain forest of the Amazon River during the flood Abstract Introduction Terrestrial arthropods were identified as secondary colonizers Survival strategies of soil arthropods in the periodically of an abandoned termite nest ( Nasutitermes tatarendae) at inundated floodplain forests (varzea) along the the beginning of the inundation period in a floodplain forest in­ Solimoes-Amazon River have long been in the focus of fluenced by whitewater in central Amazonia. About 50 indivi­ duals of non-termite meso- and macrofauna were recorded research (Ir m le r 1981, A d is 1992). Migration to tree per liter of nest volume, mainly terrestrial Arachnida, Isopoda, canopies is one possibility for soil species to survive Diplopoda, Blattodea, Staphylinidae, and Formicidae. Appa­ flooding. Termites represent a considerable part of the rently, abandoned termite nests could be important for the sur­ varzea fauna (M a r tiu s 1989), and a great part of their vival of the soil-fauna which migrates to the canopy during the arboreal nests is abandoned (M a r t iu s 1994). How­ flood period. ever, the question whether these nests are „refugia“ for the migrating soil fauna has never been analysed. Kurzfassung Terrestrische Arthropoden als Besiedler eines verlasse­To get an insight into this problem we present here nen Termitennests in einem Überschwemmungswald some data gathered during field studies on termite co­ während der Überflutungsphase lonization (M a r t iu s 1989) of llha de Marchantaria, an In einem várzea-Wald Zentralamazoniens wurden zu Beginn island in the river Amazon near Manaus, Brazil (fig. 1), der Überschwemmungsphase terrestrische Arthropoden als and a typical varzea (= white water floodplain forest Sekundärbesiedler eines verlassenen Termitennestes ( Nasu­ fig. 2) site (J u n k 1984). titermes tatarendae) identifiziert. Etwa 50 Individuen der The fauna of an abandoned termite nest of Nasutiter­ Meso- und Makrofauna (keine Termiten) wurden pro Liter Nestvolumen registriert, überwiegend terrestrische Arachnida, mes tatarendae was determined. The nest was loca­ Isopoda, Diplopoda, Blattodea, Staphylinidae und Formicidae. ted on a tree which base had been flooded the day be­ Verlassene Termitennester können offensichtlich für das fore sampling. Its lower half (with a volume of more Überleben der Bodenfauna, die während der Flutphase in die than 80 liters) was cut from the tree with a chain saw, Baumkronen wandert, wichtig sein. then transported in an aluminium container to the la­ boratory, split into small fractions of approx. 5-10 dm3 Resumo each, and extracted by heat in a Tullgren Funnel. Spe­ Colonizaçio por artrópodos terrestres de um ninho aban­ cimens were separated to orders, stored in alcohol, donado de cupim no inicio do período da enchente numa and identified. Additionally, nine abandoned arboreal floresta de várzea Nasutitermes nests (mostly N. corniger) which were Artrópodos terrestres foram identificados como colonizadores secundónos de um ninho abandonado de cupim ( Nasutiter­ found on the forest floor during the low-water period mes tatarendae) no inicio do período da enchente numa flore­ were screened for associated fauna. sta de várzea da Amazonia Central. Aproximadamente 50 in­ dividuos de meso- e macrofauna (nao cupins) foram achados Secondary colonizers of the abandoned nest of no volume de um litro de ninho, sobretudo Arachnida, Isopo­ Nasutitermes tatarendae da, Diplopoda, Blattodea, Staphylinidae, e Formicidae. Evi­ Table 1 shows the arthropods which we found in the dentemente ninhos abandonados de cupins podem ser impor­ abandoned termite nest of Nasutitermes tatarendae. tantes para a sobrevivéncia da fauna de solo que migra ao dossel da floresta durante a inundaçâo. No termites, not even of other species, were found in this nest. Due to the sampling procedure, the small Authors mesofauna (usually <1mm, e.g. Acari and Collembola) Dr. Christopher Martius, Universität Göttingen, II. Zoologi­ has not been quantitatively assessed, and its pre­ sches Institut, Abteilung Ökologie, Berliner Str. 28, D-37073 sence is only assigned with an ,,x“ in table 1. Animals Göttingen, Germany; of size classes usually >1 mm (larger meso- and Dr. Hubert Höfer & Dipl.-Biol. Manfred Verhaagh , Staatli­ macrofauna) have been quantitatively assessed. ches Museum für Naturkunde, Postfach 6209, D-76042 Karls­ At least 77 different arthropod taxa were registered in ruhe, Germany; this single abandoned nest. With a total of 4054 indivi­ Priv.-Doz. Dr. Joachim Adis, Max-Planck-Institut für Limnologie, AG Tropenökologie, Postfach 165, D-24302 Plön, Germany; duals of the fauna with a size of >1 mm the nest con­ Dr. Volker Mahnert , Muséum d’Histoire naturelle, Case tained a high density of arthropods (corresponding to postale 434, CH-1211 Genève, Switzerland. about 50 individuals per liter of nest volume). We clas- ©Staatl. Mus. f. Naturkde Karlsruhe & Naturwiss. Ver. Karlsruhe e.V.; download unter www.zobodat.at 18 andrias, 13 ( 1994) Figure 1. Aerial view of llha de Marchantaria during high water level; Photo: J. A dis. Figure 2. Edge of the study site in the varzea forest on llha de Marchantaria, where the termite nest has been found; Photo: H. Hofer. ©Staatl. Mus. f. Naturkde Karlsruhe & Naturwiss. Ver. Karlsruhe e.V.; download unter www.zobodat.at Martius et al.: Arthropods, abandoned termite nest, central Amazonia 19 Table 1. Arthropods found in an abandoned nest of Nasutiter- Crematogaster sp. (limata- group) mes tatarendae. x = no quantitative assessment; n = not de­ Formicinae 1291 2 termined. Paratrechina sp. Camponotus sp. Taxa Number of Number of Dolichoderinae 1 1 Individuals Species Tapinoma melanocephalum Chelicerata - Arachnida Bethylidae 7 4 Araneae 156 11 unident. Hymenoptera 8 4 Pseudoscorpiones 1 1 Opiliones sum >4054 >77 sp. 1 ad. 22 1 sp. 2 ad. 17 1 sp. 3 ad. 18 1 sify them mostly as opportunistic users of the nest juv. 412 structure (facultative „termitariophiles“; A rau jo 1970). ? Acari X It was particularly striking how many large scolopen- Crustacea, Chilopoda, Diplopoda ders were extracted. This shows the extension of the Isopoda internal surface area of the chambers in termite nests Dubioniscus sp. (Eubelidae) 159 1 (cf. M a r tiu s 1994). Prosekia sp. (Philosciidae) 158 1 Spider species are listed in table 2. All spider species, Chilopoda - Scolopendromorpha 72 n collected in the abandoned termite nest, have already Diplopoda - Polydesmida (juv. & adults) 185 2 been collected in the same varzea forest during an ecological survey of the spider fauna (Hofer unpubl.). Insecta Most of them can be considered ground living species. Collembola X ? The oonopid species, Anapistula sp. and Theotima cf. Orthoptera - Blattodea 104 n minutissima (P etrunkevitch ), are tiny spiders domi­ Dermaptera 2 n nant in leaf litter samples, pitfall traps and ground- Plomoptera - Stenorrhyncha 8 n photoeclectors taken in blackwater and whitewater in­ Coleóptera undation forests during the non-inundated period (Ho­ unident, larvae 7 n fer 1990). When these forests are being flooded (Fe- Carabidae 26 10 bruary-April) these spiders migrate into the trunk re­ Elateridae 3 n gion, where they have been collected in arboreal fun­ Staphylinidae 370 n nel traps. The same is true for Tricongius amazonicus Pselaphidae 3 n Platnick & H ofer , 1990, a small sized (=3 mm) gna- Tenebrionidae 27 3 phosid species. The juveniles migrate to the trunks, Scarabaeidae X n where they become adult. Adult spiders return to the Scolytidae 7 n ground when the water has receded from the forest Colydiidae 16 1 (September) and reproduce there (Platnick & H ofer Histeridae sp. 1 5 1 1990). Thereafter the spiderlings of the new genera­ sp. 2 12 1 tion spread out on the newly emerged ground. sp. 3 1 1 Orthobula sp. and Diestus sp. are medium sized spi­ Scydmaenidae 13 1 ders which were only rarely captured in traps during Scaphidiidae sp. 1 3 1 our studies in inundation forests. It seems possible sp. 2 3 1 that they live permanently in higher strata. sp. 3 1 1 Ctenus spp. are the most conspicuous large ground li­ Leptodiridae 1 1 ving spiders (>10 mm) in all central Amazon forests. In others (small mesofauna) 64 7 inundation forests they avoid being flooded by vertical Hymenoptera or horizontal movements away from the rising water. Formicidae (det. M.Verhaagh ) Consequently they also appear in arboreal funnel Ponerinae 782 1 traps during inundation. Pachycondyla constricta A wide variety of beetles was found In the nest, many workers (735) of them very small. Although Staphylinidae are among alates (13) the most abundant groups (tab. 1), no termitophilous pupae (34) species (Kistner 1969) were present among them, Myrmicinae 86 4 probably because these species only live in inhabited Solenopsis (Diploropthrum) sp. termite colonies. Cyphomyrmex rimosus Among the ants the ponerine Pachycondyla (=Meso- Pheidole sp. ponera) constricta (M a y r , 1883) was present with wor- ©Staatl. Mus. f. Naturkde Karlsruhe & Naturwiss. Ver. Karlsruhe e.V.; download unter www.zobodat.at 20 andrias, 13 (1994) Table 2. Spiders (Arachnida) collected in the abandoned nest of ground and in the vegetation, and nests are located in Nasutitermes tatarendae In the varzea forest (det. H. Hofer ). soil, or rotting wood, sometimes high on trees (V e r ­ h a a g h , unpubl.). Therefore, workers from the termite Family Genus/Species Number of nest did not necessarily come from the ground. We did Individuals not find any brood, but there might have been brood Spiders <2 mm body length: together with more workers in the upper part of the ter­ Oonopidae mite nest which was not examined.