8. Systematics and Faunistics

Poster presentation: First record of a root community in Southeast Asia: cave- dwelling from Maros karst, Sulawesi (: Fulgoromorpha: : Bennini) Andreas Wessel* 1, 2, Roland Mühlethaler 1, Kristina von Rintelen 1, Thomas von Rintelen 1, Björn Stelbrink 1, Ekkehard Wachmann 1, Hannelore Hoch 1

1 Museum für Naturkunde – Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der

Humboldt-Universität zu Berlin, Invalidenstraße 43, D-10115 Berlin, Germany 2 Institute for Evolution and Biodiversity, Westfälische Wilhelms-Universität Münster,

Hüfferstraße 1, D-48159 Münster, Germany

A systematic survey of Maros karst caves in summer 2009 revealed the first known terrestrial cave with roots in the dark zone and an associated fauna for Southeast Asia. Remarkably, this very first discovery of available resources for a root community in the region coincides with the finding of planthoppers as sap- sucking primary consumers. Planthoppers are common elements of root communities in different parts of the world. A total of more than 50 cave-dwelling species are known from Africa (incl. Madagascar), Australia, Latin America, and several oceanic islands. Two-thirds of the troglobiotic and troglophilic species belong to the Cixiidae as well as the newly discovered species from Sulawesi. The Maros cave planthoppers however, are the first representatives of the tribe Bennini ever recorded in a subterranean environment. The Bennini (about 100 species) are characterised by a unique feature – they possess very conspicuous lateral appendages each ending in a wax-covered sensillum. The precise function of these appendages and a possible role in orientation in the dark is unknown as in general the biology of this group is poorly studied. It is assumed that the ability of planthoppers to communicate by substrate vibrations is a prerequisite for the colonisation of cave environments. A well-studied example from Hawaii shows species-specific “song” patterns and revealed a complex pattern of subterranean speciation. The successful recording of vibrational signals from the Maros cave may open up a new model system for the study of the dynamics of subterranean evolution.

~ 149 ~ 20th International Conference on Subterranean Biology (ICSB) • 29 August – 3 September 2010, Postojna, Slovenia • International Society for Subterranean Biology (ISSB)

museum für naturkunde

First record of a root community in Southeast Asia: troglophilic planthoppers from Maros karst, Sulawesi (Hemiptera: Fulgoromorpha: Cixiidae: Bennini)

Andreas Wessel1,2*, Roland Mühlethaler1, Kristina von Rintelen1, Thomas von Rintelen1, Björn Stelbrink1, Ekkehard Wachmann1 & Hannelore Hoch1

1 Museum für Naturkunde – Leibniz-Institut for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstraße 43, D-10115 Berlin, Germany 2 Institute for Evolution and Biodiversity, University Münster, Hüfferstraße 1, D-48159 Münster, Germany *Correspondence: [email protected]

A systematic survey of Maros karst caves in summer 2009 revealed the first knownterrestrial cave with roots in the dark zone and an associated fauna for Southeast Asia (cf. Deharveng & Bedos 2000). Remarkably, this very first discovery of available resources for a root com- munity in the region coincides with the finding ofplant­ hoppers as sap-sucking primary consumers. (Fig. 1-4)

Planthoppers are common elements of root communities in different parts of the world. A total of more than 50 cave-dwelling species are known from Africa (incl. Ma- 1 sec dagascar), Australia, Latin America, and several oceanic

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 islands. Two-thirds of the troglobitic and troglophilic species belong to the Cixiidae, as does the newly dis- covered species from Sulawesi. (Fig. 7, Hoch & Wessel 2006)

The Maros cave planthoppers, however, are thefirst re- presentatives of the tribe Bennini ever recorded in a subterranean environment. The Bennini are characte- rised by a unique feature – they possess very conspicuous lateral appendages each ending in a wax-covered sen- sillum (Fig. 1 & 3, inset). The precise function of the- 0.1 sec se appendages and a possible role in orientation in the

Fig. 1. Maros cave planthopper, Gen. nov., spec. nov. (Cixiidae: Bennini) Fig. 5.0.1 Oscillograms0.2 0.3 of 0.4the vibrational0.5 0.6 signals0.7 of0.8 the 0.9Bennini1 planthoppers.1.1 1.2 1.3 1.4 1.5 1.6 1.7 1.8 1.9 dark2 is unknown as in general the biology of this group is poorly studied. Hitherto, about 20 Bennini species are described, but a current taxonomic revision will raise the number above 100 species (Hoch, in prep.). The Maros Bennini will be described as new species in a newly erec- ted genus (Hoch et al., in prep).

Sulawesi The reported here show no significant troglo- morphies, however, they are also not known from epi- Fig. 2. Maros karst west of Balocci, the arrow indicates the position of the Gua Assuloang (Gallery Entr.). gean samples. Planthopper nymphs feed on roots and can be defined asterrestrial epikarst fauna. Wandering into caves, the emerging adults can survive and repro- duce from the outset in the dark zone without further adaptation necessary and then develop troglomorphies PANGKAJENE in the course of an rapid adaptive shift (see Howarth & Hoch 2004, Wessel et al. 2007). We could describe them as exapted eutroglophile accidentals (cf. Sket 2008).

BALOCCI It is assumed that the ability of planthoppers to commu- Gua Assuloang nicate by substrate vibrations is a prerequisite or exap- tation for the colonisation of cave environments E W S G N • B R (Hoch & Wessel 2006). A well-studied I N E A A K K MAROS E I N R G KAPPANG example from Hawaii shows species- B • A­ N BANTIMURUNG E specific “song” patterns and re- S W W recent S vealed a complex pattern of sub- E N • town / village B cave terranean speciation (Wessel & G survey reports Bennini R N E I island Bohol. A Fig. 6. Location of the Gua Assuloang, Maros Karst near Balocci, Kepulauan Pangkajene, Sulawesi Sela- Hoch 1999, Wessel 2008). The K planthoppers in a karst cave K tan / South Sulawesi, Indonesia. A Philippine

I recording of vibra­ E successful N

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Tettigometridae Borystheninae Andini tional signals B

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Cixiidae 28 5 Bothriocerinae Bennarellini • The animals belong to a known epigean

cave planthopper (Fig. 5) may E S

Delphacidae 1 2 Bennini (1/~100) species. This find is momentous, however, W W . These are the first

Kinnaridae 3 1 Brixidiini open up a new model system S E

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as the adults were accompanied by several •

Meenoplidae 11 1 Brixiini (10/~150) N th

for the study of the dynamics instar

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Derbidae Cajetini G R

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Achilidae (15/>409) of subterranean evolution. I larval stages of Bennini ever found, A

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Achilixiidae Duiliini A

and will permit first insights intoI E

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References G

Fulgoridae Eucarpiini B the developmental biology

Deharveng, L. & Bedos, A. N

2000. The cave fauna of Sou- •

Dictyopharidae Gelastocephalini E S

theast Asia. Origin, evolution and ecology. In: Wilkens, H., of the enigmatic “sideW

S

W organs”!

E

•

N

Fig. 3. Roots in the cave: locus typicus Gua Assuloang. Inset: Bennini planthopper in situ. Issidae (2/~236) Culver, D. C. & Humphreys, W. F. (eds). Subterranean ecosystems. B R G E N

I A Acanaloniidae Pentastirini (8/~622) Elsevier, Amsterdam: 603-632. K Hoch, H., Asche, M., Burwell, C., Monteith, G. & Wessel, A. 2006. Nogodinidae Pintaliini Morphological alteration in response to endogeic habitat and ant association in Flatidae Semonini two new planthopper species from New Caledonia (Hemiptera: Auchenorrhyncha: Fulgoro-

Ricaniidae Stenophlepsiini morpha: Delphacidae). Journal of Natural History 40: 1867-1886. Hoch, H. & Wessel, A. 2006. Communication by Substrate-Borne Vibrations in Cave Plan- Hypochthonellidae 1 thoppers (Auchenorrhyncha: Hemiptera: Fulgoromorpha: Cixiidae) In: Drosopoulos, S. & Tropiduchidae Claridge, M.F. (eds). Sounds and Communication. CRC-Taylor&Francis, Boca Raton,

Lophopidae London, New York: 187-197. Howarth, F.G. & Hoch, H. 2004. Adaptive shifts. In: Culver, D.C. & White, W. (eds). Ency- Eurybrachidae clopedia of Caves. Elsevier Academic Press, Amsterdam etc.: 17-24. Gengidae Sket, B. 2008. Can we agree on an ecological classification of subterranean animals? Journal of Natural History 42: 1549-1563. Cicadomorpha Wessel, A. 2008. Incipient non-adaptive radiation by founder effect? Oliarus polyphemus Fen- Coleorrhyncha nah, 1973 – a subterranean model case. (Hemiptera, Fulgoromorpha, Cixiidae). Doctoral Thesis, Heteroptera Humboldt-Universität zu Berlin, Berlin: 144 pp. Wessel, A., Erbe, P. & Hoch, H. 2007. Pattern and process: Evolution of troglomorphy in troglobitic endogean troglophilic the cave-planthoppers of Australia and Hawai’i–Preliminary observations (Insecta: Hemipte- ra: Fulgoromorpha: Cixiidae). Acta carsologica 36: 199-206. Fig. 7. Systematic distribution of subterranean taxa in the Fulgoromorpha. For the Cixiinae tribes the Wessel, A. & Hoch, H. 1999. Remane‘s statistic species criterion applied to Hawaiian cave number of subterranean species of the total of known species is given in parentheses (modified planthoppers (Hemiptera: Auchenorrhyncha: Fulgoromorpha: Cixiidae). Reichenbachia 33: Fig. 4. Reduviid bug, member of the accompanying fauna, collected in close vicinity of the Bennini. from Hoch et al. 2006). 27-35.