An Australian immigrant in southern California: Paraphloeostiba gayndahensis (Coleoptera: Staphylinidae) Margaret K. Thayer, Field Museum of Natural History, Chicago, IL, USA

Table 1. Known habitats and microhabitats of Paraphloeostiba gayndahensis by geographical area. All data for each country are pooled (i.e., subunits are not correlated with other data). ABSTRACT COUNTRY Habitats Microhabitats or collecting methods Paraphloeostiba gayndahensis (MacLeay, SUBUNITS 1871) was described from and is believed AUSTRALIA ACT (AUSTRALIAN CAPITAL tropical & subtropical rainforest, fallen rotting fruit of various rainforest trees: native to Australia, where it is widely TERRITORY) including open edges Ficus spp. (Moraceae), Endiandra introrsa NEW SOUTH WALES subtropical rainforest- (Lauraceae), Schizomeria ovata distributed in native forests. It has been QUEENSLAND Argyrodendron (Cunoniaceae), Brachychiton sp. SOUTH AUSTRALIA littoral rainforest (Sterculiaceae), and (unidentified) purple collected in New Zealand since 1941 and VICTORIA dry sclerophyll forest fruits on rainforest floor successively in parts of Italy, France, Spain, wet sclerophyll forest fallen female cones of Bunya pine (Araucaria Eucalyptus forest bidwillii), hoop pine (Araucaria Portugal, and Germany since 1988. It is also open hardwood-Araucaria grove cunninghamii), Queensland kauri (Agathis established in southern California since at Araucaria plantation w/Ficus robusta) (all Araucariaceae) Nothofagus cunninghamii forest in male cycad cone least 1995 in the vicinity of Santa Barbara. disturbed tall forest rotten apple on gravel modified bush ripe peaches (harvested) Although its chief microhabitat appears to be in town moist rotting fruit stalk Xanthorrhoea fermenting plant matter of various kinds and (Xanthorrhoeaceae) damp patch of decaying banana tree fronds & (in Australia) inflorescences of Araceae, in detritus in & under rotten mushrooms California it has been implicated as a berlese, fine wood debris under Nothofagus welcome pollinator of cultivated Annona logs litter under flying fox [fruit bat] roost cherimola, the cherimoya. The species sieved litter breeds very abundantly in rotting fallen fruits in flowers of cunjevoi/“Arum” inflorescences (Alocasia macrorhizos Schott [Araceae]) of various trees. Laboratory rearing was on flowers Amgeboi (?) pyrethrin-fogging Araucaria cunninghamii bark successful through the full life cycle; both human dung pitfall traps adults and larvae fed on rotting cherimoya, grass pile pitfall in compost bin rotting banana and the mold growing on it, rotting food scraps Drosophila sp. adults, and yeast-based rainforest canopy (25m) intercept trap flight intercept & malaise traps Drosophila rearing medium. Fig. 1 Paraphloeostiba flying at dusk truck [net] trap 0.5-2m above ground gayndahensis, female; yellow pan traps body length ca. 2.5 mm. FRANCE ARIÈGE subalpine zone badger droppings CORS E (CORSICA) greenhouse (hothouse) in rotting vegetables/plant matter HAUTE-GARONNE (surroundings not specified) rotting papaya on ground DESCRIPTION AND RECORDS LOT traps baited with melon peels PYRÉNÉES- in banana flower Omalium gayndahense (Fig. 1, adult) was one of 550+ beetle species described by OCCIDENTALES[?] under bark fallen Eucalyptus logs (Corse) Fig. 6. European records of Paraphloeostiba gayndahensis, 2001. PYRÉNÉES-ORIENTALES car net MacLeay (1871) from the vast collections made by Masters near Gayndah, Fig. 3. Australian records of Paraphloeostiba gayndahensis, 2001. VAR Years shown are earliest known for a site (if any); some Italian Queensland, Australia (see Fig. 3), where he had been sent to collect lungfish Years shown are earliest known for a site (if any). GERMANY records date from 1988. NORDRHEIN-WESTFALEN Bruchwald [swamp forest] window trap (Monteith 2001). (Fraxinus excelsior, Alnus glutinosa, hybrid Populus) Bernhauer (1926) synonymized the species with Phloeonomus singularis (Kraatz), NATURAL HISTORY ITALY but Steel (1960) resurrected it and transferred it to his new genus Paraphloeostiba ABRUZZO [no information] in more or less rotting pomelos [citrus?] BASILICATA in decaying matter (often fruits) with many other Asian and Australasian species previously placed in Phloeonomus Steel (1960) provided the first data on the biology or habits of P. gayndahensis: specimen(s) CALABRIA pitfall traps or newly described. He provided collection records from Queensland, New South LIGURIA labeled as “bred from hoop pine cones” [Araucaria cunninghamii] in Queensland and also from LOMBARDIA Wales, and Lord Howe Island. I have subsequently collected or seen specimens Nowra, NSW, “on Datura arborea” (flowers or foliage??). PUGLIA SARDEGNA from South Australia, Victoria, and the Australian Capital Territory as well. SICILIA VENETO New Zealand Kuschel (1990) first recorded P. gayndahensis from outside Australia, Recent records (published and not) have provided far more biological information, summarized in Table 1. In Australia, P. gayndahensis occurs in a tremendous variety of habitats, but mainly NEW ZEALAND - from the Auckland area, based on specimens collected as early as 1941 (published AOTEAROA apple orchard in grass clippings native forests; elsewhere it is mostly or entirely in human-disturbed areas. Within these NORTH ISLAND kiwi fruit orchard old bracts of Musa inflorescences erroneously as 1944) up through the 1980s. Recent collections suggest its range is suburban/bush mix various decaying fruits on ground still expanding. habitats, members of the species occupy varied microhabitats, but most are in fermenting plant in compost bin on Uromycladium notabile (Acacia rust) galls material (or other decay situations) or in flowers (or have been caught in flight). These are of Acacia mearnsii Europe Zanetti (1995) published the first European occurrence, from northern and typical habits for tropical or subtropical members of the Phloeonomus group of genera, and I PORTUGAL southern continental Italy, and also Sicily, but without specific localities or dates; have reared P. gayndahensis from the rotting flowering/fruiting stem of a Xanthorrhea (“grass MADEIRA laurel forest sifted more recently it has also been found in Sardinia (A. Zanetti, email December 2001). tree”). In contrast, members of the Phloeonomus group found in the Holarctic Region live SPAIN almost exclusively under the bark of logs. The only such record for P. gayndahensis is from ISLAS CANARIAS banana plantation attracted to beer Since then, an increasing torrent of European records has appeared, suggesting a (La Gomera, La built-up area, near beach on rotting orange Palma) sifted from plant waste continuing expansion in Europe as well: Corsica, interestingly from logs of Eucalyptus, nearly all species of which are native to Australia; this, however, seems like to be coincidental since P. gayndahensis has not turned up in UNITED STATES CALIFORNIA orchard including cherimoya, in rotting cherimoya & other fruits on ground • France (Assing 1996, including Corsica; Rogé & Dauphin 1997; Brustel & Rogé extensive subcortical collecting in Australia. Tronquet (2000) reported a distinct reduction in (Santa Barbara Co.) avocado, and apricot trees leaf litter around fruit trees 1998; Mineau 1998; Tronquet 1998, 2000; Letellier & Dauphin 1999) in staminate phase cherimoya flowers (at populations of two native Proteinus species (Staphylinidae: Proteininae) in localities where P. dusk) • Germany (Assing 1996, Kohler 1997) gayndahensis has become established.

• Spain: Canary Islands (Assing 1996, Assing & Wunderle 1999) In July 2000, I visited the orchard near Santa Barbara to investigate Paraphloeostiba there. They were breeding profusely,with numerous Nitidulidae, in all the types of fallen fruits we • Portugal: Madeira (Assing 1996) REFERENCES checked, as well as in cherimoya flowers during their early-evening staminate (pollen-shedding) Assing, V. North America In 1996, botanist C. A. Schroeder of UCLA (since deceased) sent me phase. The orchard’s main crops are avocado, cherimoya, and apricot, with small numbers of 1996. Zur Kenntnis und gegenwärtigen Verbreitung von Paraphloeostiba gayndahensis (Macleay), einer nun for identification some little staphylinids found in the flowers of cultivated cherimoya orange and banana trees. Adults were abundant and very active during the day; larvae were auch für Deutschland erstmals nachgewiesenen Adventivart (Col., Staphylinidae). Entomologische Nachrichten und Berichte 40: 179-181. (Annona cherimola) near Santa Barbara, California. They appeared to be pollinating present, but less numerous at that season. I collected a promising rotting cherimoya containing Assing, V., J. Frisch, M. Kahlen, I. Löbl, G. H. Lohse, V. Puthz, M. Schülke, H. Terlutter, M. Uhlig, J. Vogel, P. the cherimoya flowers, which otherwise require hand pollination for commercially Wunderle, and L. Zerche. adults and at least a few larvae for rearing, and reared a full generation (adults to adults) in 1998. 23. Familie: Staphylinidae, pp. 119-197. In: Lucht, W. and B. Klausnitzer, Die Käfer Mitteleuropas. Band adequate fruit set. He and the grower were thus very interested in what was known about one month with little trouble. Steel (1970) reported that the two species of Phloeonomus 15. 4 Supplementband. Goecke & Evers, Krefeld. 398 pp. Assing, V. and P. Wunderle. of the species’ biology and what might be needed to maintain and increase the and one species of Phloeostiba that he reared did not feed on insect food, unlike most 1999. On the Staphylinidae of La Palma, Canary Islands, with descriptions of new species and remarks on some population. Although the beetles appeared to me indistinguishable from Australian Canarian Stenus Latreille 1796 (Coleoptera). Entomologische Zeitschrift 190: 433-443. Omaliinae (Steel 1970, Thayer 1985a, b, unpublished) Bernhauer, M. P. gayndahensis, that very unexpected identification seemed unbelievable, since 1926. Fauna Buruana. Zur Staphylinidenfauna der Molukken. Treubia 7: 311-327. Brustel, H. and J. Rogé. the (equally startling) European records were not yet available. The combination of 1998. Sur quelques Coléptères rare ou peu connus du sud-ouest de la France. Entomologiste 54: 203-209. all these records, however, makes it clear that P. gayndahensis is indeed a world Fig. 4. New Zealand records of Paraphloeostiba gayndahensis, 2001. Ciceroni, A., V. Puthz, and A. Zanetti. Years shown are earliest known for a site (if any). 1995. Coleoptera, Polyphaga III (Staphylinidae). In: Minelli A., Ruffo S. & La Posta, S., eds., Checklist delle traveler. The California record was briefly noted in Newton et al. (2000), and it specie della fauna italiana. Fasc. 48. Calderini, Bologna. 65 pp. seems likely that the species has spread some at least locally. Kohler, F. 1997. [Comments on the beetle fauna of the Rhine province 11.] Mitteilungen der Arbeitsgemeinschaft Rheinischer Koleopterologen 7: 29-51. All the records are mapped together in Fig. 2, and in more detail in Figs. 3-6. Kuschel, G. 1990. Beetles in a suburban environment: a New Zealand case study. The identity and status of Coleoptera in the natural and modified habitats of Lynfield, Auckland (1974-1989). DSIR Plant Protection Report 3: 118 pp. Letellier, Y. and P. Dauphin. 1999. Présence de Gironde de Paraphloeostiba gayndahensis (Macleay) (Coleoptera Staphylinidae). Bulletin Fig. 7. Paraphloeostiba gayndahensis, larva. de la Société Linnéenne de Bordeaux 27: 80. MacLeay, W. 1871. Notes on a collection of insects from Gayndah. Transactions of the Entomological Society of New South Rearing this species during field work in Australia in 1987, I found the beetles took about 2 weeks to Wales 2: 79-205. Mineau, A. develop from pupa to adult. Both adults and larvae ate insects and grazed on the rotting debris in 1998. Capture de plusieurs spécimens de Paraphloeostiba gayndahensis McLay dans le Var (Coleoptera which they were collected (Xanthorrhoea flowering/fruiting stalk) but I was unable to tell exactly what Staphylinidae Omaliinae). Entomologiste 54: 272. Monteith, G. B. they were eating. 2001. Gayndah, lungfish and beetles -- a fishy tale. C. T. White Memorial Lecture for 2000. Queensland Naturalist 39: 5-32. Newton, A. F., M. K. Thayer, J. S. Ashe, and D. S. Chandler. Observation of the California specimens showed that both adults and larvae (Fig. 7) will eat adult 2000. Family 22. Staphylinidae Latreille, 1802, pp. 272-418. In: Arnett, R. H., Jr. and M. C. Thomas, eds., American Beetles, Volume 1, Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Drosophila (freeze-killed, cut open), but are not restricted to insect food. In addition to the field- Press, Boca Raton, FL. xv + 443 pp. collected fruit, I also provided bits of banana (as Steel did for Phloeonomus), which both adults and Rogé, J. and P. Dauphin. 1997. Sur la présence en France de Paraphloeostiba gayndahensis (Macleay) ( Coleoptera Staphylinidae). larvae ate, or at least grazed on. Small spherical black sporulating mold structures appeared on these Bulletin de la Société Linnéenne de Bordeaux 25: 115-116. Steel, W. O. bits of fruit, and the larvae, especially, ate the black mold bodies avidly. The field-collected fruit had a 1960. Three new Omaliine genera from Asia and Australia previously confused with Phloeonomus Thomson. thriving nematode population, but not for very long – adults and larvae were both very interested in Transactions of the Royal Entomological Society of London 112: 141-172. 1970. The larvae of the genera of Omaliinae with particular reference to the British fauna. Transactions of the them. The larvae, at least, seemed to slurp down the shiny little nematodes like sucking up spaghetti Royal Entomological Society of London 122: 1-47. Thayer, M. K. strands, and reacted more strongly (moving faster), to new pieces of fruit when those contained 1985a. The larva of Brathinus nitidus LeConte and the systematic position of the genus (Coleoptera: nematodes or mold spherules. Both adults and larvae also ate yeast-based “instant” Drosophila culture Staphylinidae). Coleopterists Bulletin 39: 174-184. 1985b. Micralymma marinum (Stroem) in North America: biological notes and new distributional records medium, but preferred the other foods mentioned if those were available. Some larvae (starting at (Coleoptera: Staphylinidae). Psyche 92: 49-55. instar I or possibly II) developed successfully on the yeast medium alone. One month after the initial Tronquet, M. 1998. Staphylins intéressants ou nouveaux pour les Pyrénées-Orientales. 1re Note. Entomologiste 54: 9-16. collection, there were still first instar larvae hatching in a culture of field-collected adults isolated from 2000. A propos de Paraphloeostiba gayndahensis (MacLeay 1871) (Col. Staphylinidae). Entomologiste 55: the original fruit and any included eggs. The success with which this species is breeding on its own in 234.[1999] Fig. 5. Southern California records of Paraphloeostiba the orchard suggests that all the grower needs to do to help them is leave fallen fruit on the ground. gayndahensis, 2001. Years shown are only collections known. Fig. 2. Known World distribution of Paraphloeostiba gayndahensis, 2001. ACKNOWLEDGMENTS I thank the late Art Schroeder for contacting me about what turned out to be such interesting little beetles, providing information about cherimoya life cycles, and helping to arrange my visit to California Tropics Ranch. I am grateful to Rich Leschen (Landcare Research, Auckland) for kindly providing additional New Zealand records on short notice, and Adriano Zanetti (Museo Civico di Storia Naturale du Verona) for generously allowing me to map numerous Italian records from his own work in progress. Kris Jacques shared his discovery of the occurrence of P. gayndahensis on Alocasia inflorescences. Avis James and Oliver Betz, both then at the Field Museum, provided the idea (and provisions) to try Drosophila rearing medium as a food source. The Bottomley family of Westlake Village, CA provided a well-timed opportunity to check out the “cherimoya beetles” as part of a most enjoyable visit. My Australian fieldwork contributing to this has been supported by numerous sources in the USA and Australia, acknowledged in past and future publications, and I thank Al Newton for sharing in all the field work and much more.