We present here the cockroaches found in 55 nests of Icteridae birds in French Guiana in July 1998. Five species of cockroaches were found, Schultesia nitor (Zetoborinae), Phoetalia pallida (Blaberinae), Pelmatosilpha guianae (Blattinae), Chorisoneura n.sp. aff. gatunae (Pseudophyllodromiidae) and Epilampra grisea (Epilamprinae). The two dominant species, S. nitor and P. pallida, were found together in the same nests, and seem to be scavengers. The ecology of S. nitor was compared with those of S. lampyridiformis, a sister species found in the same type of habitat in Brazil

Deleporte, P.

Grandcolas, P.

van Baaren, J.

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AMAZONIANA XVII ( l/2): 243-248 Kiel. Dezember 2002Cockroaches ¡n French Guiana Icteridae birds nestsbyJ. van Baaren, P. Deleporte & P. GrandcolasDr. Joan van Baaren, UMR 6552, Ethologie Evolution Ecologie, Université de RennesI, Campus de Beaulieu, Avenue du Général Leclerc, 35042 Rennes Cedex" France; e-mail: joan.van-baaren@univ-rennesl.frDr. P. Deleporte, UMR 6552, Ethologie Evolution Ecologie, Station Biologique dePaimpont, 35380 Paimpont, France; e-mail: pierre.deleporte@sbp.univ-rennesl.frDr. P. Grandcolas, ESA 8043 CNRS, Laboratoire d'Entomologie, Muséum Nationald'Histoire Naturelle, 45 rue Buffon, 75005 Paris, France; e-mail: pg@mnhn"fr(Accepted for publication: July, 2001 ).AbstractWe present here the cockroaches found rn 55 nests of Icferidae birds in French Guiana in July I 998. Fivespecies of cockroaches were found, Schultesia nitor (Zetoborinae), Phoetaliu pcl1lrla (Blaberinae),Pelmatosilpha guianae (Blattinae), Chorisoneura n.sp. aff. gutunae (Pseudophyllodromiidae) and Epilum-pra griseo (Epilamprinae). The two dominant species, S nitor and P. pallida, were f'ound together in thesame nests, and seem to be scavengers. The ecology of S. nitor was compared with those o{ S. lontp.rri-difonnis, a sister species found in the same type ofhabitat in Brazil.Key words: Bird nest, cockroach, Schultesia, Zetoborinae, Phoetøliø, Blaberinae.ResumoApresentamos aqui as baratas encontradas em 55 ninhos de pássaros lcteridae na Guiana Francesa em Ji¡lhode 1998. Cinco espécies de baratas foram encontradas: Schultesia nitor (Zetoborinae), Phoetuliu pullida(Blaberinae), Pelmato.silpha guianae (Blattinae), Chorisoneut'a n.sp. aff . gotnnue (Pseudophyllodromiidae)e Epilantpra griseo (Epilamprinae). As duas espécies dominantes, S. ttitor e P. pallitlo foram encontradasjtrntas nos mesmos ninhos e parecem ser necrófagas. A ecologia de S. nitor foi comparada com a de S.lomplridiformis. uma espécie irmã encontrada no mesmo tipo de hábitat no Brasil.lntroductionThe ecology of non-domestic cockroaches is generally still poorly known. particularlyfor Neotropical families. In Neotropical rain forests, Blattaria species inhabit groundlitter where they found abundant but scattered resources, and also patchy habitats likeloose bark of dead or living trees, mould, tree holes, epiphytes, birds nests and antsnests (GRANDCOLAS 1998, 1999). There are very few records of cockroaches takenfrom nests of birds (ROTH & WILLIS 1960, ROTH 1973 and PEREIRA TORRES2001 for South America, VIDLICKA 1999 for Europa). ROTH (1973) found l0cockroach species in pendulous nests of Cacicus birds in aBraztlian tropical forest. Hedescribed a new species placed in a new getJs, Schultesia lampyridiformis, whose maleISSN 0065-675512002/2431 O MPI für Limnologie, AG Tropenökologie. Plön: INPA, Manaus243genitalia supported its belonging to the Zetoborinae subfamily, but whose nondeplanatefonn and marked resemblance to winged adult flrreflies separated it froln other generain this family. The genus Schultesia appeared to be specialised in pendulous nests ofIcteridae birds because two species were found respectively in such nests, one in Brazil(5. lampyridiþr¡ø¡s, ROTH, 1973) and the other in French Guiana (5. nitor. GRAND-COLAS 1993), and nowhere else since then. Recent investigations by Pereira Torres(2001) tend to confirm this fact, Schultesia lampyridiformrs having been found in nestsof the lcteridae bird Psarocolius decumanus but not in nests of eleven non-icterid birdspecies. The pendulous lcteridae nests are located in forest edges (HAVERSCHMIDT1968), which are typically short-lived young environments (REMSEN & PARKERl 983).The aim of this paper is to document the diversity of Blattaria species found in onecolony of pendulous Icteridae nests in French Guiana and discuss their habitat prefer-ences wilh a particular attention to Schultesia and Phoetalia.Material and methodsCollecting was carried out during July I 998. at the end of the rainy season, when reproduction of birdswas low, and in the middle of the day when cockroaches should be resting. The bird coÌony was settledon a big isolated lern¡i¡talia guyanensi.r tree EICHL., heavily covered with epiphytes and containinghundreds of nests of two species of Icteridae birds. the crested oropendola Ps¿trocolius tlecumutnts(PALLAS l7ó9) and the yellow-rumped cacique Cacicus cela (LINNE 1758). On a paltn tree thirty metersaway stood another colony of both species of birds, a much smaller and visibly newly settled group ofseventeen nests. Both trees grew in a rather open area, near the French Guiana coast. close to the Couna-mama creek, emerging high over scrubby vegetation. Thirty eight nests were collected from the biggestcolony, and 17 from the smallest colony, all hanging from the same branch or palm. The nests wereindividually packcd in closed plastic bags and were soon carefully inspected, each cockroaclr founcl wasmeasured (total length) and its species and sex identified. The content ofeach nest, besides cockroaches.was also noted.The nests could be attributed to each ofthe two bird species because oftheìr different structure. Nestsofcrested oropendola had their opening on top ofthe nest. while nests ofyellow-rumped caciques had alateral opening, about I 5 cm below the top. Nests of both species were between 60 to I 00 cm long. andhad walls about 0.5 cm thick. They consisted of long and thin woven blades of grass. The bottom wasframed with an accumulation ofshort blades ofgrass. about five cm thick. looking like a "nest in the nest".Some nests lacked this kind of thick bottom.ResultsThe five species of cockroaches found in these nests were: Epilampra grisea DE CEER1773 (Blaberidae, Epilamprinae), Schultesia ¡¿ilor GRANDCOLAS l99l (Blaberidae,Zetoborinae), Pelmatosilpha guianae HEBARD 1926 (Blattidae), Chorisoneura n.sp.aff .gatunae HEBARD l92l (Pseudophyllodrorniidae). and Phoetalia pallida BRUNNERVON WATTENWYL 1865 (Blaberidae, Blaberinae) (Table l),Among the l7 nests collected from the small colony, all were uninhabited by birds,but two of them contained a mouse nest (one with a pregnant female mouse of undeter-mined species):One yellow-rumped cacique nest contained one adult female of Epilam-pra grisea. One nest of crested oropendola contained a rotten egg, with lnaggots, andtwo adult males of Schultesia nitor.Among the 38 nests collected from the largest colony, l3 were from crested oropen-dola, five of them contained cockroaches, and two of the latter contained also fragments244of rotten eggs and young birds dead a long time ago. The 25 other nests were fromyellow-rumped cacique, ten of them contained cockroaches, and three of the lattercontained fragments of addled eggs and old corpses of young birds, a fourth onecontaining fleshless remains of a dead adult bird with two eggs, all completely dried.Four species of cockroaches were found in this sample. First, two individuals of Pelnta-tosilpha guianae were found, one last instar nymph in a crested oropendola nest and oneadult female in a yellow-rumped cacique nest, oçcurring in both cases with other speciesof cockroaches in the same nest. Second, Chorisoneura n.sp.aff. gatunae was found inone crested oropendola nest (six individuals together) and in six nests of yellow-rurnpedcacique (only one individual in each, and some oothecae attached to the walls of thenests). Although they were generally found in the same nests with other species ofcockroaches (five observations out ofseven), they did not occur on the bottom with theother species, but higher on the walls of the nests.The third species, Phoetalia pallida, was found in one nest of crested oropendolaand in four nests of yellow-rumped cacique, always in the bottom part of the nest andassociated with individuals of S. nitor. Tbe number of individuals found per nest washighly variable, respectively two nymphs; one nymph; four nymphs; one adult felnalewith her 26 newly born first instar nymphs; and two adult females together with tennymphs. In the latter case the ten nyrnphs were different sizes, ranging from ó tol7mm, and thus could not come all from the same oothecae.The fourth species, S. nitor, was found in five nests of crested oropendola and innine nests of yellow-rumped cacique (thus this species was found in 37 percent of thenests from this big bird colony, and in all but one nests containing other cockroachspecies) (Table l). The number of individuals per occupied nest was 2.6 ! 0.3, with nosignificant difference between nests of crested oropendola and yellow-rumped cacique(Student t-test: t = 1.17, P = 0.27). Thirty five nymphs were found, whose lengthsranged from 5 to 14 mm. There was no difference in cockroach sex ratios betweennests, with 1.4 ! 0.2 rnales and 1.1 t 0.3 females in each nest (Student t-test: t = 0.77,P = 0.45). In most nests with three or more larvae, the high differences in lengthindicated that all the individuals could not come frorn the same oothecae. The smallestnymphs were generally found closer to the top of the nest than were the biggest ones.One adult male was found alone in a nest, and I adult female was found with two lastinstar nymphs.The nest of yellow-rumped cacique containing remains of a dead female bird andtwo eggs contained the most cockoaches. Some S. nitor and P. pallida occuffedcompletely inside the dry skin of the dead bird, and all of them were last instar nymphs.A Chi-square test showed that the cockroaches were significantly more numerous innests containing remains of birds or eggs (Xtr,r= 13.12, P < 0.0001). Nests withoutthick bottoms were empty of cockroaches.DiscussionROTH (1973) found cockroaches belonging to ten different species (and 6 differentgenera) in pendulous nests of Cacicus birds in Brazil. In our study, we found only fìvespecies belonging to five different genera, three of thern different from those found byROTH (1973). Only two genera, Schultesict and Chorisoneura, were found in bothstudies (Table l).We found two individuals of Pelmatosilpha guianae in nests of the largest bird245cofony. P. guianae and other species of the salne genus had already been found inepiphytic plants (ROCHA E SILVA & RODRIGUES LOPES 1976: DELEPORTE,GRANDCOLAS unpubl. data; DEJEAN pers" comm.), under loose bark and in decayinglogs (ROTH & WILLIS 1960). In our study, the two P. guianae likely came frornsurrounding epiphytes, which were abundant in the Terminalia tree supporting the bigbird colony and absent frorn the palm tree supporting the smaller one.In our study, two species, P. pallída and S. nitor, seelned to be more abundant in thenests. Very little is known from other studies about P. pallida" GRANDCOLAS ( 1998)placed this species in the family Blaberinae, in a phylogenetically non resolved groupwith the genera Hormetica, Parahormetica, Brachycola and Bionoblqtta. ln this group,only the habitat of Hormetica is known and consists in epiphytes and palms, like thatof the recently described Lucihormeticafenestrqtq ZOMPRO & FRITZSCHE, 1999" Itis likely that P. pallida inhabits epiphytes and comes occasionally in Cacic¿¿s nests.Nothing is known about the social life or the anti-predator behaviour of P. pallida. Ourdata suggest that this species is gregarious, because all 46 individuals (among which 26first instar nymphs) were found in only five nests among 55 inventoried ones. More-over, more than one individual were found together even in nests lacking evidentattractive food resources like remains of birds or eggs. The facts that significantly rnorecockroaches were found in nests with remains of birds or eggs, and that sorne individu-als were found inside the remains of a dead bird indicate that this species could be ascavenger, occasionally found in Cacicus nests when they happen to offer some foodresource, VIDLICKA (1999) found young larvae of two species of Ectobiinae cock-roaches in birds nest in Slovakia and supposed that they looked for rernains ofbird foodand probably protection during winter).Little information is available about S. nitor. The description of the species wasmade from individuals collected with light traps. The only previous observations of S.nitor in its natural habitat were GRANDCOLAS (1993) and Deleporte (unpubl. data)who analysed respectively in July and September l99l the content of pendulous nests(4 nests each time) from the same abandoned colony of Cacicus haemorhou.s in FrenchGuiana. The nests were located at ten meters height on an isolated tree and were emptyof birds. No ,S. ¡¿i¿or were found on the surrounding epiphytes. Only one to two adultor larvae were found inside two of the four collected nests each time. The only otherknown species of the same genus, S. lampyridiþrtøls, is morplrologically very sirnilar(GRANDCOLAS l99l) and colonises the same habitat, which seems specific to thisgenus. ROTH (1973) analysed the S. lampyridiformis individuals found in three coloniesof Cacicus sp. birds and noted that most of the nests were apparently abandoned and afew had the remains of dead young birds. He concluded that the cockroaches wereprobably scavengers and may have inhabited the nests while the birds occupied thecolony. ROTH (1973) found in the first colony 4 males, 8 females and 29 nymphs ofS. làLmpyridiformis in one nest, one nymph alone in a second nest, four males, threefemales and seven nymphs in a third nest and six nymphs alone in a fourth nest. All ofthe nests containing S. lampyridiforø¡¡ individuals contained also Chorisoneura sp.Inthe second colony, 5 males, 2 females and l9 nymphs of S. lampyridiformis were foundin eight nests with three blattellid nymphs, one of them belonging to Chorísoneurainversa HEBARD. In the third colony (60 feet from the ground), in several nests twomales, three females and one nymph of S. lampyridiformis were found with individualsof Dendroblatta cnephaia Hebard and Chorisoneura sp. ROTH (1973) concluded that246the bird nest habitat is undoubtedly a normal habitat for the genera Schultesia and,Chorisoneura, which our study tends to confirm.In conclusion, our new data about S. nitor suggest that the whole genus Schultesiais strictly linked to pendulous bird nests. Recent srudies by PEREIRA TORRES (2001)tend to confirm this fact: non-icterid bird nests contained no SchultesÌa cockroaches butharboured different Epilampridae, Blattellidae, Pseudophyllodromiidae, Polyphagidaeand Blattidae cockroaches, while one blattellid cockroach and I I I Schultesia lampyridi-/'ormis were found in icterid bird nests. Not only this habitat presents a particularphysical sttucture (an intetwoven network of grass fibres), but the nests systematicallystand in isolated or elne[gent trees, being thus more exposed to climatic factors thancomparable habitats standing inside the continuous forest canopy. Some strong rnorpho-logical differences between schultesia and other genera of the same lineage (GRAND-COLAS l99l) could be linked to the colonisation of this particular habitat. This isparticularly likely for the slender general body shape and for the colour pattern: to thehuman eye, the light yellowish longitudinal stripes render the individuals difficult todiscem among the interwoven fibres.AcknowledgmentsWe are grateful to Roseli Pellens lor translating the abstract into Portuguese, and to the three anonymousreferees for helpful comments on an earlier version of this manuscripf. We are much indebtcd to Prol-.Joachiln Aclis for very uselul information and suggestions. The experiments cornply with the current lawsof France.ReferencesGRANDCOLAS, P. (1991): Descriptions de nouvelles Zetoborinae guyanaises avec quelques remarquessur la sous-famille. - Bull. Soc. Ent. Fr.95 241-246.GRANDCOLAS, P. (1993): Habitats of solitary and gregarious species in the Neotropical Zetoborinae(lnsecta, Blattaria). - Stud. Neotrop. Fauna & Environm. 28: 179-190.GRANDCOLAS, P. (1998): The evolutionary interplay of social behavior, resource use and anti-predatorbehavior in Zetoborinae+Blaberinae+Gyninae+Diplopterinae cockroaches: A phylogenetic analysis. -Cladistics 14: 1 17-127.GRANDCOLAS, P. ( 1999): The origin of diversity in cockroaches: a phylogenetic perspective of sociality.reproduction, communication and ecology. - In: MELIC, 4., DE HARO, J.J., MENDEZ. M. & LRIBERA (eds): Evolucion y filogenia de Arthropoda. Boletin de la S.E.A., Zaragoza26:397-420,HAVERSCHMIDT, F. (1968): Birds of Surina¡¡. - Olivier & Boyd, Edinburgh.PEREIRA TORRES, W.M. (2001): Fauna de artropodes associada a ninhos de aves e mamiferos etr umafloresta de terra firme na Amazonia Central, Brasil, Dissertação, Universidade do Amazonas, INPA,Manaus.REMSEN, J.V. & T.A. PARKER (1983): Contribution of river-created habitats to bird species richness inAmazonia. - tsiotropica 15: 223-231.ROCHA E SILVA, I. & S.M. RODRIGUES LOPES (1976): Blattaria de bromelia (Dictyoptera). - Rev.Bras. biol. 36: 873-901.ROTH, L.M. (1973): Brazilian cockroaches lound in birds'nests. with description ofnew genera andspecies. - Proc. Ent. Soc. Wash.75: l-27.ROTH, L.M. & E.R. WILLIS (1960): The biotic associations of cockroaches. -The Lord Baltirnore Press,Baltimore.VIDLICKA, L. (1999): Cockroaches (Blattaria) in bird nests. - Folia faunistica Slovaca 4:41-43.247ZOMPRO,O.&I.FRITZSCHE(1999): Lucihornetiutfènestratu n.gen.,n.sp..thefìrstrecorc{ oflunrines-cence in an orthopteroid insect (Dictyoptera: Blaberidae: Blaberinae: Brachycolini). - Amazoniana l5(3/4\:2ll-219.Table l: Cockroaches found in birds nests in our studv and in that of ROTH ( I 973).Our study ROTH's studv (1973)Cockroach genera found in birds nestsSchultesioChorisoneuroE¡tilampruPelmato,silphaPhoetalioSchultesiaChorisoneuraAlpheliriaAtttuxninaDendrohluttuLophohleÍteData concerning the genus Schultesia onlyFirst colony:Nest l: 4 adult males, 8 adult females"29 nyrnphs.Nest 2: i nymph.Nest 3: 4 adult males. 3 aclult fenrales.7 nymphs.Nest 4: 6 nymphs.Second colony (8 nests): 5 adult males.2 adult females, l9 nymphs.Third colony (several nests): 2 adult males3 adult females, I nymph.Small bird colony: 2 adull males.Large bird colony:- S. nitor was found in 37 Vo of the3[J studied nests.- Total number of indiviclLrals:I adult male, I adult female, 35 nyrlphs.- Mean number per nest: 2.57 t 0.12 ind.,I .43 ! 0.23 males and I . l4 t 0.30 females.- Maximum number in a nest: 5 individuals.248

Cockroaches in French Guiana Icteridae birds nests

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