Results of a biological inventory of the Nosy Ankao island group, Parc National de Loky-Manambato, northeastern Madagascar Steven Goodman, Yasser Anbdou, Yannie Andriamiarantsoa, Brian L Fisher, Owen Griffiths, Brad Keitt, Jean-Jacques Rafanomezantsoa, Eric Rajoelison, Jean-Claude Rakotonirina, Luc Ranaivoarisoa, et al.

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Steven Goodman, Yasser Anbdou, Yannie Andriamiarantsoa, Brian L Fisher, Owen Griffiths, et al.. Results of a biological inventory of the Nosy Ankao island group, Parc National de Loky-Manambato, northeastern Madagascar. Malagasy Nature, Association Vahatra, 2017, 11. ￿hal-01493362￿

HAL Id: hal-01493362 https://hal.archives-ouvertes.fr/hal-01493362 Submitted on 27 Mar 2017

HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Results of a biological inventory of the Nosy Ankao island group, Parc National de Loky-Manambato, northeastern Madagascar

Steven M. Goodman1,2, Yasser Anbdou3, Yannie Andriamiarantsoa4, Brian L. Fisher5,6, 13 7, Brad Keitt8, Jean Jacques Rafanomezantsoa6, Eric T. Rajoelison6, Jean Claude Rakotonirina6, Luc Ranaivoarisoa4, Patrick Ranirison4,9, Voahangy Soarimalala2,10, Michaël L. Tantely11, Pablo Tortosa12 & Achille P. Raselimanana2,13 Résumé détaillé 1 2 3 Plantes vasculaires – 7 10 11 12 Capurodendron greveanum et Eugenia aff. calciscopulasum

Malagasy Nature 2 Goodman et al

Mystroxylon aethiopicum et Diospyros erythrosperma et Mimusops coriacea et Ficus tiliifolia Moustiques – Mystrium, Xymmer, Proceratium et Tanipone ont Culex tritaentiorhynchus Aedes lambrechti Aedes Culex Anopheles et Aedeomyia ne et amphibiens – Aedes Culex Anopheles Plasmodium Anopheles arabiensis Fourmis – Oiseaux – Goodman et al 3

et al. Petits mammifères – et al. et al., et al., et al. et al., Rattus rattus et R. et al.et al., norvegicus Conclusions – Introduction et al. Goodman et al

et al., 2006 et al. et al. et al.

Figure 1. Goodman et al

& Gordon, et al. et al bay on the north-east side of Madagascar, which was Historical exploration of the biological not known to the natives by any particular name, we called Port Leven, the islands between Andrava and resources of the Nosy Ankao island group Looké [=Loky] Bays… et al et al.

Figure 2. Goodman et al

Methods Context of the 2016 biological inventory and study sites

Table 1.

Name Study group Home institution Yasser Anbdou Ants

Table 2.

Nosy Ankao Nosy Manampao Nosy Ratsy Nosiborona 7.1 Yes Some Yes Some Yes 1 2 Yes Yes Yes Yes Yes Yes 1 2

Table 3.

Taxonomic group Nosy Ankao Nosy Manampao Nosy Ratsy Nosiborona Ants Birds Goodman et al 7

Figure 3.

Study sites Islands (geographical coordinates based on centroid positions) Villages on Nosy Ankao Weather data Goodman et al

Figure 4.

Luc Ranaivoarisoa and assisted by Yannie Andriamiarantsoa and Patrick Ranirison with Data collection

Linear transects et al. Goodman et al

Figure 5.

Surface area sampling General botanical collections 10 Goodman et al

Table 4.

Group Capurodendron Group Mystroxylon Group Mimusops Floristic group greveanum and Eugenia aethiopicum and Diospyros coriacea and Ficus aff. calciscopulasum erythrosperma tiliifolia 3 2 Tr 10, Tr 11, Tr 12

Ecological Substrate parameters Florisitic 22 7 parameters Structural parameters Tr 11, Tr 12 Tr 10 110 127

Spatial parameters and Southern domains Eastern domains

Plant life-forms Data analysis Floristic richness Geographic range of plant Goodman et al 11

et al. Fisher, Jean Jacques Rafanomezantsoa, Eric Tsiriniaina Rajoelison, and Jean Claude groups Inventory methods Non-selective method

Table 5.

Distance to Latitude south Longitude east Island Elevation Period Habitat nearest point on 20 m 22 m forest 12 Goodman et al

Sample processing and data analysis ), which has up-to-date summaries of what is known about the of Malagasy ants, Selective method

Figure 6. Goodman et al 13

= the AB et al.,

Figure 7. Goodman et al

Figure 8.

Inventory methods Capture of preimaginal stages et al

Handling of mosquitoes et al. et al Aedes Skusea, Anopheles gambiae Capture of adult stages et al et al. Goodman et al

Figure 9.

Possible Plasmodium infection of mosquitoes et al Plasmodium et al et al P. falciparum and P. vivax, both

Reptiles and amphibians (Achille P. Direct observations Inventory techniques Goodman et al

Systematic searching Pandanus and Ravenala Pitfall traps

Figure 10. Goodman et al 17

et al et al et al Small mammals (Yasser Anbdou, Steven M. Goodman, Voahangy Soarimalala, and Pablo Other aspects Live traps Specimens Liophidium aff. torquatum Goodman et al

Results Weather data

Table 6. Global vegetational characteristics Floristic diversity Minimum Maximum Date temperature temperature 33 1 32 33 1 33 0 23 33 0 0 33 0 31 32 0 33 Life-forms spectrum of the vegetation

Vascular plants General aspects Goodman et al

Figure 11.

Figure 12. 20 Goodman et al

Figure 13.

Table 7

Nosy Family and species Nosy Ankao Nosy Ratsy Nosiborona Manampao Acanthaceae Hypoestes strigata Benoist Amaranthaceae Lagrezia ambrensis Anacardiaceae Poupartia gummifera Annonaceae Monanthotaxis pilosa Apocynaceae Cynanchum arenarium Landolphia gummifera Gonocrypta s Pervillea Secamone aff. Secamone capitata Bignoniaceae Rhodocolea racemosa Boraginaceae Cordia myxa L. Capparaceae Crataeva excels Celastraceae Brexia aff. humbertii Brexia madagascariensis Loeseneriella rubiginosa Maytenus undata Mystroxylon aethiopicum Loes. Combretaceae Combretum coccineum Commelinaceae Commelina aff. madagascarica Convolvulaceae Ipomoea rubens Cyperaceae Cyperus betafensis Cyperus longifolius Asparagaceae lanceolata Goodman et al 21

Table 7 Nosy Family Genus and species Nosy Ankao Nosy Ratsy Nosiborona Manampao Ebenaceae Diospyros erythrosperma Diospyros aff. ferrea Diospyros pervilleana Diospyros platycalyx Diospyros pruinosa Diospyros quercina Diospyros squamosa Erythroxylaceae Erythroxylum retusum Erythroxylum sphaeranthus Erythroxylum platyclados Erythroxylum rignyanum Euphorbiaceae Euphorbia tetraptera Margaritaria anomala Thecacoris aff. perrieri Leandri Fabaceae Albizia arenicola. Dichrostachys richardiana Senna occidentalis Loganiaceae Strychnos Lythraceae Capuronia benoistii Malvaceae Grewia aff. barorum Grewia betulaefolia Hibiscus megistanthus Meliaceae Malleastrum boivinianum Moraceae Ficus lutea Ficus tiliifolia Myrtaceae Eugenia aff. calciscopulorum Eugenia aff. oligantha Ochnaceae Ochna ciliata Lam. Oleaceae Jasminum greveanum Noronhia seyrigii Orchidaceae Angraecum aff. magdalenae Primulaceae Oncostemum Oncostemum Rhamnaceae Scutia myrtina ( Rhizophoraceae Cassipourea microphylla Rubiaceae Coffea boiviniana Coptosperma Empogona ovalifolia Ixora aff. ripicola Ixora emirnensis Pyrostria sambavensis Tarenna grevei Tricalysia ambrensis Rutaceae Vepris aff. arenicola Salicaceae Casearia nigrescens Sapindaceae Allophylus cobbe Macphersonia gracilis hildebrandtii Sapotaceae Capurodendron greveanum Mimusops coriacea Sideroxylon aff. saxorum Vitaceae Cissus microdonta 22 Goodman et al

Structural groups Floristic groups

Figure 14.

Table 8.

Indicator Group Family Taxa Group designation Island Sapotaceae Capurodendron greveanum Capurodendron greveanum 1 and Eugenia aff. calciscopulasum Myrtaceae Eugenia aff. calciscopulasum Celastraceae Mystroxylon aethiopicum 100.0 Mystroxylon aethiopicum and 2 Ebenaceae Diospyros erythrosperma Diospyros erythrosperma Sapotaceae Mimusops coriacea Mimusops coriacea and Ficus 3 Moraceae Ficus tiliifolia tiliifolia Goodman et al 23

Figure 15.

Table 9.

Transects Floristic group Structural group Vegetation group A 1 Tr 11, Tr 12 B 2 Tr 10 3 B

Table 10.

Group Name of vegetational group Structural form Island Capurodendron greveanum and Eugenia aff. calciscopulasum Mystroxylon aethiopicum and Diospyros erythrosperma Mimusops coriacea and Ficus tiliifolia

Description of vegetation groups Group and aff. calciscopulasum (Group I)

Structural characteristics of Group I Capurodendron

Structural variant 1: Low-growing forest with closed Floristic characteristics of Group I understory -- Goodman et al

Figure 16. Capurodendron greveanum and Eugenia aff. calciscopulasum Capurodendron greveanum. forest.

Figure 17.

Macphersonia gracilishildebrandtii Dracaena lanceolata Maytenus undata Strychnos Vepris aff. arenicolaEugenia aff. Diospyros pervilleana calciscopulasum Goodman et al

Figure 18.

Figure 19.

Group and (Group II) Capurodendron greveanum Eugenia aff. calciscopulasum. Structural variant 2: Low-growing forest with open canopy -- Floristic characteristics of Group II Vepris aff. arenicola and Maytenus undata. Capurodendron greveanum and Diospyros erythrosperma h Goodman et al

Figure 20. Capurodendron greveanum and Eugenia aff. calciscopulasum

Figure 21

Figure 22 Goodman et al 27

Structural variant 1: Low-growing forest with open understory -- Structural characteristics of Group II

Figure 23.

Figure 24. Mystroxylon aethiopicum and Diospyros erythrosperma Goodman et al

Figure 25

Figure 26.

Mystroxylon aethiopicum Cynanchum arenarium Grewia aff. barorum Group and (Group III) Margaritaria anomala Mystroxylon aethiopicum Structural variant 2: Low-growing forest with open canopy -- Floristic characteristics of Group III Mimusops coriacea and Ficus tiliifolia Grewia betulaefoliaGrewia aff. barorum. Margaritaria anomala Goodman et al

Figure 27

Figure 28.

Figure 29. Mimusops coriacea and Ficus tiliifolia 30 Goodman et al

Figure 30.

Structural characteristics of Group III

Landolphia gummifera

Figure 31.

Figure 32. Goodman et al 31

Cissus microdonta Mimusops coriacea Ficus tiliifolia Rattus. Ants Mimusops coriacea, Ficus tiliifolia, and Commelina Faunistics aff. madagascarica Snails

Tropidophora vittata Xymmer Tropidophora vesconis Malagidris Xymmer Omphalotropis Mystrium oberthueri Ampelita vesconis Proceratium Edentulina Edentulina intermedia Variation in species richness between sites Conulinus vesconis Microcystis Opeas soulaianus Lisachatina immaculata 32 Goodman et al

Table 11.

Dry forest Nosy Dune forest Status Nosy Ratsy Nosiborona Nosy Ankao Manampao Nosy Ankao w G w G w G W G W G Subfamily Amblyoponinae Mystrium oberthueri E Xymmer Subfamily Dolichoderinae Tapinoma E Subfamily Dorylinae Lioponera Tanipone subpilosa Tanipone Subfamily Formicinae Camponotus alamaina Camponotus E Camponotus grandidieri E Camponotus hova Camponotus repens Nylanderia madagascariensis E Paratrechina longicornis Plagiolepis Plagiolepis E Plagiolepis alluaudi Plagiolepis madecassa Tapinolepis Subfamily Myrmicinae Carebara E Carebara grandidieri E Carebara Crematogaster hafahafa E Crematogaster maina E Crematogaster tricolor E Crematogaster tavaratra Crematogaster rasoherinae Malagidris jugum Monomorium termitobium E E Monomorium platynode E Monomorium micrommaton E Monomorium lepidium E Monomorium hanneli E Nesomyrmex gracilis E Pheidole Pheidole E Pheidole megacephala Strumigenys inatos E Strumigenys fanano E Strumigenys sphera E Strumigenys olsoni E Strumigenys maxillaris Strumigenys emmae Terataner E Terataner alluaudi Tetramorium ryanphelanae E Tetramorium sericeiventre Tetramorium quasirum E Tetramorium delagoense Tetramorium trafo E Tetramorium scytalum Tetramorium Subfamily Ponerinae Anochetus madagascariensis Anochetus grandidieri Goodman et al 33

Dry forest Nosy Dune forest Status Nosy Ratsy Nosiborona Nosy Ankao Manampao Nosy Ankao w G w G w G W G W G Bothroponera wasmanii E Hypoponera Hypoponera E Hypoponera Hypoponera E Hypoponera E Leptogenys fasika Leptogenys maxillosa Plathytyrea arthuri Subfamily Proceratiinae Proceratium E Subfamily Pseudomyrmicinae Tetraponera 17 17 21 11 12 17 21 1 3 7 3 method Species richness per site 33 22 25 25 27 66 sites Number of species unique to a site 7 5 6 4 4

Figure 33.

Faunistic similarities between sites Plagiolepis alluaudi and Pheidole megacephala Abundance of introduced species Tetramorium delagoense Goodman et al

Table 12.

Dry forest Nosy Dune forest Nosy Ratsy Nosiborona Nosy Ankao Manampao Nosy Ankao Dry forest Nosy Ankao Nosy Manampao 0.10 Nosy Ratsy 0.13 0.21 Dune forest Nosy Ankao 0.20 0.07 0.12 Nosiborona

Table 13.

Dry forest Nosy Nosy Dune forest Taxa Nosiborona Nosy Ankao Manampao Ratsy Nosy Ankao Subfamily Formicinae Paratrechina longicornis Pheidole megacephala 1.00 0.20 0.20 Plagiolepis alluaudi 0.20 Subfamily Myrmicinae Tetramorium delagoense 0.32 1.00 Tetramorium sericeiventre 0.20 Tetramorium scytalum 0.32 Strumigenys emmae Strumigenys maxillaris 0.20 Subfamily Ponerinae Leptogenys maxillosa

Table 14.

Nosy Ankao island group Analabe Anala Status A M R B Subfamily Amblyoponinae Mystrium oberthueri E Xymmer Subfamily Dolichoderinae Tapinoma E Subfamily Dorylinae Lioponera Tanipone subpilosa Tanipone Subfamily Formicinae Camponotus alamaina Camponotus E Camponotus grandidieri E Camponotus hova Camponotus repens Nylanderia madagascariensis E Paratrechina longicornis Plagiolepis Plagiolepis E Plagiolepis alluaudi Plagiolepis madecassa Tapinolepis Subfamily Myrmicinae Carebara E Carebara grandidieri E Carebara Crematogaster hafahafa E Crematogaster maina E Crematogaster tricolor E Crematogaster tavaratra Goodman et al

Table 14. Nosy Ankao island group Analabe Anala Status A M R B Crematogaster rasoherinae Malagidris jugum Monomorium termitobium E E Monomorium platynode E Monomorium micrommaton E Monomorium lepidium E Monomorium hanneli E Nesomyrmex gracilis E Pheidole Pheidole E Pheidole megacephala Strumigenys inatos E Strumigenys fanano E Strumigenys sphera E Strumigenys olsoni E Strumigenys maxillaris Strumigenys emmae Terataner E Terataner alluaudi Tetramorium ryanphelanae E Tetramorium sericeiventre Tetramorium quasirum E Tetramorium delagoense Tetramorium trafo E Tetramorium scytalum Tetramorium Subfamily Ponerinae Anochetus madagascariensis Anochetus grandidieri Bothroponera wasmanii E Hypoponera Hypoponera E Hypoponera Hypoponera E Hypoponera E Leptogenys fasika Leptogenys maxillosa Plathytyrea arthuri Subfamily Proceratiinae Proceratium E Subfamily Pseudomyrmicinae Tetraponera All species shared with Analabe Forest 18 5 10 8 25 - All species shared with Analamera 10 2 3 3 - 12

Comparison of the ant fauna of the Nosy Ankao island group to the mainland sites of Daraina and Analamera

Malagidris Tanipone Goodman et al

Mosquitoes

, P Culex tritaeniorhynchus Aedes Aedes lambrechti Aedes Culex Anopheles arabiensis and Aedeomyia madagascarica. These P obtained.

Table 15. et alet al., et al

Nosy Ankao Nosy Manampao Nosy Ratsy Nosiborona Total Genus and species HLC LT HLC LT HLC LT HLC Aedeomyia madagascarica 1 1 Aedes aegypti 2 3 37 albocephalus albopictus 1 2 1 fryeri 3 7 2 7 2 21 lambrechti 3 171 20 monetus 13 1 10 tiptoni 13 13 Anopheles arabiensis 2 2 Culex quinquefasciatus tritaeniorhynchus 3 10 Total 28 398 6 27 18 50 527

Larval breeding sites

Culex tritaeniorhynchus and Aedes albopictus. Anopheles arabiensis A. aegypti.

Table 16.

Breeding sites types Species Cement Metallic Canoe Coconut Mangrove Tree holes wells container Aedes aegypti Aedes albocephalus Aedes albopictus Aedes fryeri Aedes lambrechti Aedes monetus Anopheles arabiensis Culex tritaeniorhynchus Goodman et al 37

A. lambrechti, A. albocephalus, and A. fryeri Laliostoma labrosum Reptiles and amphibians Zonosaurus madagascariensis

Figure 34. Goodman et al

Table 17.

Island IUCN Habitat and Mode of Relative Nosy Nosy Nosy Nosiborona Taxa status biotope displacement abundance Ankao Manampao Ratsy 6 ha 365 ha 29 ha 6 ha AMPHIBIANS Mantellidae Laliostoma labrosum Forest and SA Total amphibian: 1 1

REPTILES Ebenavia inunguis Forest Geckolepis maculata Forest Hemidactylus frenatus Paroedura lohatsara Forest Forest Phelsuma abbotti SA forest Phelsuma aff. Forest SA dorsivittata Uroplatus ebenaui Forest Zonosaurus Forest and A madagascariensis Scincidae Amphiglossus Forest Cryptoblepharus A boutonii Madascincus aff. Forest A miafana Paracontias aff. Forest and hildebrandti Trachylepis elegans Trachylepis gravenhorstii Forest and SA Chamaeleonidae Brookesia aff. tristis Furcifer petteri Forest Furcifer oustaleti Forest and SA Acrantophis Forest and SA madagascariensis Lamprophiidae Liophidium aff. Forest and SA torquatum Lycodryas aff. inopinae Forest Madagascarophis Forest, forest SA colubrinus Mimophis mahfalensis Forest, forest A Madatyphlops Forest SA microcephalus aff. Forest and grandidieri Total reptiles: 25 18 11 4 1 Total amphibians and 18 11 4 2 reptiles: 26 Goodman et al

Phelsuma aff. dorsivittata Bismarckia or satrana Paracontias aff. hildebrandti and Xenotyphlops aff. grandidieri Xenotyphlops aff. Mimusops grandidieri coriacea and Ficus tiliifolia X. grandidieri X. grandidieri grandidieri Xenotyphlops et al. Phelsuma aff. dorsivittata; Amphiglossus aff. miafana Madatyphlops Brookesia aff. tristis Xenotyphlops and Liophidium aff. torquatum, Lycodryas aff. inopinae, and Xenotyphlops aff. grandidieri Furcifer Paroedura petteri lohatsara and Uroplatus ebenaui Furcifer petteri Zonosaurus madagascariensis,

Figure 35. Goodman et al

Figure 36. Amphiglossus aff. miafanaBrookesia aff. tristis Phelsuma aff. dorsivittata Lycodryas aff. inopinae

Figure 37. Paroedura lohatsara Uroplatus ebenaui Furcifer petteri Goodman et al

Figure 38. Xenotyphlops aff. grandidieri Paracontias aff. hildebrandti Mimusops coriacea and Ficus tiliifolia formation (see Phelsuma aff. dorsivittata, Paracontias aff. Trachylepis elegans and T. gravenhorstii hildebrandti, Cryptoblepharus boutonii, Brookesia aff. tristis, and Xenotyphlops aff. grandidieri Madatyphlops microcephalus, Amphiglossus Madascincus aff. miafana

Table 18.

Characteristics Pitfall line 1 Pitfall line 2 Pitfall Line 3 Total Habitat type Ground cover

Taxa Amphiglossus 2 2 Madascincus aff. miafana 7 21 Trachylepis elegans 2 1 3 6 Madatyphlops microcephalus 1 1 Liophidium aff. torquatum 1 1 2 10 13 32 11 11 11 33 10 10 10 10 110 110 110 330 9.1 8.2 11.8 9.7 Goodman et al

Table 19.

Characteristics Pitfall line 1 Pitfall line 2 Total Habitat type Ground cover Taxa Paracontias aff. hildebrandti 1 1 Trachylepis gravenhorstii 3 11 Liophidium aff. torquatum 2 2 11 3 14 11 11 22 5 110 20.0 5.5 12.7

Table 20.

Characteristics Pitfall line 1 Pitfall line 2 Total Habitat type Ground cover Zonosaurus madagascariensis 3 2 5 Madagascarophis colubrinus 1 1 Liophidium aff. torquatum 1 1 3 7 11 11 22 5 110 7.3 5.5 6.4

Table 21.

Mode of capture Taxa Activity period Direct Systematic Pitfall traps observation searching Laliostoma labrosum Ebenavia inunguis Geckolepis maculata Hemidactylus frenatus Paroedura lohatsara Phelsuma abbotti Phelsuma aff. dorsivittata Uroplatus ebenaui Zonosaurus madagascariensis Amphiglossus Cryptoblepharus boutonii Madascincus aff. miafana Paracontias aff. hildebrandti Trachylepis elegans Trachylepis gravenhorstii Brookesia aff. tristis Furcifer petteri Furcifer oustaleti Acrantophis madagascariensis Liophidium aff. torquatum Lycodryas aff. inopinae Madagascarophis colubrinus Mimophis mahfalensis Madatyphlops microcephalus Xenotyphlops aff. grandidieri Total 18 8 5 Goodman et al

et al Anous stolidus and A. tenuirostris Birds Sterna caspia S. dougallii S. fuscata Thalasseus bergii Accipiter T. bergii madagascariensis and Berniera Bernieria madagascariensis Buteo brachypterus Falco peregrinus

Table 22.

Nosy Ankao Nosy Manampao Nosy Ratsy Taxa Ardea cinerea Bubulcus ibis Butorides striata Egretta dimorpha Buteo brachypterus Falco newtoni Turnix nigricollis Dryolimnas cuvieri Nesoenas picturata Centropus toulou Caprimulgus madagascariensis Cypsiurus parvus Eurystomus glaucurus Upupa marginata Hypsipetes madagascariensis Cisticola cherina Terpsiphone mutata Nectarinia souimanga Nectarinia notata Zosterops maderaspatana Corvus albus Acridotheres tristis Foudia madagascariensis Total species 25 6 4 Goodman et al

around human habitations to obtain more information R. rattus and a R. norvegicus Small mammals Suncus murinus human habitations. Felis silvestris Rattus rattus Rattus Tenrec ecaudatus or The R. rattus Setifer setosus

Table 23.

Nosy Ankao Nosy Manampao Nosy Ratsy 100 10 1000 captures Zonosaurus madagascariensis 0 0 1 Small mammal captures Rattus rattus 11 0 Rattus norvegicus 11 0 0 6.3 2.9 0.0 1 Goodman et al

Table 24. Rattus rattus

Position of trap Localization for those on ground Localization for those off ground Next to On Off On Branch or Branch or trunk In litter fallen and Other Other ground ground roots trunk > 10 cm < 10 cm rotten wood 1 10 20 Sex and age 10 10 2 7 3 13 2 1 1 2

Table 25. Rattus rattus Position of trap Localization for those on ground Localization for those off ground On Off In litter Next to On Other (mostly Branch or Branch Other ground ground fallen and roots trunk or trunk rotten > 10 cm < 10 cm wood 1 1 70 3 1 Sex and age 7 7 3 1 2 1 1

Discussion Vascular plants Ficus tiliifolia Goodman et al

Ants Ant diversity on islands in the Nosy Ankao group Snails Tropidophora thesauri Edentulina minor E. arenicola Gulella microdon

Figure 39. Goodman et al

Comparing the ant diversity on islands in the Nosy Ankao group to that of Daraina and Analamera Mystrium, Xymmer, and Proceratium, Tanipone Proceratium and Tanipone Introduced species Reptiles and amphibians Faunistic and biogeographic considerations Pheidole megacephala dominates the entire Mimusops coriacea and Ficus tiliifolia Plagiolepis alluaudi Goodman et al

et al., 2007; et al Ebenavia inunguis et al., Paracontias aff. hildebrandti Mimusops coriacea and Ficus tiliifolia Xenotyphlops aff. grandidieri Brookesia micra et al Paroedura lohatsara Taxonomic, ecological, and conservation considerations et al Liophidium aff. torquatum L. torquatum Rattus et al et al et al Seasonality of reproduction Zonosaurus madagascariensis Furcifer petteri Paracontias aff. hildebrandti Goodman et al

Birds Dryolimnas cuvieri Faunistic, biogeographic, and conservation aldabranus, Centropus toulou insularis, Hypsipetes considerations madagascariensis rostratus, Nectarinia souimanga aldabrensis, Zosterops maderaspatana aldabrensis, Dicrurus aldabranus, and Foudia (eminentissima aldabrana et al. et al., Acridotheres tristis, is as Upupa marginata Rattus Small mammals Faunistics Goodman et al

Figure 40. Sterna fuscata

Rattus Figure 41. et al. Goodman et al

Tenrec ecaudatus, Setifer setosus, and Microgale brevicaudata Rattus R. rattus Acranthophis madagascariensis, and Rattus Manambato (Goodman et al., Tenrec and Setifer Conservation priorities and recommendations Some important generalities associated with invasive invertebrates Rattus Rattus Rattus feed for Rattus Rattus Rattus Rattus, Nosy Ankao Rattus Introduced plants Zonosaurus madagascariensis, trees, Casuarina equisetifolia Goodman et al

et al., 2012 P. megacephala C. equisetifolia Mystrium, Lioponera, Tanipone, C. equisetifolia Leptogenys, and Proceratium P. megacephala Xymmer and Malagidris. et al of P. megacephala Leucaena leucocephala

Introduced ants Introduced rodents

Pheidole megacephala and Solenopsis Rattus rattus R. geminata norvegicus Pheidole megacephala et al P. megacephala P. megacephala Paratrechina longicornis, Plagiolepis alluaudi, Strumigenys maxillaris, S. emmae, Tetramorium sericeiventre, and T. delagoense Goodman et al

2011; Feare et al. Other introduced mammals – Nosy Manampao voalavo arabo Suncus murinus Introduced ants Paratrechina longicornis, Plagiolepis alluaudi, Pheidole megacephala, Strumigenys emmae, Tetramorium scytalum, and Leptogenys maxillosa – cat scats Introduced snails Lisachatina immaculata Introduced rodents Rattus Introduced birds rattus. Acridotheres tristis Ficus Upupa marginata, for state. Goodman et al

Questions of biosecurity et al Rattus R. rattus Rattus Introduced birds Acridotheres tristis Acknowledgements season. Goodman et al

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