SPtaturoceedingss of a rare of the endemic Hawaiian Marque Entomologicalsan snout butterflySociety (2018) 50:25–34 25

Scientific Note

Conservation Note on the Status of the Rare Endemic Marquesan Snout , collenettei

Akito Y. Kawahara1 and Emmanuel F.A. Toussaint1* 1Florida Museum of Natural History, University of Florida, Gainesville, Florida, 32611, USA. *Both authors contributed equally. [email protected], [email protected]

Abstract. The Marquesan snout butterfly Libythea( collenettei) also known as “Papillon à museau des Marquises,” is the only endemic butterfly from the Mar- quesas Islands, . The butterfly is known from just five historic records. We report results from an intensive two–week survey in 2018. Our survey took place on the two islands where historic collection records exist ( and ), plus and , where the species has been thought to exist. Despite visiting multiple localities including sites where the species was previously observed, we were unsuccessful at detecting this species. The larval host plant, pacifica (Cannabaceae), can still be found on the Marquesas, indicating that the butterfly might still exist in the archipelago. Because the phenology of this species is unknown, future surveys should be conducted on the same islands but during different seasons. Given the very restricted geographic range of this species and threats to its habitat, we suggest that it be listed as endangered on the IUCN Red List.

Keywords: French Polynesia, Danaus plexippus, Hypolimnas bolina otaheitae, IUCN Red List; , , ,

The Marquesas Islands are an extremely to the islands. Centipedes and lizards were remote archipelago located ca. 4000 kilo- accidental introductions that deeply altered meters southeast of Hawaii, ca. 1500 kilo- the native biota (Adamson 1939, Cheesman meters northeast of the and 1927, Lorence et al. 2016). The first major ca. 5000 kilometers southwest of Mexico. human colonizations on the largest islands This archipelago is comprised of twelve (i.e., Nuku Hiva, Hiva Oa, Ua Pou) took small volcanic islands formed over the place around 1200 AD, and their ecosys- past ca. 6 million years (Maury et al. 2016), tems have been continuously degraded by with rugged landscapes and dense natural logging, fires, and plantations (Wilmshurst forests. Although a large fraction of the et al. 2011, Lorence et al. 2016). In particu- flora is endemic (48%, Lorence et al. 2016), lar, the planting of Caribbean pine (Pinus anthropogenic disturbance, in the form of caribaea, Pinaceae) since 1974 on Nuku native ecosystem destruction (forests, lakes, Hiva and Hiva Oa (Allen et al. 2011), and etc.) has shaped a very different landscape of Leucaena leucocephala (Caesalpinioi- than what the islands were before the first deae) on Ua Pou (Butaud and Jacq 2013, Polynesian colonization 800–900 years ago Lorence et al. 2016) have triggered major (Allen 2014). Humans introduced pigs and ecosystem shifts that significantly impacted rats before bringing horses, cattle, and goats wetlands, native flora, and associated fauna. 26 Kawahara and Toussaint The Caribbean pine was planted to initiate by the Bernice Pauahi Bishop Museum a new forestry industry in the Marquesas and Hawaiian Sugar Planters’ Association in the 1970s, but due to abrupt economic allowed for the description of many new changes, mature pine plantations remained Marquesan (Adamson 1939). unharvested across the archipelago, es- diversity in the Marquesas is pecially on Nuku Hiva (Meyer 2007). depauperate in many groups but diverse Aquatic ecosystems that once prevailed on in others (Gillespie et al. 2008). For the Toovii Plateau have suffered from the instance, there are radiations of Rhynco- dessication of phreatic tables and acidifica- gonus weevils (Claridge et al. 2017) and tion of soils where these trees grow (Meyer Simulium black flies (Craig et al. 2001). 2016, de Grandmont 2018). The drying–out Among Lepidoptera (>300 species in the of phreatic tables was likely caused by Marquesas including ca. 240 endemic vegetation changes driven by enhanced species), () are soil leaching (Fossati and Marquet 1998, poorly represented, with records of only Meyer 2016). The continuous presence of five species (Holloway 1983, Adler and grazing cattle and horses on Nuku Hiva has Dudley 1994, Patrick and Patrick 2012, accentuated the demise of the mid–elevation Ramage 2017). These species are Libythea swamp “Vaihakaomeama,” which harbored collenettei, Danaus plexippus (Nym- a rich endemic diversity of and phalidae, Danainae), Hypolimnas bolina plants, including populations of the Pacific otaheitae (Nymphalidae, Nymphalinae), black duck, Anas superciliosa (Brown 1931, Phalanta marquesana (Nymphalidae, Adamson 1936, Meyer 2016). Similarly, the Heliconiinae), and Badamia exclamatio- invasion of Nuku Hiva and Ua Pou by L. leu- nis (Hesperiidae, Coeliadinae). Phalanta cocephala was fueled by grazing and fires. marquesana and B. exclamationis have This plant has now spread across these two not been found since the 1920s; the former islands and contributed to the dessication of was originally described in Atella by Riley habitats (Butaud, pers. comm.). as an endemic species to the Marquesas The Marquesan endemic insect fauna is (Poulton and Riley 1928), but was later diverse but equally threatened by human also reported from the Society Islands activities (Florence and Lorence 1997, (Holloway 1983). Badamia exclamationis Gargominy et al. 2016; but see Polhemus was found on Hiva Oa in February 1925, and Englund 2016, Roderick and Gillespie but no additional specimens have been 2016 on the exceptionally well–preserved reported since then (Poulton and Riley aquatic entomofauna of the Marquesas 1928). This skipper is currently found Islands). The insect fauna of the Marque- across the Indo–Australian archipelago sas was first documented in 1924–1925 with a geographic range stretching from during the St George Expedition that India to Japan, and from China to Aus- was inaugurated by the Scientific Expe- tralia and some Oceanian islands such ditionary Research Association. During as Fiji. (Another species of Badamia, B. that expedition, the British Natural His- atrox, is widespread in the South Pacific tory Museum entomologists Lucy Evelyn and occasionally listed as occurring in Cheesman OBE, Cyril Leslie Collenette, the Marquesas (e.g., Holloway and Peters and Cynthia “Madame Dragonfly” Long- 1976, Patrick and Patrick 2002), but these field collected insects for five weeks on the reports could not be verified.) islands of Fatu Hiva, Hiva Oa, Nuku Hiva, The Marquesan snout butterfly Lib( - and (Hornell 1924, Cheesman ythea collenettei, Nymphalidae, Liby- 1928, Adamson 1939). The Pacific Ento- theinae) is the only butterfly endemic to mological Survey (1927–1932) organized the Marquesas (Shields 1987, Kawahara Status of a rare endemic Marquesan snout butterfly 27

Figure 1. Habitus of the Marquesan snout butterfly,Libythea collenettei, known as “Pa- pillon à museau des Marquises” and its putative host plant, Celtis pacifica. Upper photo: dorsal view of L. collenettei Holotype, lower photo: ventral view of L. collenettei Holotype.

2006a). Little is known about its biology, on other days, especially in the hills, but no except that the larval host–plant might be others were seen. In flight they were often Celtis pacifica (Rosales, Cannabaceae) confused with the introduced Polystes wasp, (Shields 1987), a plant that is widespread the speed of flight and apparent size being much the same, although I do not think that in the Pacific islands (Figure 1). This tree this was a definite example of mimicry.” is found at low elevations between sea Only five specimens of this butterfly level and ca. 550 m in semi–xerophilic to have been collected to date, from the is- mesophilic forests on the six main Mar- lands of Nuku Hiva and Ua Pou (Figure 1). quesas islands (Fatu Hiva, Hiva Oa, Nuku Three female specimens are known from Hiva, Tahuatu, , and Ua Pou) and the type series from Nuku Hiva, collected (Butaud et al. 2008, Butaud on January 18, 1925, by Cyril Leslie Col- pers. comm.). The only information avail- lenette, and two specimens were sampled able on the behavior of this butterfly can in August 2001 from Ua Pou (one male, be gleaned from Cyril Leslie Collenette’s August 20, 2001), and Nuku Hiva (one notes from the 1925 St George expedition, female, August 28, 2001) by Ronald En- published in Poulton and Riley (1928): glund. The specimen collected on Ua Pou “Three specimens taken and two or three others seen in the Hooumi Valley, Nuka in 2001 is the only known male specimen Hiva Island, Marquesas, on 18.i.25. They fre- and is currently stored in the collection of quented a stream–bed near sea–level. They the Bernice P. Bishop Museum, Honolulu, were attracted by some flowering plants HI, USA (Kawahara 2003). Because this growing in the water, but flew round them species has been observed and collected only a few seconds at each visit, settling in January and August, it may be multi- very rarely. On 16.i.25, 5 or 6 specimens voltine. However, many Libythea species were seen, on a ridge above the Typee Val- are univoltine, emerging in the early sum- ley, at about 1200 ft. They flew against the mer and overwintering as adults until the wind, fairly rapidly and without settling. A considerable search was made for the insect early spring (Kawahara 2006a, b). Further 28 Kawahara and Toussaint surveys are clearly needed to shed light on by walking in fields, along trails, and on this species’ natural history. roads for 2–3 hours at each visited site Morphologically, L. collenettei lacks two (Table 1) and used butterfly nets to tap characters that are present among other Li- vegetation to allow perched butterflies bythea species: the single wide band across to fly. Although it rained occasionally in hindwing cells M1–M3, and the spine at Nuku Hiva and Hiva Oa during our field- the apex of the valva (Kawahara 2013). work (albeit never during a whole day), Based on a morphological phylogeny of most days when surveys took place were the Libytheinae, Kawahara (2009) inferred sunny. We visited sites on each island that L. collenettei to be sister to the remainder had historical collection records. We also of Libythea. However, it is possible that visited pristine patches of native forest and L. collenettei is recovered in this position open areas, including disturbed ones, such because it shares morphological characters as the one found near Mount Pouhekaei in with included outgroups although they may Ua Pou, near the site at which L. collen- be homoplasies (Kawahara 2009). Overall, ettei was found in 2001 (Figures 2 and 3). L. collenettei is morphologically similar We decided to survey the islands in early to the Southeast Asian L. geoffroyi whose February, one week after the St George easternmost distribution stretches as far as Expedition survey dates from late–Janu- the Solomon Islands and New Caledonia. ary 1928. While it would have been best It is therefore possible that L. collenettei is to survey multiple weeks during the time the Pacific Island allopatric sister species of the historic collection dates, this could of L. geoffroyi (Waterhouse 1937, Shields not be done due to limitations of funds 1987). Preliminary molecular evidence for fieldwork. We are currently seeking supports this relationship (N. Wahlberg, funding to conduct a future survey on Ua pers. comm.), although additional data will Pou in August, the island and month from be needed to firmly confirm the placement which L. collenettei was last collected. of L. collenettei within the genus. Despite Figure 2 shows a map of localities its unique morphology, distribution, and surveyed, and Table 1 provides details of rarity, this species has not been the focus sampling localities across the archipelago. of conservation initiatives, and as a result, Despite our searches, L. collenettei could is not listed on the IUCN Red List. not be found on any of the islands that we To assess the possible conservation need visited, although its putative host–plant, for L. collenettei, the authors visited Hiva Celtis pacifica, is still found in the Mar- Oa, Nuku Hiva, and Ua Pou (Marquesas quesas (Butaud et al. 2008, Butaud pers. Islands) and Tahiti (Society Islands), comm.) (Figure 1). French Polynesia, between February 3 The Marquesas Islands have been and February 12, 2018. Although Hiva Oa significantly altered by human distur- and Tahiti do not have known collection bance since the 1920s. Our observations records of L. collenettei, we visited these confirm earlier reports (e.g., Meyer 2007, islands because of their size, remaining 2016; de Grandmont 2018) of significant natural habitat, and their proximity to the anthropogenic disturbance on Nuku Hiva, islands from which the butterfly has been where Caribbean pine trees, introduced recorded. We searched extensively for from the Caribbean Islands and planted this butterfly, and visited the islands dur- ~1970s, cover most of the Toovii Plateau. ing the following dates: Tahiti: February The landscape is also altered by grazing 2; Nuku Hiva: February 3–6; Hiva Oa: cattle, goats, horses, pigs, and chickens, February 6–9; Ua Pou: February 9–12. making the understory nearly completely We conducted visual searches of habitats barren (see also Meyer 2016). Some en- Status of a rare endemic Marquesan snout butterfly 29

Figure 2. Maps highlighting sampling localities for the three islands visited in Febru- ary, 2018. demic plants remain along steep slopes area (Butaud pers. comm.). Therefore it and valleys (Figure 3), but conservation is possible that historical reports of L. regulations, despite existing, are often collenettei on the Toovii Plateau were not enforced, and the islands are isolated, of individuals that had traveled from making it difficult to protect the butterfly other parts of Nuku Hiva. Other areas and its habitat. Some sites where L. col- on this island where the plant has been lenettei was once observed (e.g., on the reported, such as the dry forest of “Terre Toovii Plateau) have been substantially déserte” between 300–500 m (Butaud affected by recent species introductions pers. comm.), could not be sampled during and land management policies. However, this expedition due to time and resource the putative host–plant Celtis pacifica is limitations. These sites and similar eco- not found on the Toovii Plateau, which is systems on other islands should be the a hydrophilic to ombrophilous forested focus of future surveys (e.g., forest patches 30 Kawahara and Toussaint Long. –149,4751 –149,4328 –140,1097 –140,1743 –140,1135 –140,1804 –140,1429 –140,1429 –140,0296 –139,0635 –139,0695 –139,0321 –139,0765 –138,9892 –138,9338 –138,928 –138,914 –138,8809 –138,9444 –140,0476 –140,05 –140,0639 –140,0791 –140,0567 –140,0472 –140,0538 –140,0963 Lat. –17,5242 –17,552 –8,9169 –8,8647 –8,8853 –8,8622 –8,8591 –8,8591 –8,8876 –9,8343 –9,8017 –9,8008 –9,799 –9,7664 –9,7624 –9,7394 –9,7401 –9,7656 –9,7726 –9,4162 –9,4104 –9,4344 –9,4458 –9,4242 –9,4308 –9,4021 –9,368 Tahiti Region Tahiti Nuku Hiva Nuku Hiva Nuku Hiva Nuku Hiva Nuku Hiva Nuku Hiva Nuku Hiva Hiva Oa Hiva Hiva Oa Hiva Hiva Oa Hiva Hiva Oa Hiva Hiva Oa Hiva Hiva Oa Hiva Hiva Oa Hiva Hiva Oa Hiva Hiva Oa Hiva Hiva Oa Hiva Pou Ua Pou Ua Pou Ua Pou Ua Pou Ua Pou Ua Pou Ua Pou Ua Locality Centre spirituel de Supermahina, Fraternité de la Sainte Famille, Archevêché de Papeete Dirt toward National road Park, near large ecosystem river and Papeno’o lowland Main road between near the lowland Plateau river with airport south a bridge, the of Toovii and Taioha’e, Main road between the slope mid with airportgrassy elevation north vegetation, the of and Taioha’e, Plateau Toovii Main road between the airport and Taioha’e, mid elevation, house «Tekeika», south of the Toovii Plateau south the of Toovii Main road between house the «Tekeika», mid airport elevation, and Taioha’e, Dirt road atoff mid the elevation main road between the north Plateau airport the of Toovii and Taioha’e, Main Plateau road between with on the muddy ponds the Toovii lowland airport and Taioha’e, Hee Tai Inn pension in Taioha’e, lowland urban lowland area, south coast Inn the of Hee island Tai pension in Taioha’e, Huemo’o near the beach, northHuemo’o coast the of island Rain puddles on a road near lowland side Taaoa, Trail to Mount Temetiu startingTrail to Mount Temetiu from the ancient cemetery , of mid elevation Temetiu Village pension in Temetiu Atuona, lowland Rainforest 1200–1300 on top Mount m Temetiu, of Trail east Atuona of airport, dirt Hibiscus road with yellow bushes Dirt road toward Nahoe, curve with road puddles Hibiscus and yellow bushes Nahoe village, edges a stream of connected to the ocean Dirt road between Nahoe and with Puama’u puddles on the edge the of cliffs Puama’u village,Puama’u on the edge the of road near the beach Road betweenRoad Nahoe and Atuona, very large puddles swampy on the road with aquatic vegetation Road betweenRoad Hakahau near and Pouhekaei Hakata’o mount,with mid ferns elevation and disturbed ecosystem Disturbed toward forest Hakahau after the junction from Hohoi for Hakata’o Summit in front Pouhekaei of the with ferns road above to Hakata’o and bushes Hakata’o village, smallHakata’o creek on the road with stream and plantations Road before junction Road from Hohoi for Hakata’o Creek near beach Hibiscus Hohoi with bushesyellow of Road afterRoad junction toward Hakahau Hohoi for Trail from Hakahetau to Manfred Ville and the waterfall Table 1. Localities 1. Table visited in the Marquesas Islands and French Polynesia, Tahiti, in 2018. Site Code FRP001 FRP002 FRP003 FRP004 FRP005 FRP006 FRP007 FRP008 FRP009 FRP010 FRP011 FRP012 FRP013 FRP014 FRP015 FRP016 FRP017 FRP018 FRP019 FRP020 FRP021 FRP022 FRP023 FRP024 FRP025 FRP026 FRP027 Status of a rare endemic Marquesan snout butterfly 31 along “La Traversière” trail on Ua Pou, habitat and unenforced environmental visited on February 11 2018). Neverthe- regulations in the Marquesas, we believe less, during the course of our fieldwork, that L. collenettei should be listed on the we rediscovered some endemic inverte- IUCN Red List. These habitats on Nuku brates such as endangered damselflies Hiva and Ua Pou are quickly disappearing, and dragonflies (Polhemus and Englund and it is possible that L. collenettei may 2016) at small pools of standing water on now be extinct. However, we are cau- Toovii Plateau, land snails (Gargominy tious to declare the species extinct, given et al. 2016), and weevils (Roderick and the limited amount of data regarding its Gillespie 2016) at the highest and most phenology and the short sampling period inaccessible parts of Mount Temetiu in of our fieldwork. Existing records are too Hiva Oa (Figure 3). A new endemic spe- scarce to infer any potential population cies of Tropidomantis praying mantis size change over the past century, but the was also discovered in Atuona Bay and total surface area of Nuku Hiva and Ua is awaiting a formal description (Sydney Pou and the total number of specimens Brannoch, pers. comm.). Endemic plants, of L. collenettei collected to date would such those in the genera Glochidion and allow for a classification of this species as Pipterus, were found along a ~1000–m “Endangered” under the IUCN Red List dirt road on the eastern slope of Nuku Criterion 3.1. According to this criterion, Hiva (Point A, Figure 2) and along trails a taxon is “Endangered” when the best near rivers of central Ua Pou (Point B, Fig- available evidence indicates that it meets ure 2), but this ecosystem is disturbed with specific criteria and is considered to be shrubs and trees of citrus, coconut, cacao, facing a very high risk of extinction in mango, and banana. During our survey, the wild (see criteria A to E in the 2001 we observed two other butterfly species, Categories and Criteria version 3.1 for Danaus plexippus and Hypolimnas bolina more details). In the case of L. collenettei, otaheitae. The only adult specimen of D. criterion B is met based on our current plexippus was observed during our survey knowledge of the species. The species’ pu- of Toovii Plateau in Nuku Hiva, and this tative range encompasses Nuku Hiva and species has historically been relatively Ua Pou for a total of <500 km2. Moreover, rare in the Marquesas and Tahiti depend- limited patches of remaining native forest ing on the availability of its host plant, exist, and remaining populations of L. col- Asclepias (Collenette 1925, Buteau, pers. lenettei may be severely fragmented. The comm.). Hypolimnas bolina otaheitae was species’ restricted distribution, our obser- relatively common in both lowland and vations, and loose environmental policies highlands, including disturbed areas on all in French Polynesia make us believe that islands visited. The sex ratio of captured a continued decline in native habitat is specimens was male–biased (27 males to inevitable. During our visit to the islands, 4 females), as previously documented for we developed a small initiative to search this species in the Marquesas (Poulton and for the species with help from locals. We Riley 1928). Badamia exclamationis and created an illustrated booklet and a poster P. marquesana were not encountered dur- with photographs of L. collenettei and its ing our fieldwork. During our survey, we host plant to help identify and report any spoke to locals on each island and many sighting of L. collenettei in Nuku Hiva, of them reported sightings of D. plexippus Hiva Oa, and Ua Pou with the hope to and H. bolina otaheitae in the past decade, find this species throughout the year. A but no one had seen L. collenettei. targeted scientific expedition should be Due to the restricted remaining natural conducted in August, the month that the 32 Kawahara and Toussaint

Figure 3. Images of the Marquesas Islands, close to localities where L. collenettei has been previously found. A. Nuku Hiva, partially preserved ecosystem near Mount Tekao. B–D. Nuku Hiva, Toovii Plateau with extensive Caribbean pine forests, feral horses, and intensive grazing damages the native ecosystem. E. Nuku Hiva, view of Ho’oumi Bay (type locality) now covered by coconut tree plantations. F. Hiva Oa, Mount Temetiu cloud forest where endangered endemic invertebrates can still be found. G. Ua Pou, site where L. collenettei was found in 2001 in front of Mount Pouhekaei. H. Ua Pou, west side of Hakahetau Valley along “La Traversière” trail with some remaining endemic vegetation. Photograph credit: Emmanuel F.A. Toussaint. Status of a rare endemic Marquesan snout butterfly 33 butterfly was last observed in 2001 on The Holocene 21:473–484. Nuku Hiva and Ua Pou (Kawahara 2003). Brown, F.B.H. 1931. Flora of Southeastern Polynesia. I. Monocotyledons. Bernice P. Bishop Museum, Honolulu. Acknowledgments Butaud, J.F., and F.A. Jacq. 2013. Flora and We thank Dan Rubinoff and three anony- vegetation on the small uninhabited islands mous reviewers for comments that greatly of the Marquesas archipelago (French improved an earlier version of this note. Polynesia): Relics of dry biota threatened The Mohamed bin Zayed (MBZ) Species by biological invasions. pp. 131–165. In: Conservation Fund #170515458 provided Larrue S, editor. Biodiversity and Societies funding that supported fieldwork. Blancas in the Pacific Islands. Chapter 7. Presses Huertas and Ryan St Laurent provided Universitaires de Provence, Aix–Marseille. the butterfly image from the NHMUK. Cheesman, L.E. 1927. A contribution towards the Insect Fauna of French Oceania: Part I. Jean–François Butaud, Ronald Englund, Ecological Entomology 75:147–161. and Daniel Polhemus provided critical Cheesman, L.E. 1928. A contribution towards information on localities that helped our the insect fauna of French Oceania. Part, survey. Jean–François Butaud also pro- II. The Annals and Magazine of Natural vided feedback on an earlier version of this History:169–194. manuscript. Jean–Yves Meyer assisted with Claridge, E.M., R.G. Gillespie, M.S. Brewer, permits and planning the expedition. Nik- and G.K. Roderick. 2017. Stepping‐stones las Wahlberg provided information on the across space and time: repeated radiation of ‐ evolution of L. collenettei. Sydney Bran- Pacific flightless broad nosed weevils (Cole- optera: Curculionidae: Entiminae: Rhynco- noch confirmed that the mantis mentioned gonus). Journal of Biogeography 44:784–796. in the text is likely a new species. We thank Collenette, C.L. 1925. The present status the local people of the Marquesas who of Danaida plexippus in the Pacific Is- provided important information, guidance, lands. Entomologists Monthly Magazine permission to explore areas of interest, and 61:198–202. offered help to publicize information on the Craig, D.A, D.C. 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