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THE TRINIDAD AND TOBAGO FIELD NATURALISTS’ CLUB LV D VHFWLRQ DOORZLQJ FRQWULEXWRUVWR GHVFULEH XQXVXDO REVHUYDWLRQVRQ RXU ÀRUD DQG IDXQD 7KH WLWOH RI

The Trinidad and Tobago Field Naturalists’ Club was founded on 10 July, 1891. Its name was incorporated by an Act of Parliament (Act 17 of 1991). The objects of the Club are to bring together persons interested in the study of natural history, the diffusion of knowledge thereof and the conservation of nature. Monthly meetings are held at St. Mary’s College on the second Thursday of every month except December. 0HPEHUVKLS LVRSHQWRDOOSHUVRQVRIDWOHDVW¿IWHHQ\HDUVRIDJHZKRVXEVFULEH WRWKHREMHFWVRIWKH Club.

Mission Statement To foster education and knowledge of natural history and to encourage and promote activities that lead to the appreciation, preservation and conservation of our natural heritage.

Management Committee 2018 -2019 Editorial Committee President Renoir Auguste Graham White (Editor) Vice President Palaash Narase Amy Deacon (Assistant Editor) Secretary Amy Deacon Yasmin Comeau Assistant Secretary Danielle Morong Luke Rostant Treasurer Selwyn Gomes Kris Sookdeo Members Dan Jaggernauth Palaash Narase (Representing Laura Baboolal the Publications Committee) Kris Sookdeo

¿OHV Enquiries All enquiries concerning the Club should be addressed to: The Honorary Secretary P.O. Box 642, Port of Spain Republic of Trinidad and Tobago E-mail: [email protected] Website: www.ttfnc.or g LIVING WORLD Journal of The Trinidad and Tobago Field Naturalists’ Club 2018

Inca clathrata quesneli Boos and Ratcliffe

Published December, 2018 Living World is published by The Trinidad and Tobago Field Naturalists’ Club. All rights reserved.

ISSN 1029-3299 Editorial

The 2018 issues of Living World was initially pub- harvestmen and cyanobacteria is highlighted by V.A. lished online as separate papers, followed by the overall Young, M.K. Moore and V. R. Townsend Jr. issue. We anticipate that this would shorten the period Representing the Amphibia, we have a Research Paper between submission and publication of articles and indeed recording the apparent absence of chytrid infection of frogs as of the next issue we plan to publish the submission and in northeast Tobago, by R. Thomson, P.A. Hoskisson, S. publication dates. Articles for Living World can now be Brozio and J. R. Downie and a Nature Note on poten- submitted online via our open access journal platform. tial case of cannibalism in the tadpoles of Mannophryne (See https://ttfnc.org/livingworld/index.php/lwj/about/sub trinitatis by R.J. Auguste and N.F. Angeli. This note missions#onlineSubmissions). highlights how much we still don’t know about this phe- We are pleased to include our Guest Editorial on the nomenon and the ecological implications within this spe- importance of and recent activities to increase our knowl- cies. Finally for the Arachnida/Amphibia R. J. Auguste, edge of our bat communities. Unlike birds, which are reg- S. Maraj and R. Deo share their observations on the cap- ularly observed and documented, our knowledge of the ture of a Leptodactylus validus by the spider Ancylometes diversity, distribution and habits of bats is poor. Yet they bogotensis. represent 70% of our mammalian fauna and are of tremen- :HKDYH¿YH1DWXUH1RWHVLQWKLVLVVXH,QDGGLWLRQWR dous ecological importance. What is particularly encour- the four already mentioned we have a report of a new spe- aging is the corporation between agencies in pushing the cies of mussel in Trinidad and new locations for a previ- bat research and conservation agenda. We are highlight- ously recorded species. ing these activities, both on our cover and within the pages The report of the Trinidad and Tobago Bird Status and of this Living World, with the expectation that work will Distribution Committee presents 152 records submitted in FRQWLQXH,QGHHGLQWKLVLVVXH&)/RXJKUH\SURYLGHVDQ 0DUW\Q.HQH¿FNSRLQWVRXWWKDWWKLVUHSUHVHQWVWKH account of the bat communities at three contrasting loca- highest number of submissions and the highest number of tions in Tobago. “reportable species” since the formation of the Committee The other papers in this issue of Living World are 23 years ago. quite variable. M.J.W. Cock provides a comprehensive This year G.A. Rivas has provided a book review of account of the sphingid moths of Trinidad, all 81 species the Reptiles and Amphibians of Trinidad and Tobago. The with illustrations of both adults and larvae. There are two Trinidad and Tobago Field Naturalists’ Club is particularly Nature Notes targeting the larvae of Saturnid moths; a cat- proud of this publication as the wide list of authors are pre- erpillar of Arsenura beebei by M. Kelly and M.J.W. Cock dominantly, if not all, members of the Club and regularly and one on Rothschildia vanschaycki by D.S. Huggins, contribute to Living World. The fruitful collaboration is K. Sookdeo and M.J.W. Cock. J.N. Sewlal concludes no doubt due to the activity of the TTFNC. her account of the orb-weaving spider diversity within the Eastern Caribbean in a summary paper which includes all Graham White previous surveys. A more intimate association between [email protected]

Cover Photograph This Common Tent-making Bat, Uroderma bilobatum is demonstrating the important role that bats play in dispersing forest fruit, in this case a Ficus fruit. The photo was taken by Merlin Tuttle, and enhanced by Edward Rooks.

ii Guest Editorial

Bats: Magnificent and Mysterious

,WLVLQGHHGDQKRQRXUWREHJUDQWHGWKHRSSRUWXQLW\WR Research Unit (also known as Trinibats) on an evening of ZULWHLQWKH/LYLQJ:RUOG-RXUQDORQDWRSLF,KROGGHDU mist netting at the Asa Wright Nature Centre in the Arima ,QWKLVJXHVWHGLWRULDO,KLJKOLJKWWKHLPSRUWDQFHRIEDWVLQ Valley on October 12 th , 2018. The bats cooperated fully RXUHFRV\VWHPVUHÀHFWRQWKHGLYHUVLW\RIEDWVLQ7ULQLGDG WKLVHYHQLQJZLWKEDWVRIVSHFLHVEHLQJFDSWXUHG and Tobago, highlight some of the research taking place in and safely released back into the night. This was the largest Trinidad and Tobago, and advocate for continued research catch rate for the entire 2018 Trinibats expedition, and we into these marvellous creatures. were able to highlight a wide array of feeding guilds to :KHQHYHU , DP JLYHQ WKH RSSRUWXQLW\ WR SUHVHQW RQ the Honourable Minister. Bats comprise over 20% of the WKHLPSRUWDQFHRIEDWV,VWDUWE\DVNLQJWKHDXGLHQFHZKDW ZRUOG¶VPDPPDOLDQIDXQDDQGORFDOO\,Q7ULQLGDG they believe to be the importance of birds. Depending on and Tobago there are roughly 70 species of bat, and this list WKHDXGLHQFH,PD\JHWDIHZSHRSOHDQVZHULQJWKDWWKHLU is likely to grow given the fact that molecular techniques plumage or songs are appealing. With some prompting, used to differentiate species have not been conducted on ¿QDOO\ PHPEHUV RI WKH DXGLHQFH VSHDN RI SROOLQDWLRQ most of the species present. Using such techniques, the seed dispersal, and insect control. Bats are the night shift only endemic species of mammal in Trinidad and Tobago for these services. One study on the pollination services was recently found in Tobago, Sir David Attenborough’s of new world phyllostomid bats found that they pollinate Myotis, Myotis attenboroughi . The rich diversity we are WKH ÀRZHUV RI  VSHFLHV RI SODQWV RI  JHQHUD DQG fortunate to have in Trinidad and Tobago can be discussed 44 families. Another study found that neotropical bats LQVHYHUDOZD\V,QWHUPVRI)DPLO\GLYHUVLW\WKHUHDUHQLQH GLVSHUVHWKHIUXLWVRIDWOHDVWVSHFLHVRISODQWVRI families of bat, with the Phyllostomidae dominating with JHQHUD DQG  IDPLOLHV 7KHLU UROH LQ PDLQWDLQLQJ RXU ¿YHVXEIDPLOLHVDQGRIWKHURXJKO\VSHFLHV(DFKRI forest ecosystems is profound, and is well demonstrated these families have their own unique characteristics. The by the cover photo where a Common Tent-making Bat, Emballonuridae, for example, are characterised by having Uroderma bilobatum is dispersing a Ficus fruit. With glandular sacs on their wings, used to mark their territories respect to fruit dispersal, bats are cited as essential for and attract mates. The Thyropteridae, consisting of one recolonisation in forest clearings since many of the species, Spix’s Disk-winged bat, Thyroptera tricolor is seeds they disperse are adapted for growth in disturbed adapted to roost in the rolled up leaves of heliconia and areas. These early successional plants include those of EDQDQDXVLQJVXFWLRQFXSVRQLWVKHHOVDQGWKXPEV,QWHUPV the Jumbie Candle, Piper sp. and Bois Canot, Cecropia of size, the smallest bat in Trinidad is the Riparian Myotis, peltata . Bats are especially good at helping to repair Myotis riparius, with a forearm between 32-38mm, while fragmented forests since they are described as mobile the largest is the Spectral Bat, Vampyrum spectrum , with a foragers, traversing these open areas and dispersing seeds IRUHDUPEHWZHHQPP into forest patches. Without insectivorous bats, the food Perhaps the best way to illustrate our bat diversity, resources we as humans depend upon would likely be however, is by feeding guild. The Spectral Bat is the PRUHH[SHQVLYH,WKDVEHHQHVWLPDWHGIRUH[DPSOHWKDW largest bat in the New World, and the largest carnivorous the value of bats to agriculture in the continental United bat in the world, feeding on birds and small mammals. 6WDWHVLVELOOLRQ\HDUZLWKUHVSHFWWRLQVHFWFRQWURO There are several gleaning animalivores, such as the Little WKHUHE\ UHGXFLQJ WKH QHHG IRU SHVWLFLGH DSSOLFDWLRQV ,Q Big-eared Bat, Micronycteris megalotis. 7KHVH FDQ ¿QG Thailand, a single species of bat has been found to prevent and consume insects perched on a leaf in the dark within WKHORVVRIDERXWWRQQHVRIULFHSHU\HDU7KHEHQH¿WV forested habitats, no easy task. Nectar feeding bats, such of bats in the control of insect populations may also extend as Geoffrey’s Hairy-legged Bat, Anoura geoffroyi , have to their consumption of mosquitoes and other vectors of VSHFWDFXODU DFUREDWLF GLVSOD\V KRYHULQJ DW ÀRZHUV IRU disease, which means they have a critical role to play in short periods while they stick out their long tongues to human health as well. consume the nectar therein. Large generalist fruit bats ,QOLJKWRIWKHPDQ\HFRV\VWHPVHUYLFHVSURYLGHGE\ like the Great Fruit-eating Bat, Artibeus lituratus , can be bats, the Minister of Agriculture, Land and Fisheries, seen dispersing a wide array of fruit, including Figs, Hog Senator the Honourable Clarence Rambarrat, has Plum, and Sapodilla. There are also several species of high recognised the importance of these creatures, going so far DQGIDVWÀ\LQJLQVHFWLYRURXVEDWVVXFKDVWKH*KRVWIDFHG as to join the Trinidad and Tobago Bat Conservation and Bat, Mormoops megalophylla WKDWÀ\DERYHWKHFDQRS\ iii IHHGLQJRQPRWKVEHHWOHVDQGÀLHV7KHUHLVRQHVSHFLHV animalivores having higher abundance in primary forests. RIEDWZKLFKVSHFLDOLVHVRQ¿VKWKH*UHDWHU)LVKLQJ%DW Throughout the world, bats continue to perform Noctilio leporinus , extending its long feet with large sharp their tasks under cover of darkness. They perform many FODZVWRJDII¿VKDWWKHZDWHU¶VVXUIDFH)LQDOO\LQ7ULQLGDG ecosystem services which we take for granted, have we possess two of the three species of vampire bat, the spectacular adaptations to perform these tasks, and Common Vampire Bat, Desmodus rotundus , which feeds continue to be important subjects of research. More on mammalian blood, and the White-winged Vampire Bat, recently, research on their behavioural and feeding ecology Diaemus youngi , which feeds on avian blood. FRQWLQXHVZLWKWKH8QLYHUVLW\RIWKH:HVW,QGLHV 8:,  The diversity of bats we see in Trinidad and Tobago LQFRQMXQFWLRQZLWK7ULQLEDWV5HVHDUFKRQWKHHI¿FDF\RI makes this a fertile ground for research. The William Beebe batbox use in Trinidad is ongoing, where the use of bat Tropical Research Centre at Simla in the Arima Valley was ER[HVLQDWWUDFWLQJEDWVDQGDOOHYLDWLQJEDWKXPDQFRQÀLFW XVHGE\'U'RQDOG*ULI¿QLQWKHVWRFRQGXFWVRPHRI is the focus. Bat boxes have been installed at the Asa his seminal work on echolocation in bats, helping uncover Wright Nature Centre, along with bat acoustic recorders, WKHIHHGLQJHFRORJ\RIWKH*UHDWHU)LVKLQJ%DW,WZDV'U in keeping with the pioneering work carried out in Trinidad *ULI¿Q ZKR FRLQHG WKH WHUP ³HFKRORFDWLRQ´ LQ  WR on echolocation. A comprehensive treatise of the bats of describe the method used by microbats navigate and forage Trinidad and Tobago has also been published by Geoffrey in the dark. The reproductive ecology of bats has also Gomes and Fiona Reid, an essential companion to novices been the subject of very interesting and ongoing research. and experts alike. Trinibats continues in our efforts to Seba’s Short-tailed Fruit Bat, Carollia perspicillata, has educate the public of the importance of bats, encouraging been used as a model organism to help understand some research and conservation. Beyond the research carried out of the serious problems found in human pregnancy. They E\WKH8:,7ULQLEDWVDQGRWKHUHQWLWLHVLQPRUHUHFHQW are ideally suited to this task since they usually carry a times we are pleased to see the removal of most bat species single embryo in a simplex uterus, similar to that seen from the vermin list in the Conservation of Wildlife Act. LQ KXPDQV ,Q XQGHUVWDQGLQJ WKH UHSURGXFWLYH HFRORJ\ Support from the Government of Trinidad and Tobago RI WKLV VSHFLHV VFLHQWLVWV VXFK DV 'U -RKQ 5DVZHLOHU ,9 and the Minister of Agriculture, Land and Fisheries in have helped us understand some of the processes which the protection of roosting habitat for bats is also vital. lead to implantation of human embryos. Some of the One such location which houses a massive population of pioneering work on paralytic rabies was carried out in EDWVLVWKHROG:RUOG:DU,,EXQNHULQ&XPXWRFDOOHGWKH Trinidad, where the link between bats and this form of ³,FHKRXVH´:HGRKRSHWKDWVWUXFWXUHVVXFKDVWKHVHZLOO WKH GLVHDVH ZDV PDGH LQ  GXULQJ D PXOWLVSHFLHV be protected in perpetuity for these bats. Bats deserve our rabies epidemic which killed 73 people, and thousands UHVSHFWDQGSURWHFWLRQ,KRSHWKDW,¶YHKLJKOLJKWHGWKHLU of livestock. Today, research continues on mimicking the importance, and the need for their continued protection. anticoagulant properties of enzymes in the saliva of the Common Vampire Bat. Though the Vampire Bat has a sordid history in Trinidad, it is hoped that in harnessing the Luke Rostant, PhD VSHFLDOSURSHUWLHVIRXQGLQWKHLUVDOLYDKXPDQVPD\EHQH¿W Department of Life Sciences, from novel drugs for stroke victims. The community of 7KH8QLYHUVLW\RIWKH:HVW,QGLHV EDWV ZH ¿QG LQ WKH IRUHVWV RI 7ULQLGDG KDYH WKHPVHOYHV St. Augustine, Trinidad been shown to be indicators of forest health, with gleaning [email protected]

iv Orb-weaving spiders of the Eastern Caribbean Jo-Anne Nina Sewlal The Caribbean Academy of Sciences, c/o Faculty of Engineering, University of the West Indies, St. Augustine, Trinidad, W.I. [email protected] ABSTRACT The Eastern Caribbean represents a poorly studied region in terms of arthropod biodiversity, in particular spiders. This paper reviews studies on the biodiversity of orb-weaving spider families on seven islands in the Eastern Caribbean, sampled between January 2006 and August 2010. Specimens were collected using two main methods; visual search and sweep-netting, and supplemented by material collected from the nests of the spider hunting wasp Sceliphron sp. Sampling HIIRUWV\LHOGHGVSHFLHVGLVWULEXWHGDPRQJVL[IDPLOLHV%LRGLYHUVLW\ZDVTXDQWL¿HGE\H[DPLQLQJWKHREVHUYHGVSHFLHV richness, diversity indices, species abundance models, and multidimensional scaling plots. Factors such as geographic ORFDWLRQDQGKDELWDWFODVVL¿FDWLRQZHUHDOVRH[DPLQHGLQWKLVVWXG\DVSRVVLEOHIDFWRUVLQÀXHQFLQJRUEZHDYHUELRGLYHUVLW\ 1RVLJQL¿FDQWGLIIHUHQFHDPRQJWKHLVODQGVZDVIRXQGLQWHUPVRIVSHFLHVULFKQHVVGLYHUVLW\HYHQQHVVDQGGRPLQDQFH6W Lucia possessed the most distinct orb-weaver species composition, while the faunas of St. Kitts and Nevis and of Grenada and Montserrat were closely associated. Key words: Anapidae, Araneidae, Mysmenidae, Symphytognathidae, Theridiosomatidae and Tetragnathidae, orb-webs

INTRODUCTION Spiders are regarded as one of the most speciose ani- et al. 2007; Stratton et al. 1979) and species composition mal orders on the planet (Coddington and Levi 1991; (Uetz 1991). This is achieved by the presence of more Foelix 1996; Nyffeler 2000) and high ranking predators diverse microclimates (Uetz 1991) produced by the vary- in multiple habitat types (Cardoso et al. 2008; Oxbrough ing amounts of rain and light that reach the lower strata et al. 2005; Sharma et al. 2010; Stratton et al. 1979; Wise of forests (Loyola and Martins 2009). Web-building spi- 1993). Spiders also make excellent candidates for bio- ders in particular are highly dependent on the vegetation diversity studies due to their abundance as well as rela- present in an area to provide them with suitable points of WLYH HDVH WR LGHQWLI\ DQG EH HI¿FLHQWO\ VDPSOHG FRP - attachment for web construction (Riechert and Gillespie pared to other invertebrates (Hsieh and Linsenmair 2011; 1986) and for protection from the elements and predators Oxbrough et al. 2005). Finally they occupy middle trophic (Enders 1974, 1976, 1977; Hodge 1988). levels in terrestrial food webs acting as prey for organisms In the Eastern Caribbean there are 15 web-building spi- in higher tropic levels (Oxbrough e t al. 2005) as well as der families recorded, out of which six construct orb-webs; predators (Cardoso et al. 2008; Stratton et al. 1979; Wise Anapidae, Araneidae, Mysmenidae, Symphytognathidae, 1993) which makes them highly regarded with respect to Theridiosomatidae and Tetragnathidae. I chose to limit ecosystem stability (Coyle 1981) and arthropod control my study to orb-weaving spiders as their characteristic (Enders 1975; Moulder and Reichle 1972; Riechert 1974; web design ensures they are relatively thoroughly sam- Turnbull 1973). pled in each locality selected. They also have a uniform The neotropics have long been declared a region rich lifestyle and literature is available to identify them, which in biodiversity but the literature contains scattered refer- makes detection of new species and changes in the ecol- ences to the spiders, particularly concerning the islands of ogy between families and species and differences between the Lesser Antilles. Many islands in the region lack spe- habitat types easier and faster. cies lists and those lists that do exist tend to be based on This paper focuses on the orb-weaving spider fami- surveys that were conducted 80 or more years ago, for lies collected on seven islands between January 2006 and example, the survey of Antigua by Bryant (1923). As a February 2008. Sampling was conducted on the islands of result, biodiversity data is either unavailable or unreliable Nevis (Sewlal and Starr 2007), St. Kitts (Sewlal 2008), for most islands and existing lists fail to document the Anguilla (Sewlal and Starr 2010), Antigua (Sewlal 2009a), relation of species composition and richness to the habitat Grenada (Sewlal 2009b), Montserrat (Sewlal 2010) and from which it is recorded. St. Lucia (Sewlal, 2011). Sampling on all of these islands Habitat has been consistently cited as one of the most was done with the aim of documenting the spider spe- LPSRUWDQW IDFWRUV DIIHFWLQJ DUWKURSRG ELRGLYHUVLW\ LQÀX - cies found in as a broad variety of habitats as possible. encing multiple facets including abundance, species rich- This paper serves to collectively examine the results of ness (Halaj et al. 1998; Hatley and MacMahon 1980; Peres the islands sampled, and looks at the relationship between 1 2 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018 habitat heterogeneity and species diversity with respect to orb-weaving spiders

METHODOLOGY Across the seven islands in this study (Fig. 1), a total of 110 localities and 27 different habitats were sampled with some habitats not found on every island sampled (Table 1). Brief descriptions of the habitat types are given Table 2. Each island in this study was sampled for a uni- form period of two weeks. Localities for sampling were selected through con- sulting with government ministries and obtaining maps or UHSRUWVGRQHWKDWFRQWDLQHGYHJHWDWLRQFODVVL¿FDWLRQV7KH FODVVL¿FDWLRQIURPWKH¿UVWLVODQGWREHVDPSOHG1HYLV – was used as a guide to maintain consistency in habitat descriptions and thus the selection of habitats for sampling in subsequent islands that were sampled. Care was taken QRW WR GDPDJH SURGXFH RU ÀRZHUV ZKHQ VDPSOLQJ VRPH habitats such as farmland used for cultivation of crops or ornamental plants. Precautions included questioning own- ers about the period between the last treatments with pes- ticides, and in determining if the habitat in some localities ZHUHWREHFODVVL¿HGDVQDWXUDORUGLVWXUEHGVXFKDVVHF - ondary forests, in terms of when was it last used for com- mercial purpose. Data from each island were collected using a combina- tion of the visual search and sweep-netting methods, both of which are effective with respect to rapid collection of the species present in an area. Visual search involved walk- ing around in the selected habitat and collecting what was seen from ground level to arm’s length above the collec- tor’s head. Microhabitats most likely to yield cryptic spe- cies were searched using this method, such as under logs, inside rotting logs, inside holes, under bark and rocks. Sweep-netting involved sweeping the vegetation with a tough canvas insect net. This method collected diurnal and nocturnal species in retreats or resting in the vegetation. Sampling concluded if no individuals of new species were found after 15 minutes of the last new species recorded Fig. 1. Map of Eastern Caribbean showing islands sampled; St for that locality. The main goal of this study was to survey Kitts (1), Nevis (2), Anguilla (3), Antigua (4), Montserrat (5), St. a single locality of each of the major vegetation types on Lucia (6) and Grenada (7). each island sampled, but where time and resources per- ZHEEXLOGLQJ VSLGHUV VSHFL¿FDOO\ RUEZHDYLQJ VSHFLHV mitted up to four localities in each major vegetation type were collected. Spiders are generally separated into web- was sampled (Table 2). After identifying specimens down builders, plant wanderers and ground wanderers. As their to species level, they were transferred and stored in glass name suggests, the web-builders depend on their webs to vials in 96% alcohol. capture prey whereas with the other two functional groups There are many different ways to separate organisms the spiders do not construct webs, but stalk and ambush into categories to allow for easier study. One such method their prey. The difference between the last two groups is is by placing them into what are known as what are known that one is predominantly found wandering on the vegeta- DV IXQFWLRQDO JXLOGV 7KHVH DUH GH¿QHG E\ :KLWPRUH tion, hence the term plant wanderers, while members of the et al. (2002) as groups that contain species that “poten- RWKHUJURXSZDONVRQWKHIRUHVWÀRRUWKHUHE\UHIHUUHGWRDV tially compete for jointly limited resources”. In this study ground wanderers. The use of functional groups can allow Orb-weaving spiders of the Eastern Caribbean 3

Table 1. Number of localities sampled in each habitat according to island for spider surveys of Eastern Caribbean. Shaded sites are KXPDQLQÀXHQFHGKDELWDWV

Islan d nests Garden In and on buildings Abandoned buildings Farmland Roadside Pastureland Sec forest Deciduous forest Lower montane Montane Palm brake Forest Valley Humid Dry Forest ponds Mangrove Littoral Woodland (O¿QZRRGODQG Vegetation Riparian Semi-evergreen forest Evergreen woodland Inland scrub Rainforest Dry evergreen forest Scelphron Coastal veg Cactus scrub Vegetation around salt around Vegetation St. Kitts 2 2 2 2 4 1 1 0 0 0 1 0 1 1 0 1 1 0 0 0 0 1 2 1 0 0 Nevis 4 3 0 1 2 0 3 0 0 1 1 0 0 0 0 2 1 0 0 0 3 0 0 0 0 0 Anguilla 1 2 3 0 2 0 0 0 0 0 0 0 2 1 0 0 0 0 0 1 0 0 0 2 0 1 Antigua 3 4 3 0 4 2 0 0 0 0 0 2 3 3 0 0 0 3 0 0 0 0 0 0 3 0 Montserrat 3 2 1 2 3 0 0 0 0 1 1 0 2 0 3 2 1 0 1 0 0 0 0 0 1 0 St. Lucia 3 1 0 2 2 0 3 2 2 0 0 0 0 0 1 2 0 1 1 0 0 0 0 0 1 0 Grenada 1 2 3 2 2 0 1 0 0 2 1 0 1 0 0 1 1 1 0 0 0 0 0 0 0 0 one to gain valuable insight into the microhabitat, vegeta- diversity between habitats. tion preferences as well as responses to habitat change of The reciprocal of the Simpson index was calculated each group (Haddad et al. 2009; Hsieh et al. 2003). as recommended by Downie et al. (1995) and Magurran Trends in biodiversity in this study were limited to (2004) to obtain a measure expressing diversity in addi- web-building species and determined by examining spe- tion to evenness. Data analysis involved determining the cies richness, diversity indices and species composition, abundance, observed species richness, species diversity, and calculating the three main diversity indices; Shannon- evenness and dominance for each island as a whole and Weiner Index ( +ƍ , reciprocal of the Simpson Index (1/ D) per habitat. However, because of the lack of independ- and Berger-Parker ( d) which looked at species diversity, ence among these indices an ANOVA using Statistix was evenness and dominance respectively (Magurran 2004), only conducted on observed species richness to determine using the following formulas; +ƍ Ȉ pi ln pi, where pi = LIWKLVYDULHGVLJQL¿FDQWO\DPRQJKDELWDWVRQWKHLVODQGV ni / N (the abundance of species i at a site as a propor- sampled. tion of the total sample at that site), 2. D = 1/y, where y The community structure of the spider fauna was

Ȉ ni (ni -1) / N(N-1)) and d = nmax / N, where nmax repre- GHWHUPLQHGE\¿WWLQJWKHUDQNDEXQGDQFHFXUYHJHQHUDWHG sents the number of individuals of the most common spe- for all the islands as a whole, to four main species abun- cies. The Shannon index has been used in numerous bio- dance models given in Magurran (2004) which give addi- diversity studies (Begossi1996) giving it the reputation as tional information on the communities in question such a benchmark measure for biological diversity (Magurran as main ecological trends. Lastly species composition was 2004), allowing the comparison of diversity with other analysed through the use of a MDS (multidimensional studies. The Shannon index is sensitive to sample size scaling) plot generated using Primer 5 software (Clarke (Magurran 2004) and to changes in the importance of rare and Warwick 2001). species (Peet 1974). Much ecological information can be obtained from the value of the Shannon index. High RESULTS Shannon values indicate high species richness and thus One way of examining the results is to split the habitats the presence of diverse communities and species that are found in more than one island into ‘naturally occurring evenly distributed (Hsieh et al. 2003; Peet 1974), while a habitats’ and ‘habitats that have developed as a result of zero value indicates the presence of a single species (Peet human activities’. Overall, the two most speciose habitat 1974). Therefore Shannon gives information on the dis- types are roadside and garden vegetation containing 17 tribution of species abundance in the community or sam- and 15 species respectively, with both being considered to ple (Peet 1974). The interpretation of the results of the belong to the latter mentioned group. Pastureland was the Shannon index with respect to comparing sites is tricky OHDVW VSHFLRVH ZLWK ¿YH VSHFLHV:LWKLQ QDWXUDO KDELWDWV due to sources of error as it is sensitive to relative species the most species-rich were littoral woodland the vegetation abundance and richness (Magurran 2004). Multiple diver- surrounding salt ponds with 11 species each. The lowest sity indices were used to better describe the differences in VSHFLHVULFKQHVVIRXQGLQSDOPEUDNHDQGHO¿QZRRGODQG 4 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

Table 2. Habitat types sampled during the course of this study and their descriptions. Habitat Description Garden Lands landscaped and maintained at private residences, hotels or for public viewing. Inside and around Inside and around buildings occupied by humans dwellings Abandoned buildings Stone and wooden structures that have not been occupied or used for a minimum of two years Farmland Land that is utilised for cultivation of crops Roadside Vegetation up to 30m from either side of the road Pasture land This habitat exists in areas once cleared for cultivation and have been abandoned. Current use of this habitat may be as pasture land for grazing livestock. Secondary vegetation Lands once cleared for agriculture disturbed for firewood or timber extraction Deciduous Seasonal This vegetation type experiences the same conditions as semi-evergreen seasonal forest. The canopy layer Forest reaching 3-9 m in height with an emergent layer 12-18 m. Buttressing is again not a feature of the trees in this vegetation type. Palms are absent while lianes and epiphytes are rare. Lower Montane This habitat type experiences a high evaporation rate. The canopy is closed and reaches heights up to 21-30 m. Rainforest The lower level consists of small tree ferns while the ground cover is extremely sparse. Again buttressing is not exhibited by the trees. Palms, lianes and epiphytes are rare in this vegetation type. Montane Forest Land has steep topography, with shallow, leached and often water-logged soils. High precipitation is characteristic of this habitat (200-254cm per annum). Palm brake This habitat type is comprised almost entirely of Prestoea montana , with a lower stratum of immature trees and palms, with ferns and heliconias comprising the understory vegetation. The palms vary from 3 m in exposed areas to 12 m in sheltered areas. Humid Valley forest This habitat type is found in small areas in the south-western volcanic regions of the island of Antigua. The dominant species in this habitat type consist of Ceiba pentandir , Ficus spp., and Delonix regia . There is a low concentration of vines, epiphytes and palms. Dry Forest Vegetation is of moderate height, with many shrubs and vines. Trees such as Ceiba pentandra and Ficus citrifolia may exceed 9 m. Vegetation around salt This includes Red Mangrove ( Rhizophora mangle ). ponds Mangrove Mangrove and vegetation under its canopy found along the banks of rivers or lakes or in dry coastal areas. Littoral woodland These areas are near the sea and confined to the north and east coasts of the island. They are exposed to sea- blast which refers to the fine salt spray deposited on the vegetation. They seldom extend inland from the coast for more than approximately 60 m. The vegetation also exhibits the mechanical effect of the wind, where they are streamlined into tables and domes. Elfin Woodland Characterised by small trees and shrubs stunted by constant wind exposure. The high humidity is ideal for epiphytes, like bromeliads, orchids, ferns and mosses. Riparian Vegetation Vegetation found along the banks of rivers or lakes Semi-evergreen This vegetation type is not exposed to violent winds. However, it does experience a drought period of five months. seasonal forest The canopy layer occurs at a height of 6-12 m, with a discontinuous emergent layer reaching 18-24 m. The lower level is dominated by palms. Buttress roots are not common in this habitat but, lianes and epiphytes are frequent. Evergreen woodland The vegetation in this habitat type seldom exceed 9m in height and consist of trees and shrubs with small leathern leaves. Native species include Pisonia fragrans , P. subcordata Burgera simaruba and Tabebuia pallida . Non- native species include Haematoxylum campechianum , Leucaena leucocephala syn. L. glauca and Acacia spp. succulent species such as Agave karatto , Cephalocereus royeni and Opuntia spp. are also found. Inland scrub This exists in areas once cleared for cultivation and have been abandoned. The areas around many of the abandoned sugar plantations exhibit this habitat. Patches of scrub vegetation usually spiny, are found dispersed in the grassland. Rainforest This vegetation type usually consists of broad-leaved trees and has a continuous canopy, commonly 30–36 m in height as well as emergent trees, 60 m or taller. The canopy layer comprises of trees of many different sizes, including pygmies, which reach only a metre. Dry Evergreen forest Bushes and gnarled little trees with hard evergreen leaves that form low woodland 3-9 m in height. Sclephron sp. nests Cells of the solitary mud wasp Sceliphron sp. Coastal Vegetation Vegetation comprises mainly of sea grapes ( Coccoloba uvifera ). Most vegetation is stunted and wind sculpted where wind exposure is high. Coastal/Cactus Scrub Found in areas of shallow and stony soil with an average annual rain-fall of 76 to 102 cm. Besides cacti, trees which grow in this habitat are stunted and wind sculpted if wind exposure is high.

Orb-weaving spiders of the Eastern Caribbean 5

Cactus scrub Cactus Coastal veg Coastal

5

nests

Scelphron Scelphron 3 Rainforest

1 Inland scrub Inland

2 3

woodland

Evergreen Evergreen 3 3 3 2

evergreen evergreen

Semi- 5 6 6

Vegetation

Riparian Riparian 6 6 Elfin woodland Elfin

7

Woodland

Littoral Littoral 5 Mangrove

6

s d n o p around salt salt around

4 4 1 4 4 4 5 1;5

Vegetation Vegetation 3;4;5 1 4;5;6 6 Dry Forest Dry

4 1;3;4 5 1;4;5;6 7 4 5 3 2 3 5;6 3 4

Forest

Humid Valley Valley Humid Palm brake Palm

2 Montane

5

Lower montane Lower

forest

Deciduous Deciduous

Forest

Secondary Secondary 6 6 4

Pastureland Roadside

6 6 1 6 2 6 266 4

4;5 Farmland

1 6 6

1;6 4

buildings

Abandoned Abandoned

Buildings Buildings

In and on on and In Garden 6 2;4 2 5 1;3;4 4 1;2;4 1;4 1;4;5 1;2;5 1;2;3;7 2;6 5 2 1;3;4;5;7 1 5 5;7 4;6 5 3 1;2 3 5 4;2;5 4 3 2 1;2;3;5 4;5;2;1 7 4;1 5 2 6 5 4 5 5;6 7 5;6 3 6 2;4;5;61;2;3;7 2;4;7 5;7 2;7 1;5;6 1;7 1;2;3;4;5;6;7 1 1;5;7 2;3;5;6 2;6 1 1;2;6;7 6 5;7 6 5 5;7 1 4 1;3;4;7 1;3;4 5;6 1;5;7 4 1;7 4;6 5 5;6 4;6;7 3 2 1 2 5 3;4;5;6 1;2;6;7 1;3;7 6;7 1;3;4;5;6;7 4 2;6 6 7 3;4;7 3;4 5;6 1;2;4;5;7 4;6;7 5 3 2 3 5;6 3 1;2;3;4;6 1;2;6 1 1;2 1;2;3;7 sp. sp A sp hamata sp. sp. cf sp. sp. 2 4 – Antigua;––– Montserrat; 6 Grenada. St. Lucia 7 5 – 4 sp. D 5 sp. C sp. B sp. A sp. . Showing species composition for the orb-weaving families, according to habitat for spider surveys of Eastern Caribbean. Key: 1 – St. Kitts; 2 – Nevis; 3 – Anguilla; – 3 Nevis; – 2 Kitts; St. – 1 Key: Caribbean. Eastern of surveys spider for habitat to according families, orb-weaving the for composition species Showing . Alpaida Araneus Araneus Chrysometa Tetragnatha nitidens Tetragnathid sp A Tetragnathid sp Theridiosomatidae A. Sp Sp B. Araneus Argiope trifasciata Metepeira compsa Mangora Neoscona neothesis Neoscona cf oaxcensis Spilasma sp. Wagneriana Symphytognathidae A Sp Tetragnathidae Mysmenidae A Sp Eustala anastera Eustala fuscovittata Gasteracanta cancriformis Kapogea Cyclosa walkenari Eriophora Cyclosa caroli Leucauge argyra Leucauge regnyi nr Araneidae Anapidae A Sp. Alcimosphenus lincinus Araneus Acacesia Argiope argentata Table 3 Table 6 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

Table 4. Showing the total island sample and mean per site for; abundance (N), observed species richness (S), species diversity ( +ƍ ), species evenness (1/ D) and dominance ( d) per habitat for Grenada, Montserrat, Nevis, St. Kitts, St. Lucia Antigua and Anguilla sampled between January 2006 and August 2010.

Island N(Act.) N(Mean) S(Act.) S(Mean) +ƍ (Act.) +ƍ (Mean) 1/D (Act.) 1/D (Mean) d(Act.) d(Mean) Grenada 110 13.8 8 2.5 1.98 0.8 5.59 3.04 0.39 0.06 Montserrat 313 24.1 13 3.3 2.38 0.9 10.67 2.9 0.35 0.06 Nevis 311 25.5 12 4.1 1.87 1 5.61 2.9 0.34 0.05 St. Kitts 380 31.7 12 3.4 2.32 0.8 7.89 2.5 0.28 0.04 St. Lucia 358 22.4 16 4.3 2.06 1 6.41 4.1 0.45 0.05 Antigua 220 14.7 15 4.5 2.09 1 7.55 3.3 0.15 0.05 Anguilla 258 21.5 12 4.1 1.48 1 3.19 3.9 0.25 0.05

KDELWDWVZLWKIRXUDQG¿YHVSHFLHVUHVSHFWLYHO\ 7DEOH  DQGHO¿QZRRGODQG7KLVUHVXOWZDVQRWVXUSULVLQJVLQFH Sampling yielded a total of 33 species from six families these low diversity habitats are found at very high alti- for this region. Grenada was found to contain the lowest tudes for example in in Nevis (Sewlal and Starr 2007), species richness (Table 4) while St Lucia and Antigua had St. Kitts (Sewlal 2008), Montserrat (Sewlal 2010) and the highest and second highest values. Notably Nevis, St. Grenada (Sewlal 2009b). Few species can withstand the Kitts and Anguilla have the same species richness values. harsh conditions present such as sea blast, salt and hur- However, single factor ANOVA test observed species ricanes. Although both littoral woodland and coastal veg- richness (F 6,72   3    ZDV QRW VLJQL¿FDQWO\ etation experience similar conditions, the coastal habitat different among the islands included in this study. contains fewer species than littoral woodland with 11 and 7KH UDQNDEXQGDQFH FXUYH SURGXFHG )LJ   ¿WWHG six species respectively. This is possibly due to the lack of only the log series model ( D = 0.151; D 0.05 = 0.154). study vegetation in coastal habitat which does not provide In terms of the similarity of species composition adequate support for webs. Most of the individuals pre- among the islands, analysis (Fig. 3) showed three distinct sent were found on the side facing inland, rather than that groups. Antigua, St Kitts and Nevis formed one cluster facing the ocean, indicating that this location provided the with Montserrat, Anguilla and Grenada forming a second spiders with protection from elements such as wind. cluster with Montserrat, with Montserrat and Anguilla The high species richness exhibited by both garden being more similar to each other than either to Grenada. St. and roadside habitats is not consistent with previous stud- Lucia contained the most distinct assemblage of species. ies, which indicate that both of these habitats provide ideal conditions for spiders. For example, they contain gaps in DISCUSSION WKHYHJHWDWLRQIRUÀ\LQJSUH\ZKLFKFDQEHEORZQLQWRRU The results revealed that in natural habitats, highest À\LQWRWKHZHEVRIRUEZHDYHUV7KHYDULHW\RIYHJHWDWLRQ species richness was found in littoral woodland and the means many suitable points of attachment for web con- vegetation around salt ponds, while lower species richness VWUXFWLRQ$OVR WKH SUHVHQFH RI DUWL¿FLDO OLJKWLQJ DWWUDFWV was exhibited in high altitude habitats such as, palmbrake

Fig. 3. Multidimensional scaling plot showing the species com- Fig. 2. Rank abundance curves for average abundances of all position similarity for Anguilla (ANG), Grenada (GRN), Montser- islands sampled (Montserrat, Nevis, St. Kitts, St. Lucia Antigua rat (MONT), Nevis (NEV), St. Kitts (SKB), St. Lucia (SLU) and and Anguilla). Antigua (ANU). Orb-weaving spiders of the Eastern Caribbean 7

À\LQJLQVHFWVWKXVSURYLGLQJDUHODWLYHO\VWHDG\IRRGVXS - to disturbance by wind and rain. However, it has more ply to nocturnal as well as diurnal species. time to recover in terms of its biodiversity. Montserrat has The low level of species richness exhibited by altered also experienced disturbance but in this case due to the habitats such as pastureland can also be explained by the constant activity of the Soufriere Hills volcano which has high level of disturbance caused by grazing, little protec- rendered approximately two-thirds of the island uninhab- tion from the elements and lack of stable structures for web itable by people and explaining why it yielded the second construction. Although farmland is considered disturbed lowest observed species richness (Table 4). habitat, like the pastureland it exhibited a higher species Sampling in St. Lucia was concentrated in the South richness, most likely as it provides a good food source and East of the island as they had the largest areas of nat- for spiders since many agricultural pests are insects. It is ural habitats to sample (Sewlal 2011). Also some of the important to note that the low species richness in certain forested habitat sampled comprised of secondary vegeta- habitats, such as dry evergreen forest, cactus scrub, low tion which accounted for the high species richness. The montane forest and rainforest, were most likely a by-prod- disturbance in this habitat caused by commercial logging uct of these habitats only being present on a single or few has created new niches, therefore possibly containing both islands. specialist and generalist species (Sewlal 2011). St. Lucia Few species were also found on buildings (both occu- is also the largest of the seven islands sampled, so this pied and abandoned). This is expected since most of the coupled with the mixture of natural and disturbed habi- species found here were also found in the neighbouring tats could account for the more habitats and microhabitats garden habitat (Table 1). As with the garden habitat, these and, in turn, the distinct species composition found on this DUHDRIWHQKDYHDUWL¿FLDOOLJKWZKLFKZLOODWWUDFWDUHOD - island. tively constant supply of insects thus providing a stable The community structure of spiders on these islands source of food. However, the exterior of these manmade ZDVIRXQGWR¿WWKHORJVHULHVDEXQGDQFHPRGHOLQGLFDWLQJ structures did not provide many suitable points of attach- that the species that make up this community arrived at ment for web placement. random intervals (May 1975), which makes sense as these The similarity among St. Kitts, Nevis and Antigua are oceanic islands. Oceanic islands have volcanic origins in terms of species composition (Fig. 3) is expected as therefore all of the fauna found there was brought over by they are relatively close to each other (Fig. 1) (Sewlal KXPDQVRUQDWXUDOO\VXFKDVRQÀRDWLQJPDWVRIYHJHWD - and Starr 2010), of similar size and share similar habi- tion or migration from continental land masses, depending tats. Montserrat, Anguilla and Grenada also have simi- on their distance from the island or nearby islands. It also lar species compositions (Fig. 3). Although these islands implies that the community has a simple ecology (May are fairly far apart geographically (Fig. 1), they have all 1975) dominated by a few factors (Magurran 2004; May experienced high disturbance events. Hurricanes have 1975). Some of the advantages of this model are that it is profound effects on web-building spiders, for example QRWLQÀXHQFHGE\VDPSOHVL]HRUWKHDEXQGDQFHRIWKHPRVW causing a decrease in population densities (Spiller and common species (Taylor 1978). Schoener 2007). Grenada has experienced two hurricanes Orb-weavers occupy a wide range of habitats, and successively in recent years; Hurricanes Ivan and Emily Araneidae was consistently the most ecologically diverse hit the island in 2005 and 2006 respectively resulting in a family in all islands sampled (Sewlal and Starr 2007; change in the understory vegetation structure. For exam- Sewlal 2008, 2009a, 2009b, 2010, 2011). It was followed ple, in the montane forest on Mount Qua Qua the natural by Tetragnathidae in terms of family diversity, except when vegetation has been replaced with dense stands of razor it was replaced by Oxyopidae in St Lucia (Sewlal 2011) grass ( Scleria secans ). This indicates that this habitat is and Salticidae in Montserrat (Sewlal 2010). However, the going through primary succession, providing a habitat for SUHIHUHQFH RI FHUWDLQ VSHFLHV IRU VSHFL¿F KDELWDWV LV QRW generalist species (Sewlal 2009b), which may explain the unusual especially with respect to orb-weavers. lowest observed species richness here (Table 4). Hurricane Two big orb-weaver species ( Araneus and Eriophora ) GDPDJH WR YHJHWDWLRQ LV D PDMRU LQÀXHQFH RQ WKH RUE were recorded from the islands. However, Nephila clavi- weaver fauna of this island, as indicated by the high spe- pes belonging to Nephilidae was absent from despite being cies richness found in the dry forest habitat which exhib- present on nearby islands like Trinidad and Tobago (Sewlal ited little damage from the hurricanes (Sewlal 2009b). 2013) and the Greater Antilles (World Spider Catalogue Anguilla has also experienced hurricane damage with the 2016). For future work, continental islands like Trinidad most recent being Hurricane Lenny in 1999. This island is and Tobago can be compared with that of the females of DOVRTXLWHÀDWZLWKDPD[LPXPHOHYDWLRQRIP 6HZODO this species are quite large, measuring between 24 to 40 and Starr 2010); most of the vegetation is short and subject mm in length, producing webs often spanning a metre in 8 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018 width. In the available habitats it was observed that mem- Coddington, J.A. and Levi, H.W. 1991. Systematics bers of Leucauge constructed web up to a metre in width and evolution of spiders. Annual Review of Ecology and so that they vegetation can support and facilitate webs of Systematics,  this size. Therefore the noted absence of very large orb- Coyle, F.A. 1981. Effects of clearcutting on the spider weaving species on the islands of the Eastern Caribbean community of a Southern Appalachian forest. Journal of may be related to their size in that the habitats available Arachnology.  on these islands cannot facilitate webs of that size and pro- Downie, I.S., Butterfield, J.E.L. and Caulson, J.S. 1995. vide enough prey to sustain spiders of this size. Habitat preferences of sub-montane spiders in Northern For future work, I would recommend that similar stud- England. Ecography,  ies be carried out on other spider groups, such as plant Clarke, K.R. and Warwick, R.M. 2001. Change in marine and ground wandering species. Also the results from these FRPPXQLWLHVDQDSSURDFKWRVWDWLVWLFDODQDO\VLVDQGLQWHU - studies can be compared with continental islands such as SUHWDWLRQQGHG3O\PRXWK35,0(5( Trinidad and Tobago. Enders, F. 9HUWLFDOVWUDWL¿FDWLRQLQRUEZHEVSLGHUV (Araneidae, Araneae) and a consideration of other meth- ACKNOWLEDGEMENTS ods of coexistence. Ecology,  I would like to thank Adrian Hailey for comments on Enders, F   7KH LQÀXHQFH RI KXQWLQJ PDQQHU RQ an earlier draft of this paper and advice on data analysis, prey size, particularly in spiders with long attack dis- *LUDOGR$OD\yQ*DQG%UXFH&XWOHUZLWKLGHQWL¿FDWLRQRI tances (Araneidae, Linyphiidae, and Salticidae). American material. This project was partially funded by grants from Naturalist,  the National Science and Technology Council (NSTC) Enders, F.A . 1976. 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Living World, Journal of the of Arachnology,  Trinidad and Tobago Field Naturalists’ Club,  World Spider Catalog. 2016. World Spider Catalog. ver- Sewlal, J.N. 2010. A Preliminary Survey for Spiders on sion 17.0, Natural History Museum Bern, [Online] avai- Montserrat, West Indies. Living World, Journal of the ODEOHDWKWWSZVFQPEHFK $FFHVVHG)HEUXDU\ Hawk-moths (Lepidoptera: Sphingidae) of Trinidad, West Indies: an illustrated and annotated list Matthew J.W. Cock CABI, Bakeham Lane, Egham, Surrey TW20 9TY, UK. [email protected] / [email protected] ABSTRACT $QLOOXVWUDWHGDQGDQQRWDWHGFDWDORJXHLVSUHVHQWHGRIVSHFLHVRI6SKLQJLGDHFRQ¿UPHGWRRFFXULQ7ULQLGDG Adhemarius daphne (Boisduval) , Aleuron chloroptera (Perty), Nyceryx stuarti (Rothschild) and Erinnyis impunctata Rothschild & Jordan DUHQHZUHFRUGVIRU7ULQLGDG,QDGGLWLRQ¿YHSULRUVSHFLHVUHFRUGVKDYHQRWEHHQFRQ¿UPHGDQGVHYHQUHSRUWHGVSHFLHV DUHFRQVLGHUHGWREHLQHUURU,PDJHVRIOLYLQJDGXOWVRIVSHFLHVDQGWKHHDUO\VWDJHVRIVL[VSHFLHVDUHLQFOXGHG9HU\ OLWWOHKDVEHHQUHFRUGHGUHJDUGLQJIRRGSODQWVDQGHDUO\VWDJHVLQ7ULQLGDGEXWNQRZQIRRGSODQWVIURPHOVHZKHUHLQWKH 1HRWURSLFVDUHWDEXODWHGDVDUHGDWDRIWKHPRQWKVRIFDSWXUHDQGORFDOLWLHVLQVXEXUEDQDQGIRUHVWHGORFDOLWLHVLQ7ULQLGDG Key words: FDWDORJXHGLVWULEXWLRQIRRGSODQWVKRUQZRUPVSKHQRORJ\7REDJR INTRODUCTION Of all the moth families in Trinidad, Sphingidae, pop- Accordingly species names follow the original spelling ularly known as hawk-moths and hornworms have been and do not show gender agreement in the case of Latin and most studied. They are large, colourful and conspicuous, Greek adjectival species names. Some authors have recog- easily recognised and frequently seen by naturalists and nised forms in two of the Trinidad species ( Erinnyis las- the public, most frequently when attracted to house lights. sauxii (Boisduval), Erinnyis obscura (Fabricius)). These Globally, their taxonomy is well understood (at least com- DUHLQIUDVSHFL¿FKDYHQRWD[RQRPLFVWDQGLQJDQGDUHV\Q - pared to nearly all other moth families), and they have been onyms of the given species (see e.g. Kitching and Cadiou used as model groups for biological and ecological studies 2000). However, inasmuch as the names of forms can be in several areas of the tropics (Moss 1912, 1920; Janzen conveniently used to refer to particular phenotypes of pol- 1986, Kawahara et al . 2013; Schreiber 1978; Stradling et ytypic species, they are useful. Thus, it is easier to refer al . 1983; Tuttle 2007). to ‘ Erinnyis obscura f. domingonis (Butler)’ and thereaf- As discussed by Cock (2003), the moths of Trinidad ter ‘f. domingonis ’ than to repeatedly refer to ‘a form of have been documented by W.J. Kaye in a preliminary cat- Erinnyis obscura with dark forewing markings’. alogue (Kaye 1901) and a catalogue (Kaye and Lamont Kawahara et al . (2009) examined the phylogeny of 1927), to which there is a short supplement (Lamont and WKH6SKLQJLGDHEDVHGRQ¿YHQXFOHDUJHQHVGHPRQVWUDW - Callan 1950). Kaye (1904) was reprinted in the Proceedings ing that more work is needed to clarify the natural groups of the Agricultural Society of Trinidad and Tobago of 1914 of Sphingidae, particularly below the subfamily level. (Kaye 1914a), with occasional supplementary informa- Although their work indicates some problems relating to tion mainly relating to common names, probably added by VXEIDPLOLHVDVSUHVHQWO\GH¿QHGWKLVLVQRWDQLVVXHZKHQ P.L. Guppy. There has been relatively little published on applied to the Trinidad fauna, and so the three subfamilies the Trinidad hawk-moths since then, apart from an inten- Smerinthinae, Sphinginae and Macroglossinae are used sive light trapping study on Sphingidae by Stradling et al . here without reservations. The tribes they recognised can (1983), which provides an important resource for what fol- EHDSSOLHGWRWKHOLPLWHG7ULQLGDGIDXQDZLWKOLWWOHGLI¿ - lows. I have recently published a checklist of the moths culty (Table 1). However, since then Kitching (2017) has of Tobago (Cock 2017b), which included 26 hawk-moth built on their phylogeny and used subtribes to encompass species. Treatments are available for Trinidad Castniidae the diversity Kawahara et al . (2009) showed, although this (González and Cock 2004), Mimallonidae (St Laurent and has not been formally published yet. Here I use Kitching’s Cock 2017) and Sematuridae (Cock 2016), but these are tribes and subtribes, and incorporate the species groups all relatively small families in Trinidad (Cock 2003). This evident in Kawahara et al. (2009) as shown in Table 2. illustrated catalogue of the Sphingidae of Trinidad is the ¿UVWRIWKHPHGLXPVL]HGIDPLOLHVWREHWUHDWHGDQGLWLV ,GHQWL¿FDWLRQ hoped that other families will be tackled in the future. When working with pinned specimens in good condition, DOO RI WKH 7ULQLGDG VSHFLHV FDQ EH LGHQWL¿HG E\ FDUHIXO 1RPHQFODWXUHDQGFODVVL¿FDWLRQ . examination and comparison with the images provided Nomenclature follows Kitching and Cadiou (2000) here. Diagnostic features are pointed out in the text which and Kitching (2017), and where this has changed since should help with similar species. Many naturalists are Kitching and Cadiou (2000) an explanation is provided. now taking photographs rather than specimens in support 10 +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 11

Table 1. 3URYLVLRQDOFODVVL¿FDWLRQRIWKH7ULQLGDG6SKLQJLGDHEDVHGRQ.DZDKDUD et al   7KHJURXSQDPHVZLWKLQ'LORSKRQRWLQL DUHQRWXVHGLQ.DZDKDUD et al   EXWDUHEDVHGRQWKHLUSK\ORJHQHWLFWUHH Subfamily Tribe Group Genera reported from Trinidad (and number of Trinidad species) 6PHULQWKLQDH $PEXO\FLQL Adhemarius    Protambulyx   Sphinginae ? Cocytius group Cocytius    Amphonyx    Neococytius   Sphingini Sphingini sensu stricto Manduca   Acherontiini Agrius   Macroglossinae 'LORSKRQRWLQL Pachylia group Pachylia   Nyceryx group Callionima (4), Nyceryx (4), Perigonia    Eupyrrhoglossum  DQG Aellopos (5) Erinnyis group Oryba*    Pachylioides    Madoryx    Hemeroplanes    Erinnyis    Isognathus    Pseudosphinx    Phryxus   Aleuron group Aleuron   Unzela   Enyo and Eumorpha group* Enyo   Eumorpha  Pachygonidia group Pachygonidia   &KRHURFDPSLQL Hyles    Xylophanes   :HDNO\VXSSRUWHGJURXSVRUSODFHPHQWVLQ.DZDKDUD et al    Table 2. &ODVVL¿FDWLRQRIWKH7ULQLGDG6SKLQJLGDHDIWHU.LWFKLQJ  LQFOXGLQJWKHJHQHUDJURXSVRI.DZDKDUD et al.   Subfamily Tribe Subtribe Genera reported from Trinidad (and number of Trinidad species) 6PHULQWKLQDH $PEXO\FLQL Adhemarius    Protambulyx   Sphinginae Sphingini Cocytiina Cocytius    Amphonyx    Neococytius   Sphingina Manduca   Acherontiina Agrius   Macroglossinae 'LORSKRQRWLQL 'LORSKRQRWLQD Pachylia group : Pachylia   Nyceryx group: Aellopos (5), Callionima (4), Eupyrrhoglossum   Nyceryx (4), Perigonia   Erinnyis group: Erinnyis    Hemeroplanes    Isognathus   Madoryx    Oryba    Pachylioides    Phryxus    Pseudosphinx   3KLODPSHOLQD Aleuron group: Aleuron   Unzela   Enyo and Eumorpha group*: Enyo   Eumorpha  Pachygonidia group: Pachygonidia   Macroglossini &KRHURFDPSLQD Hyles    Xylophanes   :HDNO\VXSSRUWHGJURXSVRUSODFHPHQWVLQ.DZDKDUD et al    Sources of records of their observations. With experience, images of living The basis of this checklist is the published work of PRWKVFDQDOVREHUHDGLO\LGHQWL¿HGDQG,KDYHLQFOXGHG Kaye and Lamont (1927), Lamont and Callan (1950), such images as are available to me to facilitate this (Figs. Schreiber (1978) and Stradling et al . (1983), together with 1-38). However, some of the better diagnostic features of my collecting while resident in Trinidad, 1978-1982 and the hindwings and abdomen may be obscured in images of on subsequent visits. I have supplemented these sources living specimens in natural resting positions. This can be with records and images from numerous colleagues and resolved by taking additional images to show these features contacts, and records from museums and collections as (e.g. Fig. 5 Protambulyx strigilis (Linnaeus) female, Fig. 15 follows: Agrius cingulata (Fabricius)), or more careful examination and comparison of the visible features. Further images of MGCL - McGuire Center for Lepidoptera and Biodiversity, pinned adults, genitalia, diagnostic details, and literature which includes W.J. Kaye’s collection (only selected references can be found on the Sphingidae Taxonomic UHFRUGVFRQ¿UPHGDQGLQFOXGHG Inventory website (Kitching 2017) and images of adults MJWC - the private research collection of M.J.W. Cock, and early stages can be found in the websites of Janzen UK (records from all specimens and unpublished notes and Hallwachs (2017) and Oehlke (2017). compiled and collated) 12 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

NHMUK - Natural History Museum, London, UK, which NMSE - National Museum of Scotland, Edinburgh, contains much historical material collected by F. Birch, UK, which includes the balance of the collections of S.M. Klages and others (only selected records con- Sir Norman Lamont and D.J. Stradling (records from ¿UPHGDQGLQFOXGHG Lamont’s specimens compiled and collated, but only selected records from Stradling’s material are included).

Figs. 1-7. /LYLQJDGXOWVRI7ULQLGDG6SKLQJLGDH   1, Adhemarius gannascus PDOH0RUQH%OHX'HFHPEHU .6RRNGHRSKRWR  2, Adhemarius daphne PDOH%UDVVR9HQDGR-XO\ .6RRNGHRSKRWR  3, Adhemarius palmeri PDOH$VD:ULJKW1DWXUH&HQWUH 0DUFK -0RUUDOOSKRWR  4, Protambulyx strigilis PDOH0RUQH%OHX-DQXDU\ .6RRNGHRSKRWR  5, Protambulyx strigilis IHPDOH3HQDO6HSWHPEHU .6RRNGHRSKRWR  6, Cocytius lucifer PDOH%ULJDQG+LOO0DUFK 0-:&RFNSKRWR  7 Neococytius cluentius IHPDOH&DW¶V+LOO-XQH .6RRNGHRSKRWR  +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 13

OUNHM - Oxford University Natural History Museum, UWIZM - University of the West Indies Zoological which includes material collected by R.M. Farmborough Museum, St. Augustine, Trinidad and Tobago, which and others (records from many, but perhaps not all, includes part of the Sir Norman Lamont collection, F.D. specimens compiled and collated). Bennett’s collection and part of D.J. Stradling’s collec- tion (records from all specimens compiled and collated).

Figs. 8-14. /LYLQJDGXOWVRI7ULQLGDG6SKLQJLGDH   8, Neococytius cluentius PDOH$VD:ULJKW1DWXUH&HQWUH0DUFK 61DQ] SKRWR  9, Manduca albiplaga PDOH%UDVVR9HQDGR-XO\ .6RRNGHRSKRWR  10, Manduca huascara %UDVVR9HQDGR-XO\  .6RRNGHRSKRWR  11, 0DQGXFDÀRUHVWDQ 6LPOD1DWXUH/RGJH-XQH 06%RWKDPSKRWR  12 , Manduca rustica rustica , 0RUQH%OHX'HFHPEHU .6RRNGHRSKRWR  13 , 0DQGXFDGLI¿VVDWURSLFDOLV 6RXWK2URSRXFKH$XJXVW 730DKDUDM SKRWR  14 , Agrius cingulata PDOH%ULJDQG+LOO0DUFK 0-:&RFNSKRWR  14 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Images of adults Images of pinned adults are provided life size under Figs. 1-38 of living adult moths show typical resting each species entry. Recognising that these may not be used SRVLWLRQVWRIDFLOLWDWHWKHLGHQWL¿FDWLRQRILPDJHVRIOLY - in hard copy, a 5cm scale bar is included on each page. ing moths. These are deliberately not representative, but Male and female specimens are both shown where sexual are simply images that I have accumulated from my own GLPRUSKLVPLVVLJQL¿FDQWDQGDWOHDVWRQHVSHFLHVIRUHDFK work and images generously shared by resident and visit- genus is shown as both male and female (where material ing naturalists. For none of these images is a scale avail- is available). Ventral (underside) views are shown where able, although these can be derived from the images pro- there are useful diagnostic features, or space would other- vided of pinned specimens. wise be wasted.

Figs. 15-22. /LYLQJDGXOWVRI7ULQLGDG6SKLQJLGDH   15 , Agrius cingulata IHPDOH0RUQH%OHX6HSWHPEHU .6RRNGHRSKRWR  16 , 3DFK\OLD¿FXV 3HQDO1RYHPEHU .6RRNGHRSKRWR  17, Callionima pan pan 0RUQH%OHX'HFHPEHU .6RRNGHR SKRWR  18, Perigonia lusca lusca PDOH%UDVVR6HFR-DQXDU\ .6RRNGHRSKRWR  19 , Aellopos ceculus 7REDJR(QJOLVKPDQ¶V %D\0DUFK 0.HOO\SKRWR  20 , Aellopos clavipes clavipes 7REDJR(QJOLVKPDQ¶V%D\)HEUXDU\ 0.HOO\SKRWR  21 , Pseudosphinx tetrio PDOH0W6W%HQHGLFW¶V2FWREHU 0-:&RFNSKRWR  22 , Isognathus scyron $ULSR6DYDQQD 6HSWHPEHU .6RRNGHRSKRWR  +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 15

Figs. 23-30. /LYLQJDGXOWVRI7ULQLGDG6SKLQJLGDH   23 , Enyo ocypete PDOH3HQDO-XQH .6RRNGHRSKRWR  24, Eumorpha satellitia licaon 3HQDO-XO\ .6RRNGHRSKRWR  25, Eumorpha vitis vitis 6RXWK2URSRXFKH'HFHPEHU 730DKDUDM SKRWR  26, Eumorpha anchemolus IHPDOH%ULJDQG+LOO0DUFK 0-:&RFNSKRWR  27, Eumorpha fasciatus fasciatus PDOH 3HQDO0DUFK .6RRNGHRSKRWR  28, Eumorpha capronnieri 0RUQH%OHX-XQH 06%RWKDPSKRWR  29, Eumorpha labruscae PDOH6RXWK2URSRXFKH-XO\ 730DKDUDMSKRWR  30 , Xylophanes chiron nechus PDOH0RUQH%OHX'HFHPEHU  .6RRNGHRSKRWR  16 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Figs. 31-38. /LYLQJDGXOWVRI7ULQLGDG6SKLQJLGDH   31, Xylophanes loelia 3HQDO)HEUXDU\ .6RRNGHRSKRWR  32, Xylophanes pluto  3HQDO  -XQH  . 6RRNGHR SKRWR  33, Xylophanes pistacina  3HQDO  1RYHPEHU  . 6RRNGHR SKRWR  34-35, Xylophanes tersa tersa 3HQDO1RYHPEHU .6RRNGHRSKRWR  36, Xylophanes tersa tersa 7REDJR(QJOLVKPDQ¶V%D\0DUFK  0.HOO\SKRWR  37 , Xylophanes titana PDOH6LPOD1DWXUH/RGJH-XQH 06%RWKDPSKRWR  38 , Xylophanes tyndarus tyndarus PDOH0RUQH%OHX0D\ 730DKDUDMSKRWR  +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 17

Figs. 39-49.  (DUO\ VWDJHV RI 7ULQLGDG 6SKLQJLGDH 39, Manduca sexta paphus PDWXUH FDWHUSLOODU $UDQJXH] *DUGHQV RQ WRPDWR 6RODQDFHDH XQGDWHG 0-:&RFNSKRWR  40-41, Manduca sexta paphus PDWXUHFDWHUSLOODU QRWUHDUHG 7REDJR(QJOLVKPDQ¶V%D\ RQRUQDPHQWDO Brugmansia VS 6RODQDFHDH -DQXDU\ 0.HOO\SKRWR  42 , Agrius cingulata FDWHUSLOODU$ULPD9DOOH\6LPOD on Ipomoea VS &RQYROYXODFHDH SDUDVLWL]HGE\XQLGHQWL¿HG0LFURJDVWHULQDH FRFRRQV -DQXDU\ 0-:&RFNSKRWR  43-45 , Erinnyis alope alope FDWHUSLOODU QRWUHDUHG 7REDJR(QJOLVKPDQ¶V%D\RQSDSD\D-DQXDU\ 0.HOO\SKRWR  46 , Isognathus scyronPDWXUHFDWHUSLOODU6RXWK2URSRXFKH$XJXVW 730DKDUDMSKRWR  47-48 , Isognathus scyron SXSDUHDUHGIURP Allamanda cathartica $SRF\QDFHDH &XUHSHXQGDWHG 0-:&RFNSKRWR  49 , Pseudosphinx tetrio PDWXUHFDWHUSLOODURQ Allamanda cathartica , 7REDJR$UJ\OO%D\)HEUXDU\ &'$GDPVSKRWR  18 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Vagility are not treated here, as information can be found on-line Some species of hawk-moths are well known to dis- HJKWWSVEUHHGLQJEXWWHUÀLHVFRPJHQHUDOLQIRUPDWLRQ SHUVHE\À\LQJJUHDWGLVWDQFHVRIWHQDVUHJXODUPLJUDQWV http://sphingidae-haxaire.com/index.php/study/live- (Hodges 1971, Tuttle 2007), and in Costa Rica many spe- stock/). There is considerable scope for documenting the cies move long distances into drier forests during the rainy life histories of Trinidad and Tobago’s hawk-moths (and season (Janzen 1986). However, there is little informa- indeed other Lepidoptera), e.g. Cock (2007, 2008). tion regarding this behaviour in Trinidad, and it may be that some species recorded from the island are not resi- Collecting dent, e.g. Hyles lineata (Fabricius) (see below). Writing Most collecting of hawk-moths in Trinidad, at least about the Caribbean species, Cary (1951) lists Agrius cin- in the last 50 years, has been by using lights, particularly gulata , Manduca rustica (Fabricius), Cocytius antaeus mercury vapour lights (MVL) with a strong ultra-violet (Drury), Protambulyx strigilis , Erinnyis ello (Linnaeus), component, to attract nocturnal adults. Based on this E. alope (Drury), 3DFK\OLD¿FXV (Linnaeus), Pachylioides collecting method, Stradling et al . (1983) recorded all resumens (Walker), Eumorpha satellitia (Linnaeus), E. Sphingidae for nearly 4000 light trap nights at Curepe and vitis (Linnaeus), E. fasciatus (Sulzer) and E. labruscae XVHGWKHVHGDWDWRH[DPLQHSDWWHUQVRIÀLJKWLQUHODWLRQWR (Linnaeus) as likely to disperse over wide stretches of rainfall and the lunar cycle, and how species frequency water. In passing, Cock and Boos (2007) reported that and diversity has varied over time. Julius O. Boos had collected specimens of Callionima fal- As an aside, it may be noted that CAB International cifera (Gehlen) and Enyo lugubris (Linnaeus) ‘at light on (CABI) staff (including F.D. Bennett, R.E. Cruttwell, the an oilrig in the sea between Venezuela and the island of author, etc.) have labelled material as from Curepe, when Trinidad’. I checked these specimens, now in UWIZM, actually it is from St. Augustine (based on the outlines while preparing the present account. They are labelled shown for Curepe and St. Augustine on Google maps). ‘Trintes Platform B, East of Mayaro Beach’ August 1978, The reasons for this are twofold: (1) although the part of i.e. to the east of Trinidad. Trintes B was deployed in the Gordon Street where the CABI premises are situated is Galeota Marine Field, east of Mayaro, from June 1978 LQ 6W$XJXVWLQH WKH &$%, RI¿FH XVHV WKH &XUHSH SRVW (MPM 1981), and is 8-9km offshore, northeast of Point RI¿FHDQGKHQFHWKHSRVWDODGGUHVVLV&XUHSHDQG  LQ Galeota (Petroleum Economist 2015), so Cock and Boos those days, data labels were hand written or hand printed (2007) referring to the platform as between Trinidad using a small hand-printing press, and the attraction of and Venezuela is misleading. Further records from off- writing or preparing hundreds of data labels with the short shore rigs would add to our insight regarding the species ‘Curepe’ rather than the long ‘St. Augustine’ is obvious. involved, and frequency and scale of the movement of This body of data from Curepe (= St. Augustine) is com- these moths. plemented by records from collecting at light in forested areas, most notably at the Morne Bleu Textel Installation Immature stages and at Simla in the Arima Valley, but with observations There is very little recorded regarding the life histories from many parts of Trinidad (Cock 2003). However, of the Sphingidae in Trinidad, so an overview is provided these represent dozens of evenings collecting compared of food plant records from other parts of the Americas to the thousands of nights trapping in St. Augustine. (Appendix Table 1). Several of the sources listed include The intensive collecting at Curepe (St. Augustine) has images of caterpillars and pupae, notably Moss (1912, UHVXOWHGLQ¿YHVSHFLHVEHLQJRQO\UHFRUGHGIURPWKHUH 1920), Dyer and Gentry (2002), Dyer et al . (2017), Janzen Erinnyis impunctata Rothschild & Jordan, Pachylia syces & Hallwachs (2017), and Oehlke (2017). In the following syces (Hübner), Callionima inuus (Rothschild & Jordan), VSHFLHVDFFRXQWVRQO\OLWHUDWXUHDQGLQIRUPDWLRQVSHFL¿F Aleuron chloroptera (Perty), and Hyles lineata (Appendix to Trinidad and Tobago is included. The few images avail- Table 2). However, when other habitats are better col- able of early stages of Trinidad hawk-moths are shown as lected, this exclusivity to Curepe is expected to disappear; Figs. 39-49. Hawk-moth caterpillars have a distinctive indeed, two further species were removed from this list as and characteristic horn or tail on the posterior end of the WKLVSDSHUZDVEHLQJ¿QDOL]HG body (hence the common name hornworms). Although In Appendix Table 2, I have attempted to classify spe- this tail may be greatly reduced in mature caterpillars of cies as being commonly (+++), occasionally (++), rarely some species, its presence will normally serve to distin- (+), or effectively never (-) associated with (i.e., collected guish hawk-moth caterpillars from those of other families in) suburban habitats and forested areas. However, the for the Trinidad fauna (but not necessarily elsewhere). aforementioned intensive collecting in Curepe is expected Pupation is generally in a cell in the soil. Rearing methods to have led to a corresponding overemphasis on suburban +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 19 habitats in this analysis. the individual records are listed. I also give an indica- The hummingbird hawk-moths, Aellopos VSSÀ\E\ tion of the habitats in which species have been found, but GD\DQGIHHGDWÀRZHUVUHVHPEOLQJVPDOOKXPPLQJELUGV UDUHO\ GLIIHUHQWLDWH PRUH WKDQ VXEXUEDQ DV H[HPSOL¿HG DQGKDYHEHHQRFFDVLRQDOO\FDXJKWXVLQJQHWVE\EXWWHUÀ\ by Curepe / St. Augustine) and forested, although where FROOHFWRUVRUSKRWRJUDSKHGKRYHULQJDWÀRZHUV:KLOH, there seems to be a pattern, the latter is characterised by was resident in Trinidad, malaise traps were run at several altitude or the part of the island. Additional information ORFDWLRQVWRFROOHFWÀ\LQJLQVHFWVLQWRHWKDQRO,VRUWHG on months and localities of captures is summarised in some of these catches from Curepe and the Aripo Valley to Appendix Tables 2 and 4. record the Sphingidae, but found only small numbers of a 6SHFLHVHQWULHVIRUVSHFLHVZKLFK,FDQFRQ¿UPDV few species by this collecting method. occurring in Trinidad follow below. These are followed by There is a group of genera that are poorly documented sections on species recorded from Trinidad which I have from Trinidad, because they are not readily attracted QRWEHHQDEOHWRFRQ¿UP VSHFLHV DQGVSHFLHVUHFRUGHG to light: Nyceryx Boisduval (four species), Perigonia from Trinidad, which I consider to be errors (7 species). Herrich-Schaffer (two species), Aellopos Hübner (four species), Oryba Walker (two species), Aleuron Boisduval The Sphingidae of Trinidad (two, possibly four species), Unzela Walker (one, possi- bly two species), and Pachygonidia Fletcher (one, pos- Family: Sphingidae sibly two species) (Haxaire 1992, 1996a, 1996b, 2005). Subfamily: Smerinthinae As noted above, Aellopos  VSS DUH SULPDULO\ GD\À\LQJ Tribe: Ambulycini though they continue to feed until dusk; the remaining 7KH¿YH7ULQLGDGVSHFLHVRIWZRJHQHUDDUHHDVLO\UHF - VSHFLHV SULPDULO\ À\ DW GXVN DQG GDZQ DQG DUH KDUGO\ ognised by the elongate forewings with an apical notch, attracted to lights, apart from Perigonia lusca (Fabricius), and the coloured dorsal hindwings: orange in Protambulyx which is regularly caught at light. All these species are Rothschild and Jordan and pink in Adhemarius Oiticica PRVWOLNHO\WREHVHHQDQGFROOHFWHGZKHQFRPLQJWRÀRZ - Filho. HUV7KHÀRZHUVWKDWDUHPRVWDWWUDFWLYHLQFOXGHVSHFLHVRI Calliandra and Inga (Fabaceae, Mimosoideae) (Haxaire 1. Adhemarius gannascus (Stoll, 1790) (Figs. 1, 50-51) 1992, 1996b). Eupyrrhoglossum Grote (one species) and Amplypterus gannascus (Stoll): Kaye (1914b), Kaye and Enyo +EQHU WKUHHVSHFLHV DUHDOVRDWWUDFWHGWRÀRZHUV Lamont (1927), Cary (1951) at dusk, but then come to light later in the night (Haxaire Amplypterus gannascus gannascus (Stoll): Schreiber F 7KXVIRUWKHVSHFLHVWKDWRQO\À\DWGXVN (1978), Stradling et al . (1983) there are very few records and specimens, and several spe- Adhemarius gannascus (Stoll): Vaglia & Haxaire (2005) cies of this group have been recorded from Trinidad by ,GHQWL¿FDWLRQ . Three species of Adhemarius are recog- Schreiber (1978) for which I have not seen specimens. It nised from Trinidad. Adhemarius palmeri (Boisduval) is seems likely that before collecting at light came to pre- easily separated from the other two, for example, by the dominate, collectors in the late 19 th and early 20 th centu- dark diagonal line beyond the basal band of the forewing; ries were aware of this behaviour and exploited it to col- when seen in life, the lines of the two wings are aligned. lect specimens of these genera. Certainly, it would merit Adhemarius gannascus and A. daphne (Boisduval) below investigation in Trinidad today to better understand the are similar, but the dark mark on the forewing costa just hawk-moth fauna. before the apex is consistently narrow in A. gannascus and wider in A. daphne . SPECIES ACCOUNTS Biology in Trinidad . According to Haxaire and Rasplus In the checklist that follows, the currently accepted (1987a), the food plants are Ocotea sp. (Lauraceae), name of each species (and where appropriate subspecies) based on the work of D.H. Janzen and colleagues (Janzen recorded from Trinidad is followed by a list of all names and Hallwachs 2017). Recently, Janzen and Hallwachs used for the species in the literature relating to Trinidad. (2016) clarify that they have not reared A. gannascus There follows brief notes on (1) taxonomic issues (if any), LWVHOIDOWKRXJKLWÀLHVLQWKHLUVWXG\DUHDDQGWKHVHHDUOLHU  LGHQWL¿FDWLRQ  ELRORJ\LQ7ULQLGDG LIDQ\WKLQJLV records are all referable to cryptic species that had been known), and (4) status in Trinidad. For the last, an indica- confused with A. gannascus . Thus, at this time, the food tion is given as to how rare or common the species is in plants and biology of A. gannascus are unknown. Trinidad, based on the number of records known to the Status in Trinidad . A fairly common forest species. author: rare <5, uncommon 5-10, occasional 11-19, quite or Most records are from the north, but Kaye and Lamont rather common 21-49, and common >50. For rare species, (1927) report it from Fyzabad (R.M. Farmborough). 20 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

2. Adhemarius daphne daphne (Boisduval, [1875]) ,GHQWL¿FDWLRQ . Adhemarius palmeri is easily separated (Figs. 2, 52) from the other two species of Adhemarius spp., for Taxonomic issues. 7KLVVSHFLHVLVLGHQWL¿HGEDVHGRQWKH example, by the dark diagonal line beyond the basal band treatment by Vaglia and Haxaire (2005). There may be a of the forewing. further cryptic species based on DNA barcoding (Janzen Status in Trinidad . An occasional species, found in and Hallwachs 2017; I.J. Kitching pers. comm. 2017). suburban areas and forested areas, but not recorded from Adhemarius daphne has not hitherto been separated from higher areas A. gannascus in publications relating to Trinidad, but the author has long been aware that two species are present 4. Protambulyx eurycles (Herrich-Schäffer, [1854]) with different genitalia, and in the past I have incorrectly (Fig. 54) referred to the present species as A. dentoni (Clark). Based Protambulyx eurycles (Herrich-Schäffer): Schreiber RQPDWHULDOLQ8:,=0,¿QGWKDWVRPHVSHFLPHQVZHUH (1978), Stradling et al . (1983) treated as A. gannascus by Stradling et al . (1983). ,GHQWL¿FDWLRQ . This distinctive species might only be ,GHQWL¿FDWLRQ . As discussed under A. gannascus above. confused with the much commoner P. strigilis below, but Status in Trinidad . An occasional forest species. that species lacks the strong wavy parallel lines on both wings, the dark forewing margin and the strong dark 3. Adhemarius palmeri (Boisduval, [1875]) (Figs. 3, 53) marking on the basal part of the forewing dorsum. Amplypterus palmeri (Boisduval): Kaye (1914b), Kaye Status in Trinidad . I only know of three records, two and Lamont (1927), Schreiber (1978), Stradling et al . from lowland forest, and one from a suburban situation. (1983)

Fig. 50. Adhemarius gannascus gannascus  6WROO PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@

Fig. 51. Adhemarius gannascus gannascus  6WROO IHPDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW6HSWHPEHU 0-:&RFN  >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 21

&XUHSH 09/ ƃ  6HSWHPEHU  0-: &RFN  9DOHQFLD)RUHVW09/ƃ$XJXVW 0-:&RFN  [UWIZM CABI.7042] [MJWC] (Fig. 54) 3DUU\ODQGV2LO¿HOG09/ƃ-XO\ 0-:&RFN  [UWIZM CABI.7043]

Fig. 52. Adhemarius daphne daphne (Boisduval) PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW0DUFK 0-:&RFN >0-:&@

Fig. 53. Adhemarius palmeri %RLVGXYDO PDOH+ROOLV5HVHUYRLUDWSXPSKRXVHOLJKWV1RYHPEHU 0-:&RFN >0-:&@

Fig. 54. Protambulyx eurycles  +HUULFK6FKlIIHU PDOH9DOHQFLD)RUHVW09/$XJXVW 0-:&RFN >0-:&@ 22 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

5. Protambulyx strigilis (Linnaeus, 1771) treated below. They are large species with yellow basally (Figs. 4-5, 55-56) on the hindwing and yellow lateral spots on the anterior Ambulyx strigilis (Linnaeus): Kaye (1901), Kaye (1914a) abdomen. Several species of Manduca Hübner have the Protambulyx strigilis (Linnaeus): Rothschild and Jordan latter character, but not the former. All Trinidad species (1903), Kaye (1914b), Kaye and Lamont (1927), Cary feed on Annonaceae and are notable for their long (1951) proboscides. Protambulyx strigilis strigilis (Linnaeus): Schreiber (1978), Stradling et al . (1983), Cock (2017b) 6. Cocytius duponchel (Poey, 1832) (Figs. 57-58) Taxonomic issues. Janzen and Hallwachs (2017) indicate Cocytius duponchel (Poey): Schreiber (1978), Stradling et that there may be two cryptic species present under this al . (1983) name in Costa Rica, but this possibility has not been Taxonomic issues. Eitschberger (2006) placed this investigated elsewhere. species in the genus Amphonyx , but this has not been ,GHQWL¿FDWLRQ . A distinctive species readily separated widely accepted (e.g. Kitching 2017). from its congener, P. eurycles , above. ,GHQWL¿FDWLRQ . See under C. antaeus below. Status in Trinidad . A common and widespread species, Biology in Trinidad. Reported Annonaceae food plants found in suburban and forest habitats. include Annona muricata (soursop, introduced), A. reticulata (custard apple, introduced) and A. squamosa Subfamily: Sphinginae (sugar apple, introduced), but there are no reports of this Tribe: Sphingini species as a pest in Trinidad. Subtribe: Cocytiina Status in Trinidad . An occasional but widespread species This group of four Trinidad species in two genera is in Trinidad, found in both suburban and forest habitats. basal to the two other recognised subtribes of Sphingini

Fig. 55. Protambulyx strigilis /LQQDHXV PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@

Fig. 56. Protambulyx strigilis /LQQDHXV IHPDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 23

7. Cocytius lucifer Rothschild and Jordan, 1903 8. Cocytius antaeus (Drury, 1773) (Fig. 60) (Figs. 6, 59) Cocytius antaeus medon [ sic ] (Stoll): Kaye (1914b) Cocytius lucifer Rothschild and Jordan: Lamont and [misspelling of medor , a synonym of antaeus ] Callan (1950), Schreiber (1978) Cocytius antaeus medor (Stoll): Kaye and Lamont (1927), Cocytius lucifer lucifer Rothschild and Jordan: Stradling Cary (1951) [ medor is a synonym of antaeus ] et al . (1983) Cocytius antaeus antaeus (Drury): Schreiber (1978) Amphonyx lucifer (Rothschild and Jordan): Eitschberger Cocytius antaeus (Drury): Eitschberger 2006 (2006) ,GHQWL¿FDWLRQ . This large species resembles the previous Taxonomic issues. As for the last species, Eitschberger two, C. duponchel and C. lucifer . Pinned specimens are (2006) placed this species in the genus Amphonyx , but this easy enough to distinguish: the hyaline discal area of the has not been widely accepted (e.g. Kitching 2017). There hindwing has the veins dark in all three species, but only in are two cryptic DNA barcode species under this name in C. antaeus is there a parallel dark line in the distal part of Costa Rica (Janzen and Hallwachs 2011), but this has not each hyaline space. This character is hidden when living been investigated elsewhere. adults are photographed, and so only characters of the ,GHQWL¿FDWLRQ . See under C. antaeus below. forewing can be used: the forewings of C. antaeus and C. Status in Trinidad . An occasional species in forest duponchel are more mottled than the rather evenly coloured habitats. forewing of C. lucifer ; and furthermore C. antaeus has a

Fig. 57. Cocytius duponchel 3RH\ PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@

Fig. 58. Cocytius duponchel 3RH\ IHPDOH9DOVD\Q3DUNDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@ 24 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE single black streak in the discal area, C. lucifer has two, Biology in Trinidad. The Annonaceae food plants and C. duponchel has at most a weak streak. reported include A. muricata (soursop, introduced), A. Biology in Trinidad. As for C. duponchel , although the reticulata (custard apple, introduced), but there are no Annonaceae food plants include A. muricata (soursop, reports of this species as a pest in Trinidad. introduced), A. reticulata (custard apple, introduced) and Status in Trinidad . An uncommon species, primarily in A. squamosa (sugar apple, introduced), there are no reports forest habitats. of this species as a pest in Trinidad. Status in Trinidad . An occasional species in suburban Subfamily: Sphinginae habitats, rarely found in forested areas. Tribe: Sphingini Subtribe: Sphingina 9. Neococytius cluentius (Cramer, 1775) This tribe is represented by the genus Manduca, with ten (Figs. 7-8, 61-62) species in Trinidad. Kawahara et al . (2013) conducted Cocytius cluentius (Cramer): Kaye and Lamont (1927) a multi-gene phylogenetic analysis of Manduca, and Neococytius cluentius (Cramer): Stradling et al . (1983) recognised (1) a basal group, the lefeburii complex, and ,GHQWL¿FDWLRQ . The females of this species probably have three well supported groups referred to here as: (2) the the greatest wing spans of any Trinidad hawk-moth, up ÀRUHVWDQ complex, (3) the sexta complex, and the (4) WRFP$GXOWVDUHVXSHU¿FLDOO\VLPLODUWRWKHODVWWKUHH occulta complex. species, but the forewing is much darker, and the lateral yellow spots on the abdomen extend further distally. The lefeburii complex of Manduca Photographs of living adults with the yellow markings This is the basal group of the genus in the phylogeny of of the hindwing and abdomen potentially hidden may Kawahara et al . (2013). be confused with the smaller Erinnyis alope (and size is not always evident in an image). When at rest, the adult 10. Manduca lefeburii (Guérin-Méneville, [1844]) of N. cluentius holds its wings more tightly against the (Figs. 63) body (Fig. 7) like Erinnyis spp. but unlike Cocytius spp. Protoparce lefeburei [sic ] (Guérin-Méneville): Kaye (e.g. Fig 6). The weak pale marking of the forewing of N. (1914b), Kaye and Lamont (1927) cluentius are closer to the base of the wing, and E. alope Protoparce lefeburei lefeburei [sic ] (Guérin-Méneville): has a dark stripe dorsally the length of the thorax, and a Schreiber (1978), Stradling et al . (1983) strong crest that can be seen in lateral view and sometimes ,GHQWL¿FDWLRQ . A very distinctive species in Trinidad, in dorsal view. although there are similar species in South America.

Fig. 59. Cocytius lucifer 5RWKVFKLOGDQG-RUGDQPDOH%ULJDQG+LOOOLJKWKRXVHVHFXULW\09/OLJKWV0DUFK 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 25

Status in Trinidad . An uncommon species, found in and apical bands of the forewing. suburban and disturbed habitats, but not forested. Status in Trinidad . An occasional species, predominantly in forested areas, especially on the ridges of the Northern 11. Manduca albiplaga (Walker, 1856) (Figs. 9, 64) Range. Protoparce albifuga [ sic ] (Walker): Kaye (1914b) Protoparce albiplaga (Walker): Kaye and Lamont (1927) The ÀRUHVWDQ complex of Manduca Manduca albiplaga (Walker): Cary (1951) The three species treated in this group do not have yellow Manduca albiplaga albiplaga (Walker): Schreiber (1978), markings laterally on the abdomen. Manduca franciscae Stradling et al . (1983) (Clark) and M. huascara (Schaus) were not included in ,GHQWL¿FDWLRQ . This is the largest Manduca species in Kawahara et al .’s (2013) study, but are placed here based Trinidad, and can be further recognised by the white discal on their obvious similarity.

Fig. 60. Cocytius antaeus 'UXU\ IHPDOH6W$XJXVWLQH09/6HSWHPEHU )'%HQQHWW >0-:&@

Fig. 61. Neococytius cluentius  &UDPHU PDOH3DUU\ODQGV2LO¿HOG09/-XO\ 0-:&RFN >0-:&@ 26 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

12. Manduca franciscae (Clark, 1916) (Fig. 65) 13. Manduca huascara (Schaus, 1941) (Figs. 10, 66-67) Manduca franciscae (Clark): Stradling et al . (1983) Protoparce corallina : Kaye (1914b) [probable mis- ,GHQWL¿FDWLRQ . This rare species might easily be LGHQWL¿FDWLRQ@ overlooked as a pale 0ÀRUHVWDQ , but it is slightly smaller, Manduca lichenea (Burmeister): Cary (1951) WKHFRORXULQJLVVLJQL¿FDQWO\SDOHUDQGWKHKLQGZLQJRI >PLVLGHQWL¿FDWLRQ@ 0ÀRUHVWDQ (Stoll) is more uniformly brown. Separation Manduca fosteri (Rothschild and Jordan): Stradling et al. RILPDJHVRIWKHWZRFRXOGEHGLI¿FXOWZLWKRXWYRXFKHU  >PLVLGHQWL¿FDWLRQ@ material. Taxonomic issues. This medium sized Manduca species Status in Trinidad . I know of just two records, from KDVEHHQPLVLGHQWL¿HGLQ7ULQLGDGFROOHFWLRQVLQWKHSDVW Arima Valley and Palmiste. as M. fosteri (Rothschild and Jordan) and M. lichenea $ULPD9DOOH\6LPOD09/ƃ0D\ 0-:&RFN  (Burmeister), which are not Trinidad species. Based on [MJWC] (Fig. 65) appearances, I suspect that this is also the species which 3DOPLVWH Ƃ  -XQH  >1 /DPRQW@ >8:,=0 Kaye (1914b) referred to as M. corallina , although 2013.13.2346] curiously, he does not refer to this species in Kaye and

Figs. 62. Neococytius cluentius  &UDPHU IHPDOH3DUU\ODQGV2LO¿HOG09/-XO\ 0-:&RFN >0-:&@

Fig. 63. Manduca lefeburii PDOH0DQ]DQLOODDWOLJKW-XQH 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 27

Fig. 64. Manduca albiplaga :DONHU PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW0D\ 0-:&RFN >0-:&@

Fig. 65. Manduca franciscae &ODUN PDOH GLVVHFWHG $ULPD9DOOH\6LPOD09/0D\ 0-:&RFN >0-:&@

Fig. 66. Manduca huascara 6FKDXV PDOH$ULPD9DOOH\6LPOD09/)HEUXDU\ 0-:&RFN >0-:&@ 28 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Lamont (1927). It was confused with the common M. 14. 0DQGXFDÀRUHVWDQ (Stoll, 1782) (Figs. 11, 68) ÀRUHVWDQ (below) by Stradling et al . (1983). 'LOXGLDÀRUHVWDQ (Stoll): Kaye (1901, 1914a) ,GHQWL¿FDWLRQ . This is a reasonably distinctive species within 3URWRSDUFHÀRUHVWDQ (Stoll): Rothschild and Jordan (1903), the Trinidad fauna: 0ÀRUHVWDQ is a more evenly coloured Kaye (1914b), Kaye and Lamont (1927) and greener species, with a diagnostic contrasting light brown 0DQGXFDÀRULVWDQ [sic ] (Stoll): Cary (1951) patch at the end of the forewing cell (although care is needed 0DQGXFD ÀRUHVWDQ ÀRUHVWDQ (Stoll): Schreiber (1978), as this patch can be weakly or very weakly expressed in Stradling et al . (1983) some individuals). Images of living adult M. huascara 0DQGXFDÀRUHVWDQ (Stoll): D’Abrera ([1987]), Cock (2017b) might be confused with Agrius cingulata or Pseudosphinx ,GHQWL¿FDWLRQ . This species might be confused with the tetrio (Linnaeus), but careful examination of the details of last two as discussed above. the forewing markings should separate them. Status in Trinidad . A common and widespread species Status in Trinidad . An occasional widespread species, in forested areas. more common in forested areas, but not from the higher parts of the Northern Range.

Fig. 67. Manduca huascara 6FKDXV IHPDOH%ULJDQG+LOOOLJKWKRXVHVHFXULW\09/OLJKWV0DUFK 0-:&RFN >0-:&@

Fig. 68. 0DQGXFDÀRUHVWDQ 6WROO PDOH0RUQH%OHXDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 29

The sexta complex of Manduca 16. Manduca sexta (Linnaeus, 1763) paphus (Cramer, 1779) (Figs. 39-40, 70) 15. Manduca rustica rustica (Fabricius, 1775) Protoparce paphus (Cramer): Kaye (1901), Guppy (1914), (Figs. 12, 69) Kaye (1914a) Macrospila sp.: Guppy (1893) Protoparce sexta (Linnaeus): Guppy (1911a, 1911b) Protoparce rustica (Fabricius): Kaye (1901, 1914a, 1914b) Protoparce sexta paphus (Cramer): Rothschild and Jordan Protoparce rustica rustica (Fabricius): Rothschild and (1903), Kaye (1914b), Kaye and Lamont (1927) , Cock Jordan (1903), Kaye and Lamont (1927) (2017b) Manduca rustica rustica (Fabricius): Schreiber (1978), Protoparce sp.: Topper (1943) Stradling et al . (1983), Cock (2017b) Manduca sexta paphus (Cramer): Cary (1951), Schreiber ,GHQWL¿FDWLRQ   7KH ¿QH JUDLQHG PRWWOLQJ RI WKH ZLQJV (1978), Stradling et al . (1983) and strong white spot in the forewing cell should serve to Manduca sexta jamaicensis (Butler): Schreiber (1978), recognise this species in Trinidad. Stradling et al   >PLVLGHQWL¿FDWLRQRUHUURU@ Biology in Trinidad . Guppy (1893) describes rearing a Manduca sexta (Linnaeus): CABI (2002c) ‘Macrospila sp.’ which he found on ‘heliotrope’, which Taxonomic issues. There appears to be at least two cryptic may well have been the polyphagous Manduca rustica . taxa under this name in Costa Rica (Janzen and Hallwachs Status in Trinidad . A fairly common and widespread 2017), but only a single species in South America based species in both suburban and forest habitats. on the barcodes in BOLD (http://boldsystems.org/index.

Fig. 69. Manduca rustica rustica )DEULFLXV PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@

Fig. 70. Manduca sexta paphus  &UDPHU PDOH&XUHSH09/-XO\ 0-:&RFN >0-:&@ 30 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE php, 24 October 2017). Schreiber (1978) records both M. GLI¿VVD and not a South American species . sexta jamaicensis and M. sexta paphus from Trinidad; the ,GHQWL¿FDWLRQ 7KLVVSHFLHVLVGDUNHUWKDQWKHVXSHU¿FLDOO\ former is restricted the Caribbean islands, including the similar M. sexta above. In living specimens the slightly Lesser Antilles, but I have seen no evidence that it also paler postdiscal lines align across the two wings. occurs in Trinidad. Status in Trinidad . A fairly common, widespread species ,GHQWL¿FDWLRQ . This medium sized hawk-moth has the found in both suburban and forested habitats. forewings rather uniformly grey, and although there are other similar species in South America, this is the only one 18. Manduca hannibal hannibal (Cramer, 1779) (Fig. 72) of this appearance known from Trinidad. Photographs of Protoparce hannibal (Cramer): Rothschild and Jordan living adults might be confused with Agrius cingulata as (1903), Kaye (1914b), Kaye and Lamont (1927) discussed above, or the darker 0GLI¿VVD (Butler) below. Manduca hannibal (Cramer): Cary (1951) Biology in Trinidad. In Trinidad, Guppy (1911b) notes Manduca hannibal hannibal (Cramer): Schreiber (1978), caterpillars on tobacco, tomato and peppers, and Kaye and Stradling et al . (1983) Lamont (1927) refer to this species as the tobacco hawk-moth. Taxonomic issues. There seems to be at least two cryptic Status in Trinidad . A fairly common species in suburban species in Costa Rica (Janzen and Hallwachs 2017) but habitats, but hardly recorded from forests. this has not been evaluated elsewhere. ,GHQWL¿FDWLRQ . A distinctive species; note the orange The occulta complex of Manduca dorsal markings on the abdomen, and the relatively This group is based on M. occulta (Rothschild and Jordan), uniform brown dorsal forewing, with a broad blackish a species closely related to 0GLI¿VD Butler, which does post-discal band and narrow submarginal white not occur in Trinidad. Kawahara et al . (2013) did not Biology in Trinidad. Kaye and Lamont (1927) give include M. ochus (Klug) in their study, and its placement ‘Frombeta (Solanaceae)’ as a food plant, but this is an error is uncertain. for Trombeta , incorrectly transcribed from Moss (1912). Status in Trinidad . An uncommon species found in 17. 0DQGXFDGLI¿VVD(Butler, 1871) tropicalis (Rothschild suburban and forested situations. and Jordan, 1903) (Figs. 13, 71) Protoparce diffusa [ sic ] tropicalis (Rothschild and Jordan): 19. Manduca ochus (Klug, 1836) (Fig. 73) Kaye (1914b) [misspelling] Protoparce ochus (Klug): Kaye (1901), Rothschild and Jordan 3URWRSDUFH GLI¿VVD WURSLFDOLV (Rothschild and Jordan): (1903), Kaye (1914a, 1914b), Kaye and Lamont (1927) Kaye and Lamont (1927), Cary (1951), Schreiber Manduca ochus (Klug): Cary (1951), Schreiber (1978), (1978), Stradling et al . (1983) Stradling et al . (1983) Manduca occulta occulta Rothschild and Jordan: Schreiber ,GHQWL¿FDWLRQ . A very distinctive species; the combination  >PLVLGHQWL¿FDWLRQRUHUURU@ of rich brown and blackish costal markings on the forewing Taxonomic issues. Schreiber (1978) lists Manduca cannot be mistaken. occulta occulta Rothschild and Jordan from Trinidad, Status in Trinidad . A rare species, from both suburban but this is the Central American equivalent of Manduca and forest areas.

Fig. 71. 0DQGXFDGLI¿VVDWURSLFDOLV  5RWKVFKLOGDQG-RUGDQ PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW0D\ 0-:&RFN  >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 31

Subfamily: Sphinginae markings with similar species such as Pseudosphinx tetrio Tribe: Sphingini and Erinnyis spp. Subtribe: Acherontiina Biology in Trinidad . The Convolvulaceae food plants include sweet potato ( Ipomoea batatas ), on which the This mostly Old World subtribe is represented by just one caterpillar are an occasional, minor pest in Trinidad species in Trinidad: Agrius cingulata . It is unique in the (Pollard 1984). Trinidad fauna in having pink markings on the abdomen. Status in Trinidad . An occasional species found in both disturbed and forest habitats. 20. Agrius cingulata (Fabricius, 1775) (Figs. 14-15, 41, 74-75) Subfamily: Macroglossinae Herse cingulata (Fabricius): Kaye (1914b), Kaye and Tribe: Dilophonotini Lamont (1927), Pollard (1984) Subtribe Dilophonotina Agrius cingulata (Fabricius): Cary (1951) Agrius cingatulus [sic ] (Fabricius): Schreiber (1978), For ease of comparison, the sequence in Kawahara et al Stradling et al . (1983) (2009) is followed here, using group names for convenience ,GHQWL¿FDWLRQ . The pink markings on the dorsal hindwing based on the obvious clades on their phylogeny. Thus, and abdomen are distinctive. Slight sexual dimorphism, I refer to the Pachylia group (two species of Pachylia the male typically has more contrasting forewing markings. Walker), the Nyceryx  JURXS  VSHFLHV RI ¿YH JHQHUD  Images of resting adults do not show the pink markings the Erinnyis group (17 species of seven genera), and the QRUPDOO\DQGWKH\PXVWEHLGHQWL¿HGE\FRPSDULVRQRIWKH Aleuron group (three species of two genera).

Fig. 72. Manduca hannibal hannibal &UDPHU IHPDOH&XUHSH09/1RYHPEHU 0-:&RFN >0-:&@

Fig. 73. Manduca ochus .OXJ PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@ 32 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Pachylia group Mimosoideae ( Inga sp., Callliandra sp., etc.) at dusk in Two species of Pachylia are reported from Trinidad. Both French Guiana (Haxaire and Rasplus 1987a). It may be feed on Ficus (Moraceae). WKDW WKLV LV SULPDULO\ D GXVN RU FUHSXVFXODU ÀLHU UDUHO\ attracted to light. 21. 3DFK\OLD¿FXV (Linnaeus, 1758) (Figs. 16, 76-77) ,GHQWL¿FDWLRQ . The dorsal forewings are fairly uniformly 3DFK\OLD¿FXV (Linnaeus): Rothschild and Jordan (1903), dark brown with pale brown patches on the costa and apex. Kaye (1914b), Kaye and Lamont (1927), Cary (1951), Status in Trinidad . A rare species in Trinidad that I have Schreiber (1978), Stradling et al . (1983), Cock (2017b) not personally encountered. Stradling et al . (1983) note ,GHQWL¿FDWLRQ . This species can be recognised by the just three records, and I have seen images of two further mottled brown dorsal forewings with a distinct pale brown females from Tobago (Cock 2017b). apical patch. The rare P. syces is darker and has a pale &XUHSHOLJKWWUDSƂ1RYHPEHU )'%HQQHWW  brown patch on the forewing costa as well. >8:,=0&$%,@ )LJ ƃ6HSWHPEHU Status in Trinidad . Kaye and Lamont (1927) considered [F.D. Bennett] [NMSE, ex D.J. Stradling collection] 3 ¿FXV one of the commonest of the Sphingidae, but observations since the 1970s show that it is only an Nyceryx group occasional species, mostly recorded from suburban areas. This group comprises the following genera from Trinidad: Callionima Lucas (four species), Nyceryx (four species), 22. Pachylia syces syces (Hübner, [1819]) (Fig. 78) Perigonia (two species), Eupyrrhoglossum (one species), Pachylia syces syces (Hübner): Schreiber (1978), Stradling and Aellopos ¿YHVSHFLHV  Callionima spp. can be readily et al . (1983), Cock (2017b) recognised by the silver distorted Y-shape on the dorsal Biology   $GXOWV KDYH EHHQ FROOHFWHG DW ÀRZHUV RI forewing; Madoryx spp. and Hemeroplanes triptolemus

Fig. 74. Agrius cingulata )DEULFLXV PDOH&DXUD9DOOH\1U&DXUD09/6HSWHPEHU 0-:&RFN >0-:&@

Figs. 75. Agrius cingulata )DEULFLXV IHPDOH%ULJDQG+LOOOLJKWKRXVHVHFXULW\09/OLJKWV0DUFK 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 33

Fig. 76. 3DFK\OLD¿FXV  /LQQDHXV PDOH6W$XJXVWLQH09/$XJXVW )'%HQQHWW >0-:&@

Fig. 77. 3DFK\OLD¿FXV  /LQQDHXV IHPDOH%ULJDQG+LOOOLJKWKRXVHVHFXULW\09/OLJKWV0DUFK 0-:&RFN >0-:&@

Fig. 78. Pachylia syces syces  +EQHU IHPDOH&XUHSHOLJKWWUDS1RYHPEHU )'%HQQHWW >8:,=0&$%,@ 34 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

(Cramer) also have one or two silver spots, but not in a 25. Callionima inuus (Rothschild and Jordan, 1903) Y-shape. The inclusion of Callionima is weakly supported, (Figs. 83-84) but the remaining genera appear to be a monophyletic Callionima inuus Rothschild and Jordan: Stradling et al . group. (1983) ,GHQWL¿FDWLRQ   7KLV VSHFLHV LV VXSHU¿FLDOO\ VLPLODU WR 23. Callionima pan pan (Cramer, 1779) (Figs. 17, 79-80) C. falcifera (below), but can be recognised by the more Callionima pan (Cramer): Stradling et al . (1983) elongate, and less falcate forewing and the more contrasting ,GHQWL¿FDWLRQ . The slightly truncate forewing apical dorsal forewing subapical reticulate pale orange patch, margin should help separate C. pan from the other which has a curved outer margin. Callionima spp. found in Trinidad. Status in Trinidad . An occasional species, recorded only Status in Trinidad . An uncommon species in Trinidad from a suburban area (Curepe). with records from a suburban area (Curepe, four records including three noted by Stradling et al . (1983)) and a 26. Callionima falcifera (Gehlen, 1943) (Figs. 85-86) forested area (Morne Bleu four records). Callionima parce parce (Fabricius): Schreiber (1978), Stradling et al   >PLVLGHQWL¿FDWLRQ@ 24. Callionima calliomenae (Schaufuss, 1870) Callionima falcifera (Gehlen): Stradling et al . (1983), (Figs. 81-82) Cock and Boos (2007) Hemeroplanes calliomenae (Schaufuss): Kaye (1914b), Taxonomic issues. A population from Mexico was Kaye and Lamont (1927), recognised as a separate subspecies in Kitching and Cadiou Callionima calliomenae (Schaufuss): Schreiber (1978), 2000), but this has now been sunk, so no subspecies name Stradling et al . (1983) LVQHHGHG .LWFKLQJ 7KLVVSHFLHVZDVLGHQWL¿HGE\ ,GHQWL¿FDWLRQ . The variable olive-grey-brown forewing dissection and comparison with the illustrations in Soares dorsal colouring and denticulate margin, and the yellow (1993). Trinidad specimens are quite variable in colour, rather than orange dorsal hindwing when visible, will some being orange-brown in general colour (Fig. 85) and distinguish this common species from others of the genus. others darker (Fig. 86). I have dissected an example of Status in Trinidad . A common species in suburban areas, ERWKDQGIRXQGQRVLJQL¿FDQWGLIIHUHQFHV but not recorded from forested areas. ,GHQWL¿FDWLRQ . The sharply falcate forewing, variable

Figs. 79-80. Callionima pan pan &UDPHU PDOH&XUHSH09/)HEUXDU\ 0-:&RFN >0-:&@ 79 GRUVDO 80 YHQWUDO

Figs. 81-82. Callionima calliomenae 6FKDXIXVV  81  PDOH &XUHSH 09/ 6HSWHPEHU  0-: &RFN  >0-:&@ 82  IHPDOH &XUHSH09/6HSWHPEHU 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 35 orange-brown dorsal forewing and pale subapical dorsal hindwing (diagnostic for this species in Trinidad), reticulated patch with a straight outer margin should serve where P. lusca has only one. to distinguish this species. Status in Trinidad . An uncommon species, all records Status in Trinidad . A common species in suburban areas, are from a suburban area (Curepe). but only occasionally caught in lowland forested areas. 28. Nyceryx maxwelli (Rothschild, 1896) (Figs. 89-90) 27. Nyceryx coffaeae (Walker, 1856) (Figs. 87-88) Taxonomic issues. This species has not previously been Nyceryx coffeae [ sic ] (Walker): Kaye and Lamont (1927), UHSRUWHGIURP7ULQLGDG0\LGHQWL¿FDWLRQZDVFRQ¿UPHG Schreiber (1978), Stradling et al . (1983) by I.J. Kitching (pers. comm. 2004) from dorsal and ,GHQWL¿FDWLRQ . This small hawk-moth is readily ventral images, and from Haxaire & Cadiou (1999). distinguished from the larger N. maxwelli (Rothschild) ,GHQWL¿FDWLRQ . The relatively large size, forewing shape and the other two Nyceryx spp., which have denticulate and markings, and extensive yellow basal-discal area of wing margins, but might be confused with Perigonia the dorsal hindwing will help to identify this species. lusca . Nyceryx coffaeae has two yellow patches on the Status in Trinidad . Nyceryx maxwelli is only known

Figs. 83-84. Callionima inuus 5RWKVFKLOGDQG-RUGDQ PDOH&XUHSHOLJKWWUDS2FWREHU )'%HQQHWW >0-:&@ 83 GRUVDO 84 , YHQWUDO

Figs. 85-86. Callionima falcifera falcifera *HKOHQ PDOH 85 &XUHSH09/0D\ 0-:&RFN >0-:&@ 86 &XUHSH09/ 6HSWHPEHU 0-:&RFN >0-:&@

Figs. 87-88. Nyceryx coffaeae  :DONHU PDOH&XUHSH09/-DQXDU\ 0-:&RFN >0-:&@ 87 GRUVDO 88 YHQWUDO 36 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE from Trinidad from one record, collected near Chatham, 30. Nyceryx stuarti (Rothschild, 1894) (Figs. 91, 95-97) southwest Trinidad, by Mary Alkins-Koo. Taxonomic issues. This species has not previously been &KDWKDP ƃ  -DQXDU\  0 $ONLQV  >8:,=0 reported from Trinidad. The two specimens reported below 2014.9.1289] (Figs. 89-90) were confused with N. riscus  DERYH  XQWLO , LGHQWL¿HG them from Haxaire (1996b, 2005) when preparing this 29. Nyceryx riscus (Schaus, 1890) (Figs. 91, 93-94) catalogue. I have subsequently checked all material in Nyceryx riscus (Schaus): Schreiber (1978), Stradling et al . MJWC, NMSE and UWIZM as N. riscus ZLWKRXW¿QGLQJ (1983) additional specimens of N. stuarti . Taxonomic issues . Haxaire (2013) reports that there is ,GHQWL¿FDWLRQ See under N. riscus above. DVLJQL¿FDQWEDUFRGHGLIIHUHQFHEHWZHHQWKH&HQWUDO Status in Trinidad . To date, this species in Trinidad American and South American populations of N. riscus . is only known from a pair taken at light in Parrylands, The South American population may require a new name February 1980, by the late Julius O. Boos. It seems likely as N. riscus was described from Mexico. that N. stuarti is associated with lowland forest, perhaps ,GHQWL¿FDWLRQ The reddish brown dorsal forewing with restricted to the south or southwest of Trinidad. a dark marginal band and the denticulate margin of both wings will separate this species and N. stuarti (Rothschild) 31. Perigonia pallida Rothschild and Jordan, 1903 (below) from other Trinidad hawk-moths. The dark (Figs. 98-99) marginal area is angled towards the tornus in N. stuarti Perigonia pallida Rothschild and Jordan: Kaye and (Fig. 91), but almost straight in N. riscus (Fig. 92). Lamont (1927) Status in Trinidad 7KLVVSHFLHVZDV¿UVWUHSRUWHGIURP Perigonia pallida pallida Rothschild and Jordan: Schreiber Trinidad by Schreiber (1978) and Stradling et al . (1983). (1978), Stradling et al . (1983) It is uncommon, but occurs in both suburban and forested ,GHQWL¿FDWLRQ . This hawk-moth is unlikely to be confused areas. There are no records from the south of Trinidad, from with any other Trinidad species, being recognisable by its where a very similar species, N. stuarti is reported below. small size and the large yellow discal area of the dorsal hindwing.

Figs. 89-90. Nyceryx maxwelli  5RWKVFKLOG PDOH&KDWKDP-DQXDU\ 0$ONLQV >8:,=0@ 89 GRUVDO 90 YHQWUDO

Figs. 91-92 'LDJQRVWLFFKDUDFWHUVIRU Nyceryx stuarti   DQG N. riscus    1RWWRVFDOH +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 37

Status in Trinidad . An uncommon species, only recorded [synonym] from suburban areas (Port of Spain, Curepe). Perigonia lusca tenebrosa (C. Felder and R. Felder): Stradling et al . (1983) [synonym] 32. Perigonia lusca lusca (Fabricius, 1777) Perigonia lusca lusca (Fabricius): Cock (2017b) (Figs. 18, 100-103) Taxonomic issues. This is a variable species with several Perigonia lusca (Fabricius): Kaye (1914b), Kaye and named forms, which have sometimes been treated as subspecies Lamont (1927) in the Trinidad literature. Here they are treated as a single Perigonia interrupta (Walker): Schreiber (1978), Stradling subspecies which is variable in the extent of yellow dorsal et al . (1983) [synonym] hindwing markings. This interpretation is supported by Perigonia lusca restituta (Walker): Stradling et al . (1983) DNA barcoding (I.J. Kitching pers. comm. 2017).

Figs. 93-94. Nyceryx riscus 6FKDXV PDOH%ULJDQG+LOOOLJKWKRXVHVHFXULW\09/OLJKWV0DUFK 0-:&RFN >0-:&@ 93 , GRUVDO 94 YHQWUDO

Figs. 95-97. Nyceryx stuarti 5RWKVFKLOG  95 PDOH3DUU\ODQGVDWOLJKW)HEUXDU\ -2%RRV >0-:&@ 96 DV)LJYHQWUDOYLHZ 97 IHPDOH3DUU\ODQGVDWOLJKW)HEUXDU\ -2%RRV >0-:&@

Figs. 98-99. Perigonia pallida 5RWKVFKLOGDQG-RUGDQPDOH&XUHSHDWOLJKW-XO\'-6WUDGOLQJ>8:,=0@ SKRWR 8:,=0  98 GRUVDO 99 YHQWUDO 38 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

,GHQWL¿FDWLRQ . This is a fairly distinctive, small species. Aellopos ceculus (Cramer), which follows. The yellow It might be confused with Nyceryx coffaeae (above), under hindwing markings are narrower in E. sagra , the detail which species differences are highlighted. Perigonia ilus of the dorsal forewing markings differ, and the ventral Boisduval is a confusingly similar species found on the forewing has some areas mottled with orange, whereas adjacent mainland that could well occur in Trinidad. It that of A. ceculus is uniformly dark apart from the white can be most readily distinguished by the dorsum of the subterminal spot. ventral hindwing being yellow rather than dull brown as in Biology in Trinidad . F.D. Bennett has collected E. sagra P. lusca (Fig. 103). In preparing this paper, I re-examined FRPLQJ WR ÀRZHUV RI Lantana montevidensis at 18.00 h VSHFLPHQVSUHYLRXVO\LGHQWL¿HGDV P. lusca , and there (Curepe, January 1977), but this species is most often were no P. ilus amongst them. collected when attracted to light. Status in Trinidad . A fairly common species found in Status in Trinidad . An uncommon species, with records suburban and forested areas. from suburban and lowland forest areas.

33. Eupyrrhoglossum sagra (Poey, 1832) (Figs. 104-106) Aellopos Hübner, [1819] Eupyrrhoglossum sagra (Poey): Kaye and Lamont (1927), 7KLVLVDJHQXVRIGD\À\LQJKXPPLQJELUGKDZNPRWKV Schreiber (1978), Stradling et al . (1983), Cock (2017b) RIZKLFK¿YHVSHFLHVKDYHEHHQUHFRUGHGIURP7ULQLGDG ,GHQWL¿FDWLRQ . This is one of two Trinidad species of EXWRQO\IRXUDUHDFFHSWHGKHUH7KH¿UVW $FHFXOXV has hummingbird hawk-moths that have yellow markings on a yellow band on the dorsal hindwing and yellow laterally the dorsal hindwing and adjacent thorax, the other being RQ WKH DEGRPHQ ZKLOH WKH WKUHH UHPDLQLQJ FRQ¿UPHG

Figs. 100-103. Perigonia lusca lusca  )DEULFLXV  100 PDOH&XUHSH09/'HFHPEHU 0-:&RFN >0-:&@ 101 IHPDOH &XUHSH09/-XQH 0-:&RFN >0-:&@ 102 IHPDOH&XUHSH09/-XQH 0-:&RFN >0-:&@ 103 DV)LJ YHQWUDOYLHZ

Figs. 104-106. Eupyrrhoglossum sagra  3RH\  104 /HIWPDOH$ULSR9DOOH\5DSVH\(VWDWHPDODLVHWUDS$XJXVW 50 %DUDQRZVNL >0-:&@ 105 IHPDOH&XUHSH09/6HSWHPEHU 0-:&RFN >0-:&@ 106 DV)LJYHQWUDO +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 39 species are dark with a white band on the abdomen: A. 34. Aellopos ceculus (Cramer, 1777) (Figs. 19, 107-109) clavipes (Rothschild and Jordan), A. titan (Cramer) and A. Eupyrrhoglossum ceculus (Cramer): Kaye (1901, 1914a) fadus &UDPHU 5HFRUGVRIWKH¿IWKVSHFLHV A. tantalus Sesia ceculus (Cramer): Kaye (1914b), Kaye and Lamont (Linnaeus) (Kaye 1901, Kaye and Lamont 1927), are here (1927), Cary (1951) FRQVLGHUHGDPLVLGHQWL¿FDWLRQRI A. clavipes , as discussed Aellopos ceculus (Cramer): Schreiber (1978), Stradling et below. However, A. tantalus zonata (Drury) occurs in the al . (1983), Cock (2017b) Caribbean as far south as Grenada, and so could turn up ,GHQWL¿FDWLRQ . With its yellow markings, A. ceculus can in Tobago or Trinidad. Hodges (1971), D’Abrera ([1987]) only be confused with Eupyrrhoglossum sagra , above, and Tuttle (2007) present diagnostic characters of the under which differences are discussed. dorsal and ventral forewing surfaces to separate the four Biology in Trinidad   7KLV LV QRUPDOO\ D GD\À\LQJ species. Aellopos fadus and A. titan both have on the species, although it comes rarely to light. F.D. Bennett ventral forewing, a double row of white spots from the KDVFROOHFWHGWZRVSHFLPHQVFRPLQJWRÀRZHUVRI Lantana inner margin to the cell; they also have an outer band of montevidensis at 18.00 h (Curepe, January 1977). white spots from the tornus to about two-thirds along costa, Status in Trinidad . An occasional species found in both single in A. titan and partially double in A. fadus ; further, suburban and forest habitats. on the dorsal surface, A. titan has a strong dark spot at the end of the cell that is obscure or missing in A. fadus . In 35. Aellopos clavipes clavipes (Rothschild and Jordan, contrast, A. tantalus and A. clavipes lack the double row 1903) (Figs. 110-111) of white spots from the inner margin to the cell, and the Aellopus [sic ] sisyphus (Burmeister): Kaye (1901), Kaye outer band is reduced to three irregular spots in M 3-Cu1 to (1914a) [synonym of A. tantalus PLVLGHQWL¿FDWLRQ@

M2-M 3 (spaces 3-5), and on the dorsal surface both have a Sesia tantalus (Linnaeus): Kaye (1914b), Cary (1951) dark spot at the end of the cell. Aellopos tantalus and A. >PLVLGHQWL¿FDWLRQ@ clavipes can be most readily separated by the presence in Sesia tantalus tantalus (Linnaeus): Kaye and Lamont A. clavipes of two (sometimes one) narrow streaks on the  >PLVLGHQWL¿FDWLRQ@ underside in Cu 1-Cu2 (space 2), although with complete Aellopos tantalus (Linnaeus): Schreiber (1978) [assumed specimens to hand, the swollen foretarsus of A. clavipes PLVLGHQWL¿FDWLRQ@ compared to the normal foretarsus of A. tantalus reliably Aellopos clavipes clavipes (Rothschild and Jordan): distinguishes the two. Schreiber (1978), Stradling et al . (1983), Kelly (2011), Cock (2017b)

Figs. 107-109. Aellopos ceculus  &UDPHU  107 PDOH*UDQGH5DYLQH5HVHUYH -%RRV >0-:&@ 108 IHPDOH&XUHSHEODFNOLJKWWUDS 0DUFK )'%HQQHWW >0-:&@ 109 DV)LJYHQWUDOYLHZ

Figs. 110-111. Aellopos clavipes clavipes  5RWKVFKLOGDQG-RUGDQ IHPDOH)RUW*HRUJHIW-DQXDU\ ):-DFNVRQ >0*&/@ SKRWR56W/DXUHQW0*&/  110 GRUVDO 111 YHQWUDO 40 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

,GHQWL¿FDWLRQ . See under Aellopos above. ,GHQWL¿FDWLRQ . See under Aellopos above. A specimen in Status in Trinidad . A rare species, with just one specimen /DPRQW¶VFROOHFWLRQLQ8:,=0KDGEHHQPLVLGHQWL¿HGDV located from Trinidad, although another is known from A. titan . the Gulf of Paria, 3½ miles offshore (I.J. Kitching pers. Biology in Trinidad. Usually seen and collected feeding comm. 2017). The early records of A. tantalus are thought DWÀRZHUV WR EH PLVLGHQWL¿FDWLRQV RI A. clavipes , as set out in the Status in Trinidad . An uncommon species with records section on species not recognised from Trinidad. There from both suburban (Curepe) and forest areas. are two recent photographic records from Tobago (Cock &XUHSH OLJKW WUDS Ƃ  $XJXVW  )' %HQQHWW  2017b). [UWIZM CABI.2161] )RUW *HRUJH  IW > P@ Ƃ -DQXDU\  ): &XUHSH &,%& 6WDWLRQ Ƃ  -XO\  7 &DVVLH  [Jackson]) [MGCL] (Figs. 110-111) [UWIZM CABI.2162] &XUHSHPDODLVHWUDSƂ-XQH>0-:&@ 36. Aellopos titan titan (Cramer, 1777) (Figs. 112-113) *UDQGH 5DYLQH UHVHUYH DW ÀRZHUV ƃ 6HSWHPEHU  Sesia titan (Cramer): Kaye and Lamont (1927) (J.O. Boos) [MJWC] (Figs. 114-115) Aellopos titan titan (Cramer): Schreiber (1978), Stradling 6DQ )HUQDQGR Ƃ  $SULO  >1 /DPRQW@ et al . (1983) [UWIZM.2013.13.2301] ,GHQWL¿FDWLRQ . See under Aellopos above. 7ULQLGDGƃ 0LOOHUFROO >0*&/@ Status in Trinidad . Known from just one record, a female captured by Sir Norman Lamont at Morne Diable, 1 July Erinnyis group 1917 – not 1918 as stated by Kaye and Lamont (1927). In Trinidad, the Erinnyis group of Kawahara et al . 0RUQH@ 'LDEOH Ƃ  -XO\  >1 /DPRQW@ >106(@ (2009) comprises Oryba (two species), Pachylioides (Figs. 112-113) Hodges (one species), Madoryx Boisduval (three species), Hemeroplanes Hübner (one species), Erinnyis Hübner 37. Aellopos fadus (Cramer, 1775) (Figs. 114-115) (seven species), Isognathus C. Felder and R. Felder (two Sesia fadus (Cramer): Kaye and Lamont (1927), Cary (1951) species) and Pseudosphinx Burmeister (one species). It Aellopos fadus fadus (Cramer): Schreiber (1978), is assumed that Phryxus Hübner (one species), which Stradling et al . (1983) Kawahara et al . (2009) do not include, also belongs here

Figs. 112-113. Aellopos titan titan &UDPHU IHPDOH0>RUQH@'LDEOH-XO\>1/DPRQW@>106(@ 112 GRUVDO 113 YHQWUDO 3KRWRV $:KLI¿Q106( 

Figs. 114-115. Aellopos fadus &UDPHU PDOH*UDQ5DYLQHUHVHUYHDWÀRZHUV6HSWHPEHU -2%RRV >0-:&@ 114 GRUVDO 115 , YHQWUDO +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 41 given the similarity to Erinnyis and Isognathus . The are available (Kitching 2017, Oehlke 2017). inclusion of Oryba in the clade is not strongly supported, Status in Trinidad . A rarely collected species, very although the remaining genera seem to be a monophyletic occasionally attracted to light. Haxaire (1996a) indicates clade, with Erinnyis , Isognathus , Pseudosphinx and WKDWDGXOWVÀ\H[WUHPHO\UDSLGO\DWGXVNZKLFKPD\UHÀHFW Phryxus forming a tight group, and so the group is referred why it is seldom collected in Trinidad. The few records to to by its most species-rich genus, Erinnyis . date are from Curepe. Erinnyis , ,VRJQDWKXV3KU\[XV and Pseudosphinx are &XUHSH09/Ƃ$XJXVW 0-:&RFN >0-:&@ noteworthy in that the adults rest with their forewings St. Augustine: ? December 1979 (Stradling et al . 1983) almost parallel with the body, rather than at an acute angle 6RXWKRI9DOHQFLD1:PLVWQHWWLQJƂ to each other as is the case with the other tribes found in September 2014 (M.G. Rutherford photo) [https:// 7ULQLGDG7KLVVKRXOGKHOSVHSDUDWHLPDJHVRIVXSHU¿FLDOO\ www.inaturalist.org/observations/11142177] similar species, apart from Neococytius cluentius which Trinidad: ? (F.W. Urich) (Kaye and Lamont 1927) also holds its wings almost parallel with the body (Fig. 7). 39. Oryba achemenides (Cramer, 1779) (Figs. 117-118) 38. Oryba kadeni (Schaufuss, 1870) (Fig. 116) Oryba achemenides (Cramer): Kaye and Lamont (1927), Oryba kadeni (Schaufuss): Kaye and Lamont (1927), Schreiber (1978), Stradling et al . (1983), Cock (2017b) Schreiber (1978), Stradling et al . (1983) ,GHQWL¿FDWLRQ . See under O. kadeni above; this is a ,GHQWL¿FDWLRQ . This species might only be confused with generally darker species. O. achemenides (Cramer) below. The two are obviously Status in Trinidad . An uncommon species in Trinidad GLIIHUHQW DV VKRZQ LQ WKH ¿JXUHV EXW WKLV LV EDVHG RQ and Tobago, with scattered records at light in suburban relative width, colour and contrast of the markings, rather and forested areas. than a single diagnostic character. Rothschild and Jordan (1903) highlighted that the distal marginal area of the 40. Pachylioides resumens (Walker, 1856) dorsal forewing is much narrower than the space between (Figs. 119-120) this and the discal line beyond the cell in O. achemenides , Pachylia resumens Walker: Kaye and Lamont (1927), but equally broad in O. kadeni . However, they are best Cary (1951) separated by the conspicuous narrow green discal band Pachylioides resumens (Walker): Schreiber (1978), bounded by brown lines or bands on the dorsal hindwing Stradling et al . (1983), Cock (2017b) in O. kadeni that is missing in O. achemenides . There ,GHQWL¿FDWLRQ   7KLV VSHFLHV LV VXSHU¿FLDOO\ VLPLODU WR is quite strong sexual dimorphism in O. kadeni , along Pachylia spp., but lacks the pale costal marking at the the lines shown for O. achemenides (Figs. 117-118), but apex of the dorsal forewing of the two Trinidad species. I have not seen any male O. kadeni from Trinidad to Status in Trinidad . An occasional species, recorded from illustrate this, although published images from elsewhere both suburban and forested areas.

Figs. 116. Oryba kadeni  6FKDXIXVV IHPDOH&XUHSH09/$XJXVW 0-:&RFN >0-:&@ 42 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

41. Madoryx oiclus oiclus (Cramer, 1779) Trinidad. The spot closest to the costa is an elongate (Figs. 121-122) angled dash in M. oiclus whereas that of M. plutonius Madoryx oiclus (Cramer): Kaye (1914b), Kaye and Lamont (Hübner) is reduced to a dot and that of M. bubastus (1927), Cary (1951), Schreiber (1978), Stradling et al . (Cramer) is round. (1983), D’Abrera ([1987]), Cock (2017b) Status in Trinidad . A common species, primarily in ,GHQWL¿FDWLRQ . The two silvery discal spots of the dorsal suburban situations. forewing separate Madoryx spp. from other genera in

Fig. 117. Oryba achemenides &UDPHU PDOH%ULJDQG+LOOOLJKWKRXVHVHFXULW\09/OLJKWV-DQXDU\ 0-:&RFN >0-:&@

Figs. 118. Oryba achemenides &UDPHU IHPDOH&XUHSH09/2FWREHU 0-:&RFN >0-:&@

Fig. 119. Pachylioides resumens :DONHU PDOH9DOVD\Q3DUNDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 43

42. Madoryx plutonius plutonius (Hübner, [1819]) Status in Trinidad . An uncommon species, with records (Fig. 123) from both suburban and forested areas. Madoryx pluto pluto (Cramer): Schreiber (1978), Stradling et al . (1983) [unavailable homonym] 44. Hemeroplanes triptolemus (Cramer, 1779) ,GHQWL¿FDWLRQ . See under M. oiclus above. (Figs. 126-127) Status in Trinidad . A rare species, only recorded from Leucorhampha triptolemus (Cramer): Rothschild and forested areas of the Northern Range. Jordan (1903), Kaye (1914b), Kaye and Lamont &XPDFD5RDGPLOHV09/ƃ-XO\ 0-: (1927), Cary (1951) &RFN >0-:&8:,=0&$%,@ƃ2FWREHU Hemeroplanes triptolemus (Cramer): Schreiber (1978), 1982 (M.J.W. Cock) [UWIZM CABI.2037] Stradling et al . (1983), Cock (2017b) /DODMD5LGJH09/ ƃ  6HSWHPEHU K ,GHQWL¿FDWLRQ . The elongate white discal spot and the (M.J.W. Cock) [MJWC notes] narrow yellow bands on the abdomen should serve to recognise this species. 43. Madoryx bubastus bubastus (Cramer, 1777) Status in Trinidad . A fairly common species, primarily (Figs. 124-125) recorded from suburban areas. Madoryx bubastus (Cramer): Kaye (1914b) Madoryx bubastus bubastus (Cramer): Kaye and Lamont 45. Phryxus caicus (Cramer, 1777) (Figs. 128-129) (1927), Cary (1951), Schreiber (1978), Stradling et al . Phryxus caicus (Cramer): Lamont and Callan (1950), (1983) Schreiber (1978), Stradling et al . (1983) ,GHQWL¿FDWLRQ . See under M. oiclus above. ,GHQWL¿FDWLRQ . The long white and black line across the

Fig. 120. Pachylioides resumens :DONHU IHPDOH)\]DEDG$SULO-XQH 50)DUPERURXJK >281+0@

Figs. 121-122. Madoryx oiclus oiclus  &UDPHU  121 PDOH9DOVD\Q3DUNDWOLJKW$XJXVW 0-:&RFN >0-:&@ 122 IHPDOH &XUHSH%/7)HEUXDU\ )'%HQQHWW >0-:&@ 44 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE dorsal forewing, orange-brown dorsal hindwing with dark on the abdominal markings: Erinnyis alope , E. lassauxii marginal lines along the veins, and black and grey banded (Boisduval), E. impunctata and E. ello have the abdomen abdomen make this a distinctive species. banded with black and light grey, whereas E. oenotrus Status in Trinidad . An uncommon species, recorded (Cramer), E. crameri (Schaus) and E. obscura (Fabricius) from suburban and disturbed areas, but not from primarily KDYHWKHDEGRPHQXQLIRUPO\EURZQ,QWKH¿UVWJURXS forested areas. E. alope has the basal and discal dorsal hindwing yellow, E. impunctata and E. ello have it dull orange (brighter Erinnyis Hübner, [1819] in female E. ello ) and E. lassauxii has just the basal Seven species of Erinnyis are recognised from area orange, almost completed obscured in the typical Trinidad. They can be divided into two groups based form (although there is another form of E. lassauxii that

Fig. 123. Madoryx plutonius plutonius +EQHU PDOH&XPDFD5RDGPLOHV09/-XO\ 0-:&RFN >0-:&@

Figs. 124-125. Madoryx bubastus bubastus &UDPHU  124 PDOH3DUU\ODQGV2LO¿HOGDWOLJKW)HEUXDU\ -2%RRV >0-:&@ 125 IHPDOH&XUHSH09/'HFHPEHU 0-:&RFN >0-:&@

Figs. 126-127. Hemeroplanes triptolemus  &UDPHU  126 PDOH&XUHSH09/2FWREHU 0-:&RFN >0-:&@ 127 IHPDOH &XUHSH09/-XQH 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 45 closely resembles E. impunctata but has not been found in hindwing yellow, with a minimal dark margin and dark Trinidad); E. impunctata and E. ello can be separated by veins extending at least halfway to the base of the wing. the dark brown dorsal forewing of E. impunctata (similar Given that several of these diagnostic characters relate to male E. crameri ), the uniformly pale grey forewing of WRWKHGRUVDOKLQGZLQJLGHQWL¿FDWLRQRILPDJHVRIOLYLQJ the female of E. ello and the irregular dark streak from PRWKVZLWKWKLVIHDWXUHFRYHUHGZLOOEHPRUHGLI¿FXOWDQG base to apex on the grey forewing of male E. ello . In the rely on a careful comparison of the forewing characters. second group, E. obscura is much smaller than the others, +HQFHLGHQWL¿FDWLRQZLOOEHIDFLOLWDWHGLILPDJHVDUHDOVR the forewing reminiscent of male E. ello , and the hindwing taken to show the hindwing. dull orange, only narrowly darker at the margin; the female of E. oenotrus is grey with two darker patches on the 46. Erinnyis alope alope (Drury, 1773) costa, the one just before mid-costa extending irregularly (Figs. 42-44, 130-131) to just above tornus; male E. oenotrus and both sexes of E. Erinnyis alope (Drury): Kaye and Lamont (1927), Cary crameri resemble each other, but the hindwing margin of (1951), CABI (2002b) E. oenotrus is even with a diffuse internal border, whereas Erinnyis alope alope (Drury): Schreiber (1978), Stradling that of E. crameri is uneven, extending basally along the et al . (1983), Cock (2017b) veins and with a sharp internal border. Erinnyis spp. might Biology in Trinidad . Although food plants include also be confused with Isognathus spp., of which two are papaya, Carica papaya , this species has not been reported known from Trinidad: the common I. scyron (Cramer) has as a pest in Trinidad. similar colouring to E. alope , but the wings are broader, ,GHQWL¿FDWLRQ . See under Erinnyis above. and the grey bands on the abdomen are narrower and not Status in Trinidad . A common species in both suburban interrupted dorsally; the rare I. caricae (Linnaeus) has the and forested areas.

Figs. 128-129. Phryxus caicus  &UDPHU  128 PDOH&XUHSH09/1RYHPEHU 0-:&RFN >0-:&@ 129 IHPDOH0DUDYDO -XQH -2%RRV >0-:&@

Fig. 130. Erinnyis alope alope  'UXU\ PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@ 46 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

47. Erinnyis lassauxii (Boisduval, 1859) (Figs. 132-134) [1988] treat as a fourth form of E. lassauxii ), but E. Erinnyis lassauxi [ sic ] (Boisduval): Kaye (1914b), Kaye impunctata lacks the dark spots on the ventral surface of and Lamont (1927) the abdomen (Fig. 136), which are found in all forms of E. Erinnyis lassauxi [ sic ] (Boisduval) ssp.: Cary (1951) lassauxii (Fig. 133). In the material that I have seen from Erinnyis lassauxii (Boisduval): Schreiber (1978), Trinidad, males of E. lassauxii are usually f. omphalae Stradling et al . (1983), Cock (2017b) (Fig. 132) and females are usually f. lassauxii (Fig. 134). ,GHQWL¿FDWLRQ . See under Erinnyis above. Three forms of Status in Trinidad . A fairly common species in both E. lassauxii have been recognised (Rothschild and Jordan suburban and forested areas. 1903, D’Abrera [1988]): f. lassauxii (dorsal hindwing dark with the orange colour much reduced; Fig. 134), f. 48. Erinnyis impunctata Rothschild and Jordan, 1903 omphaleae (Boisduval) (dorsal hindwing with extensive (Fig. 135-136) orange area and a broad dark margin; Fig. 132) and f. Erinnyis oenotrus (Cramer): Stradling et al . (1983) [partial merianae Grote (similar to f. omphaleae , but the orange PLVLGHQWL¿FDWLRQVHHEHORZ@ more extensive and the dark margin correspondingly Taxonomic issues. This species not previously been narrower). These names have no taxonomic standing, recorded from Trinidad. When compiling records for this EHLQJLQIUDVSHFL¿FEXWWKH\DUHDXVHIXOWRROWRUHIHUWR catalogue, two specimens in F.D. Bennett’s series of E. the forms of a polytypic species . I have seen no specimens oenotrus in UWIZM were found. They would have been of the last-named form from Trinidad. Were f. merianae combined with that species in the Stradling et al . (1983) to occur, specimens might be confused with E. impunctata dataset. below (which Rothschild and Jordan (1903) and D’Abrera ,GHQWL¿FDWLRQ . See under Erinnyis above. This species

Fig. 131. Erinnyis alope alope  'UXU\ IHPDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW$XJXVW 0-:&RFN >0-:&@

Figs. 132. Erinnyis lassauxii %RLVGXYDO I omphaleae  %RLVGXYDO PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 47 differs from E. oenotrus and E. crameri in having grey Taxonomic issues . Barcoding of the material reared by bands on the abdomen, like those of E. ello , E. lassauxii Janzen and Hallwachs (2017) suggests that caterpillars of and some other species of Erinnyis . It is closest in what appears to be E. ello reared from Manilkara chicle appearance to E. lassauxii f. merianae as noted under that (Sapotaceae) in Costa Rica are a separate species, with species, but this form has not been reported from Trinidad. matching barcodes from Mexico to the Amazon (Belém) Status in Trinidad . Just two records from Curepe. This (I.J. Kitching pers. comm. 2017). Manilkara chicle does seems to be a rare species, but would have been overlooked. not occur in Trinidad, but M. bidentata (known locally as &XUHSH09/ƃ-DQXDU\ )'%HQQHWW >8:,=0 balata) is occasional in Trinidad forests and more common &$%,@ ƃ  )HEUXDU\  )' %HQQHWW  in coastal areas (Hill and Sandwith 1947). It would be [UWIZM CABI.1933] worth looking for caterpillars of E. ello on this tree in Trinidad, and experimental studies could clarify the status 49. Erinnyis ello ello (Linnaeus, 1758) (Figs. 137-138) and biology of the two barcode groups. Dilophonota ello (Linnaeus): Kaye (1901), Guppy (1911a, Biology in Trinidad . In Trinidad, Guppy (1911b) notes 1911b), Kaye (1914a) caterpillars on rubber and cassava. Urich (1915) notes Erinnyis ello (Linnaeus): Rothschild and Jordan (1903), it as a pest of cassava, usually kept in check by natural Kaye (1914b), Urich (1915), Kaye and Lamont (1927), enemies: Telenomus sp. (Scelionidae) attacking eggs, and Cary (1951), Winder (1976), Des Vignes (1986), CABI at least one species of Microgasterinae parasitic on the (2002a), MFPLMA (2010) caterpillars. Kaye and Lamont (1927) refer to this species Erinnyis ello ello (Linnaeus): Schreiber (1978), Stradling as the cassava hawk-moth. In recent decades there have et al . (1983), Cock (2017b) been periodic outbreaks reported on cassava (Des Vignes

Fig. 133. Erinnyis lassauxii %RLVGXYDO I omphaleae  %RLVGXYDO PDOHDV)LJYHQWUDO

Fig. 134. Erinnyis lassauxii %RLVGXYDO I lassauxi  %RLVGXYDO IHPDOH%ULJDQG+LOOOLJKWKRXVHVHFXULW\09/OLJKWV-DQXDU\ 0-:&RFN >0-:&@ 48 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

1986, MFPLMA 2010). While living in Trinidad, 1978- had descended to the ground and were walking on the road. 1982, I also observed occasional outbreaks on planted It was the popping noise made by inadvertently driving patches of introduced rubber trees, Hevea brasiliensis over them that drew the outbreak to my attention – in those (Euphorbiaceae) at Fishing Pond, Arima Valley (between days, as was the case with most cars, we did not have air- Simla and Asa Wright), and Southern Main Road (not conditioning! far from Pitch Lake). These scattered patches of planted ,GHQWL¿FDWLRQ . See under Erinnyis above. rubber trees are not obvious when the trees are mature, Status in Trinidad . One of the commonest species as in Arima Valley and on Southern Main Road, and the everywhere, occasionally abundant. RXWEUHDNVZHUH¿UVWQRWLFHGEHFDXVHWKHPDWXUHFDWHUSLOODUV

Figs. 135-136. Erinnyis impunctata 5RWKVFKLOGDQG-RUGDQPDOH&XUHSH09/)HEUXDU\ )'%HQQHWW >8:,=0&$%,@ 135 GRUVDO 136 YHQWUDO

Fig. 137. Erinnyis ello ello /LQQDHXV PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@

Fig. 138. Erinnyis ello ello /LQQDHXV IHPDOH&XUHSHFDWHUSLOODURQFDVVDYDHPHUJHG-XO\ 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 49

50. Erinnyis oenotrus ( Cramer, 1780) (Figs. 139-140) validity (Kitching and Cadiou 2000, Haxaire and Herbin Erinnyis oenotrus (Cramer): Kaye (1914b), Kaye and 2000). Tuttle (2007) showed that both species could be Lamont (1927), Cary (1951), Schreiber (1978), reared from the same female and accordingly made E. Stradling et al . (1983) domingonis a junior subjective synonym of E. obscura . ,GHQWL¿FDWLRQ . See under Erinnyis above. The name domingonis may be useful to refer to the dark Status in Trinidad . A fairly common species, recorded form, i.e. E. obscura f. domingonis . Stradling et al . (1983) from both suburban and forested areas record two males as E. domingonis from 3,767 trap nights in the St. Augustine area, 1969-1977. These specimens 51. Erinnyis crameri (Schaus, 1898) (Figs. 141-142) are now in NMSE and one is shown here as Fig. 144. Erinnyis crameri (Schaus): Rothschild and Jordan (1903), ,GHQWL¿FDWLRQ . See under Erinnyis above. Kaye (1914b), Kaye and Lamont (1927), Cary (1951), Status in Trinidad . A common species in suburban areas, Schreiber (1978), Stradling et al . (1983) with rather few records from forested areas. ,GHQWL¿FDWLRQ . See under Erinnyis above. Status in Trinidad . A common species in both suburban 53. Isognathus scyron (Cramer, 1780) and forested situations. (Figs. 22, 45-46, 145-146) Unnamed caterpillar: Guppy (1893) 52. Erinnyis obscura obscura (Fabricius, 1775) Anceryx scyron (Cramer): Kaye (1901), Kaye (1914a) (Figs. 143-144) Isognathus scyron (Cramer): Rothschild and Jordan Erinnyis obscura obscura (Fabricius): Cary (1951), (1903), Kaye (1914b), Cary (1951), Schreiber (1978), Schreiber (1978), Stradling et al . (1983) Stradling et al . (1983), D’Abrera ([1987]) Erinnyis domingonis (Butler): Stradling et al . (1983) [synonym] ,GHQWL¿FDWLRQ . See under Erinnyis above. There is slight Taxonomic issues . For many years, E. domingonis was sexual dimorphism: the male has two dark dashes on the considered a valid species, resembling E. obscura but discal cell of the dorsal forewing; these are absent in the darker, although doubts had been expressed about its female, which is darker than the male but has a more

Figs. 139-140. Erinnyis oenotrus &UDPHU  139 PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@ 140 , IHPDOH&XUHSH09/-DQXDU\ 0-:&RFN >0-:&@

Figs. 141-142. Erinnyis crameri 6FKDXV  141 PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@ 142 , IHPDOH9DOVD\Q3DUNDWOLJKW-XQH 0-:&RFN >0-:&@ 50 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE conspicuous light brown area beyond the end of the cell Macrospila tetrio (Linnaeus): Caracciolo (1890) of the dorsal forewing, and the grey bands on the abdomen Pseudosphinx tetrio (Linnaeus): Kaye (1901), Rothschild are more obvious. and Jordan (1903), Kaye (1914a, 1914b), Kaye and Biology in Trinidad . The caterpillars are a common Lamont (1927), Cary (1951), Schreiber (1978), sight on the introduced ornamental Allamanda cathartica Stradling et al . (1983), Cock (2003, 2008), Cooper and (Apocynaceae), often noticed by naturalists and the public. Cooper (2009), Cock (2017b) Status in Trinidad . Very common in suburban areas, but ,GHQWL¿FDWLRQ . The mottled grey dorsal forewings, dark also regularly found in forested areas, indicating that other dorsal hindwings and grey banded abdomen should food plants must also be used. distinguish this species in Trinidad. There is modest sexual dimorphism: the dorsal forewing markings of males are 54. Isognathus caricae (Linnaeus, 1758) (Fig. 147) more contrasting, with an extensive dark area adjacent to Isognathus caricae (Linnaeus): Lamont and Callan (1950), the tornus, whereas the larger females are more uniformly Schreiber (1978), Stradling et al . (1983) pale grey. ,GHQWL¿FDWLRQ . See under Erinnyis above. The yellow Biology in Trinidad . Kaye and Lamont (1927) refer to dorsal hindwing with dark veins is unique in Trinidad. this species as the frangipani hawk-moth and note that Status in Trinidad . A rare species, with records from it is a ‘constant pest on frangipani ( Plumeria ); the large Palmiste (Lamont and Callan 1950), and Curepe (Stradling voracious larvae completely stripping a tree of its leaves et al . 1983). in a few days’. Cock (2008) illustrates the early stages in &XUHSHOLJKWWUDSƂ6HSWHPEHU )'%HQQHWW  Trinidad and discusses this species in Trinidad and Tobago. >8:,=0&$%,@ƃ-XO\ '-6WUDGOLQJ  The caterpillars in Trinidad have been observed almost [NMSE] exclusively on the naturalised frangipani, Plumeria rubra 3DOPLVWH Ƃ $SULO  >1 /DPRQW@ >106(@ Ƃ  (Apocynaceae), but may also use the garden ornamental April 1934 [N. Lamont] [NMSE] (Fig. 148) Allamanda cathartica (Apocynaceae) occasionally. Status in Trinidad . A fairly common species that could turn 55. Pseudosphinx tetrio (Linnaeus, 1771) up anywhere. Caterpillars are more conspicuous than adults, (Figs. 21, 47, 148-149) and more frequently observed by members of the public.

Figs. 143-144. Erinnyis obscura obscura )DEULFLXV  143 PDOH&XUHSH09/-DQXDU\ 0-:&RFN >0-:&@ 144 PDOHI domingonis 6W$XJXVWLQHDWOLJKW)HEUXDU\ '-6WUDGOLQJ >106(@ SKRWR$:KLI¿Q106( 

Figs. 145-146. Isognathus scyron  &UDPHU  145  PDOH &XUHSH 09/ 6HSWHPEHU  0-: &RFN  >0-:&@ 146  IHPDOH 6W $XJXVWLQHIURPFDWHUSLOODUVRQ Allamanda cathartica HPHUJHG-XO\ 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 51

Aleuron group more common in Trinidad than light trap records indicate. In the Trinidad fauna, Aleuron (two species) and Arima Valley, Asa-Wright Nature Centre: ? 6 October Unzela (one species) form a small, isolated group that is 2017 (tlaloc27, poor photo) [https://www.inaturalist. rarely collected. org/observations/8552081] &XUHSH OLJKW WUDS ƃ -DQXDU\  )' %HQQHWW  56. Aleuron carinata (Walker, 1856) (Figs. 150-151) >1+08.@ ƃ  -DQXDU\  )HEUXDU\  )' Aleuron carinata (Walker): Schreiber (1978), Stradling et Bennett) [MJWC] al . (1983) ,GHQWL¿FDWLRQ . A small, rather plain species, generally 57. Aleuron chloroptera (Perty, [1833]) (Fig. 152) brown, but with distinctive banding on the anterior part of ,GHQWL¿FDWLRQ . Th greenish colour of this distinctive small the abdomen. hawk-moth rapidly fades to olive-brown (as in Fig. 152); Status in Trinidad . A rare species in Trinidad collections it has a broad dark margin to the dorsal hindwing, and the with only three records. Haxaire (1992) states that the forewing is distinctly hooked at mid-termen. JHQXV FDQ EH FDXJKW FRPLQJ WR ÀRZHUV DW GXVN  Status in Trinidad . There is just one record of this 19.00h in French Guiana) and very rarely come to light, so species from Trinidad. Although this specimen was taken different collecting methods should show this species to be at light, A. chloroptera is more likely to be found feeding

Fig. 147. Isognathus caricae /LQQDHXV IHPDOH3DOPLVWH$SULO>1/DPRQW@>106(@ SKRWR$:KLI¿Q106( 

Fig. 148. Pseudosphinx tetrio  /LQQDHXV PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@ 52 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

DWÀRZHUVDWGXVNRUDVFDWHUSLOODUV Schreiber (1978), Stradling et al . (1983) &XUHSH EODFN OLJKW WUDS ƃ  6HSWHPEHU  )' ,GHQWL¿FDWLRQ . The white ventral body, narrow white Bennett) [UWIZM CABI.2094] band on the abdomen and wing markings will all help to recognise this small hawk-moth. 58. Unzela japix japix (Cramer, 1776) (Figs. 153-155) Status in Trinidad . This species has been rarely collected Enyo japix japix (Cramer): Rothschild and Jordan (1903), in Trinidad. There is an old, undated specimen in NHMUK,

Figs. 149. Pseudosphinx tetrio (Linnaeus) IHPDOH3RUWRI6SDLQ-XO\ :GH9RRJG >0-:&@

Figs. 150-151. Aleuron carinata  :DONHU PDOH&XUHSHEODFNOLJKWWUDS-DQXDU\)HEUXDU\ )'%HQQHWW >0-:&@ 150 , GRUVDO 151 YHQWUDO

Fig. 152. Aleuron chloroptera  3HUW\ PDOH&XUHSHEODFNOLJKWWUDS6HSWHPEHU )'%HQQHWW >8:,=0@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 53 and recently one was photographed attracted to light in the 59. Enyo lugubris lugubris (Linnaeus, 1771) Arima Valley. (Figs. 156-158) $ULPD9DOOH\$VD:ULJKW1DWXUH&HQWUH"ƃ0DUFK 6 Enyo lugubris (Linnaeus): Kaye (1901), Kaye (1914a), Nanz photo , http://www.stevenanz.com/Main_Directory/ Cock and Boos (2007) Trips/2015_Trinidad_Tobago/Trinidad_Tobago_Leps/ Epistor lugubris lugubris (Linnaeus): Rothschild and source/img_3286.htm) (Figs. 153-155). Jordan (1903), Kaye and Lamont (1927) 7ULQLGDGƃ>1+0@ Epistor lugubris (Linnaeus): Kaye (1914b), Cary (1951) Enyo lugubris lugubris (Linnaeus): Schreiber (1978), Subfamily: Macroglossinae Stradling et al . (1983) Tribe: Dilophonotini ,GHQWL¿FDWLRQ . A sexually dimorphic species. The male Subtribe Philampelina might only be mistaken for E. ocypete (Linnaeus) (below), Based on the phyllogentic tree in Kawahara et al . but that species has a very conspicuous yellow-white patch (2009, Figure 3), this subtribe comprises two groups: the on the dorsal hindwing dorsum that is absent in E. lugubris . Enyo and Eumorpha group (three species of Enyo and nine However, this feature is not visible in images of the living of Eumorpha Hübner), and the Pachygonidia group (one moths, and the discal spot at the end of the cell should be species of Pachygonidia ). examined – dark in E. lugubris , faint or pale in E. ocypete . The female of E. lugubris is less colourful than that of E. Enyo – Eumorpha group ocypete , and the dorsal forewing discal line extends to The three Trinidad species of Enyo form a weakly the dorsum as a weakly distinguished line, whereas in E. supported group with Eumorpha . ocypete it extends to the dorsum as the sharply contrasting inner border of a broad dark band.

Figs. 153-155. Enyo japix japix  &UDPHU $ULPD9DOOH\$VD:ULJKW1DWXUH&HQWUH0DUFK SKRWRV61DQ]  153 GRUVDOYLHZ 154 ODWHUDOYLHZ 155 YHQWUDOYLHZ 1RWWRVFDOH

Figs. 156-158. Enyo lugubris lugubris  /LQQDHXV  156 PDOH&XUHSH09/0D\ 0-:&RFN >0-:&@ 157 DV)LJ YHQWUDOYLHZ 158 IHPDOH&XUHSH09/6HSWHPEHU 0-:&RFN >0-:&@ 54 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Status in Trinidad . A common species almost exclusively Status in Trinidad . An uncommon species in Trinidad, found in suburban areas. with all records from suburban areas.

60. Enyo ocypete (Linnaeus, 1758) (Figs. 23, 159-160) Eumorpha Hübner, [1807] Enyo ocypete (Linnaeus): Schreiber (1978), Stradling et This genus has nine Trinidad species. It was al . (1983), Cock (2017b) previously been placed in a separate tribe Philampelini, ,GHQWL¿FDWLRQ . A strongly sexually dimorphic species. but is clearly nested within Dilophonotini in Kawahara et Diagnostic features to distinguish E. ocypete from E. al .’s (2009) phylogeny. Ponce et al . (2015) discuss the lugubris are discussed under that species. phylogeny of the genus and the development of a dorsal Status in Trinidad . Not as common as E. lugubris in posterior eye spot on the caterpillars, grouping species Trinidad, but E. ocypete is also a common species. It is into six provisional groups (referred to as clades I–VI). found in both suburban and lowland forested areas. Their phylogeny indicates that E. phorbas (Cramer), E. capronnieri (Boisduval) and E. labruscae form the basal 61. Enyo gorgon (Cramer, 1777) (Figs. 161-162) clade I of the genus, although the inclusion of the last- Enyo gorgon (Cramer): Kaye (1901), Kaye (1914a) named is not strongly supported. The remaining Trinidad Epistor gorgon (Cramer): Rothschild and Jordan (1903), species form two groups based on adult markings, E. Kaye (1914b), Kaye and Lamont (1927) vitis and E. fasciatus in one and the remaining species, E. Epistor gorgon gorgon (Cramer): Schreiber (1978), obliquus (Rothschild and Jordan), E. megaeacus (Hübner), Stradling et al . (1983) E. satellitia and E. anchemolus (Cramer) in the other. ,GHQWL¿FDWLRQ . A strongly sexually dimorphic species. However, the phylogeny of Ponce et al . (2015) indicates The male is distinctive due to the arrangement of three that although the last two species are very close, the others shades of brown on the dorsal forewing, and the bulging are spread across separate clades that do not match adult FRVWD7KHIHPDOHVXSHU¿FLDOO\UHVHPEOHVWKRVHRIRWKHU markings; thus, E. fasciatus and E. megaeacus are in clade members of the genus, but the contrasting shades of brown IV whereas E. vitis is in clade VII with E. anchemolus , and the distinct discal eye spot of the dorsal forewing make while E. obliquus is in clade II and E. satellitia in clade VI. it easily recognisable.

Figs. 159-160. Enyo ocypete /LQQDHXV  159 PDOH&XUHSH09/-DQXDU\ 0-:&RFN >0-:&@ 160 IHPDOH&XUHSH09/ 6HSWHPEHU 0-:&RFN >0-:&@

Figs. 161-162. Enyo gorgon &UDPHU  161 PDOH9DOVD\Q3DUNDWOLJKW$XJXVW 0-:&RFN >0-:&@ 162 IHPDOH&XUHSH OLJKWWUDS-XO\ )'%HQQHWW >8:,=0&$%,@ SKRWR8:,=0  +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 55

62. Eumorpha anchemolus (Cramer, 1779) E. obliquus are distinctly larger than E. satellitia and E. (Figs. 26, 163-164) megaeacus (which is not helpful when working from an Pholus anchemolus (Cramer): Rothschild and Jordan image with no scale) . Characters of the dorsal forewing (1903), Kaye (1914b), Kaye and Lamont (1927) can be used as follows to separate the four species. There Eumorpha anchemolus (Cramer): Cary (1951) is a darker area running from the base of the forewing Eumorpha anchemola [sic ] (Cramer): Schreiber (1978), along the dorsum to about the mid-point where it either Stradling et al . (1983), D’Abrera ([1987]) ends in a sharply demarcated, pointed wedge ( E. satellitia , Eumorpha triangulum (Rothschild and Jordan): Cock E. obliquus , E. megaeacus ), or a diffuse area spreading  >PLVLGHQWL¿FDWLRQ@ further into the forewing disc ( E. anchemolus ). In E. ,GHQWL¿FDWLRQ 7KH¿UVWIRXUVSHFLHVRI Eumorpha treated satellitia , the basal side of this wedge is also sharply here are all rather similar: E. anchemolus , E. satellitia , E. demarcated and pointed, in E. obliquus , a dark subdiscal obliquus and E. megaeacus . Eumorpha anchemolus and line runs from the upper inner corner of the wedge to the

Fig. 163. Eumorpha anchemolus  &UDPHU PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW$XJXVW 0-:&RFN >0-:&@

Figs. 164. Eumorpha anchemolus  &UDPHU IHPDOH%ULJDQG+LOOOLJKWKRXVHVHFXULW\09/OLJKWV0DUFK 0-:&RFN >0-:&@ 56 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE costa, and in E. megaeacus the basal side of the wedge is 64. Eumorpha obliquus (Rothschild and Jordan, 1903) barely differentiated from the wedge and forms part of a (Figs. 166-167) continuous dark band from the base of the forewing to the Pholus obliquus Rothschild and Jordan: Kaye and Lamont end of the wedge, the band being sharply demarcated by a (1927) pale stripe above it from the base of the wing. Eumorpha obliquus (Rothschild and Jordan): Cary (1951), Males of E. anchemolus from Trinidad generally Cock (2017b) have a reddish tone (Fig. 163), whereas the three females Eumorpha obliquus obliquus (Rothschild and Jordan): H[DPLQHGODFNWKLV )LJ &RFN  PLVLGHQWL¿HG Schreiber (1978), Stradling et al . (1983) two females from Brigand Hill as E. triangulum. Taxonomic issues. Kitching & Cadiou (2000) treated Eumorpha anchemolus and E. triangulum can be separated E. obliquus as having three subspecies, but Eitschberger by the shape of the subapical costal patch on the forewing (2011) raised E. orientis (Daniel) and E. guadelupensis upperside: in E. anchemolus this is cut off by Rs4 (vein (Chalumeau & Delplanque) to species level, leaving E. 6), making it quadrate as in E. satellitia and the other obliquus with no subspecies. species, whereas in E. triangulum the inner, basal corner ,GHQWL¿FDWLRQ . See under E. anchemolus (above). is elongated downwards across about half the wing width Status in Trinidad . An uncommon species in Trinidad, (Kitching 2017). recorded from Morne Bleu and Curepe, but rare in Status in Trinidad . An occasional species in Trinidad, suburban areas. rarely found in suburban areas and normally found in forested areas, most commonly at Morne Bleu. 65. Eumorpha megaeacus (Hübner, [1819]) (Fig. 168) Eumorpha eacus (Cramer): Stradling et al . (1983) 63. Eumorpha satellitia (Linnaeus, 1771) licaon (Cramer, [unavailable homonym] 1775) (Figs. 24, 165) ,GHQWL¿FDWLRQ . See under E. anchemolus (above). Pholus satellitia licaon (Cramer): Kaye (1914b), Kaye Status in Trinidad . A rare species in Trinidad with just and Lamont (1927) two records. The food plants are particular species of Pholus satellitia lichaon [ sic ] (Cramer): Cary (1951) Ludwigia spp. (Onagraceae) (Appendix Table 1). Janzen & Eumorpha satellitia satellitia (Linnaeus): Schreiber Hallwachs (2017) found it almost entirely on L. leptocarpa (1978), Stradling et al . (1983) [Jamaican subspecies] and L. octovalvis , rather than other Ludwigia spp. in Costa Eumorpha satellitia licaon (Cramer): Cock (2017b) Rica. These two species are particularly associated with ,GHQWL¿FDWLRQ . See under E. anchemolus (above). freshwater swamps and riparian habitats (CABI 2015, Status in Trinidad . A rather common and widespread University of Florida 2017), which are under-collected species in Trinidad, more frequent in forested areas than habitats in Trinidad, and may explain why there are not suburban areas more records of E. megaeacus .

Fig. 165. Eumorpha satellitia licaon &UDPHU PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 57

Cushe Village, Cunapo Southern Main Road, approx- ,GHQWL¿FDWLRQ . The broad white stripe from the base (or LPDWHO\PLGZD\EHWZHHQ%LFKHDQG5LR&ODUR"Ƃ near) to the apex of the dorsal forewing separates E. vitis , September 2017 (K. Mahabir photo) E. fasciatus and Hyles lineata (below) from other Trinidad +ROOLV5HVHUYRLUDWSXPSKRXVHOLJKWVƃK hawk-moths. Both Eumorpha spp. have an additional September 1978 (M.J.W. Cock) [MJWC] transverse stripe from mid-dorsum to costa, just before apex. The margin of the dorsal hindwing is broadly 66. Eumorpha vitis vitis (Linnaeus, 1758) (Figs. 25, 169) pink in E. fasciatus and narrowly pale brown in E. vitis Pholus vitis vitis (Linnaeus): Kaye and Lamont (1927) making these species easy to separate. However, when Eumorpha vitis vitis (Linnaeus): Cary (1951), Schreiber this character is not visible in images of living moths, (1978), Stradling et al . (1983), Cock (2017b) they can be separated by the double discal spot in E. vitis ,

Fig. 166. Eumorpha obliquus 5RWKVFKLOGDQG-RUGDQ PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW$XJXVW 0-:&RFN  >0-:&@

Figs. 167. Eumorpha obliquus 5RWKVFKLOGDQG-RUGDQ IHPDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW6HSWHPEHU 0-: &RFN >0-:&@ 58 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE which is single in E. fasciatus , and by the narrow, pale line 67. Eumorpha fasciatus fasciatus (Sulzer, 1776) from just before mid-dorsum to the main white stripe at (Figs. 27, 170) right angles to the dorsum in E. fasciatus and the less well Pholus fasciatus (Sulzer) ( vitis ): Kaye (1914b), marked, broader, double line in the same area, but nearer Pholus fasciatus (Sulzer): Kaye and Lamont (1927) the wing base and at an angle to the dorsum in E. vitis . Eumorpha fasciata fasciata [ sic ] (Sulzer): Schreiber Status in Trinidad . A common species, particularly in (1978), Stradling et al . (1983) suburban and disturbed areas. ,GHQWL¿FDWLRQ . See under E. vitis above.

Fig. 168. Eumorpha megaeacus +EQHU PDOH+ROOLV5HVHUYRLUDWSXPSKRXVHOLJKWVK6HSWHPEHU 0-:&RFN  >0-:&@

Fig. 169. Eumorpha vitis vitis /LQQDHXV PDOH6W$XJXVWLQH09/WUDS$XJXVW )'%HQQHWW >0-:&@

Fig. 170. Eumorpha fasciatus fasciatus  6XO]HU PDOH&XUHSH09/'HFHPEHU 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 59

Status in Trinidad . An occasional species, recorded from 69. Eumorpha capronnieri (Boisduval, [1875]) suburban and disturbed areas, but not forests. (Figs. 28, 172) Pholus capronnieri (Boisduval): Kaye and Lamont (1927) 68. Eumorpha phorbas (Cramer, 1775) (Fig. 171) Eumorpha capronnieri (Boisduval): Schreiber (1978), Pholus phorbas (Cramer): Rothschild and Jordan (1903), Stradling et al . (1983), Cock (2017b) Kaye (1914b), Kaye and Lamont (1927) ,GHQWL¿FDWLRQ . The brown and green dorsal forewings are Eumorpha phorbas (Cramer): Schreiber (1978), Stradling very distinctive. et al . (1983) Status in Trinidad . An occasional species in forested areas. ,GHQWL¿FDWLRQ . The mottled dark green dorsal forewing should serve to recognise this species. Eumorpha 70. Eumorpha labruscae (Linnaeus, 1758) (Figs. 29, 173) capronnieri is similar in markings, but most of the Pholus labruscae (Linnaeus): Kaye (1914b), Kaye and forewing is pale brown. Eumorpha labruscae is uniformly Lamont (1927), Cruttwell (1974) green, but lighter in colour and with straight discal lines Eumorpha labruscae labruscae (Linnaeus): Schreiber across the wing, and a small brown patch centrally; when (1978), Stradling et al . (1983) the hindwings are visible, the unusual blue markings are Taxonomic issues. Kitching & Cadiou (2000) included a very distinctive. subspecies from Galapagos, but this is sunk by Kitching Status in Trinidad . A fairly common species in all (2017). forested areas, but rare in suburban areas. ,GHQWL¿FDWLRQ . The red, yellow and blue markings of the

Fig. 171. Eumorpha phorbas &UDPHU PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW$XJXVW 0-:&RFN >0-:&@

Fig. 172. Eumorpha capronnieri %RLVGXYDO PDOH5LR&ODUR*XD\DJXD\DUH5RDGPLOHVWRQH09/6HSWHPEHU 0-: &RFN >0-:&@ 60 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE dorsal hindwings are very unusual, but the rather uniform ,GHQWL¿FDWLRQ . This medium sized, brown species, with green dorsal forewing with straight discal lines is also two discal lines on the dorsal hindwing can only be distinctive. confused with P. subhamata (Walker), which has the apex Status in Trinidad . A common species in Stradling et of the forewings truncate rather than falcate. Pachygonidia al .’s (1983) study, but it seems to have been much less subhamata is recorded from Trinidad by Schreiber (1978) common in subsequent years. Most records are from but I have not seen any specimens. suburban areas. Status in Trinidad . Just one poorly documented record IURPHDUO\ODVWFHQWXU\7KLVLVDQRWKHUJHQXVWKDWÀLHV Pachygonidia group at dusk (and probably also dawn) and is best captured at 7KLVJHQXVZLWKRQHUHFRUGRIRQHFRQ¿UPHG7ULQLGDG ÀRZHUV +D[DLUHE  species forms a separate group, most closely related to >3DOPLVWH@ƂXQGDWHG>1/DPRQW@>8:,=0 several Old World genera (Kawahara et al. 2009, Fig. 3). Subfamily: Macroglossinae 71. Pachygonidia caliginosa (Boisduval, 1870) (Fig. 174) Tribe Macroglossini Pachygonia [sic ] caliginosa (Boisduval): Schreiber Subtribe: Choerocampini (1978), Stradling et al . (1983) Two genera of Macroglossini are found in Trinidad: There is a specimen in Lamont’s collection in UWIZM the Old World genus Hyles with a single representative, with no data, erroneously labelled Eupyrrhoglossum and the New World genus Xylophanes with 12 species. sagra (Poey). It is probably a Trinidad specimen, taken The latter are quite easy to recognise by virtue of the by Lamont prior to 1915, most likely at Palmiste (I have resting position with the wings held horizontal at an angle observed that material of this vintage from Palmiste listed of about 60° and the long, relatively thin body. by Kaye and Lamont (1927) is unlabelled in UWIZM).

Fig. 173. Eumorpha labruscae /LQQDHXV IHPDOH9DOVD\Q3DUNDWOLJKW6HSWHPEHU 0-:&RFN >0-:&@

Fig. 174. Pachygonidia caliginosa  %RLVGXYDO IHPDOH>7ULQLGDG@>1/DPRQW@>8:,=0@ SKRWR8:,=0  +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 61

72. Hyles lineata (Fabricius, 1775) (Fig. 175) &XUHSHOLJKWWUDSƃ-XO\ )'%HQQHWW >8:,=0 Celerio lineata (Fabricius): Cary (1951) CABI.4988] Hyles lineata lineata (Fabricius): Stradling et al . (1983) Taxonomic issues. Hyles lineata is a New World species 73. Xylophanes anubus (Cramer, 1777) (Fig. 176) (introduced in Hawai’i) that could be confused with the Xylophanes anubus (Cramer): Kaye and Lamont (1927), very similar Old World species, H. livornica Esper, which Cary (1951), Schreiber (1978), Stradling et al . (1983) has been reported from French Guiana (Haxaire 1993). ,GHQWL¿FDWLRQ . The larger size and relatively uniform 7KH LGHQWL¿FDWLRQ RI WKH VLQJOH NQRZQ VSHFLPHQ IURP beige coloured dorsal forewings should distinguish this 7ULQLGDG EHORZ  ZDV FRQ¿UPHG XVLQJ WKH GLDJQRVWLF species from other Xylophanes spp. characters set out in Haxaire (1993) and Eitschberger and Status in Trinidad . This species is quite common in Steiniger (1976). forested areas, particularly in the Northern Range, but rare ,GHQWL¿FDWLRQ . The broad white stripe from base to apex in suburban and disturbed areas. RIWKHIRUHZLQJLVVXSHU¿FLDOO\VLPLODUWR Eumorpha vitis and E. fasciatus , but both those species have a transverse 74. Xylophanes ceratomioides (Grote and Robinson, stripe from mid-termen to costa just before apex. Any 1867) (Fig. 177) future Trinidad specimens should be checked against H. Xylophanes ceratomioides (Grote and Robinson): livornica . Schreiber (1978), Stradling et al . (1983) Status in Trinidad . Only known from a single specimen ,GHQWL¿FDWLRQ . A very distinctive species in the Trinidad taken by light trap at Curepe. Given that this is a vagile fauna; the arrangement of the brown and pinkish areas of species, it may be that it is not normally resident in Trinidad. the dorsal forewing are like no other species, although

Fig. 175. Hyles lineata )DEULFLXV PDOH&XUHSHOLJKWWUDS-XO\ )'%HQQHWW >8:,=0&$%,@ SKRWR8:,=0 

Fig. 176. Xylophanes anubus &UDPHU PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKWHDUO\1RYHPEHU 0-:&RFN >0-:&@ 62 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

WKHUHDUHRWKHUVXSHU¿FLDOO\VLPLODU Xylophanes spp. on the forewings, the other two being X. neoptolemus (Cramer) mainland (D’Abrera 1987). and X. tersa (Linnaeus). The latter has yellow markings Status in Trinidad . A fairly common species in forested on the dorsal hindwing whereas the other two are pinkish areas, but rare in suburban and disturbed areas. red, but this character is not evident in images of the living adults. Quite subtle differences can be seen comparing the 75. Xylophanes chiron (Drury, 1773) nechus (Cramer, dorsal forewings, but the body colouring and markings are 1777) (Figs. 30, 178-179) also helpful. In X. tersa , the base of the dorsal forewing Xylophanes chiron nechus (Cramer): Kaye and Lamont (1927), is pale, sharply contrasting with the adjacent thorax (Figs. Cary (1951), Schreiber (1978), Stradling et al . (1983) 34-36), whereas there is little colour difference between ,GHQWL¿FDWLRQ . A very distinctive species in the Trinidad the wing base and adjacent thorax for X. loelia and X. fauna. neoptolemus . In addition, X. tersa has golden dorso-lateral Status in Trinidad . This is a common, sometimes very longitudunial bands on the abdomen (Figs 34, 190), not common, species in forested areas, and is also occasionally present on the other two species. The dorsal thorax and found in suburban and disturbed areas. body of X. loelia is paler than in X. neoptolemus , and X. loelia has a faint thin brown dorsal line on the abdomen but 76. Xylophanes loelia (Druce, 1878) (Figs. 31, 180-181) not the thorax, whereas X. neoptolemus has three parallel Xylophanes loelia (Druce): Schreiber (1978), Stradling et brown dorsal lines on the abdomen and one on the thorax. al . (1983), Vaglia et al . (2008) Status in Trinidad . This is an occasional species in ,GHQWL¿FDWLRQ . This is the least common of three medium suburban and disturbed areas. It may have been overlooked sized species with slightly contrasting brown dorsal as the common X. neoptolemus by Stradling et al . (1983).

Fig. 177. Xylophanes ceratomioides *URWHDQG5RELQVRQ PDOH7RFRDWOLJKW-XQH 0-:&RFN >0-:&@

Figs. 178-179. Xylophanes chiron nechus &UDPHU  178 PDOH&XPDFD5RDGPLOHV09/-XO\ 0-:&RFN >0-:&@ 179 IHPDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XQH 0-:&RFN >0-:&@ +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 63

77. Xylophanes neoptolemus (Cramer, 1780) 79. Xylophanes pluto (Fabricius, 1777) (Figs.182-183) (Figs. 32, 186-187) Choerocampa neoptolemus (Cramer): Kaye (1901), Kaye Xylophanes pluto (Fabricius): Kaye (1914b), Kaye and (1914a) Lamont (1927), Schreiber (1978), Stradling et al . Xylophanes neoptolemus (Cramer): Rothschild and Jordan (1983), Cock (2017b) (1903), Kaye (1914b), Kaye and Lamont (1927), ,GHQWL¿FDWLRQ . The dorsal green forewing, infused with Kitching and Cadiou (2000), Vaglia et al . (2008) white and multiple discal lines should separate this species Xylophanes neoptolemus trinitatis Closs: Closs (1917), from others even if the vivid orange hindwing is not visible. Schreiber (1978), Stradling et al . (1983) Status in Trinidad . A fairly common species in both Xylophanes neoptolemus trinitatas [sic ] Closs: Cary (1951) suburban and forested areas. ,GHQWL¿FDWLRQ . See under X. loelia (above). Status in Trinidad . A common and widespread species. 80. Xylophanes porcus (Hübner [1823]) continentalis Rothschild and Jordan, 1903 (Figs. 188-189) 78. Xylophanes pistacina (Boisduval, [1875]) Xylophanes porcus continentalis Rothschild and Jordan: (Figs. 33, 184-185) Schreiber (1978), Stradling et al . (1983) Xylophanes pistacina pistacina (Boisduval): Schreiber ,GHQWL¿FDWLRQ . This plain brown species is unlikely to be (1978), Stradling et al . (1983) confused with any other at present, but there are cryptic ,GHQWL¿FDWLRQ . The mixture of pastel green-grey colours species under this name in mainland South America (I.J. and a simple discal line on the dorsal forewings make this Kitching pers. comm. 2017). species distinct. Status in Trinidad . An occasional species in forested Status in Trinidad . An occasional species in suburban areas. and forested areas.

Figs. 180-181. Xylophanes loelia 'UXFH PDOH&XUHSH09/6HSWHPEHU 0-:&RFN >0-:&@ 180 GRUVDO 181 YHQWUDO

Figs. 182-183. Xylophanes neoptolemus &UDPHU PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW-XO\ 0-:&RFN >0-:&@ 182 GRUVDO 183 YHQWUDO 64 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

81. Xylophanes tersa tersa (Linnaeus, 1771) 82. Xylophanes thyelia thyelia (Linnaeus, 1758) (Figs. (Figs. 34-36, 190-191) 192-195) Choerocampa tersa (Linnaeus): Kaye (1901, 1914a) Xylophanes thyelia (Linnaeus): Kaye (1914b), Kaye and Xylophanes torsa [sic ] (Linnaeus): Kaye (1914b) Lamont (1927), Cary (1951) Xylophanes tersa (Linnaeus): Rothschild and Jordan Xylophanes thyelia thyelia (Linnaeus): Schreiber (1978), (1903), Kaye and Lamont (1927) Stradling et al . (1983) Xylophanes tersa tersa (Linnaeus): Schreiber (1978), ,GHQWL¿FDWLRQ . A very distinctive species in the Trinidad Stradling et al . (1983), Cock (2017b) fauna, due to its small size, pale line from base to apex of ,GHQWL¿FDWLRQ . See under X. loelia (above). the dorsal forewing, and rich colouring ventrally. Status in Trinidad . The commonest hawk-moth in Status in Trinidad . An uncommon species, primarily Trinidad, occurring in all habitats sampled. found in forested areas, but occasionally in suburban areas.

Figs. 184-185. Xylophanes pistacina %RLVGXYDO>@  184 , PDOH&XUHSH09/-XQH 0-:&RFN >0-:&@ 185 IHPDOH $ULPD9DOOH\6LPOD09/0D\ 0-:&RFN >0-:&@

Figs. 186-187. Xylophanes pluto  )DEULFLXV PDOH6W$XJXVWLQH097UDS$XJXVW )'%HQQHWW >0-:&@ 186 GRUVDO 187 , YHQWUDO

Figs. 188-189. Xylophanes porcus continentalis 5RWKVFKLOGDQG-RUGDQPDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKWXQWLOK 6HSWHPEHU 0-:&RFN >0-:&@ 188 GRUVDO 189 YHQWUDO +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 65

83. Xylophanes titana (Druce, 1878) (Figs. 37, 196-197) ,GHQWL¿FDWLRQ . Images of living adults of this species Xylophanes titana (Druce): Stradling et al . (1983), Cock might be confused with X. tersa , but X. titana is larger, (2007) the dorsal forewing is darker and more contrasting and the Biology in Trinidad. Cock (2007) illustrates the adult abdomen and thorax have two pale dorsal lines and early stages from a caterpillar collected on Endlichera Status in Trinidad . An occasional species in forested umbellata (Rubiaceae) on Morne Catherine. areas.

Figs. 190-191. Xylophanes tersa tersa /LQQDHXV PDOH&XUHSH09/$XJXVW 0-:&RFN >0-:&@ 190 GRUVDO 191 YHQWUDO

Figs. 192-195. Xylophanes thyelia thyelia /LQQDHXV PDOHV 192 0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKWK0DUFK 0-:&RFN >0-:&@ 193 DV)LJYHQWUDOYLHZ 194 &XPDFD5RDGPLOHV09/2FWREHU 0-:&RFN >0-:&@ 195 DV)LJYHQWUDOYLHZ

Figs. 196-197. Xylophanes titana 'UXFH PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKWXQWLOK6HSWHPEHU 0-:&RFN  >0-:&@ 196 GRUVDO 197 YHQWUDO 66 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

84. Xylophanes tyndarus tyndarus (Boisduval, [1875]) LQLGHQWL¿FDWLRQLQWKHLQWHUSUHWDWLRQRIODEHOV µ7ULQLGDG¶ (Figs. 38, 198-199) occurs in several Neotropical collecting localities in diverse Choerocampa tyndarus (Boisduval): Kaye (1901, 1914a) countries), and even transcription errors. Although Stradling Xylophanes tyndarus (Boisduval): Rothschild and Jordan et al . (1983) also list these species, this is purely on the (1903), Kaye (1914b), Kaye and Lamont (1927), Cary basis of the distribution reported by Schreiber (1978) and (1951), Cock (2017b) QRWRQQHZREVHUYDWLRQV1HYHUWKHOHVVWKH¿UVWIRXURI Xylophanes tyndarus tyndarus (Boisduval): Schreiber WKHVHVSHFLHVDUHGXVNDQGGDZQÀRZHUIHHGHUVXQOLNHO\WR (1978), Stradling et al . (1983) EHFDSWXUHGDWOLJKW VHHLQWURGXFWLRQ DQGWKH¿IWKFRXOGEH Taxonomic issues. Haxaire (2013) described a new overlooked for the common X. tersa , so in principle there subspecies for Central America, whereas the nominotypical is no reason to say that their presence in Trinidad will not subspecies is restricted to South America, including EHFRQ¿UPHGLQWKHIXWXUH Trinidad. ,GHQWL¿FDWLRQ . This plain green species, slightly darker Aleuron iphis (Walker, 1856) beyond the discal line, should not be confused with any Aleuron iphis (Walker): Schreiber (1978), Stradling et al . other Trinidad species. (1983) Status in Trinidad . An occasional species in forested 7KLVUDUHGXVNÀRZHUIHHGHURFFXUVLQ)UHQFK*XLDQD areas, rare in suburban areas. (Haxaire and Rasplus 1987b) and so might be a Trinidad species. 6SHFLHVQRWFRQ¿UPHGIURP7ULQLGDG The following species have been recorded as occurring Aleuron neglectum Rothschild and Jordan, 1903 in Trinidad by Schreiber (1978) and Stradling et al . (1983), Aleuron neglectum neglectum Rothschild and Jordan: EXW,KDYHEHHQXQDEOHWRFRQ¿UPWKHPIURPVSHFLPHQV Schreiber (1978), Stradling et al . (1983) photographs or recent records, and there are no specimens Same comments as for the preceding A. iphis . known to I.J. Kitching (pers. comm. 2017) in the NHMUK or elsewhere. Schreiber (1978) based his distribution table Pachygonidia subhamata (Walker, 1856) on a review of 17 museums and private collections in Europe Pachygonia [sic] subhamata (Walker): Schreiber (1978), and USA containing nearly 75,000 labelled Neotropical Stradling et al . (1983) specimens. At the time, Schreiber (1978) lists 22,815 *LYHQWKDWWKLVLVDGXVNÀRZHUIHHGHUDQGKDVEHHQ Neotropical records from the NHMUK, but the NHMUK recorded from coastal Venezuela (I.J. Kitching pers. Sphingidae holdings have been doubled by the addition of comm. 2017), it may yet be found to occur in Trinidad. the J.-M. Cadiou collection. Of the collections checked by Schreiber, I have only checked the NHMUK (partly in Unzela pronoe pronoe Druce, 1894 person, but completely by consultation with I.J. Kitching), Enyo pronoe pronoe (Druce): Schreiber (1978), Stradling and the implication is that voucher specimens for some of et al . (1983) Schreiber’s records are held in one or more of the museums 7KLV GXVN ÀRZHUIHHGHU RFFXUV LQ )UHQFK *XLDQD or collections that I have not seen. Having said that, there (Haxaire and Rasplus 1987b), so could be a Trinidad is also the possibility of errors – in labelling, in curation, species.

Figs. 198-199. Xylophanes tyndarus tyndarus %RLVGXYDO PDOH0RUQH%OHX7H[WHO,QVWDOODWLRQDWOLJKW0D\ 0-:&RFN  >0-:&@ 198 GRUVDO 199 YHQWUDO +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 67

Xylophanes turbata (Edwards, 1887) [synonym of A. tantalus ] Xylophanes turbata (Edwards): Schreiber (1978), Sesia tantalus (Linnaeus): Kaye (1914b), Cary (1951) Stradling et al . (1983) Sesia tantalus tantalus (Linnaeus): Kaye and Lamont (1927) Schreiber (1978) includes this species from Trinidad Aellopos tantalus (Linnaeus): Schreiber (1978), Stradling in his distribution table, but not in his distribution map. It et al . (1983) has been recorded from coastal Venezuela (I.J. Kitching Kaye (1901) included this species in his provisional list pers. comm. 2017) and so it is possible that it could occur of the Trinidad moths as its synonym as Aellopus sisyphus in Trinidad. It resembles X. tersa , but the dorsal hindwing (Burmeister), referring to a specimen in ‘Coll. Schaus’. is pale brown (D’Abrera [1987]), so it is unlikely to have The bulk of William Schaus’s collection is now held in been overlooked in Stradling et al .’s (1983) comprehensive the USNM although some is in NHMUK and other major light trapping at Curepe. It just might be worth watching collections (Heinrich and Chapin 1942). This specimen out for this species in the dry northwest peninsular or has not been found in USNM (J.W. Brown pers. comm. Bocas Islands. 2017) or NHMUK (I.J. Kitching pers. comm. 2017). Kaye and Lamont (1927) list this species from Trinidad based Species recorded from Trinidad, but considered to be on a record from Fort George (1,000 ft.) [305 m] January errors 1922 (F.W. Jackson). The specimen was located in 6HH FRPPHQWV XQGHU µ6SHFLHV QRW FRQ¿UPHG IURP MGCL and found to be A. clavipes . Early last century, A. Trinidad:’ above. However, in contrast to the species clavipes was considered to be a subspecies of A. tantalus , treated in that section, there are arguments for thinking following the treatment of Rothschild and Jordan (1903), that the following species are unlikely to occur in Trinidad which probably explains the use of the name A. tantalus as set out below. from Trinidad. Schreiber (1978) included records from Kaye and Lamont (1927) so it is reasonable to suggest Adhemarius ypsilon (Rothschild and Jordan, 1903) that his inclusion of this species from Trinidad is based on Amplyterus ypsilon Rothschild and Jordan: Schreiber that source. I conclude that these early records refer to A. (1978), Stradling et al . (1983) clavipes and not A. tantalus . Although recorded from Mexico to Peru, this species is not known from northern Venezuela (I.J. Kitching, pers. Eumorpha cissi (Schaufuss, 1870) comm. 2017) and is unlikely to occur in Trinidad. Eumorpha cissi (Schaufuss): Stradling et al . (1983) Stradling et al . (1983) list one specimen of this species Manduca occulta (Rothschild and Jordan, 1903) from Trinidad, but I have failed to locate a specimen in Manduca occulta occulta (Rothschild and Jordan) : UWIZM or NMSE. It is not impossible that they were Schreiber (1978), Stradling et al . (1983) misled by the slight sexual dimorphism shown by E. Schreiber (1978) lists Manduca occulta occulta from anchemolus , and mistook the darker, less red female for a Trinidad, but this is the Central American equivalent of separate species (as the present author also did initially). 0DQGXFDGLI¿VVD and not a South American species . Eumorpha cissi is an Andean species extending into western Venezuela, and found above 1500m (5,000ft) Manduca sexta jamaicensis (Butler, 1875) (D’Abrera [1987]), and unlikely to occur in Trinidad, and Manduca sexta jamaicensis (Butler): Schreiber (1978), so this record is not accepted. Stradling et al . (1983) Schreiber (1978) records both M. sexta jamaicensis Xylophanes crotonis (Walker, 1856) and M. sexta paphus from Trinidad; the former is restricted Xylophanes crotonis (Walker): Schreiber (1978), Stradling the Caribbean islands, including the Lesser Antilles, and I et al . (1983) have seen no specimens to indicate that it also occurs in Schreiber (1978) includes this species from Trinidad Trinidad. in his distribution table, but not in his distribution map. It appears to be an Andean species (I.J. Kitching pers. comm. Isognathus rimosa (Grote, 1865) 2017) and unlikely to occur in Trinidad. Isognathus rimosa (Grote): Cary (1951) Cary (1951) lists this species as perhaps occurring in DISCUSSION Trinidad (‘Trinidad (?)’), but I know of no records. I have recorded 84 species of Sphingidae from 7ULQLGDG SOXV  XQFRQ¿UPHG UHFRUGV RI ZKLFK ¿YH Aellopos tantalus tantalus (Linnaeus, 1758) are considered possible and seven are considered likely Aellopus [sic ] sisyphus (Burmeister): Kaye (1901) to be errors. This compares with 54 recorded by Kaye 68 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE and Lamont (1927), 81 recorded by Schreiber (1978), collecting effort in each month is uneven, so more detailed and 77 recorded by Stradling et al . (1983) out of a total DQDO\VLVLVQRWMXVWL¿HG2IWKHFROOHFWLQJVLWHVVKRZQLQ possible list of 94 including published records that they Appendix Table 3, the area lumped together under ‘Morne had not substantiated. Comparing my list of 84 species Bleu’ is on the ridge tops of the northern range at the head with Stradling et al ¶V $SSHQGL[   FRQ¿UPHG of the Arima Valley at about 600–700 m (2,000–2,300 records, they treat Erinnyis domingonis as a separate IW ZKHUHDVWKHUHPDLQGHUDUHDOODWVLJQL¿FDQWO\ORZHU species whereas here it is treated as a synonym of E. elevations. There is no hint that any species are only found obscura , I reject their record of E. cissi , and they do not DWWKHKLJKHUHOHYDWLRQVLWHVDOWKRXJKDVLJQL¿FDQWQXPEHU include $GKHPDULXVGDSKQH the four species of Aellopos , of less common species have yet to be recorded from the Aleuron chloroptera , Erinnyis impunctata , Pachygonidia high elevation sites. On balance, there is little evidence of caliginosa , Nyceryx stuarti and Unzela japix . Of these, species being restricted by elevation rather than by habitat. $GKHPDULXVGDSKQH Aleuron chloroptera , N. stuarti and This illustrated checklist of Trinidad hawk-moths Erinnyis impunctata are here recorded from Trinidad for VKRXOGHQDEOHDGXOWVRIDOONQRZQVSHFLHVWREHLGHQWL¿HG WKH¿UVWWLPH and it is my hope that this will stimulate interest in this 2IWKHVSHFLHVDFFHSWHGVL[  DUHEDVHGRQVLQJOH charismatic group in Trinidad and Tobago. Using this records ( $HOORSRVFODYLSHV$WLWDQ , Aleuron chloroptera , FKHFNOLVW LGHQWL¿FDWLRQV FDQ QRZ EH UHOLDEO\ PDGH DQG Hyles lineata , Nyceryx maxwelli , Pachygonidia further studies on distribution and frequency, adult feeding caliginosa ), a slightly higher rate than for Lycaenidae in habits, food plants and life history studies, etc., can all be the recent analysis by Cock & Robbins (2016), where six readily taken up. RXWRI/\FDHQLGDH  DUHNQRZQRQO\IURPVLQJOH specimens. This percentage of records based on single ACKNOWLEDGEMENTS specimens suggests that further collecting will reveal more I would like to thank the following: Ian Kitching new records for Trinidad. (NHMUK) who shared his encyclopaedic knowledge of Comparing the records from suburban habitats and Sphingidae, answered my questions about material in forest habitats (Appendix Tables 2 and 3), it can be seen the NHMUK and reviewed an advanced draft helping that despite the totally disproportionate investment in me to avoid several errors; Pauline Geerah and Mike G. collecting in suburban habitats (Curepe / St. Augustine), Rutherford (UWIZM) who shared images and data of there are many species that were collected as frequently all Sphingidae in UWIZM, some of which are included or more frequently in forest habitats (Appendix Table here: C. Dennis Adams (deceased), Marc S. Botham, Matt 2). I have categorised 23 species as relatively frequent in Kelly, Tarran P Maharaj, John Morrall, Steve Nanz, and suburban habitats and 23 as relatively frequent in forest Kris Sookdeo for sharing their images and allowing me habitats. Six species are frequently collected in both: to use those shown in Figs. 1-49; Fred D. Bennett for Erinnyis alope , E. ello , E. crameri , Eumorpha satellitia , encouraging and facilitating my interest in Sphingidae in Xylophanes neoptolemus and X. tersa . There has been an Trinidad; the staff of the Morne Bleu Textel Installation even larger bias towards collecting in the northern half of (especially Mike Dookie), Simla (Jack Price), Hollis Trinidad than in the south, and not surprisingly, although Reservoir, Brigand Hill Lighthouse, etc. who years ago all bar one of the 84 Trinidad species have been recorded welcomed moth collectors on occasional evenings and from the north, only 51 have been recorded from the south facilitated my collecting; Bob Ramnanan (CABI, Trinidad (Appendix Table 2). Further collecting and observations and Tobago) and Sarah Maharaj (Central Experiment in the south is likely to show that most or all species also Station, Centeno, Trinidad and Tobago) who assisted with occur there. ¿QGLQJ HDUO\ SXEOLFDWLRQV -HDQ +D[DLUH IRU DGYLFH DQG Cock (2017b) listed 26 Sphingidae from Tobago help with literature; Ryan St. Laurent who photographed $SSHQGL[7DEOH UHSUHVHQWLQJRIWKHVSHFLHV the Trinidad specimen of Aellopos clavipes in MGCL, UHFRUGHGKHUHIURP7ULQLGDG7KLVLVVLJQL¿FDQWO\KLJKHU included here; John W. Brown (USNM) who checked for WKDQ  IRU DOO EXWWHUÀLHV &RFN D  UHÀHFWLQJ WKH Trinidad specimens of Aellopos spp. and other selected greater mobility of adult hawk-moths compared to other species in the USNM (and found none); and those who families of Lepidoptera. All species recorded from Tobago have facilitated access to the collections in their care, DUHNQRZQIURP7ULQLGDGDQGFDQEHLGHQWL¿HGXVLQJWKH including staff of the NHMUK (Alessandro Giusti, Martin images provided here. Honey, Geoff Martin), USNM (Marc Epstein), OUNHM The table of captures in forested areas (Appendix *HRUJH0F*DYLQ 106( .HLWK%ODQG$VKOHLJK:LI¿Q  Table 4) indicates that hawk-moths are collected in and UWIZM (various staff of the UWI Department of VLJQL¿FDQW QXPEHUV LQ HYHU\ PRQWK EXW WKH DPRXQW RI Zoology over the years). +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 69

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Ithaca, USA and place: what determines the saturniid and sphingid moth London, UK: Cornell University Press. viii + 227 p. fauna of Santa Rosa National Park, Costa Rica, and what Lamont, N. and Callan, E.McC. 1950. Moths new to does it mean to conservation biology? Brenesia , 25/26: Trinidad, B.W.I. Zoologica , 35: 197-207. 51-87. MFPLMA (Ministry of Food Production, Land and Janzen, D.H. and Hallwachs, W. 2016. DNA barcoding Marine Affairs, Trinidad and Tobago) 2010. Cassava the Lepidoptera inventory of a large complex tropical hornworm Erinnyis ello. Fact Sheet #1. Research Division, conserved wildland, Area de Conservacion Guanacaste, Ministry of Food Production, Land and Marine Affairs, northwestern Costa Rica. Genome , 59: 641-660. Trinidad and Tobago, 2 p. Janzen, D. H. and Hallwachs, W. 2017. Dynamic database MPM (Ministry of Petroleum and Mines, Trinidad and for an inventory of the macrocaterpillar fauna, and its Tobago) 1981. Annual Report for 1978. Government of food plants and parasitoids, of Area de Conservacion Trinidad and Tobago, vi + 26 p. Guanacaste (ACG), northwestern Costa Rica. [Online] Moss, A.M. 1912. On the Sphingidae of Peru. Transactions Available at http://janzen.sas.upenn.edu (Accessed: 14 of the Zoological Society of London , 20: 73-135. April, 2017). Moss, A.M. 1920. Sphingidae of Para, Brazil. Novitates Kawahara, A.Y., Mignault, A.A., Regier, J.C., Kitching, Zoologicae , 27: 333-424, plates i-xi. +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 71

Oehlke, B. 2017. Sphingidae of the Americas. [Online] St Laurent, R.A. and Cock, M.J.W. 2017. Annotated Available at http://www.silkmoths.bizland.com/Sphinx/ list of Mimallonidae (Lepidoptera, Mimallonoidea) from danjansphinx.htm (Accessed: 14 April, 2017). Trinidad and Tobago. Zootaxa , 4268: 53-70. Petroleum Economist 2015. Energy Map of Trinidad and Stradling, D.J., Legg, C.J. and Bennett, F.D. 1983. Tobago. 2015 Edition. [Online] Available at http://ngc. Observations on the Sphingidae (Lepidoptera) of Trinidad. co.tt/wp-content/uploads/pdf/publications/energy-map- Bulletin of Entomological Research , 73: 201-232. of-tnt-2015-ed.pdf (Accessed: 14 April, 2017). Topper, B.F. 1943. Tomato cultivation on the College Pollard, G. 1984. Insect pests of sweet potato in the Farm. Tropical Agriculture , 19(9) (1942): 180-182. Caribbean with particular reference to the borer ( Megastes Tuttle, J.P. 2007. The Hawk Moths of North America. grandalis Guen.). p. 147-152. In: Dolly, D. ed., Root Crops Washington DC: The Wedge Entomological Research in the Caribbean. Proceedings of the Caribbean Regional Foundation. xviii + 253 p. Workshop on Tropical Root Crops. St. Augustine, Trinidad University of Florida 2017. Ludwigia leptocarpa . and Tobago: Faculty of Agriculture, University of the West Available at KWWSVSODQWVLIDVXÀHGXSODQWGLUHFWRU\ Indies. 269 p. ludwigia-leptocarpa/ (Accessed 22 July, 2017). Ponce, F., Breinholt, J.W., Hossie, T., Barber, J.R., Urich, F.W. 1915. Cassava insects. Bulletin of the Janzen, D.H., Hallwachs, W. and Kawahara, A.Y. 'HSDUWPHQWRI$JULFXOWXUH7ULQLGDGDQG7REDJR , 14(2): 2015. A molecular phylogeny of Eumorpha (Lepidoptera: 38-40. Sphingidae) and the evolution of anti-predator larval Vaglia, T. and Haxaire, J. 2005. Revision des Adhemarius eyespots. Systematic Entomology , 40(2): 401-408. du complex gannascus (Stoll, 1790). Lambillionea , Rothschild, W. and Jordan, K. 1903. A revision of the 105(supplement 3): 3-[45]. lepidopterous family Sphingidae. Novitates Zoologicae , Vaglia, T., Haxaire, J., Kitching, I.J., Meusnier, I. and 9(supplement): i-cxxxv, 1-972, 67 pl. Rougerie, R. 2008. Morphology and DNA barcoding Schreiber, H. 1978. Dispersal centres of Sphingidae reveal three cryptic species within the Xylophanes (Lepidoptera). Biogeographica, Volume 10. The Hague: neoptolemus and loelia species-groups (Lepidoptera: Dr W. Junk Publishers. 195 p. Sphingidae). =RRWD[D 1923: 18-36. Soares, A. 1993. Nota sôbre o gênero Callionima Lucas, 1856 Winder, J.A. 1976. Ecology and control of Erinnyis ello (Lepidoptera, Sphingidae, Macroglossinae, Dilophonotini). and E. alope , important insect pests in the New World. Revista Brasileira de Entomologia , 37: 35-39. PANS , 22(4): 449-466. 72 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Appendix Table 1. 2YHUYLHZRIWKHPDLQIRRGSODQWVRI6SKLQJLGDHIRXQGLQ7ULQLGDGEDVHGRQUHFRUGVIURPHOVHZKHUH Species Food plants Sources 1 Adhemarius gannascus $SSDUHQWO\XQNQRZQ 2 Adhemarius daphne Lauraceae ( Ocotea ) -DQ]HQDQG+DOOZDFKV  2HKONH  3 Adhemarius palmeri $SSDUHQWO\XQNQRZQ 4 Protambulyx eurycles 3UREDEO\$QDFDUGLDFHDHDQG6LPDURXEDFHDH 2HKONH  5 Protambulyx strigilis Anacardiaceae ( Anacardium , Astronium , '\HUDQG*HQWU\  7XWWOH  -DQ]HQDQG Spondias , Schinus , Tapirira , Simarouba ) +DOOZDFKV  2HKONH  6 Cocytius duponchel Annonaceae ( Annona , Rollinia, Guatteria , 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG Xylopia ) +DOOZDFKV  2HKONH  7 Cocytius lucifer Annonaceae ( Annona , Desmopsis ) -DQ]HQDQG+DOOZDFKV  2HKONH  8 Cocytius antaeus Annonaceae ( Annona , Rollinia ) 0RVV  7XWWOH  -DQ]HQDQG +DOOZDFKV  2HKONH  9 Neococytius cluentius Annonaceae ( Annona ), Piperaceae ( Piper ) 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG +DOOZDFKV  10 Manduca lefeburii Salicaceae ( Casearia , Zuelania ) +D[DLUHDQG5DVSOXV D -DQ]HQDQG +DOOZDFKV 2HKONH  11 Manduca albiplaga Annonaceae ( Annona , Rollinia ) and 0RVV  -DQ]HQDQG+DOOZDFKV   /DPLDFHDH Aegiphila ); perhaps also 2HKONH  Cordiaceae 12 Manduca franciscae $SSDUHQWO\XQNQRZQ 13 Manduca huascara $SSDUHQWO\XQNQRZQ 14 0DQGXFDÀRUHVWDQ 3RO\SKDJRXVRQDWOHDVWIDPLOLHV 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG SDUWLFXODUO\%LJQRQLDFHDHDQG/DPLDFHDHEXW +DOOZDFKV  2HKONH  DOVR%RUDJLQDFHDH&RUGLDFHDH5XELDFHDH 9HUEHQDFHDHHWF 15 Manduca rustica Polyphagous, including Bignoniaceae, 0RVV  -DQ]HQDQG+DOOZDFKV   &RQYROYXODFHDH&RUGLDFHDH/DPLDFHDH 2HKONH  0DOYDFHDHDQG9HUEHQDFHDH 16 Manduca sexta Mainly Solanaceae ( Capsicum , Datura , 0RVV  7XWWOH  '\HUDQG*HQWU\ Nicotiana , Solanum DOVR/DPLDFHDH  -DQ]HQDQG+DOOZDFKV 2HKONH (Aegiphila ), Martyniaceae ( Martynia ,  Proboscidia ) and Petiveriaceae ( Rivina ) 17 0DQGXFDGLI¿VVD Solanaceae ( Nicotiana , Solanum ) 0RVV  18 Manduca hannibal Solanaceae ( Cestrum , Datura , Solanum ), 0RVV  '\HU et al   -DQ]HQDQG /DPLDFHDH Aegiphila, Clerodendrum ) +DOOZDFKV 2HKONH  19 Manduca ochus Solanaceae ( Cestrum ) -DQ]HQDQG+DOOZDFKV 20 Agrius cingulata Convolvulaceae ( Convolvulus , Ipomoea , 0RVV  7XWWOH  -DQ]HQDQG Merremia ) +DOOZDFKV  2HKONH  21 3DFK\OLD¿FXV Moraceae ( Brosimum , Castilla , Ficus , 0RVV  7XWWOH  '\HUDQG*HQWU\ Maclura)  -DQ]HQDQG+DOOZDFKV  2HKONH  22 Pachylia syces Moraceae ( Ficus , Artocarpus 8UWLFDFHDH 0RVV  '\HUDQG*HQWU\   (Cecropia ) -DQ]HQDQG+DOOZDFKV  2HKONH  23 Callionima pan $SSDUHQWO\XQNQRZQ 24 Callionima calliomenae $SSDUHQWO\XQNQRZQ 25 Callionema inuus Apocynaceae ( Ambelania ) 0RVV  26 Callionima falcifera Apocynaceae ( Stemmadenia ) -DQ]HQDQG+DOOZDFKV  2HKONH  27 Nyceryx coffaeae 5XELDFHDH Calycophyllum , Uncaria ) -DQ]HQDQG+DOOZDFKV  2HKONH  28 Nyceryx maxwelli $SSDUHQWO\XQNQRZQ 29 Nyceryx riscus Bignoniaceae ( Xylophragma ) -DQ]HQDQG+DOOZDFKV  2HKONH  30 Nyceryx stuarti 5XELDFHDH Uncaria ) -DQ]HQDQG+DOOZDFKV  +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 73

Appendix Table 1 Continued. 2YHUYLHZRIWKHPDLQIRRGSODQWVRI6SKLQJLGDHIRXQGLQ7ULQLGDGEDVHGRQUHFRUGVIURPHOVHZKHUH Species Food plants Sources 31 Perigonia pallida $SSDUHQWO\XQNQRZQ 32 Perigonia lusca 5XELDFHDH Calycophyllum , Coffea , Guettarda , 0RVV  7XWWOH  -DQ]HQDQG+DOOZDFKV Ourouparia , Uncaria ), Aquifoliaceae ( Ilex )  2HKONH  33 Eupyrrhoglossum sagra 5XELDFHDH Chomelia , Guettarda , Sabicea ) 0RVV  -DQ]HQDQG+DOOZDFKV   2HKONH  34 Aellopos ceculus 5XELDFHDH Ourouparia , Sabicea ) 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG +DOOZDFKV  35 Aellopos clavipes 5XELDFFDH Randia ) 0RVV  7XWWOH  -DQ]HQDQG+DOOZDFKV  2HKONH  36 Aellopos titan 5XELDFHDH Alibertia , Genipa , Guettarda , 0RVV  7XWWOH  -DQ]HQDQG+DOOZDFKV Randia )  2HKONH  37 Aellopos fadus 5XELDFHDH Alibertia , Augusta , Genipa ) 0RVV  -DQ]HQDQG+DOOZDFKV   2HKONH  38 Oryba kadeni 5XELDFHDH Cosmibuena , Isertia , Palicourea , 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG Remijia ) +DOOZDFKV  2HKONH  39 Oryba achemenides 5XELDFHDH Ourouparia , Uncaria ) 0RVV  -DQ]HQDQG+DOOZDFKV  40 Pachylioides resumens Apocynaceae ( Echites , Forsteronia , 0RVV  -DQ]HQDQG+DOOZDFKV   Zschokkea ) 2HKONH  41 Madoryx oiclus Bignoniaceae ( Tecoma 9HUEHQDFHDH +D[DLUHDQG5DVSOXV D -DQ]HQDQG (Rehdera ) +DOOZDFKV  2HKONH  42 Madoryx plutonius Aquifoliaceae ( Ilex 0HODVWRPDWDFHDH 0RVV  -DQ]HQDQG+DOOZDFKV   (Conostegia , Miconia 9RFK\VLDFHDH Erisma, 2HKONH  Vochysia ) 43 Madoryx bubastus 5XELDFHDH Calycophyllum , Guettarda ) -DQ]HQDQG+DOOZDFKV  2HKONH  44 Hemeroplanes Apocynaceae ( Amblyanthera , Echites , 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG triptolemus Fischeria , Mesechites , Zschokkea ) +DOOZDFKV  2HKONH  45 Phryxus caicus Apocynaceae ( Mesechites ) -DQ]HQDQG+DOOZDFKV  2HKONH  46 Erinnyis alope (XSKRUELDFDH Hevea , Manihot ), Caricaceae 0RVV  +D[DLUHDQG5DVSOXV D  (Carica ) 7XWWOH  '\HUDQG*HQWU\  -DQ]HQDQG +DOOZDFKV  2HKONH  47 Erinnyis lassauxii Apocynaceae ( Gonolobus , Morrenia ) 0RVV  -DQ]HQDQG+DOOZDFKV   2HKONH  48 Erinnyis impunctata $SSDUHQWO\XQNQRZQ 49 Erinnyis ello (XSKRUELDFHDH Euphorbia, Manihot , 0RVV  7XWWOH  '\HUDQG*HQWU\ Poinsettia, Sebastiania ), Caricaceae ( Carica );  -DQ]HQDQG+DOOZDFKV  2HKONH also Apocynaceae, Sapotaceae ( Bumelia ,  Chrysophyllum , Manilkara ) 50 Erinnyis oenotrus Apocynaceae ( Forsteronia , Echites , 0RVV  -DQ]HQDQG+DOOZDFKV   Zschokkea ) 2HKONH  51 Erinnyis crameri Apocynaceae ( 5DXYRO¿D , Stemmadenia ) -DQ]HQDQG+DOOZDFKV  2HKONH  52 Erinnyis obscura Apocynaceae ( Blepharodon , Cynanchum , 0RVV  7XWWOH  -DQ]HQDQG+DOOZDFKV Gonolobus , Sarcostemma )  2HKONH  53 Isognathus scyron Apocynaceae ( Allamanda 6HHWH[W 2HKONH  54 Isognathus caricae Apocynaceae ( Allamanda , Himatanthus ) 2HKONH  55 Pseudosphinx tetrio Apocynaceae ( Plumeria ) 0RVV  -DQ]HQDQG+DOOZDFKV   2HKONH  56 Aleuron carinata 'LOOHQLDFHDH Curatella , Doliocarpus ) -DQ]HQDQG+DOOZDFKV  2HKONH  57 Aleuron chloroptera 'LOOHQLDFHDH Curatella , Davilla ) -DQ]HQDQG+DOOZDFKV  2HKONH  58 Unzela japix 'LOOHQLDFHDH Curatella , Davilla , Doliocarpus , 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG Pinzonia , Tetracera ) +DOOZDFKV  2HKONH  59 Enyo lugubris 9LWDFHDH Cissus 'LOOHQLDFHDH Doliocarpus ) 0RVV  7XWWOH  -DQ]HQDQG+DOOZDFKV  2HKONH  74 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Appendix Table 1 Continued. 2YHUYLHZRIWKHPDLQIRRGSODQWVRI6SKLQJLGDHIRXQGLQ7ULQLGDGEDVHGRQUHFRUGVIURPHOVHZKHUH

Species Food plants Sources 60 Enyo ocypete 'LOOHQLDFHDH Curatella , Davilla , Doliocarpus , 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG Tetracera ), Onagraceae ( Ludwigia 9LWDFHDH +DOOZDFKV  2HKONH  (Cissus ) 61 Enyo gorgon 9LWDFHDH Cissus , Vitis ) 0RVV  -DQ]HQDQG+DOOZDFKV   2HKONH  62 Eumorpha anchemolus 'LOOHQLDFHDH Davilla , Doliocarpus 9LWDFHDH 0RVV  -DQ]HQDQG+DOOZDFKV   (Ampelopsis , Cissus ) 2HKONH  63 Eumorpha satellitia 9LWDFHDH Ampelopsis , Cissus ) 0RVV  7XWWOH  -DQ]HQDQG +DOOZDFKV  2HKONH  64 Eumorpha obliquus Marcgraviaceae ( Schwartzia 9LWDFHDH 0RVV  -DQ]HQDQG+DOOZDFKV  (Cissus ) 65 Eumorpha megaeacus Onagraceae ( Ludwigia ) 0RVV  -DQ]HQDQG+DOOZDFKV   2HKONH  66 Eumorpha vitis 9LWDFHDH Cissus , Vitis ) 0RVV  7XWWOH  '\HUDQG*HQWU\  -DQ]HQDQG+DOOZDFKV  2HKONH  67 Eumorpha fasciatus Onagraceae ( Ludwigia ) 0RVV  7XWWOH  -DQ]HQDQG +DOOZDFKV  2HKONH  68 Eumorpha phorbas Marcgraviaceae ( Schwartzia ) -DQ]HQDQG+DOOZDFKV  69 Eumorpha capronnieri $SSDUHQWO\XQNQRZQ 70 Eumorpha labruscae 9LWDFHDH Ampelopsis , Cissus ) 0RVV  7XWWOH  '\HUDQG*HQWU\  -DQ]HQDQG+DOOZDFKV  2HKONH  71 Pachygonidia $SSDUHQWO\XQNQRZQRWKHUVSHFLHVRI -DQ]HQDQG+DOOZDFKV  caliginosa Pachygonidia IHHGRQ'LOOHQLDFHDH 5XELDFHDH9LWDFHDH 72 Hyles lineata 3RO\SKDJRXVLQFOXGLQJ0HODVWRPDWDFHDH 7XWWOH  -DQ]HQDQG+DOOZDFKV   Nyctaginaceae, Onagraceae, Portulacaceae 2HKONH  73 Xylophanes anubus 5XELDFHDH Palicourea , Psychotria ) 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG +DOOZDFKV  2HKONH  74 Xylophanes 5XELDFHDH Hamelia , Margaritopsis , '\HUDQG*HQWU\  -DQ]HQDQG+DOOZDFKV ceratomioides Psychotria )  2HKONH  75 Xylophanes chiron 0DQ\5XELDFHDH Coussarea , Faramea , 0RVV  '\HUDQG*HQWU\  '\HU et Margaritopsis , Morinda , Palicourea , al   -DQ]HQDQG+DOOZDFKV  2HKONH Psychotria , Rudgea , Spermacoce ),  0HODVWRPDWDFHDH Miconia 9RFK\VLDFHDH (Vochysia ) 76 Xylophanes loelia 5XELDFHDH Spermacoce ); also Onagraceae 0RVV  -DQ]HQDQG+DOOZDFKV  (Ludwigia ) 77 Xylophanes $SSDUHQWO\XQNQRZQ neoptolemus 78 Xylophanes pistacina 5XELDFHDH Psychotria ) 2HKONH  79 Xylophanes pluto (U\WKUR[\ODFHDH Erythroxylon 5XELDFHDH 0RVV  +D[DLUHDQG5DVSOXV E 7XWWOH (Augusta , Hamelia , Manettia , Margaritopsis ,  '\HUDQG*HQWU\  -DQ]HQDQG Morinda ) +DOOZDFKV  2HKONH  80 Xylophanes porcus 5XELDFHDH Hamelia , Margaritopsis , 0RVV  '\HUDQG*HQWU\  -DQ]HQDQG Palicourea , Psychotria ) +DOOZDFKV  2HKONH  81 Xylophanes tersa 5XELDFHDH Diodia , Hamelia , Margaritopsis , 0RVV  7XWWOH  '\HUDQG*HQWU\ Pentas , Richardia , Spermacoce )  -DQ]HQDQG+DOOZDFKV  2HKONH  82 Xylophanes thyelia $SSDUHQWO\XQNQRZQ 83 Xylophanes titana 5XELDFHDH Manettia ) -DQ]HQDQG+DOOZDFKV  2HKONH  84 Xylophanes tyndarus 5XELDFHDH Faramea ) -DQ]HQDQG+DOOZDFKV  2HKONH  +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 75

Appendix Table 2 2YHUYLHZRIWKHIUHTXHQF\RIFDSWXUHGLVWULEXWLRQDQGKDELWDWRI6SKLQJLGDHLQ7ULQLGDGDQG7REDJR Curepe 1969- 1976 LVWKHWRWDOQXPEHURIHDFKVSHFLHVUHFRUGHGE\6WUDGOLQJ et al.   ZLWKWKHWKUHHOLJKWWUDSVLQ&XUHSH 6W$XJXVWLQH RYHU WUDSQLJKWV Curepe 1978-1982 LVEDVHGRQP\LUUHJXODUOLJKWWUDSUHFRUGVRYHUWKHVH\HDUVLQP\JDUGHQLQ&XUHSH 6W$XJXVWLQH  Other records HQFRPSDVVHVDOOUHFRUGVQRWIURP&XUHSH QRWHWKDWUHFRUGVIURP&XUHSH6W$XJXVWLQHWKDWDUHQRWLQFOXGHGLQWKH¿UVW WZRGDWDVHWVDUHQRWLQFOXGHGLQWKHVHQXPEHUV  Suburban species and Forest species JLYHVDVXEMHFWLYHLQGLFDWLRQRIWKHUHODWLYH IUHTXHQF\LQWKHVHKDELWDWVEDVHGRQVLJQVRU±IRUDOPRVWRUFRPSOHWHO\DEVHQW Recorded from South Trinidad: the south of 7ULQLGDGLVWUHDWHGDVVRXWKRIWKH&HQWUDO5DQJHWKHUHDUHQRUHFRUGVIURPWKHVWULSVRXWKRI7DEDTXLWHDQGQRUWKRI6DQ)HUQDQGR Recorded from Tobago LVWDNHQIURP&RFN E  Species Curepe Curepe Other Suburban Forest Recorded Recorded 1969-1976 1978-1981 records species species from South from Trinidad Tobago  Adhemarius gannascus    + +++  Adhemarius daphne   5  +++ X  Adhemarius palmeri   5 ++ ++ X 4 Protambulyx eurycles     ++ X 5 Protambulyx strigilis    ++ ++ X X  Cocytius duponchel 44   ++ + X  Cocytius lucifer    + +++ X 8 Cocytius antaeus    ++ + X  Neococytius cluentius   4 + + X  Manduca lefeburii   4 ++   Manduca albiplaga    + +++  Manduca franciscae     +  Manduca huascara ?   + +++  0DQGXFDÀRUHVWDQ    44 + +++ X X  Manduca rustica    ++ ++ X X  Manduca sexta    +++  XX  0DQGXFDGLI¿VVD    ++ ++ X  Manduca hannibal    + + X  Manduca ochus    + ++ X  Agrius cingulata    X  3DFK\OLD¿FXV    ++ ++ X X  Pachylia syces    +  X  Callionima pan   4  +  Callionima calliomenae    +++   Callionema inuus    +++   Callionima falcifera    +++ ++ X  Nyceryx coffaeae    +++   Nyceryx maxwelli    X  Nyceryx riscus    ++ +  Nyceryx stuarti    ++ X  Perigonia pallida    +++  Perigonia lusca 58   +++ + X X  Eupyrrhoglossum sagra 8   ++ ++ X X  Aellopos ceculus    ++ ++ X X  Aellopos clavipes    X  Aellopos titan    + X  Aellopos fadus    + +  Oryba kadeni    ++ +  Oryba achemenides    ++ ++ X  Pachylioides resumens   4 ++ + X X  Madoryx oiclus    +++ + X X  Madoryx plutonius   5 ++ 76 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Appendix Table 2 Continued 2YHUYLHZRIWKHIUHTXHQF\RIFDSWXUHGLVWULEXWLRQDQGKDELWDWRI6SKLQJLGDHLQ7ULQLGDGDQG7REDJR Species Curepe Curepe Other Suburban Forest Recorded Recorded 1969-1976 1978-1981 records species species from South from Trinidad Tobago  Madoryx bubastus    ++ ++ 44 Hemeroplanes triptolemus    +++ + X X 45 Phryxus caicus    ++ X  Erinnyis alope    +++ +++ X X  Erinnyis lassauxii 44   ++ ++ X X 48 Erinnyis impunctata    +  Erinnyis ello    +++ +++ X X  Erinnyis oenotrus     ++ +  Erinnyis crameri    +++ +++  Erinnyis obscura  4  5 +++  Isognathus scyron    +++ + X 54 Isognathus caricae   4 + X 55 Pseudosphinx tetrio  5  ++ ++ X X  Aleuron carinata    +  Aleuron chloroptera    + 58 Unzela japix    +  Enyo lugubris    +++ X  Enyo ocypete 84 8  +++ + X X  Enyo gorgon 5  4 ++  Eumorpha anchemolus    + +++ X  Eumorpha satellitia  8  +++ +++ X X  Eumorpha obliquus 4   +++ X  Eumorpha megaeacus    +  Eumorpha vitis  5  +++ + X X  Eumorpha fasciatus  8 5 +++ + X  Eumorpha phorbas 5   + +++ X  Eumorpha capronnieri    +++ X X  Eumorpha labruscae    +++ X  Pachygonidia caliginosa   "  Hyles lineata    +  Xylophanes anubus    +++ X  Xylophanes ceratomioides 5   + +++ X  Xylophanes chiron   84 ++ +++ X  Xylophanes loelia   4 +  Xylophanes neoptolemus    +++ +++ X  Xylophanes pistacina    + + X  Xylophanes pluto    +++ + X X  Xylophanes porcus    +++  Xylophanes tersa    +++ +++ X X  Xylophanes thyelia   8 + +++ X  Xylophanes titana    +++ 84 Xylophanes tyndarus    + +++ X X 7RWDO 8848   1XPEHURIVSHFLHV          7KLVLVWKHQXPEHURIVSHFLHVDOORFDWHGVL[VSHFLHVDUHUDWHGLQERWKKDELWDWV Manduca huascara and 0ÀRUHVWDQ ZHUHPL[HGWRJHWKHUDV 0ÀRUHVWDQ 7ZRVSHFLPHQVRI Erinnyis impunctata were included under E. oenotrus. 4,QFOXGLQJWZRVSHFLPHQVLGHQWL¿HGE\6WUDGOLQJ et al   DV E. domingonis. +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 77

Appendix Table 3. 7KHPDLQFROOHFWLQJVLWHVIRU7ULQLGDG6SKLQJLGDHIURPQRQVXEXUEDQVLWHV0RUQH%OHXLQFOXGHVWKH0RUQH%OHX 7H[WHO,QVWDOODWLRQDVZHOODVVLWHVDORQJWKH$ULPD%ODQFKLVVHXVH5RDGIURPWKHEDVHRIWKHURDGWR0RUQH%OHX7H[WHO,QVWDOODWLRQWR WKHHQGRI/DV/DSSDV7UDFHDQG/DODMD5LGJH3DOPLVWHZDV6LU1RUPDQ/DPRQW¶V(VWDWHQHDU6DQ)HUQDQGRDOWKRXJKLWLVDKRXVLQJ GHYHORSPHQWQRZZKHQ/DPRQWFROOHFWHGIURPDURXQGWRWKHVWKHVSHFLHVKHFROOHFWHGLQGLFDWHVLWPXVWKDYHEHHQPL[HG HVWDWHZLWKVRPHJDUGHQDQGVRPHIRUHVWIUDJPHQWV$ULPD9DOOH\LQFOXGHVREVHUYDWLRQVIURPWKH$VD:ULJKW1DWXUH&HQWUH6LPODDQG 6LU1RUPDQ/DPRQW¶V6W3DWUFN¶V(VWDWHDGMDFHQWWR6LPOD&XPDFDUHSUHVHQWVRQHVSHFL¿FVLJKWRQWKH&XPDFD5RDGZKHUH,UDQD PHUFXU\YDSRXUOLJKWRQWZRRFFDVLRQVµ2WKHU¶LQFOXGHVPDWHULDOIURPGLYHUVHSDUWVRI7ULQLGDGLQFOXGLQJ,QQLVV)LHOG%DODQGUD6DQJUH *UDQGH+ROOLV5HVHUYRLU5LR&ODUR±*XD\DJXDUH5RDG0DWXULWDDQG3HQDOIRUHDFKRIZKLFKDWOHDVWWHQUHFRUGVDUHLQFOXGHGDVZHOO DVVLWHVZLWKIHZHUUHFRUGV

Species St. Hill Bleu Aripo Other Arima Valley Valley Morne Brigand Cumaca Palmiste Benedicts Parrylands  Adhemarius gannascus    8  Adhemarius daphne 4     Adhemarius palmeri  5   4 4 Protambulyx eurycles   5 Protambulyx strigilis  5    Cocytius duponchel       Cocytius lucifer    8 Cocytius antaeus     Neococytius cluentius      Manduca lefeburii    Manduca albiplaga  4    Manduca franciscae    Manduca huascara 4  8   0DQGXFDÀRUHVWDQ         Manduca rustica      Manduca sexta    0DQGXFDGLI¿VVD  5      Manduca hannibal    Manduca ochus     Agrius cingulata      3DFK\OLD¿FXV      Pachylia syces  Callionima pan 4  Callionima calliomenae   Callionema inuus  Callionima falcifera      4  Nyceryx coffaeae  Nyceryx maxwelli   Nyceryx riscus    Nyceryx stuarti   Perigonia pallida  Perigonia lusca       Eupyrrhoglossum sagra      Aellopos ceculus     Aellopos clavipes   Aellopos titan   Aellopos fadus   Oryba kadeni   Oryba achemenides     Pachylioides resumens   78 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Appendix Table 3. &RQWLQXHG7KHPDLQFROOHFWLQJVLWHVIRU7ULQLGDG6SKLQJLGDHIURPQRQVXEXUEDQVLWHV

Species St. Hill Bleu Aripo Other Arima Valley Valley Morne Brigand Cumaca Palmiste Benedicts Parrylands  Madoryx oiclus   Madoryx plutonius    Madoryx bubastus   44 Hemeroplanes triptolemus    45 Phryxus caicus    Erinnyis alope       Erinnyis lassauxii 5     48 Erinnyis impunctata  Erinnyis ello        Erinnyis oenotrus      Erinnyis crameri  4   Erinnyis obscura    Isognathus scyron    54 Isognathus caricae 4 55 Pseudosphinx tetrio     Aleuron carinata   Aleuron chloroptera 58 Unzela japix   Enyo lugubris    Enyo ocypete     Enyo gorgon  Eumorpha anchemolus      Eumorpha satellitia 8        Eumorpha obliquus   Eumorpha megaeacus   Eumorpha vitis     Eumorpha fasciatus 4   Eumorpha phorbas   5   Eumorpha capronnieri 4    Eumorpha labruscae     Pachygonidia caliginosa  Hyles lineata  Xylophanes anubus  4   Xylophanes ceratomioides 8 8     Xylophanes chiron        Xylophanes loelia   Xylophanes neoptolemus          Xylophanes pistacina    Xylophanes pluto  5     Xylophanes porcus      Xylophanes tersa    8 5  Xylophanes thyelia 4    Xylophanes titana   84 Xylophanes tyndarus 5    1XPEHURIUHFRUGV          1XPEHURIVSHFLHV       5   +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 79

Appendix Table 4 7KHPRQWKVRIFDSWXUHIRU7ULQLGDG6SKLQJLGDHIURPQRQVXEXUEDQVLWHV

Species May July April Total June March August October January February December November September  Adhemarius gannascus      4    4  60  Adhemarius daphne      9  Adhemarius palmeri         12 4 Protambulyx eurycles   2 5 Protambulyx strigilis       4 4 34  Cocytius duponchel 4     12  Cocytius lucifer   5  8 8 Cocytius antaeus    4  Neococytius cluentius     5  Manduca lefeburii  2  Manduca albiplaga     5  33  Manduca franciscae   2  Manduca huascara        22  0DQGXFDÀRUHVWDQ  5  8    51  Manduca rustica    4    13  Manduca sexta     7  0DQGXFDGLI¿VVD  4        21  Manduca hannibal     5  Manduca ochus     4  Agrius cingulata        4   25  3DFK\OLD¿FXV   4      13  Pachylia syces  Callionima pan     4  Callionima calliomenae  1  Callionema inuus  Callionima falcifera   5  10  Nyceryx coffaeae  Nyceryx maxwelli  1  Nyceryx riscus   2  Nyceryx stuarti  2  Perigonia pallida  Perigonia lusca      6  Eupyrrhoglossum sagra    4  Aellopos ceculus     5  Aellopos clavipes  2  Aellopos titan  1  Aellopos fadus  1  Oryba kadeni  1  Oryba achemenides    3  Pachylioides resumens   2  Madoryx oiclus  1  Madoryx plutonius    4  Madoryx bubastus   2 80 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Appendix Table 4 Continued 7KHPRQWKVRIFDSWXUHIRU7ULQLGDG6SKLQJLGDHIURPQRQVXEXUEDQVLWHV

Species May July April Total June March August October January February December November September

44 Hemeroplanes triptolemus     4 45 Phryxus caicus   2  Erinnyis alope       19  Erinnyis lassauxii     4  11 48 Erinnyis impunctata  Erinnyis ello         4   57  Erinnyis oenotrus      7  Erinnyis crameri       11  Erinnyis obscura   2  Isognathus scyron           15 54 Isognathus caricae 4 4 55 Pseudosphinx tetrio  4    10  Aleuron carinata  1  Aleuron chloroptera 58 Unzela japix  1  Enyo lugubris        8  Enyo ocypete     8  Enyo gorgon  Eumorpha anchemolus    4 4 4  18  Eumorpha satellitia         27  Eumorpha obliquus 4  7  Eumorpha megaeacus  1  Eumorpha vitis      5 13  Eumorpha fasciatus     5  Eumorpha phorbas    5     28  Eumorpha capronnieri       9  Eumorpha labruscae      6  Pachygonidia caliginosa  Hyles lineata  Xylophanes anubus      5  26  Xylophanes ceratomioides    4  8   26  Xylophanes chiron   4   5 4  73  Xylophanes loelia    3  Xylophanes neoptolemus   5        4 58  Xylophanes pistacina  2  Xylophanes pluto  5     20  Xylophanes porcus      17  Xylophanes tersa     4   33  Xylophanes thyelia       8  Xylophanes titana    8   16 84 Xylophanes tyndarus 5   8 1XPEHURIUHFRUGV             960 1XPEHURIVSHFLHV         44    75 +DZNPRWKV 6SKLQJLGDH RI7ULQLGDG 81

Annex Table 5. ,QGH[RIVSHFLHVE\QXPHULFDOVHTXHQFHE\JHQXVDQGE\VSHFLHV 1XPHULFVHTXHQFH $OSKDEHWLFDOVHTXHQFHE\JHQXV $OSKDEHWLFDOVHTXHQFHE\VSHFLHV  Adhemarius gannascus  Adhemarius daphne  achemenides, Oryba  Adhemarius daphne  Adhemarius gannascus  albiplaga, Manduca  Adhemarius palmeri  Adhemarius palmeri  alope, Erinnyis 4 Protambulyx eurycles  Aellopos ceculus  anchemolus, Eumorpha 5 Protambulyx strigilis  Aellopos clavipes 8 antaeus, Cocytius  Cocytius duponchel  Aellopos fadus  anubus, Xylophanes  Cocytius lucifer  Aellopos titan  bubastus, Madoryx 8 Cocytius antaeus  Agrius cingulata 45 caicus, Phryxus  Neococytius cluentius  Aleuron carinata  caliginosa, Pachygonidia  Manduca lefeburii  Aleuron chloroptera  calliomenae, Callionema  Manduca albiplaga  Cocytius duponchel  capronnieri. Eumorpha  Manduca franciscae  Cocytius lucifer 54 caricae, Isognathus  Manduca huascara  Callionema inuus  carinata, Aleuron  0DQGXFDÀRUHVWDQ  Callionima calliomenae  ceculus, Aellopos  Manduca rustica  Callionima falcifera  ceratomioides, Xylophanes  Manduca sexta  Callionima pan  chiron, Xylophanes  0DQGXFDGLI¿VVD 8 Cocytius antaeus  chloroptera, Aleuron  Manduca hannibal  Enyo gorgon  cingulata, Agrius  Manduca ochus  Enyo lugubris  clavipes , Aellopos  Agrius cingulata  Enyo ocypete  cluentius, Neococytius  3DFK\OLD¿FXV  Erinnyis alope  coffaeae, Nyceryx  Pachylia syces  Erinnyis crameri  crameri, Erinnyis  Callionima pan  Erinnyis ello  daphne, Adhemarius  Callionima calliomenae 48 Erinnyis impunctata  GLI¿VVD0DQGXFD  Callionema inuus  Erinnyis lassauxii  duponchel, Cocytius  Callionima falcifera  Erinnyis obscura  ello, Erinnyis  Nyceryx coffaeae  Erinnyis oenotrus 4 eurycles, Protambulyx  Nyceryx maxwelli  Eumorpha anchemolus  fadus, Aellopos  Nyceryx riscus  Eumorpha capronnieri  falcifera, Callionema  Nyceryx stuarti  Eumorpha fasciatus  fasciatus, Eumorpha  Perigonia pallida  Eumorpha labruscae  ¿FXV3DFK\OLD  Perigonia lusca  Eumorpha megaeacus  ÀRUHVWDQ0DQGXFD  Eupyrrhoglossum sagra  Eumorpha obliquus  franciscae, Manduca  Aellopos ceculus  Eumorpha phorbas  gannascus, Adhemarius  Aellopos clavipes  Eumorpha satellitia  gorgon, Enyo  Aellopos titan  Eumorpha vitis  hannibal, Manduca  Aellopos fadus  Eupyrrhoglossum sagra  huascara, Manduca  Oryba kadeni 44 Hemeroplanes triptolemus 48 impunctata, Erinnyis  Oryba achemenides  Hyles lineata  inuus, Callionema  Pachylioides resumens 54 Isognathus caricae 58 japix, Unzela  Madoryx oiclus  Isognathus scyron  kadeni, Oryba  Madoryx plutonius  Madoryx bubastus  labruscae, Eumorpha  Madoryx bubastus  Madoryx oiclus  lassauxii, Erinnyis 44 Hemeroplanes triptolemus  Madoryx plutonius  lefeburii, Manduca 45 Phryxus caicus  Manduca albiplaga  lineata, Hyles  Erinnyis alope  0DQGXFDGLI¿VVD  loelia, Xylophanes  Erinnyis lassauxii  0DQGXFDÀRUHVWDQ  lucifer, Cocytius 48 Erinnyis impunctata  Manduca franciscae  lugubris. Enyo  Erinnyis ello  Manduca hannibal  lusca, Perigonia  Erinnyis oenotrus  Manduca huascara  maxwelli, Nyceryx  Erinnyis crameri  Manduca lefeburii  megaeacus, Eumorpha  Erinnyis obscura  Manduca ochus  neoptolemus, Xylophanes  Isognathus scyron  Manduca rustica  obliquus, Eumorpha 54 Isognathus caricae  Manduca sexta  obscura, Erinnyis 55 Pseudosphinx tetrio  Neococytius cluentius  ochus, Manduca  Aleuron carinata  Nyceryx coffaeae  ocypete. Enyo  Aleuron chloroptera  Nyceryx maxwelli  oenotrus, Erinnyis 58 Unzela japix  Nyceryx riscus  oiclus, Madoryx  Enyo lugubris  Nyceryx stuarti  pallida, Perigonia  Enyo ocypete  Oryba achemenides  palmeri, Adhemarius  Enyo gorgon  Oryba kadeni  pan, Callionima  Eumorpha anchemolus  Pachygonidia caliginosa  phorbas, Eumorpha  Eumorpha satellitia  3DFK\OLD¿FXV  pistacina, Xylophanes  Eumorpha obliquus  Pachylia syces  pluto, Xylophanes  Eumorpha megaeacus  Pachylioides resumens  plutonius, Madoryx  Eumorpha vitis  Perigonia lusca  porcus, Xylophanes  Eumorpha fasciatus  Perigonia pallida  resumens, Pachylioides  Eumorpha phorbas 45 Phryxus caicus  riscus, Nyceryx  Eumorpha capronnieri 4 Protambulyx eurycles  rustica, Manduca  Eumorpha labruscae 5 Protambulyx strigilis  sagra, Eupyrrhoglossum  Pachygonidia caliginosa 55 Pseudosphinx tetrio  satellitia, Eumorpha  Hyles lineata 58 Unzela japix  scyron, Isognathus  Xylophanes anubus  Xylophanes anubus  sexta, Manduca  Xylophanes ceratomioides  Xylophanes ceratomioides 5 strigilis, Protambulyx  Xylophanes chiron  Xylophanes chiron  stuarti, Nyceryx  Xylophanes loelia  Xylophanes loelia  syces, Pachylia  Xylophanes neoptolemus  Xylophanes neoptolemus  tersa, Xylophanes  Xylophanes pistacina  Xylophanes pistacina 55 tetrio, Pseudosphinx  Xylophanes pluto  Xylophanes pluto  thyelia, Xylophanes  Xylophanes porcus  Xylophanes porcus  titan, Aellopos  Xylophanes tersa  Xylophanes tersa  titana, Xylophanes  Xylophanes thyelia  Xylophanes thyelia 44 triptolemus, Hemeroplanes  Xylophanes titana  Xylophanes titana 84 tyndarus, Xylophanes 84 Xylophanes tyndarus 84 Xylophanes tyndarus  vitis, Eumorpha Apparent lack of chytrid infection in northeast Tobago’s frogs Robyn Thomson 1, Paul A. Hoskisson 2, Sarah Brozio 2 and J. Roger Downie 1 1. School of Life Sciences, Graham Kerr Building, University of Glasgow, Glasgow G12 8QQ 2. Strathclyde Institute of Pharmacy and Biomedical Sciences, University of Strathclyde, 161 Cathedral Street, Glasgow G4 0RE Corresponding author: [email protected] ABSTRACT Six species of Tobago frogs (including the stream frog Mannophryne olmonae ) from 11 sites across the northeast of the island were tested in 2016 for the presence of the pathogenic chytrid fungus Batrachochytrium dendrobatidis . No chytrid was detected in any of the 176 samples (20-39 frogs per species). These results contrast with a 2006 survey which found chytrid in 25% of the M. olmonae individuals sampled, though none in smaller samples of four other species. Together ZLWKVLPLODUUHVXOWVIURP7ULQLGDGRXU¿QGLQJVLQGLFDWHWKDWWKHLQIHFWLRQPD\KDYHGLHGRXWRUEHDWDYHU\ORZOHYHOLQ Trinidad and Tobago. However, there is a need to sample from the south of Tobago and from two threatened species not yet assessed. Key words: amphibians, Caribbean, chytridiomycosis, Tobago

INTRODUCTION The infectious disease chytridiomycosis whose north-east of the island. At two sites, only Mannophryne aetiological agent is the fungus Batrachochytrium olmonae was sampled. At the other eight sites 1-4 species dendrobatidis  %G  KDV EHHQ LGHQWL¿HG DV RQH FDXVH RI were sampled. Table 1 lists only the species that were WKHZLGHVSUHDGSRSXODWLRQGHFOLQHVDIÀLFWLQJWKH:RUOG¶V sampled at each site, not the complete list of species amphibians (Stuart et al . 2004; Kilpatrick et al . 2010). detected at each site. M. olmonae is a day active frog, so In recent years it has become apparent that amphibian VDPSOLQJRFFXUUHGGXULQJGD\OLJKWKRXUV7KHRWKHU¿YH species vary widely in their susceptibility to the disease, VSHFLHV DUH DFWLYH DQG ZHUH VDPSOHG DW QLJKW 6FLHQWL¿F with some acting as carriers while themselves remaining names follow Frost (2017). asymptomatic, whilst others suffer high mortality rates (Venesky et al . 2014). Sampling methods In Trinidad and Tobago, Alemu et al . (2008,2013) Frogs were caught by hand or with the aid of a small reported the occurrence of Bd infection in both of the hand-net, and were transferred to individual polythene bags. endemic aromobatid species, Mannophryne olmonae 7KHFROOHFWLRQWHDPZHUHDOOWUDLQHGLQIURJLGHQWL¿FDWLRQ (Tobago) and M. trinitatis (Trinidad) although they saw by JRD and had Murphy (1997) available to check any no signs of clinical disease in either species. Greener et al . GRXEWIXOLGHQWL¿FDWLRQV:KHUHSRVVLEOHFK\WULGVDPSOLQJ (2017) and Shepherd et al . (2016) carried out new surveys took place at the capture site and frogs were released once for chytrid in Trinidad, re-sampling M. trinitatis but also all had been swabbed. On occasion, when the number testing other species, both from single-species sites and of frogs captured was very large, or the weather too from multi-species assemblages. Greener et al. found no wet for reliable swabbing, the frogs in their bags were chytrid infected individuals in a sample of 116 M. trinitatis transferred to our base in Charlotteville, swabbed there from six sites, including some sites where Alemu et al. and returned to their capture site next morning. Swabbing (2013) collected chytrid-positive samples. Shepherd et al . was all carried out by RT to ensure uniform technique, and found no chytrid in 245 frogs from a further 15 species followed the standard protocol (Brem et al . 2007), as used sampled across nine widely separated sites, both single and also in Trinidad by Greener et al . (2017) and Shepherd et PXOWLVSHFLHVDVVHPEODJHV,QWKHOLJKWRIWKHVH¿QGLQJVLW al . (2016). Gloves were routinely used for frog handling VHHPHGZRUWKZKLOHWRUHWHVW7REDJR¶VIURJVIRUFK\WULG and discarded after each use, as were the polythene bags, WHQ\HDUVDIWHUWKHSUHYLRXVDVVHVVPHQW:HUHSRUWRQWKH to avoid any cross contamination. In addition, nets were results of these tests here. disinfected with bleach after use. Clinical grade sterile Deltalab swabs were used to sample the skin of each frog. METHODS They were stored in sterile 1ml vials with 0.5 ml ethanol Site and species selection added to preserve the collected DNA, and sealed with a Chytrid swabbing was conducted in Tobago between screw cap. Vials were stored in our base freezer and later June and August 2016. A total of eleven sites were visited transferred to the UK by air in a cool bag. and six species sampled (Table 1; Figure 1), all in the Each site was sampled only once per species, to avoid 82 Lack of chytrid infection in northeast Tobago 83

Fig. 1. Map of northeast Tobago showing the sites sampled for this study. any risk of sampling the same individuals more than lacked extractable DNA, ten were from the Tobago glass once. The exception to this was Dead Bay pond where frog, Hyalinobatrachium orientale (reducing the number ten Leptodactylus validus were sampled one night early of that species that could be tested to 20). During the during the study, and then a further 25 were sampled six ¿HOGZRUNQRIURJVZHUHREVHUYHGZLWKFOLQLFDOV\PSWRPV weeks later; the intention was to test whether there was of chytrid infection. A few frogs did look unhealthy e.g. a any evidence of a change in chytrid occurrence over time, few Boana xerophylla (previously Hypsiboas crepitans ) given the multi-species breeding assemblages at this site. had green growths on their hands and feet, but this is not a symptom normally associated with chytridiomycosis. Sample analysis DNA was extracted using Bioline genomic extraction DISCUSSION kit and standard PCR carried out as described by Shepherd Alemu et al . (2008) carried out their survey during et al . (2016). The positive control DNA for Bd was June-September 2006. They sampled 84 Mannophryne supplied by Professor Andrew Cunningham, Institute of olmonae  IURP ¿YH QRUWKHDVW ULYHUV $UJ\OH %ORRG\ Zoology, London. %D\ 'RFWRU¶V .LQJ¶V %D\ DQG /RXLV G¶2U  DQG WKUHH Northside Road streams, but caught most of these frogs RESULTS  DW'RFWRU¶V$W'RFWRU¶VWKH\IRXQGSRVLWLYH A total of 188 frogs were sampled, with the smallest for chytrid; chytrid was also detected at two other sites species sample size being 27 (Table 2). DNA could not (Argyle and one of the Northside Road streams), giving be extracted from 12 samples, so the total number of  SRVLWLYHV IRU WKH ZKROH VDPSOH 7KH\ DOVR WHVWHG analysed samples was 176. No sample tested positive for four other species: two each of Leptodactylus validus , chytrid. As insurance that we had performed the assay Pristimantis charlottevillensis and Hyalinobatrachium correctly, our negative control showed no DNA and our orientale , and 34 Rhinella marina, all of which were positive controls with low and higher amounts of chytrid negative for chytrid . None of the chytrid-positive frogs DNA both showed positive. Since no Tobago sample showed clinical signs of disease. gave positive results, there was no basis for a comparison Our survey, ten years after that of Alemu et al ., between the two Leptodactylus validus samples collected FRYHUHG VHYHUDO RI WKH VDPH VLWHV $UJ\OH /RXLV G¶2U from Dead Bay pond at different times, so they are all 'RFWRU¶V DQGWKHUHIRUHSURYLGHVDGLUHFWFRPSDULVRQ7KH presented together in Table 2. Of the 12 samples which difference in results could relate to sample size (Alemu 84 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

Table 1. Descriptions of the eleven sites, with the frog species sampled from each. Site names are as on local maps, except where such names are lacking and have been given by University of Glasgow expeditions e.g ‘Mystery’ river. Abbreviations for frog names: OL, Mannophryne olmonae ; VA, Leptodactylus validus ; PU, Engystomops pustulosus ; BX, Boana xerophylla ; CV, Pristimantis charlottevillensis ; HO, Hyalinobatrachium orientale . GPS co-ordinates differed slightly at different locations within each site; for simplicity, we give only one set of co-ordinates for each site. Site name and Species Site characteristics GPS co-ordinates collected Hermitage HO,CV,BX, VA Medium to high vegetation with Heliconia and ferns.Canopy closed N11.31435, W060.57454 where glass frogs were located, and stream running constantly. Elsewhere, canopy open, stream intermittent. Substrate rocks and pebbles. Cambleton CV,VA Medium to high vegetation with Heliconia and some bamboo. N11.31667, W060.55733 Canopy closed, stream intermittent. Substrate: leaf litter. Merchiston OL, BX, CV Medium height vegetation, mostly small plants. Canopy open, stream N11.28638, W060.54179 intermittent. Substrate: leaf litter. Louis d’Or ‘original’ CV High vegetation with Heliconia and ferns at some points, elsewhere, N11.27180, W060.56355 bamboo, ferns and open canopy. Stream constant. Substrate: leaf litter, rocks, pebbles. Louis d’Or ‘new’ OL, CV Medium height vegetation with Heliconia , ferns and small plants. N11.27049, W060.56311 Closed canopy, stream intermittent. Substrate: leaf litter. Main Ridge HO,CV High palms and ferns. Canopy closed, stream constant. Substrate: N11.28667, W060.59545 rocks, silt, mud. ‘Mystery’ OL High ferns and small plants with canopy partly closed. Stream N11.31566, W060.62614 constant. Substrate: leaf litter, rocks, pebbles. Argyle Waterfall OL Low ferns, canopy open. Major river. Substrate: rocks and pebbles. N11.25953, W060.58602 Doctor’s river OL, HO, CV High Heliconia and ferns where glass frogs were found. Elsewhere, N11.31104, W060.53991 smaller plants including low ferns, canopy open, stream constant. Substrate: leaf litter, rocks, pebbles. Dead Bay river HO, CV, VA High Heliconia and ferns where glass frogs were found, canopy open, N11.29070, W060.63354 stream constant. Substrate: leaf litter, rocks, pebbles. Dead Bay pond BX,VA,PU Pond surrounded by low vegetation; open canopy. Edges part N11.29152, W060.63214 concrete, part grass. Bottom of pond muddy with rotting timber.

Table 2. The number of individual frogs of each species sampled at each site. *= no DNA extracted. Species name abbreviations as LQ7DEOHa DGGLWLRQDOVDPSOHZHHNVODWHUWKDQWKH¿UVW Numbers of each species Site HO OL VA PU BX CV Hermitage 6(*1) 0 2 0 1 6(*1) Cambleton 0 0 1 0 0 2 Merchiston 0 18 0 0 5 2 Louis d’Or original 0 0 0 0 0 2 Louis d’Or new 0 2 0 0 0 2 Main Ridge 11(*5) 0 0 0 0 5 Mystery 0 4 0 0 0 0 Argyle Waterfall 0 4 0 0 0 0 Doctor’s 4 3 0 0 0 3 Dead Bay river 9(*4) 0 1 0 0 5 Dead Bay pond 0 0 10(~25) 30(*1) 25 0 TOTALS 30 31 39 30 31 27 Lack of chytrid infection in northeast Tobago 85 et al FDXJKWDW'RFWRU¶VDORQHZKLOHRXU M. olmonae PDSV RI FK\WULG¶V GLVWULEXWLRQ XS WR  &DULEEHDQ VDPSOH ZDV  LQ WRWDO IURP  ¿YH VLWHV  +RZHYHU LI LVODQGV SRVLWLYH IRU FK\WULG ZHUH &XED +LVSDQLROD chytrid was going to take hold in Tobago, it would be (Dominican Republic), Puerto Rico, Dominica and the H[SHFWHGWRVSUHDGLQWKHWHQ\HDUVVLQFHWKH¿UVWVXUYH\ British Virgin Islands, whilst those showing no chytrid and possibly to affect other species. Instead, we found no were Jamaica, Montserrat, Barbados and Grenada. Other chytrid in M. olmonae RU LQ VXEVWDQWLDO VDPSOHV RI ¿YH islands appeared not to have been assessed, although Olson other species. Of these, three ( P. charlottevillensis , B. et al. ¶VGDWDVHWRPLWWHG$OHPX et al ¶V  7REDJRVWXG\ xerophylla and H. orientale ) were found at the same sites Since then, Greenhawk et al . (2017) and Sabino-Pinto et as M. olmonae and therefore might have been expected to al   KDYHFRQ¿UPHGWKHSUHVHQFHRIFK\WULGDWORZ have become infected. Research elsewhere (Scheele et al. levels of prevalence in Puerto Rico and Cuba respectively. 2017) shows that asymptomatic infections in one species ,QFRQWUDVW+XGVRQ et al. (2016) reported the devastating can amplify the harmful effects on a susceptible species effects on the Critically Endangered endemic mountain where the two co-occur. The Dead Bay pond we surveyed chicken ( Leptodactylus fallax ) of the arrival of chytrid is used as a breeding site by several species in very large ¿UVW LQ 'RPLQLFD DQG ODWHU LQ 0RQWVHUUDW 5RGULJXH] numbers ( Trachycephalus typhonius and Dendropsophus Brenes et al. (2016) discussed the importance of studying minutus in addition to those we surveyed: JRD, personal chytrid prevalence in low altitude tropical locations where observations) and would therefore be expected to act as a the infection can be asymptomatic but act as a reservoir place where chytrid, if present, could easily spread. Our for spread to cooler, wetter, often montane habitats conclusion therefore is that chytrid, while present in at ZKHUHPRUWDOLW\RFFXUV2XU¿QGLQJVJLYHVRPHJURXQGV least one species ten years ago, is now absent or at very IRU RSWLPLVP FRQFHUQLQJ 7REDJR¶V IURJV EXW UHJXODU low levels or in very restricted locations. monitoring is advisable. 2QH GH¿FLHQF\ LQ RXU UHVXOWV ZDV WKH ODFN RI '1$ in 12 of our samples. Ten of these cases were from ACKNOWLEDGEMENTS H.orientale (Table 2), suggesting that the failures may :HWKDQNPHPEHUVRIWKH8QLYHUVLW\RI*ODVJRZ relate more to the species than to our technique overall. H[SHGLWLRQIRUDVVLVWDQFHZLWK¿HOGZRUNWKH8QLYHUVLW\RI These are very small delicate frogs and it may be that in *ODVJRZ¶V &KDQFHOORU¶V )XQG *ODVJRZ 1DWXUDO +LVWRU\ some individuals, our swabbing was too gentle to extract 6RFLHW\¶V%ORGZHQ/OR\G%LQQV%HTXHVWDQGVHYHUDORWKHU an adequate sample. WUXVWVIRU¿QDQFLDOVXSSRUW$QJHOD5DPVH\'HSDUWPHQW One of Alemu et al. ¶VFRQFHUQVZDVWKDWWKH\KDGIRXQG RI1DWXUDO5HVRXUFHVDQG(QYLURQPHQW7REDJR+RXVHRI FK\WULGLQRQHRI7REDJR¶VHQGHPLFVSHFLHV$WWKDWWLPH Assembly provided permits for our work. IUCN rated M. olmonae as Critically Endangered, so the discovery of chytrid in the population was particularly REFERENCES worrying. Since then, following work by Lehtinen et al. Alemu,J.B., Cazabon, M.N.E, Dempewolf, L., Hailey, (2016), the conservation status of M. olmonae has been A., Lehtinen, R.M., Mannette, R.P., Naranjit, K.T. VRIWHQHG WR 9XOQHUDEOH :KHQ DGGHG WR RXU ¿QGLQJV RQ and Roach, A.C.J. 2008. Presence of the chytrid fungus chytrid, Lehtinen et al ¶VUHVXOWVSURYLGHDPRUHKRSHIXO Batrachochytrium dendrobatidis in populations of the view of the future of this species. Critically Endangered frog Mannophryne olmonae in %RWK RI WKH LQYHVWLJDWLRQV LQWR FK\WULG¶V VWDWXV LQ 7REDJR:HVW,QGLHV Ecohealth , 5:34-39. 7REDJR¶VIURJVKDYHEHHQFRQGXFWHGLQWKHQRUWKHDVWRI Alemu, J.B., Cazabon-Manette, M.N.E., Cunningham, WKH LVODQG 7KLV LV MXVWL¿DEOH VLQFH WKH QRUWKHDVW LV WKH A.A., Dempewolf, L., Hailey, A., Mannette, R.P., SULQFLSDO ORFDWLRQ RI 7REDJR¶V HQGHPLF DQG WKUHDWHQHG Naranjit, K.T., Perkins, M.W. and Schmidt- VSHFLHV 0XUSK\ +RZHYHUQRWDOORIWKHWKUHDWHQHG Roach, A.C.J. 2013. Presence of the chytrid fungus species (IUCN 2016) have been assessed for chytrid: Batrachochytrium dendrobatidis in a Vulnerable frog in Pristimantis turpinorum is a Tobago endemic with a 7ULQLGDG:HVW,QGLHV Endangered Species R esearch, 20: very restricted range, an IUCN rating of Vulnerable, 131-136. and should be assessed; )OHFWRQRWXV ¿W]JHUDOGL occurs Brem, F., Mendelson III, J.R. and Lips, K.R. 2007. Field- in Tobago, Trinidad and Venezuela, but has an IUCN sampling protocol for Batrachochytrium dendrobatidis rating of Endangered- the problem is that its habits make from living amphibians using alcohol preserved swabs LWH[WUHPHO\GLI¿FXOWWRFDSWXUHLQDGHTXDWHQXPEHUVWR (version 1.0). Conservation International, Arlington, assess properly. Virginia, USA, p.1-10.[On-line] Available at http://www. The occurrence and impact of chytrid in the Caribbean amphibians.org (Accessed May, 2017). has been patchily reported. Olson et al. (2013) provided Frost, D.R.  $PSKLELDQ 6SHFLHV RI WKH :RUOG 86 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018 an online reference, version 6.0 American Museum of Murphy, J.C. 1997. Amphibians and Reptiles of Trinidad 1DWXUDO+LVWRU\1HZ2QOLQH@$YDLODEOHDW and Tobago. Krieger Publishing Company, Malabar, http://research.amnh.org/herpetology/amphibia/index. Florida. html. (Accessed 4 October, 2017) Olson, D.H., Aanensen, D.M., Walker, S.F., Bielby, Greener, M.S., Shepherd, R., Hoskisson, P.A., Asmath, J., Garner, T.W.J., Weaver, G., Group T.B.M. and H. and Downie, J.R. +RZPDQ\7ULQLGDGVWUHDP Fisher, M.C. 2013. Mapping the global emergence of the frogs ( Mannophryne trinitatis ) are there, and should they amphibian chytrid fungus. PLoS ONE, 8: e56802. be regarded as Vulnerable to extinction? Herpetological Rodriguez-Brenes, S., Rodriguez, D., Ibanez, R. and Journal, 27:5-11. Ryan, M.J. 2016. Spread of amphibian chytrid fungus Greenhawk, N., Zlotnik, S., Billy, L.M., Boas, S. and across lowland populations of tungara frogs in Panama. Gabel,S. 2017. Baseline amphibian survey and sampling PLoS ONE , 11: e0155745. RI %G LQ WKH ,FDFR DQG +RUPLJD YDOOH\V 3XHUWR 5LFR Sabino-Pinto,J., Bletz, M.C., Iturriaga, M., Vences, M. Phyllomedusa , 16: 63-69. and Rodriguez, A. 2017. Low infection prevalence of the Hudson, M.A., Young, R.P., D’Urban Jackson, J., amphibian chytrid fungus Batrachochytrium dendrobatidis Orozco ter Wegel, P., Martin, L., James, A., Sulton, (Chytridiomycetes: Rhizophydiales) in Cuba. Amphibia- M., Garcia, G., Griffiths, R.A., Thomas, R., Magin, C., Reptilia , 38:243-249. Bruford, M.W. and Cunningham, A.A. 2016. Dynamics Scheele, B.C., Hunter, D.A, Brannelly, L.A., Skerratt, and genetics of a disease driven species decline to near L.F. and Driscoll, D.A. 5HVHUYRLUKRVWDPSOL¿FDWLRQ extinction: lessons for conservation. 6FLHQWL¿F 5HSRUWV , of disease impact on an endangered amphibian. 6:e30772. Conservation Biology , 31: 592-600. IUCN. 2016. The IUCN red list of threatened species. Shepherd, R., Hoskisson, P.A., and Downie, J.R. 2016. International Union for the Conservation of Nature and $SSDUHQWDEVHQFHRIFK\WULGLQIHFWLRQLQ7ULQLGDG¶VIURJV natural Resources. [On-line] Available at www.iucnredlist. Living World: Journal of the Trinidad and Tobago Field com (Accessed 4 October, 2017) Naturalists’ Club , 2016: 19-22. Kilpatrick, A.M., Briggs, C.J., and Daszak, P. 2010. Stuart, S., Chanson, J.S., Cox, N.A., Young, B.E., The ecology and impact of chytridiomycosis: an emerging Rodrigues, A.S.L., Fishman, D.L. and Waller, R.W. disease of amphibians. Trends in Ecology and Evolution , 2004. Status and trends of amphibian declines and 25: 109-118. extinctions worldwide. Science , 306: 1783-1786. Lehtinen,R.M., Calkins, T.L., Novick,A.M. and Venesky, M.D., Raffel, T.R., McMahon, T.A. and Rohr, McQuigg, J.L. 2016. Reassessing the conservation J.R. 2014. Confronting inconsistencies in the amphibian- status of an island endemic frog . Journal of Herpetology , chytridiomycosis system; implications for disease 50:249-255. management. Biological Reviews , 89: 477-483. Influence of forest disturbance on bat community diversity in northeast Tobago, West Indies Ciara F. Loughrey [email protected]

ABSTRACT Bat assemblages were sampled at three habitats of varied disturbance within northeast Tobago over a 10-week period. Overall 995 individuals, representing 15 species, were caught through the use of mist nets. Community diversity, evenness, abundance, and feeding guild composition across sites were recorded. Our analysis showed lower species diversity and richness within the site experiencing ongoing disturbance. Results from the secondary forest suggest diversity can recover to natural levels if this disturbance ends. However, some species were only located within primary forest, which may be a UHVXOWRIVSHFL¿FHFRORJLFDOUHTXLUHPHQWV

Key words : Chiroptera, mist-netting, diversity

INTRODUCTION Anthropogenic disturbance has reduced the quality, and diversity has been found within deforested areas of size, and number of forest habitats within the Neotropics, tropical forests (Brosset et al. 1996, Fenton et al. 1992). consequently impacting the animal species residing Selective logging in Trinidad was determined to have within them (Chapin III et al. 2000, Gibson et al. 2011). altered feeding guild abundance in comparison to primary Different bat species show different levels of sensitivity IRUHVW &ODUNH et al. 2005), while local-scale fragmentation to environmental change, in some cases leading to of Neotropical forests was found to reduce species richness local extinctions and consequently shifts in community in Panama (Meyer et al. 2008). This paper presents the composition (Fenton et al. 1992, Medellín et al. 2000). results of mist-net sampling of bat communities from three This response to disturbance can differ between feeding sites in northeast Tobago; an urban area, secondary forest guilds, for example gleaning animalivore numbers may and primary forest. GHFUHDVH ZKLOH IUXJLYRUHV EHQH¿W &ODUNH et al. 2005). It is important to assess the effect of disturbance on bat METHODS communities within the tropics as they provide a variety 6XUYH\VZHUHFRQGXFWHGRYHUDWHQZHHNSHULRGIURP of important ecosystem services such as pollination, seed 21 June to 17 August 2016 using mist nets at ground and dispersal, and pest control (Medellín et al. 2000). VXEFDQRS\ OHYHO$ WRWDO RI  QLJKWV RI VDPSOLQJ WRRN The country of Trinidad and Tobago has recorded 68 place at three locations within northeast Tobago, six species of bats, 24 of which are found in Tobago (Gomes sample nights for each site. The sites consisted of an urban and Reid 2015). While studies have previously been carried area experiencing frequent anthropogenic disturbance out in Tobago to assess the bat fauna present, there is (Charlotteville Village, N11.32148, W060.55398) a currently no literature assessing the composition of habitat- secondary forest undergoing succession following previous VSHFL¿FEDWFRPPXQLWLHVRQWKHLVODQG7REDJR¶VQDWXUDO disturbance (Dead Bay River, N11.29115, W060.63352), landscape supports a variety of forest habitats, containing and a primary forest containing the mature climax lower montane, lowland and xerophytic rainforest, semi- vegetation of the island (Main Ridge Forest Reserve, evergreen and deciduous seasonal forest, dry evergreen N11.27832, W060.58253) (Fig.1). Detailed forest maps of IRUHVWDQGIRUHVWHGZHWODQG,QUHFHQWKLVWRU\DÀXFWXDWLQJ 7REDJRDOORZHGDFFXUDWHLGHQWL¿FDWLRQRIIRUHVWW\SHDQG agricultural industry has left secondary forest covering the land cover (Helmer et al. 2012). majority of the landscape, as sites have been left to recover To determine the community composition at each site, and regenerate when plantations were abandoned (Helmer LQGLYLGXDOV ZHUH FDXJKW DQG LGHQWL¿HG WR VSHFLHV OHYHO et al. 2012). Within the centre of the landmass, a smaller WKURXJKWKHXVHRI¿HOGNH\V *RPHVDQG5HLG5HLG area of the primary montane forest remains, protected 2009). Three nylon mist nets were placed at ground level (2 since 1776 as part of the Main Ridge Forest Reserve. [P[P  PRQR¿ODPHQWPPKHLJKWP DQG Small towns surround the forested area, situated along the two nets at sub-canopy level (2 x 9m) to intercept potential coastal road (Helmer et al. 2012, UNESCO). ÀLJKWSDWKVDORQJQDWXUDOFRUULGRUV1HWVZHUHRSHQHGDW The impact of landscape disturbance has been observed sunset for four hours each night, typically from 1830h to to affect multiple aspects of bat diversity and community KDQGFKHFNHGHYHU\PLQXWHV8SRQFDSWXUHHDFK structure within the Neotropics. Decreased species richness individual was weighed using Pesola Spring Balances 87 88 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

chosen against Chao 1 and Chao 2, as both Chao estimators calculated species richness to be lower than our observed species richness. Beta diversity was calculated using the :KLWWDNHU ,QGH[ ȕZ  DVVHVVHG SDLUZLVH RYHU WKH WKUHH VLWHV :KLWWDNHU :KLWWDNHU¶VLQGH[RIEHWDGLYHUVLW\ was used as it is an effective index of species turnover (Koleff et al.    6LPSVRQ¶V 5HFLSURFDO ,QGH[ DQG 6LPSVRQ¶V (YHQQHVV ZHUH XVHG WR HYDOXDWH WKH VSHFLHV diversity and species dominance respectively within each test site (Simpson 1949). Simpsons Index (1/D) ZDV GLYLGHG E\ VSHFLHV ULFKQHVV 6  WR ¿QG 6LPSVRQV Evenness. One-way analysis of variance (ANOVA, F) was conducted on these two measures to determine if the GLIIHUHQFHV DFURVV KDELWDWV ZDV VWDWLVWLFDOO\ VLJQL¿FDQW Power analysis of this ANOVA calculation (f=1.6, p= 0.01, power = 0.9) recommended a minimum of 4.013 Fig. 1. Map of Tobago showing location of study sites (Urban Site: red, Secondary Site: blue, Primary Site: yellow) T VDPSOHVQLJKWVSHUVLWHZKLFKFRQ¿UPHGRXUVL[VDPSOH QLJKWVSHUVLWHDVDGHTXDWH:KHUHVWDWLVWLFDOVLJQL¿FDQFH and forearm size was measured using 0-150mm Dialmax ZDVGHWHFWHGDGGLWLRQDOSRVWKRF7XNH\7HVWVZHUHFDUULHG Vernier Dial Callipers (to the nearest 0.1mm) to assist in out to assess the sites pairwise. Analysis of individual VSHFLHVLGHQWL¿FDWLRQ *RPHVDQG5HLG5HLG  abundance per feeding guild were compared between sites 'DWDZHUHDQDO\VHGXVLQJSURJUDPPH5DQGSDFNDJH XVLQJ3HDUVRQ¶V&KL6TXDUHG7HVW ³YHJDQ´ 2NVDQHQ et al . 2018). Calculation of mist net hours (MNH) assessed the capture rates between sites, and RESULTS 3HDUVRQ¶V&KLVTXDUH7HVWZDVXVHGWRHYDOXDWHVWDWLVWLFDO Sampling Effort: A total of 375.5 mist net hours were VLJQL¿FDQFH 7KH QXPEHU RI LQGLYLGXDOV FDXJKW RI HDFK carried out over 18 sample nights. The overall capture VSHFLHV ZDV UHFRUGHG DEXQGDQFH  :KLWWDNHU¶V 5DQN rate was found to be 2.8 bats net -1 h -1 . Capture rate was Abundance Plots were applied to represent the relative 1.9 bats net -1 h -1 at the urban site and 2.4 bats net -1 h -1 at DEXQGDQFHV SHU VLWH DJDLQVW VSHFLHV UDQN 7KLV DOORZHG the secondary site. The most successful capture rate was comparison between sites, despite differences in size and found at the primary site at 4.1 bats net -1 h -1 (Table 1). species composition (Magurran and McGill 2011). To This difference in individuals caught was shown to be allow improved comparison of species richness between VWDWLVWLFDOO\VLJQL¿FDQW Ȥ S   VLWHV VDPSOH EDVHG DFFXPXODWLRQ FXUYHV ZHUH ¿WWHG IRU Species Composition & Abundance: A total of species richness at each site. An asymptote for abundance 995 bats were sampled, representing 15 species from GDWDZDVHVWLPDWHGXVLQJ¿UVWRUGHU-DFNNQLIHHVWLPDWRUV four families (Table 2). The Phyllostomidae accounted Macgurran and McGill (2011), found that non-parametric for the majority of bats caught (94.7%), followed by PHDVXUHV &KDR  &KDR  -DFNNQLIH  SHUIRUPHG EHWWHU Vespertilionidae (3.3%), then Molossidae (1.5%) and in comparison to parametric measures as more precise, Emballonuridae (0.5%). Individual species counts were HI¿FLHQW DQG OHVV ELDVHG ,Q WKLV VWXG\ -DFNNQLIH ZDV dominated by species from the Phyllostomidae family

Table 1. 6XPPDU\RIWRWDOQXPEHURILQGLYLGXDOVVDPSOHG Q PLVWQHWKRXUVDQGUDWHRIFDSWXUHSHUQHWSHUKRXUDVGH¿QHGE\WKHGLI - ferent nets used for each site. Site Ground Nets Triple High Nets Total

n MNH Bats net -1 h -1 n MNH Bats net -1 h -1 n MNH Bats net -1 h -1

Urban 64 48.5 0.8 158 72 2.2 223 120.5 1.9

Secondary 138 47 2.9 146 72 2 284 119 2.4

Primary 221 44.5 5 267 73.5 3.6 488 118 4.1

Total 424 140 3 572 217.5 2.6 995 357.5 2.8 Bat Diversity in Northeast Tobago 89

- Artibeus jamaicensis (60.3%), Carollia perspicillata out slightly, no clear asymptote was reached (Fig. 3). (15.6%), and Artibeus cinereus (11.8%). All other species Regression analysis for total species richness (Fig. 4) were present at relatively low numbers (2.6% - 0.2%). estimated the asymptote would occur at 8.1 for the urban The urban site contained 46.7% of the total species caught, site, 15.5 for the secondary site, and 14.8 for the primary the secondary site contained 86.7%, and the primary site. This indicates sampling detected close to the total site contained 93.3%. Six species were shared among estimated species for the urban and primary site, while at all habitats sampled ( C. perspicillata, Glossophaga the secondary site 3 - 4 species remained unrecorded. longirostris, Artibeus lituratus, A. jamaicensis, A. cinerus The beta diversity across all sites was 0.36. The lowest and Myotis attenboroughi ). Three species were found beta diversity value was found between the secondary exclusively within the primary site ( Saccopteryx leptura, DQGSULPDU\VLWH ȕZ  IROORZHGE\WKHXUEDQDQG Micronycteris megalotis, and Centurio senex ). One VHFRQGDU\VLWH ȕZ  7KHKLJKHVWEHWDGLYHUVLW\YDOXH individual of the species Anoura geoffroyi was captured ZDVIRXQGEHWZHHQWKHXUEDQDQGSULPDU\VLWH ȕZ   at the secondary site, and one individual at the primary Bat species diversity as calculated through Simpsons Index VLWH:KLWWDNHU UDQN DEXQGDQFH SORWV )LJ   GHSLFW WKH ' VKRZHGDVLJQL¿FDQWGLIIHUHQFHEHWZHHQWKHVWXG\ relative abundance of species within each sample site. At sites (one-way ANOVA, F (2, 15) =15.52, p=0.0002). The all three sample sites, the majority of species are found to XUEDQVLWHVKRZHGVWDWLVWLFDOO\VLJQL¿FDQWGLIIHUHQFHVIURP be rare while a small number of species are abundant in both secondary and primary sites (p=0.0001, p=0.005 high numbers, indicating a low species evenness. respectively). As p>0.05 between the secondary site and Species Richness & Diversity: Species accumulation WKH SULPDU\ VLWH WKHUH ZDV QR VLJQL¿FDQFH GLIIHUHQFH curves achieved a near asymptote for both the urban and EHWZHHQEDWVSHFLHVGLYHUVLW\DWWKHWZRVLWHV1RVLJQL¿FDQW primary site. A slightly lower rate of accumulation is GLIIHUHQFHZDVSUHVHQWIRU6LPSVRQ¶V(YHQQHVVEHWZHHQ present for the secondary site and while the curve levels any of the survey sites (Table 3).

Table 2. Bat species captured within the study, along with their respective family and sub-family, feeding guild, total sample abundance and sample abundance found within each Site. See table 4 for feeding guilds.

Taxon Guild Site Total Family Subfamily Species Urban Secondary Primary Emballonuridae Saccopteryx leptura AI 0 0 5 5 Molossidae Molossus molossus AI 13 2 0 15 Phyllostomidae Carolliinae Carollia perpicillata F 14 52 79 145 Glossophaginae Anoura geoffroyi N 0 1 1 2 Glossophaga longirostris N 1 4 2 7 Phyllostomianae Micronycteris megalotis GA 0 0 2 2 Phyllostomus hastatus O 0 3 6 9 Stenodermatinae Artibeus lituratus F 3 11 8 22 Artibeus jamaicensis F 185 131 284 600 Artibeus cinereus F 6 44 66 116 Chiroderma villosum F 0 9 2 11 Centurio senex F 0 0 2 2 Sturnira lilium F 0 9 17 26 Vespertilionidae Eptesicus brasiliensis AI 0 11 11 22 Myotis nigricans AI 1 7 3 11 Total 223 284 488 995 90 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

Fig. 2. Log relative species abundance for a) urban site, b) secondary site, c) primary site

Fig. 4.  (VWLPDWHGVSHFLHVULFKQHVVXVLQJ¿UVWRUGHUMDFNNQLIH for A. urban site, B. secondary forest site, C. primary forest site. Fig. 3. Sample-based species accumulation curve for each site sampled.

Feeding Guilds: Trophic analysis for number of to three species not detected in either of the other two individuals captured showed frugivores dominated all habitats. Higher bat species turnover occurs between the habitats, representing over 90% of individuals at any site urban site and the primary site, where habitat variation sampled. Aerial insectivores were the next most common lowers the proportion of species shared between two sites guild, averaging over 5% of total number captured. (Estrada and Coates-Estrada 2002). Both the beta diversity Nectarivores, omnivores and gleaning animalivores and alpha diversity values agree with our assumption of a all fell beneath 1% of captures. (Table 4). Gleaning disturbance gradient affecting the habitats, from the urban animalivores and omnivores were represented by just one to secondary site then primary site. species each, Micronycteris megalotis and Phyllostomus One trait that appears to be shared by all bat hastatus respectively (Table 1). In addition, the gleaning assemblages sampled in Tobago is domination of animalivorous guild was only present within the primary community composition by a small number of species, as VLWH1RVWDWLVWLFDOVLJQL¿FDQFHZDVIRXQGIRUGLIIHUHQFHV VHHQ LQ WKH:KLWWDNHU UDQN DEXQGDQFH SORWV DQG VSHFLHV in feeding guild abundance between sample sites. evenness calculations. This is a typical characteristic of island habitats and tropical rainforest communities (Meyer DISCUSSION DQG.DONR6WHYHQVDQG:LOOLJ ,QWKLVVWXG\ The three bat communities sampled were found to A. jamaicensis is the predominant species found at all differ in terms of species abundance, species diversity, VLWHV $V D JHQHUDOLVW IUXJLYRUH FDSDEOH RI XQGHUWDNLQJ and species richness. Both diversity and richness were SURORQJHGÀLJKWV A. jamaicensis is able to exploit human VLJQL¿FDQWO\ORZHULQWKHXUEDQVLWHZKLFKVXIIHUVIURP PRGL¿HG ODQGVFDSHV DYRLGHG E\ PRUH VHQVLWLYH VSHFLHV continuous anthropogenic disturbance. The secondary site (Ortega and Castro-Arellano 2001). was indicated to contain the highest species diversity and The urban site used within our study suffers from light estimated richness. The highest abundance of individuals pollution, sound pollution, human activity and reduced was recorded at the primary site, which was also home vegetation, and subsequently provides fewer ecological Bat Diversity in Northeast Tobago 91 niches (Brosset et al. 1996, Hourigan et al . 2006). The captured within the study, and the resulting seed dispersal low species diversity and richness recorded at this site carried out by various species allows multiple stages of corresponds with other studies noting lower diversity succession to occur. Bats are responsible for the dispersal of LQ GHIRUHVWHG KDELWDWV &ODUNH et al. 2005, Fenton et al. SLRQHHUÀRUDVSHFLHVIROORZLQJKDELWDWGLVWXUEDQFH8SRQ 1992). Four out of the seven species caught within the maturing, this initial vegetation provides more favourable XUEDQ DUHD KDG SUHYLRXVO\ EHHQ LGHQWL¿HG DV WKH ¿UVW conditions for specialised bat species that frequent later species to colonise a disturbed area ( A. jamaicensis , A. VWDJHVRIVXFFHVVLRQXQWLOWKHVHHGVRIFOLPD[ÀRUDFDQ cinereus , C. perspicillata , M. molossus) (Brosset et al. be dispersed (Gomes and Reid 2015). However, as is the 1996). The disturbances affecting this habitat may prevent case with M. megalotis , the absence of C. senex within the reforestation from occurring, thus reducing the ability of secondary site may show conditions are still not suitable other, less resilient, bat species to persist here. for rarer species that require climax vegetation. C. senex The Phyllostominae sub-family of bats, represented has adapted to feed upon mature canopy fruits, and so here by M. megalotis and P. hastatus , show sensitivity is less involved in the reforestation of disturbed areas to disturbance and are often only present in pristine than species such as C. perspicillata and A. jamaicensis landscapes (Bobrowiec and Gribel 2010, Medellín et al. (Gomes and Reid 2015). 2000). M. megalotis was located solely within the primary The interaction of multiple frugivorous species with site. This habitat restriction may be a consequence of their environment can create a more heterogeneous dietary specialisation as a gleaning animalivore, hunting habitat. Species diversity was highest in the secondary arthropods and vertebrates from surfaces in narrow space, site, and our analysis estimates that this site contains the rendering the species unable to adapt to anthropogenic highest number of species. The species accumulation GLVWXUEDQFH &ODUNH et al. 2005). P. hastatus is a large curve (Fig. 3) indicates there may still be a small number omnivorous predator that avoids areas where habitat of species to be discovered given further sampling effort. PRGL¿FDWLRQLVSUHVHQW 6FKXO]H et al. 2000). The presence The primary site contained more of the specialist species, of P. hastatus within the secondary site indicates the but with increased disturbance in the secondary forest, the VLWH KDV UHFRYHUHG WR D VLJQL¿FDQW GHJUHH IURP SUHYLRXV secondary site may be able to support a wider range of disturbance. species. Both forest specialists and generalists, and edge Frugivores accounted for over 90% of individuals DQGRSHQHFRV\VWHPVSHFLHVFDQ¿QGVXLWDEOHKDELWDWKHUH

Table 3. Diversity analysis. Species richness (S), total number of individuals caught, Simpsons Index (1/D, and Simpsons evenness index (1/D/S) between each sample site. Site Number of Species (S) Number of Individuals Simpson Index (1/D) Simpsons Eveness Index 1/D/S) Urban 7 223 1.435 0.205 Secondary 12 285 3.623 0.302 Primary 14 488 1.592 0.185

Table 4. Feedings guilds present at each site, by individuals captured and percentage of total sample

Site Urban Secondary Primary Total

Feeding Guild Number Percent Number Percent Number Percent Number Percent Captured Captured Captured Captured Frugivore 208 93.27% 256 90.14% 458 93.85% 922 92.66%

Nectarivore 1 0.45% 5 1.76% 3 0.61% 9 0.90%

Aerial Insectivore 14 6.28% 20 7.04% 19 3.89% 53 5.33%

Gleaning 0 0.00% 0 0.00% 2 0.41% 2 0.20% Animalivore Omnivore 0 0.00% 3 1.07% 6 1.23% 9 0.90% 92 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

(Castro-Luna et al. 2007a, Connell 1978). , WKDQN WKH WHDP PHPEHUV RI WKH  ([SHGLWLRQ ZKR The primary site showed the highest capture rate, FDUULHGRXWWKH¿HOGZRUN FRQWUDVWLQJ ZLWK UHVXOWV IURP 7ULQLGDG &ODUNH et al. 2005), French Guiana (Brosset et al. 1996), and Mexico REFERENCES (Medellín et al.  7REDJR¶VSULPDU\IRUHVWKDVEHHQ Bobrowiec, P. and Gribel, R . 2010. Effects of different protected since 1776 when the Main Ridge was established secondary vegetation types on bat community composition in as a Forest Reserve, preventing rainforest fragmentation, Central Amazonia, Brazil. Animal Conservation , 13: 204-216. which has affected abundance in other Neotropical sites Brosset, A., Charles-Dominique, P., Cockle, A., (Meyer et al. 2008). The importance of maintaining Cosson, J.F. and Masson, D. 1996. Bat communities conservation efforts and protecting areas such as the Main and deforestation in French Guiana. 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E  0LVW QHWV DUH H[WUHPHO\ HI¿FLHQW IRU VDPSOLQJ Chapin III, F.S., Zavaleta, E.S., Eviner, V.T., Naylor, the Phyllostomidae family, which is the most species-rich R.L., Vitousek, P.M., Reynolds, H.L., Hooper, bat family present in the Neotropics. However, additional D.U., Lavorel, S., Sala, O.E. and Hobbie, S.E. 2000. sampling methods such as acoustic sampling or harp traps Consequences of Changing Biodiversity. Nature , 405: could identify previously undetected species (Ochoa et al. 234-242. 2000) and build a more complete community inventory Clarke, F., Pio, D. and Racey, P. 2005. A comparison for Tobago. The secondary site has an elevation of around of logging systems and bat diversity in the Neotropics. 600m, double the height of both the urban and primary Conservation Biology , 19: 1194-1204. sites. Previous publications in the tropics have found bat Connell, J.H. 1978. Diversity in tropical rainforests and species diversity to typically decrease with increasing coral reefs. Science 199: 1302–1310. HOHYDWLRQ DQG D SHDN ULFKQHVV RFFXUV DW DURXQG P Estrada, A. and Coates-Estrada, R., 2002. Bats in elevation (McCain 2007). We can therefore exclude this continuous forest, forest fragments and in an agricultural factor as an alternative explanation to our results. Future mosaic habitat-island at Los Tuxtlas, Mexico. Biological VWXGLHVPD\VWLOOEHQH¿WIURPVDPSOLQJPXOWLSOHIRUHVWVLWHV Conservation , 103: 237-245. at differing levels of elevation to identify any relationship Gibson, L., Lee, T.M., Koh, L.P., Brook, B.W., between elevation and species diversity within Tobago. Gardner, T.A., Barlow, J., Peres, C.A., Bradshaw, C.J., Tobago has large expanses of secondary forest, and the Laurance, W.F., Lovejoy, T.E. and Sodhi, N.S., 2011. ability of the forest to recover from previous disturbance Primary forests are irreplaceable for sustaining tropical has allowed bat community diversity to recover. As the biodiversity. Nature , 478(7369): 378. majority of bat species within our study are frugivores, a Gomes, G.A. and Reid, F.A. 2015. Bats of Trinidad and suggestion for further research would be to assess the role Tobago: A Field Guide and Natural History, Trinibats . 286p. of bats in forest regeneration on the island. Helmer, E.H., Ruzycki, T.S., Benner, J., Voggesser, S.M., Scobie, B.P., Park, C., Fanning, D.W. and Ramnarine, ACKNOWLEDGEMENTS S. 2012. Detailed maps of tropical forest types are within ,ZRXOGOLNHWRWKDQNWKHIXQGLQJERGLHVDQGVSRQVRUV reach: Forest tree communities for Trinidad and Tobago WKDW VXSSRUWHG WKH 8QLYHUVLW\ RI *ODVJRZ¶V 7REDJR PDSSHGZLWKPXOWLVHDVRQ/DQGVDWDQGPXOWLVHDVRQ¿QH 2016 Expedition, including the University of Glasgow resolution imagery. Forest Ecology and Management , &KDQFHOORU¶V )XQG *ODVJRZ 1DWXUDO +LVWRU\ 6RFLHW\ 279: 147 - 166. DQG WKH *LOFKULVW (GXFDWLRQDO7UXVW , WKDQN P\ DGYLVRU Hourigan, C.L., Johnson, C. and Robson, S.K.A., 2006. Dr. Stewart A. White for his help and guidance during this The structure of a micro-bat community in relation to SURMHFW,WKDQN'U/XNH5RVWDQWDQG0DUN*UHHQHUIRU gradients of environmental variation in a tropical urban SURYLGLQJ WUDLQLQJ DQG H[SHUWLVH , DOVR WKDQN 7ULQLEDWV area. Urban Ecosystems, 9: 67-82. and the Trinidad 2016 Expedition for loaning equipment. Fenton, M., Acharya, L., Audet, D., Hickey, M., Bat Diversity in Northeast Tobago 93

Merriman, C., Obrist, M., Syme, D. and Adkins, B. insectivorous bat diversity in protected areas from northern 1992. Phyllostomid bats (Chiroptera: Phyllostomidae) Venezuela. Acta Chiropterologica , 2: 171-183. as indicators of habitat disruption in the Neotropics. Oksanen, J., Blanchet, F.G, Friendly, M., Kindt, R., Biotropica , 440-446. Legendre, P., McGlinn, D., Koleff, P., Gaston, K.J., and Lennon, J.J. 2003. Minchin, P.R., O’Hara, R.B., Simpson, G.L., Solymos, Measuring beta diversity for presence–absence data. P., Stevens, M.H.H., Szoecs, E., and Wagner, H. 2018. Journal of Animal Ecology 72: 367-382. YHJDQ&RPPXQLW\(FRORJ\3DFNDJH5SDFNDJHYHUVLRQ Magurran, A.E. and McGill, B.J. 2011. Biological 2.5-1. [Online]. Available at https://CRAN.R-project.org/ diversity: frontiers in measurement and assessment, SDFNDJH YHJDQ (Accessed 27 th October 2018) Oxford University Press. 345p. Ortega, J. and Castro-Arellano, I. 2001. Artibeus McCain, C.M., 2007. Could temperature and water jamaicensis . Mammalian species , 662:1-9. availability drive elevational species richness patterns? Reid, F.A. 2009. A Field Guide to the Mammals of A global case study for bats. Global Ecology and Central America and South-East Mexico, Oxford, Oxford biogeography , 16: 1-13. University Press . 346p. Medellín, R.A., Equihua, M. and Amin, M.A. 2000. Bat Schulze, M.D., Seavy, N.E. and Whitacre, D.F. 2000. diversity and abundance as indicators of disturbance in A Comparison of the Phyllostomid Bat Assemblages in Neotropical rainforests. Conservation Biology , 14: 1666-1675. Undisturbed Neotropical Forest and in Forest Fragments Meyer, C.F., Fründ, J., Lizano, W.P. and Kalko, of a Slash-and Burn Farming Mosaic in Petén, Guatemala E.K. 2008. Ecological correlates of vulnerability to 1. Biotropica , 32: 174-184. fragmentation in Neotropical bats. Journal of Applied Simpson, E.H. 1949. Measurement of diversity. Nature , Ecology , 45: 381-391. 163:688. Meyer, C.F. and Kalko, E.K . 2008. Assemblage- Stevens, R.D. and Willig, M.R. , 2002. Geographical level responses of phyllostomid bats to tropical forest ecology at the community level: perspectives on the fragmentation: land-bridge islands as a model system. diversity of New World bats. Ecology , 83:545-560. Journal of Biogeography , 35: 1711-1726. Whittaker, R.H.  9HJHWDWLRQ RI WKH 6LVNL\RX Ochoa, J., O’Farrell, M.J. and Miller, B.W. 2000. mountains, Oregon and California. Ecological Contribution of acoustic methods to the study of monographs , 30, 279-338. Epizoic cyanobacteria associated with harvestmen (Arachnida: Opiliones) from Tobago, West Indies

Virginia A. Young 1, Michael K. Moore 2 and Victor R. Townsend, Jr. 3 1. Department of Biology, Mercer University, Macon, Georgia 31207 USA [email protected] 2. Department of Biology, Mercer University, Macon, Georgia 31207 USA [email protected] 3. Department of Biology, Virginia Wesleyan College, 5817 Wesleyan Drive, Virginia Beach, Virginia 23455, USA [email protected]

ABSTRACT On Tobago, there is a species of sclerosomatid harvestman ( Prionostemma sp.) that serves as a host to a variety of cryp- togams. In the living condition, the exterior of the harvestmen is light blue in appearance, which quickly fades to a bluish- green following preservation in ethanol. Observations of specimens with scanning electron microscopy (SEM) revealed the occurrence of several types of epizoic organisms, including prokaryotic cells (several sizes of bacilli and cocci) and at OHDVWWZRVSHFLHVRIIXQJL0RUHRYHUXVLQJ3&5ZLWKVSHFL¿FSULPHUVZHFRQ¿UPHGWKHSUHVHQFHRIDWOHDVWRQHVSHFLHV of epizoic cyanobacterium occurring on the tegument of the harvestmen. The tegument of the carapace and abdomen of Prionostemma sp. is full of numerous small pits that may facilitate colonisation and survival by epizoic organisms.

Key words &DPRXÀDJH&U\SWRJDP)XQJL6FOHURVRPDWLGDH7HJXPHQW INTRODUCTION Epizoic organisms attach to the external surfaces epizoic cryptogams in different types of forests and also of animals, living as either commensals or mutualists noted several similarities shared by arthropod hosts that are (reviewed by Machado and Vital 2001). Among terrestrial inhabited by cryptogams. In both weevils and harvestmen, vertebrates, examples include lichens that dwell on the shells the surfaces of the exoskeletons feature pits, tubercles and of giant tortoises in the Galapagos Islands (Hendrickson granulations, traits that may facilitate colonisation and and Weber 1964), bryophytes and algae growing on the survival of the epizoites at early life stages. In addition, heads of lizards in Mexico (Gradstein and Equihua 1995) arthropod hosts tend to be slow-moving, survive for and algae occurring on the hairs of Neotropical sloths multiple years and inhabit moist habitats. Machado and (Thompson 1972). Epizoic organisms have also been Vital (2001) also observed that colonising cryptogams observed on terrestrial arthropods, including bryophytes are generally common and widespread taxa, rather than growing upon millipedes in Colombia (Martínez-Torres et specialists, and also have the ability to colonise ephemeral al. 2011), liverworts and lichens that dwell on mantids in habitats and grow quickly. Observations of epizoic Costa Rica (Lücking et al. 2010), and cryptogams and their organisms have generally been made only for harvestmen associated microfaunas (e.g., oribatid mites, rotifers and that occur in moist, tropical environments. nematodes) occurring on weevils in New Guinea (Gressitt In this study, we used scanning electron microscopy and Sedlacek 1967; Gressitt et al. 1968). Among arachnids, and molecular techniques to investigate further the in particular, interactions involving epizoic organisms are relationship between epizoic organisms on the carapace known to occur in harvestmen (Order Opiliones), which DQGDEGRPHQRIDQXQLGHQWL¿HGVSHFLHVRIVFOHURVRPDWLG may carry cyanobacteria, liverworts, and nonpathogenic KDUYHVWPDQ IURP 7REDJR :HVW ,QGLHV 6SHFL¿FDOO\ ZH fungi (Machado et al. 2000; Machado and Vital 2001; used SEM to survey the diversity of microorganisms on Proud et al. 2012; Townsend et al. 2012). Only three species the external surface of the exoskeleton and molecular of harvestmen representing two families, Gonyleptidae WHFKQLTXHV WR FRQ¿UP WKH SUHVHQFH RI F\DQREDFWHULD (suborder Laniatores) and Sclerosomatidae (suborder among the observed types of cryptogams. Eupnoi), have been observed to have relationships with epizoic cyanobacteria or bryophytes (Machado and Vital METHODS 2001; Proud et al. 2012; Townsend et al. 2012). $GXOW VFOHURVRPDWLG KDUYHVWPHQ LGHQWL¿HG DV Machado and Vital (2001) noted variation in the Prionostemma sp. 6 by Townsend et al. (2012), were colonisation frequencies of gonyleptid harvestmen by collected by hand from the leaf litter and vegetation along 94 Epizoic Cyanobacteria Associated with Harvestmen 95

the trail connecting the Speyside Overlook to Pigeon Peak RESULTS from 10-14 August 2010 in Tobago (Townsend et al. 2012). The tegument of the carapace and abdomen of The primary habitat in this area (11N 17.845 60W 32.934 Prionostemma sp. 6 are covered in numerous small pits at 405 m in elevation) is lower montane rainforest, and (Figs. 1, 2A-B). Using SEM, we observed that many of Prionostemma sp. 6 is one of the most commonly species, WKHVHSLWVZHUH¿OOHGZLWKDPL[WXUHRIVRLOSDUWLFOHVDQG with individuals often observed moving across the forest other detritus (Fig. 2C). In addition, we observed the ÀRRUDWQLJKW 7RZQVHQG et al. 2012). In contrast to their occurrence of prokaryotic organisms (Fig. 2C) in many of syntopic congeners, the individuals of Prionostemma sp. 6 the pits on the carapace and abdomen. Bacteria included with epizoic cyanobacteria that we observed had a distinct several types of cocci (Fig. 2D) and bacilli (Fig. 2D-F). light denim blue colour on their dorsum that faded to We also observed the occurrence of larger, eukaryotic cells bluish-green following preservation in 70% ethanol. including leaf-shaped structures (Fig. 2E) that resemble We prepared multiple adults (n = 5) for SEM by the thalli of Laboulbeniales fungi as well as those that dehydrating them in an ethanol ladder and chemically may represent fungal hyphae (Fig. 2F). The results of our drying them with hexamethyldisilazane. Individuals were DQDO\VLV RI WKH VXUIDFH ¿OPV WKDW ZH UHPRYHG IURP WKH mounted on aluminium stubs using carbon adhesive strips, harvestmen revealed the presence of a distinct band of 700 sputter coated with gold for 2 min and examined with a EDVHSDLUVDSRVLWLYHUHVXOWWKDWFRQ¿UPVWKHSUHVHQFHRI Hitachi S-3400 VP SEM at accelerating voltages of 5-15 cyanobacteria (Fig. 3). kV in the SEM laboratory on the campus of Virginia Wesleyan University. Voucher specimens are deposited in the American Museum of Natural History (AMNH) arachnological collection. Several techniques were used to remove cyanobacterial cells from the preserved individuals in order to isolate the JHQRPLF'1$IURPWKHEDFWHULD6XUIDFH¿OPVRIHSL]RLF cyanobacteria were removed from the harvestmen by applying a piece of tape to their body, and then either scraping or washing the tape into a test tube (following the protocol described by Proud et al. 2012). In addition to the scotch tape technique, cyanobacterial cells were also scraped off the harvestmen into a 70% ethanol solution and entire bodies of the harvestmen were added to a 70% ethanol solution and agitated with forceps and scissors to dislodge the bacterial cells. Dislodged cells from these preparations were pelleted in a microcentrifuge tube at 13,000 rpm for 2 min. DNA was isolated using the GE Healthcare Illustra Bacteria Genomic Mini Spin Kit   IROORZLQJ 3URWRFRO  IRU SXUL¿FDWLRQ of genomic DNA from gram-positive bacteria. The cyanobacteria Oscillatoria (Carolina Biological Supply) was used as a positive control, as it contains the phycocyanin operon from which the IGS primers can amplify a ~700 base pair PCR product. Both an IGS forward primer (5’-GGCTGCTTGTTTACGCGACA-3’) and an IGS reverse primer (5’-CCAGTACCACCAGCAACTAA-3’) were used to amplify the IGS with PureTaq RTG PCR beads (GE Healthcare; Proud et al. 2012). The PCR Fig. 1. Photograph of the dorsal habitus of the sclerosomatid o settings were: 94 C for 5 min followed by 40 cycles of harvestman Prionostemma sp. 6 with bluish-green cyanobacteria 94 o C for 20 seconds, 60 o C for 30 seconds, and 72 o C for 1 on the dorsal surface. a = abdomen; c = carapace; cx = coxa; e min. PCR products were analysed by gel electrophoresis. = eye mound (ocularium); t = free tergite. Scale bars = 1.5 mm. 96 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE

Fig. 2. SEM micrographs showing the mixture of detritus and microorganisms on the tegument of the sclerosomatid harvestman Prionostemma sp. 6. Lateral (A) and anterior (B) views of the dorsal scutum revealing the density and distribution of pits on the tegument. C). Detritus from a pit on the posterior region of the body. D) Arrows indicate bacteria (bacilli and cocci) from a pit on the FDUDSDFH( %DFWHULD DUURZV DQGDIXQJDOWKDOOXV W IURPDSLW) /DUJHHXNDU\RWLFFHOOV HF ZLWKVPDOOHUSURNDU\RWLFFHOOV DUURZV  6FDOHEDUV —PIRU$±%DQG—PIRU&±) Epizoic Cyanobacteria Associated with Harvestmen 97

colonisation by epizoic organisms. One striking difference between the cyanobacteria observed by Machado and Vital (2001) and Proud et al. (2012) and the cyanobacteria that occurs on the tegument of the harvestman studied here is colour. The specimens that we studied were not green, but rather a distinct blue. We believe that this difference LQ FRORUDWLRQ SUREDEO\ UHÀHFWV LQWHUVSHFL¿F YDULDWLRQ LQ the pigments used by different species of cyanobacteria. In addition, the epizoic cyanobacteria observed by Proud et al. (2012) were generally similar in size to those that we observed on the specimens studied here. However, the epizoites that Proud et al. (2012) observed were aggregated in dense patches across most of the dorsal surface of the carapace and abdomen. In contrast, the epizoic cyanobacteria that we observed here occurred generally in DWKLQ¿OPZLWKWKHPRVWRIWKHVPDOOSURNDU\RWLFFHOOV occurring predominantly as dense clusters in the pits on Fig. 3. *HOVKRZLQJ3&5SURGXFWVIROORZLQJDPSOL¿FDWLRQRIWKH intergenic sequence of the PC operon within cyanobacteria. the tegument of the host. '1$ IURP VFUDSLQJ WDSH DSSOLHG WR WKH ERG\ RI DQ LQGLYLGXDO The nature of the epizoic relationship that exists of Prionostemma  VS  /DQH %  DQG '1$ UHPRYHG IURP between cryptogams and harvestmen (i.e., commensalism washing the tape applied to the harvestmen body (Lane C) were or mutualism) remains to be empirically investigated. FRPSDUHG ZLWK SRVLWLYH FRQWURO '1$ IURP Oscillatoria (Lane The cyanobacteria on the dorsum of the harvestmen are ' DQGDQHJDWLYHFRQWURORIQR'1$ /DQH( $OOODQHVZHUH FRPSDUHG ZLWK WKH  EDVH SDLU '1$ /DGGHU 1HZ (QJODQG photosynthetic and relatively small. They are probably %LR/DEV16 7KHVLPLODUEDQGVRQ/DQH%DQG/DQH' unable to penetrate the cuticle of the host and, thus, are DWWKHEDVHSDLUUHJLRQ LQGLFDWHGE\DUURZ FRQ¿UPWKDWWKH FHUWDLQO\QRWSDUDVLWLF2QWKHEDVHVRIRXU¿HOGREVHUYDWLRQV cells collected from the tegument of the Prionostemma sp. 6 of harvestmen with cryptogams, the hosts do not appear to were cyanobacteria. be encumbered by the biomass of epizoic organisms that DISCUSSION they carry. To the contrary, our observations indicate that 2XU FRQ¿UPDWLRQ RI F\DQREDFWHULD OLYLQJ RQ WKH individuals with epizoic organisms move normally and are dorsal tegument of the carapace and abdomen of adults not lethargic. of the harvestmen Prionostemma sp. represents only the From the perspective of the epizoic organisms, the third documented case of a epizoic interaction between body of the host is an unusual microenvironment that photosynthetic prokaryotes and harvestmen. Interestingly, supports few, if any, potential competitors for light or all three instances involve host species that live in moist other limiting resources. Thus, monotypic colonies of Neotropical environments (Fig. 4). In addition, the species epizoic cyanobacteria can thrive on the tegument of their of harvestmen serving as hosts have a tegument dominated KRVWV+RZHYHUWKHEHQH¿WLIDQ\WRWKHKRVWLVQRWDV by small pits or tubercles, providing suitable habitats for clear. Harvestmen do not appear to be capable of feeding

Fig. 4. $UURZVLQGLFDWHWKHWKUHHORFDWLRQVIURPZKLFKKDUYHVWPHQDVVRFLDWHGZLWKF\DQREDFWHULDKDYHEHHQFDSWXUHG0DSPRGL¿HG from Olson et al.  98 /LYLQJ:RUOG-7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE upon epizoic organisms that live on their dorsum. While Gressitt, J.L. and Sedlacek, J. 1967. Papuan weevil genus harvestmen pass their distal leg segments between the Gymnopholus : Supplement and further studies in epizoic chelicerae in a behaviour known as “grooming”, they symbiosis. 3DFL¿F,QVHFWV , 9: 481-500. simply cannot reach the surfaces of the dorsal scutum nor Grether, G.F., and Donaldson, Z.R. 2007. Communal do they have the anatomy necessary (i.e., chelate tarsi) to roost site selection in a Neotropicalharvestman: habitat remove epizoic organisms from the dorsal surfaces of their limitation vs. tradition. Ethology, 113: 290–300. bodies. Hendrickson, J.R. and Weber, W.A. 1964. Lichens on Gressitt et al. (1968) hypothesised that the presence Galapagos giant tortoises. Science , 144: 1463. of mosses on the dorsal surfaces of weevils in New Lücking, R., Mata-Lorenzen, J. and Dauphin L.G. 2010. Guinea conferred crypsis and thus enabled hosts to avoid Epizoic liverworts, lichens and fungi growing on Costa predators, especially diurnal predators that rely upon visual Rican shield mantis (Mantodea: Choeradodis ). Studies on FXHVWR¿QGWKHLUSUH\+DELWDWXVHDQGFLUFDGLDQDFWLYLW\ Neotropical Fauna and Environment , 45: 175-186. patterns of several species of Neotropical Prionostemma Machado, G., Raimundo, R.L.G. and Oliveira, P.S. spp. have been studied (Donaldson and Grether 2007; 2000. Daily activity schedule, gregariousness, and defen- Grether and Donaldson 2007; Wade et al. 2011). In these sive behaviour in the Neotropical harvestman Goniosoma harvestmen, adults generally are more active in the leaf longipes (Arachnida: Opiliones: Gonyleptidae). Journal litter at night, spending the day, occupying perches on tree of Natural History , 34: 587-596. trunks, buttresses and the exposed surfaces of leaves in Machado, G., and Vital. D.M. 2001. On the occur- the understory. If host species with epizoic cyanobacteria rence of epizoic algae and liverworts on the harvestmen exhibit similar habits, it may be that individuals with Neosadocus aff. variabilis (Opiliones: Gonyleptidae). FU\SWRJDPVKDYHEHWWHUFDPRXÀDJHDQGDUHOHVVREYLRXVWR Biotropica, 33: 535-538. visually oriented predators. Unfortunately, the behavioural Martínez-Torres, S.D., Flórez Daza, A.E. and Linares- ecology of Prionostemma p. 6 or that of its congeners in Castillo, E.L. 2011. Meeting between kingdoms: Discovery Costa Rica has not yet been examined. of a close association between Diplopoda and Bryophyta in a WUDQVLWLRQDO$QGHDQ3DFL¿FIRUHVWLQ&RORPELD ,QWHUQDWLRQDO ACKNOWLEDGEMENTS Journal of Myriapodology , 6: 29-36. We thank S. Broadbridge (Caribbean Discovery Tours) Olson, D.M., Dinerstein, E., Wikramanayake, E.E., IRU KHOSLQJ ZLWK WKH ¿HOGZRUN DQG IRU DVVLVWDQFH ZLWK Burgess, N.D., Powell, G.V.N., Underwood, E.C., ground transportation. We are grateful to R. Shirley, S. D’amico, J.A., Itoua, I., Strand, H.E., Morrison, 'XQQDQG$.D\HIRUDVVLVWDQFHZLWK¿HOGFROOHFWLRQVDQG J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, molecular work and T. Pesacreta and D. Proud (University of Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P., and Louisiana at Lafayette) for helping with the scanning electron Kassem, K.R. 2001. Terrestrial ecoregions of the world: microscopy. We thank G. Machado and an anonymous A new map of life on earth: A new global map of terrestrial reviewer for constructive comments on an earlier version ecoregions provides an innovative tool for conserving bio- of our manuscript. This research was supported by a faculty diversity, BioScience , 51: 933-938. development grant from the CLA at Mercer University Proud, D.N., Wade R.R., Rock, P., Townsend, VR. (MKM). We also thank the Forestry Division of the Ministry Jr., and Jiménez Chavez, D. 2012. Epizoic cyano- of Agriculture, Land and Marine Resources of Trinidad and bacteria associated with a Neotropical harvestman Tobago. All specimens were legally collected and exported (Opiliones: Sclerosomatidae) from Costa Rica. Journal of under permit number 001414 (to MKM in 2010). Arachnology, 40: 259-261. Thompson, R.H. 1972. Algae from the hair of the sloth REFERENCES Bradypus . Journal of Phycology , 8: 8. Donaldson, Z.R. and Grether, G.F. 2007. Tradition with- Townsend, V.R. Jr., Moore, M.K., Proud, D.N. and out social learning: Scent-mark based communal roost for- Young, V.A. 2012. A preliminary survey of the Harvestmen mation in a Neotropical harvestman ( Prionostemma sp.). (Arachnida: Opiliones) of Tobago, West Indies . Living Behavioral Ecology and Sociobiology, 61: 801–809. World, Journal of the Field Naturalist Club of Trinidad Gradstein, S.R. and Equihua, C. 1995. An epizoic bryo- and Tobago, 2012: 41-53. phyte and algae growing on the lizard Corythophanes cris- Wade, R.R., Loaiza Phillips, E., Townsend, V.R. Jr. tatus in Mexican rain forest. Biotropica , 27: 265-268. and Proud, D.N. 2011. Activity patterns of two spe- Gressitt, J.L., Samuelson, G.A. and Vitt, D.H. 1968. cies of Neotropical harvestmen (Arachnida, Opiliones) Moss growing on living Papuan moss-forest weevils. from Costa Rica. Annals of the Entomological Society of Nature , 217: 765-767. America, 104: 1360-1366. Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018 99 Nature Notes Is this the caterpillar of Arsenura beebei (Lepidoptera: Saturniidae)? We report here on a very distinctive caterpillar and width, and in the final instar the subdorsal scoli are absent suggest its likely identity. M.K. and Dr Feroze Omardeen (Lampe 2010). The Central American population, now encountered it on 11 March 2018 along Martinez Trace, known as A. arianae Brechlin and Meister, has the head just north of Brasso Seco in the Northern Range of and anal segment brown and a pale, broad, diffuse lateral Trinidad, West Indies. The approximate coordinates were: line, linked by irregularly diffuse pale transverse bands 10°44’00.00” N, 61°18’08.08” W. The caterpillar was (Costa et al. 2001, Janzen and Hallwachs 2018). Both stationary on a rotting leaf stalk of Cecropia peltata L. these reports are of gregarious species, although it is not (Urticaceae) on the ground, so there was no evidence as out of the question that the apparently solitary blue cater- to what the food plant was. The caterpillar was striking. pillar had strayed from a social group and/or was looking It was black with 10 bright blue bands and 7.5-8.0cm in for a pupation site. We note that there are very few records length (Fig. 1). of A. armida from Trinidad, all from the southern part of Although the caterpillar was not reared, we can deduce the island (Kaye and Lamont 1927), and there has been its identity with some confidence. Its large size indicates only one record since 1918 (M.J.W. Cock unpublished). an uncommonly large adult moth. The cuticle surface is The caterpillar of A. beebei has not previously been smooth apart from small subdorsal scoli on the three tho- reported, as far as we are aware. In contrast to A. armida, racic segments and segments 8–10 of the abdomen, and A. beebei has been regularly collected in the Northern it has very large, conspicuous spiracles. These charac- Range of Trinidad, with more than 30 records extending ters suffice to place it in only one genus among the large from Port of Spain to Toco (M.J.W. Cock unpublished) moths recorded from Trinidad: Arsenura (Saturniidae, (Fig. 2). Given this geographic pattern and the lack of any Arsenurinae) (Lampe 2010, Janzen and Hallwachs 2018). close resemblance of the specimen in question to known The young caterpillars of Arsenura spp. have long, thin caterpillars of the A. armida complex, we can consider it scoli, often 2–3 times the body width, which become rela- very likely that the caterpillar of A. beebei is now known tively shorter as the caterpillars grow until they resemble and is strikingly distinct in colouration. those seen in Fig. 1, or are absent altogether (Lampe 2010, Janzen and Hallwachs 2018). Two species of Arsenura are known from Trinidad: A. armida (Cramer) and A. beebei (Fleming) (Kaye and Lamont 1927, Cock 2003). Arsenura armida has been treated as a widespread polytypic species (Lemaire 1980), but it is now clear that it is more appropriate to treat it as a complex of species, most of which is yet to be worked out (e.g. Brechlin and Meister 2010). The caterpillars of at least some populations of this complex are known. When mature, caterpillars of a population from Carabobo Province, northern Venezuela, are dark brown-black with irregular yellow-brown bands, strongest on abdomi- Fig. 2. Arsenura beebei male from high in the central part of nal segments 1–7, and no wider than 1/3 of the segment the Northern Range (Morne Bleu Textel installation, 20 January 2007). Wingspan 14 -15 cm. Photo by J. Muddeman (Spainbirds Nature Tours, www.spainbirds.com). REFERENCES Brechlin, R. and Meister, F. 2010. Zwei neue Arten der Gattung Arsenura Duncan, 1841 (Lepidoptera: Saturniidae). Entomo-Satsphingia, 3(4): 39-43. Cock, M.J.W. 2003. On the number of species of moths (Lepidoptera) in Trinidad and Tobago. Living World, Journal of the Trinidad and Tobago Field Naturalists’ Fig. 1. Final instar caterpillar of presumed Arsenura beebei. Club, 2003: 49-58. 100 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

Costa, J.T., Fitzgerald, T.D. and Janzen, D.H. 2001. Trail- Lampe, R.E.J. 2010. Saturniidae of the World. following behavior and natural history of the social caterpillar Pfauenspinner der Welt. München, Germany: Verlag Dr. Arsenura armlda in Costa Rica (Lepidoptera: Saturniidae: Friedrich Pfeil. 368 p., 336 pl. Arsenurinae). Tropical Lepidoptera, 12(1-2): 17-23. Lemaire, C. 1980. Les Attacidae americains. The Janzen, D. H. and Hallwachs, W. 2018. Dynamic data- Attacidae of America (=Saturniidae) Arsenurinae. C. base for an inventory of the macrocaterpillar fauna, and Lemaire, Neuilly-sur-Seine, France: 199 p., 76 pl. its food plants and parasitoids, of Area de Conservacion Guanacaste (ACG), northwestern Costa Rica. [Online]. Available at http://janzen.sas.upenn.edu (Accessed: 21 Matt Kelly March 2018) [email protected] Kaye, W.J. and Lamont, N. 1927. A catalogue of the Trinidad Lepidoptera: Heterocera (Moths). Memoirs of the Department Matthew J.W. Cock of Agriculture of Trinidad and Tobago, 3: 144 p. [email protected]; [email protected]

The caterpillar of Rothschildia vanschaycki (Lepidoptera, Saturniidae), a little known silk moth from Trinidad, W.I.

On 9 March 2018, one of us (DSH) found and photo- published images of Saturniidae caterpillars (e.g. Lampe 2010, graphed a large, strikingly colourful caterpillar (Fig. 1) at Janzen & Hallwachs 2018) showed that Rothschildia is the Guapo Beach, near Point Fortin (approximately 10.19N, only genus from Trinidad which shows this configuration of 61.67W), on a species of mangrove growing on land. The short-spined, small subdorsal, dorsolateral and lateral scoli caterpillar was large, at least 6 cm long so probably in the on all thoracic and abdominal segments. Rothschildia is a final instar. It was stationary on a stem about 1.5m off genus of New World atlas moths, with at least 25 species the ground, and we presume that it was on its food plant distributed primarily in the Neotropical Region (Lemaire since there were few other potential food plants nearby, 1978, 1996). In recent years, several new species have and it wasn’t actively searching for a pupation site. Based been described, and some subspecies have been revised to on this image, the food plant is either white mangrove species (e.g. Brechlin and Meister 2012). There are only (Laguncularia racemosa (L.) Gaertn. f., Combretaceae) or three species of Rothschildia known from Trinidad: R. red mangrove (Rhizophora mangle L., Rhizophoraceae), erycina erycina (Shaw), R. aurota aurota (Cramer) and R. but key diagnostic features are not visible (Y. Baksh- vanschaycki Brechlin and Meister (Kaye & Lamont 1927, Comeau pers. comm.). Brechlin and Meister 2012, M.J.W. Cock unpublished data). The size and arrangement of spined scoli indicate that it The mature caterpillar of R. erycina erycina is black with is a species of Saturniidae. Comparison of the image with white bands and partial orange-red bands, and that of R. aurota aurota is yellow-green dorsally, blue green ventrally, with a pale lateral line, small red scoli, red spiracles and no bands (Lampe 2010). As the specimen in question is very different from both of these, we infer that it is the hitherto unknown caterpillar of R. vanschaycki. Caterpillars of some subspecies of the closely related R. lebeau (Guerin- Méneville) are known (Lampe 2010, Janzen & Hallwachs 2018). Lemaire (1978, 1996) treated the widespread R. lebeau as having six subspecies, some of which are now treated as valid species (Brechlin and Meister 2012). Two subspecies might be found in Trinidad: R. lebeau lebeau is found in northern Venezuela to the west of Trinidad, while R. lebeau amacurensis Lemaire is known from the Orinoco Delta to the south of Trinidad, with one doubtful record from Trinidad (Lemaire 1978) which may prove to be the recently described R. vanschaycki. The caterpillar Fig. 1. Presumed final instar caterpillar of Rothschildia of the latter is unknown, but that of R. lebeau lebeau from vanschaycki. Carabobo Province, northern Venezuela, is illustrated by Nature Notes 101

/DPSH SODWH ,QWKHSHQXOWLPDWHLQVWDUWKHERG\ $OWKRXJK ZH DUH FRQ¿GHQW RI RXU LGHQWL¿FDWLRQ RI is fairly uniformly green darker ventrally, with small red this striking caterpillar, it would be desirable to locate scoli, yellow-brown spiracles and bands of grey (anteriorly) DGGLWLRQDOVSHFLPHQVWRUHDUWKURXJKWRFRQ¿UPWKLVDQG and white (posteriorly) on the dorsal and lateral parts of the WKHIRRGSODQW,WVGLVWLQFWLYHQHVVVXSSRUWV%UHFKOLQDQG DEGRPLQDOVHJPHQWV7KH¿QDOLQVWDULVVLPLODUEXWWKHUHG Meister’s (2012) decision to treat R. vanschaycki as a VFROLDUHPLVVLQJ,QVRPHFKDUDFWHUVWKH*XDSR%HDFK separate species rather than as a subspecies of R. lebeau . caterpillar is close to those of R. lebeau lebeau : the rather uniform green ground colour, small red scoli, absence of a REFERENCES lateral line and the presence of transverse bands. However, Brechlin, R. and Meister, F. 2012. Neue Taxa der Gattung LWDOVRGLIIHUVVLJQL¿FDQWO\DVWKHWUDQVYHUVHEDQGVRIWKH Rothschildia Grote, 1896 (Lepidoptera: Saturniidae). abdomen are much more striking, being white anteriorly in (QWRPR6DWVSKLQJLD± the dorsal half, red posteriorly from the subdorsal scolus Janzen, D. H. and Hallwachs, W. 2018. Dynamic data- on one side dorsally to the other side, and black from the base for an inventory of the macrocaterpillar fauna, and subdorsal scolus to the bottom of the prolegs, which have its food plants and parasitoids, of Area de Conservacion a red spot laterally, and the spiracles in the black band are Guanacaste (ACG), northwestern Costa Rica. [Online]. concolourous. Unfortunately the head of the Guapo Beach Available at http://janzen.sas.upenn.edu (Accessed: 14 caterpillar is not visible in the photo. March 2018). Rothschildia vanschaycki was recently described from Kaye, W.J. and Lamont, N. 1927. A catalogue of the Trinidad based on a single specimen collected at Port of Trinidad Lepidoptera: Heterocera (Moths). Memoirs of the Spain (Brechlin and Meister 2012). MJWC had been Department of Agriculture of Trinidad and Tobago, 3: 144 p. DZDUHRIWKLVDVDUDUHVSHFLHVLQ7ULQLGDGEXWLGHQWL¿HG Lampe, R.E.J. 2010. Saturniidae of the World. Pfauenspinner it as R. lebeau ,QDGGLWLRQWRWKHW\SHVSHFLPHQWKHUH der Welt. Verlag Dr. Friedrich Pfeil, 368 p, 336 plates. are two further specimens known to us from Trinidad: a Lemaire, C. 1978. Les Attacidae americains. The male collected at Valpark Shopping Plaza, Valsayn South Attacidae of America (=Saturniidae) Attacinae. Neuilly- (10.633N 61.418E, 26 June 1978, M.J.W. Cock) and a sur-Seine, France: C. Lemaire. 238 p., 49 pl. female from ‘Trinidad’ (no other data) in the Natural Lemaire, C. 6DWXUQLLGDHS± In : Heppner, History Museum, London. Further, KS has photographed J.B. ed. Atlas of Neotropical Lepidoptera. Checklist: an adult male about 3km from Valsayn South at the Caroni 3DUW % 'UHSDQRLGHD ± %RPE\FRLGHD  6SKLQJRLGHD Swamp Visitor Centre, on the eastern side of Caroni Gainesville, Florida: Association for Tropical Lepidoptera 6ZDPS 1 : 1RYHPEHU  )LJ   6FLHQWL¿F3XEOLVKHUVS and at least three images of males from the same area have rainingrainers [Deo, R.] 2017. [Online]. Available at https:// been posted on the internet: late June 2011 (What’s that www.inaturalist.org/observations/11377399 (Accessed 14 Bug? 2011), 30 May 2016 (Rutherford 2016), 4 November March 2018)  UDLQLQJUDLQHUV    ,W LV D UDUHO\ HQFRXQWHUHG Rutherford, M.G. 2016. [Online]. Available at https:// species in Trinidad, and the places where it has been seen www.inaturalist.org/observations/12069611 (Accessed 14 suggest an association with coastal swamps, which this March 2018) provisional mangrove food plant association supports. What’s that Bug? 2011. Giant Silkmoth from Trinidad: Rothschildia vanschaycki . [Online]. Available at https:// www.whatsthatbug.com/2011/07/10/giant-silkmoth-from- trinidad-rothschildia-lebeau-amacurensis/ (Accessed 14 March 2018)

David S. Huggins [email protected]

Kris Sookdeo [email protected]

Fig. 2 . Living Rothschildia vanschaycki male, &DURQL6ZDPS Matthew J.W. Cock Visitor Centre, 2 November 2014 (K. Sookdeo). [email protected]; [email protected] 102 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

First report of the freshwater mussel Mytilopsis leucophaeata (Family Dreissenidae) in Trinidad W.I. and a new locality record of the native Anodontites crispata (Family Unionidae)

Freshwater mussels have been documented within ballast water discharge. Trinidad since the 1800s (Guppy 1866) with two species Whilst there is no documentation of this species of the Unionidae family; Anadontitis crispata (Bruguiere forming dense colonies that out compete native bivalves 1792) and Mycetopoda VSLGHQWL¿HG,QWKLVQRWHZHUHFRUG in the other Caribbean countries, monitoring of the spread a new location for A. crispata and a new species record of this species and any negative impact on other species Mytilopsis leucophaeata (Conrad 1831) for Trinidad. in Trinidad, is important. Mytilopsis leucophaeata could During a survey of freshwater biota in water courses potentially compete with native mollusc species such as the in northeast Trinidad in August 2015, a population of Neritina sp., which was also found in low densities at the the mussel Mytilopsis leucophaeata (Conrad 1831) was site. No predation of M. leucophaeata was observed at the discovered in the small river immediately north east of FROOHFWLRQVLWH,QFDSWLYLW\KRZHYHUWKHQDWLYHIUHVKZDWHU the Rio Seco/Salybia River bridge on the Toco Main Road prawn, Macrobrachium crenulatum did actively attack (UTM 20N 716200, 1184191). The density of mussels and feed on it. We suspect this prawn could be a natural was estimated at between 20 to 100 individuals per square predator as they also occur at the collection site. metre. They were clustered on both rock and driftwood 2XU VHFRQG QHZ ¿QGLQJ UHIHUV WR WKH QDWLYH VSHFLHV substrate. No individuals were observed in the Rio Seco Anodontites crispata. 7KLVVSHFLHVZDV¿UVWUHFRUGHGE\ River or any other watercourse that was surveyed. *XSS\IURPWKH&DURQLÀRRGSODLQV *XSS\ %DFRQ Specimens were collected and sent to Miramar recorded the species at Cuche River and Nariva Swamp =HHPXVHXP 1HWKHUODQGV IRU FRQ¿UPDWLRQ RI WKH in 1978 (Bacon 1979) and since then shells of A. crispata LGHQWL¿FDWLRQ /LYH VSHFLPHQV ZHUH DOVR KRXVHG LQ D (=A. cf. irisans ) were collected from Poole and Caparo IUHVKZDWHUDTXDULXPDW7KH8QLYHUVLW\RIWKH:HVW,QGLHV drainages in 2007 (Mohammed et al.   7DEOH ,Q St. Augustine for up to 24 months after collection. addition mussels (> 2cm in length) were observed by Dr Mytilopsis leucophaeata (Conrad 1831), (Dark false Mary Alkins-Koo in freshwater areas of the Carlisle River, mussel or Conrad’s false mussel) is a member of the Dreis- south Chatham in the early 1980s. Although these were senidae family native to North America. M. leucophaeata QRWLGHQWL¿HGLWLVVXVSHFWHGWKH\FRXOGEHPHPEHUVRIWKH has a thick, usually dark periostracum covering the shell 8QLRQLGDHIDPLO\+HUHZHGRFXPHQWDQHZORFDWLRQIRU A. and a maximum length of 2cm. (Fig.1). Juveniles may also crispata in southern Trinidad within the South Oropuche be striped. The ventral margin of the shell is fairly straight. drainage where a tributary crosses Mulchan Road (UTM ,QWHUQDOO\WKHUHLVDVHSWXPQHDUWKHXPERDQGSDUWO\FRY - 20N 672341, 1119862). The site has a muddy substrate, ered by this is a small triangular apophysis. This charac- with slow moving turbid water and riparian vegetation, a teristic septum is absent in the Mytilidae or true mussels. KDELWDWNQRZQWREHIDYRXUHGE\WKHVSHFLHV +XHKQHU This species has also been reported within suitable habitats Gagnon et al. 2006) and similar to that where Mohammed in Martinique (Delannoye et al. 2015), Guadeloupe in 1980 et al. (2008) documented shells in 2007. The species is also DQG *%,) 0DUJDULWD1XHYD(VSDUWDDQG%RFD known to be sensitive to low dissolved oxygen concentrations &KDYH]$Q]RDWHJXLLQ9HQH]XHOD *%,) DQGDVIDU DQGWKHIDPLO\GRHVQRWWROHUDWHDQ\VDOLQLW\ +XHKQHU south as French Guiana (Massemin et al.  ,WVUDQJH Gagnon et al. 2006). is suspected to be underestimated (Pointier 2015). 7KHXQLRQLGVIXO¿OLPSRUWDQWHFRV\VWHPIXQFWLRQVDQG There have been several instances of the species being are one of the most threatened freshwater taxa globally UHVSRQVLEOHIRUELRIRXOLQJLQ(XURSHDQDQGWHPSHUDWHUHJLRQV (Zieritz et al. 2016). Unlike M. leucophaeata , which has ']LXELĔVND+ROHFN et al. 2004, Therriault et al. 2004), planktonic free swimming stage in their life cycle, the and the initial expansion to these more southerly locations 8QLRQLGVSODQNWRQLFVWDJHUHOLHVRQ¿VKJLOOVDVSULPDU\ are thought to be the result of transport by watercraft and host for both support and nutrition (Williams et al. 1993). QRWQHFHVVDULO\QDWXUDOGLVSHUVDO +ROHFN et al. 2004). -XYHQLOHVPRYHXSVWUHDPYLDWKHLU¿VKKRVW6XEWOHFKDQJHV ,WLVSRVVLEOHWKDWVXEVHTXHQWQDWXUDOVSUHDGWR7ULQLGDG LQKDELWDWDQGRUH[WUDFWLRQRIWKHLU¿VKKRVWFDQOHDGWR PD\ KDYH RFFXUUHG WKURXJK SODQNWRQLF GULIW LQÀXHQFHG loss of this taxon. This species may be a candidate as an by the Orinoco discharge plume, or transport via birds or indicator species indicating well aerated slow moving possibly a direct introduction from its natural range via QXWULHQWSRRUVWUHDPVZLWKLQWDFW¿VKSRSXODWLRQV Nature Notes 103

Figure 1. Mytilopsis leucophaeata collected in North East near Rio Seco River, Trinidad. Photo credits- Robert Vink, The Netherlands.

Table 1 Occurrences of freshwater mussels in Trinidad. *See location description in text

Species Location Sighting Date Publication Anodontites crispata &DURQLÀRRGSODLQV na Guppy, 1866

Cuche River May 1978 Bacon et al. 1979

Nariva August 1978 Bacon et al. 1979

Nariva August 1988 6PLWKVRQLDQYLD,:5DPQDULQH 1988 Poole River May 2007 Mohammed et al. 2008

Caparo River November 2007 Mohammed et al. 2008

Mulchan Trace, Penal Rock September 2014 current paper Road Mycetopoda sp. Nariva August 1988 Bacon et al. 1979

Ortoire May 2007 Mohammed et al. 2008

Mytilopsis leucophaeata 1(RI5LR6HFR5LYHU August 2015 current paper

REFERENCES Guppy R .J.L. 1866. Additions to the catalogue of the land Bacon, P.R., Jaikaransingh, E. and Seeberan, G . 1979. and freshwater mollusca of Trinidad, 1:135-139. Notes on some freshwater molluscs from Nariva Swamp, Holeck, K.T., Mills, E.L., MacIsaac, H.J., Dochoda, Trinidad. Living World, Journal of Trinidad and Tobago M.R., Colautti, R.I. and Ricciardi, A . 2004. Bridging Field Naturalists’ Club, 1978-1979: 14-15. troubled waters: biological invasions, transoceanic ship- Delannoye R., Charles L., Pointier J. and Massemin D. SLQJ DQG WKH /DXUHQWLDQ *UHDW /DNHV AIBS Bulletin , 1RQPDULQH0ROOXVFRI0DUWLQLTXH/HVVHU$QWLOOHV 54:919-929. &ROOHFWLRQ ,QYHQWDULHV DQG %LRGLYHUVLWH 9RO 0XVHH Huehner, M.K., 1987. Field and laboratory determina- 1DWLRQDOG¶+LVWRLUH1DWXUHOOH3DULVS tion of substrate preferences of unionid mussels. The Ohio Dziubińska, A. 2011. Mytilopsis leucophaeata , an alien dre- Journal of Science , 87: 29-32. LVVHQLGELYDOYHGLVFRYHUHGLQWKH*XOIRI*GDĔVN VRXWKHUQ Gagnon, P., Michener, W., Freeman, M. and Box, J.B., Baltic Sea). Oceanologia , 53:651-655. 2006. Unionid habitat and assemblage composition in GBIF.org  $XJXVW *%,)2FFXUUHQFH'RZQORDG coastal plain tributaries of Flint River (Georgia). South- https://doi.org/10.15468/dl.fpwlzt eastern Naturalist, :31-52. 104 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

Massemin, D., Clavier, S. and Pointier, J.P., 2011. First of freshwater mussels of the United States and Canada. record of Pisidium punctiferum (Guppy, 1867) and Eupera Fisheries, 18:6-22. viridans (Prime, 1865) (Mollusca: Sphaeriidae) from Zieritz, A., Lopes-Lima, M., Bogan, A.E., Sousa, R., French Guiana. Novapex, 12:109-118. Walton, S., Rahim, K.A.A., Wilson, J.J., Ng, P.Y., Mohammed, R.S., Nieweg, D.C., Rostant, W.G. and Froufe, E. and McGowan, S., 2016. Factors driving Badal, P ., 2008. Collections of freshwater mussel shells changes in freshwater mussel (Bivalvia, Unionida) diver- of Anodontites sp. and Mycetopoda sp. in rivers of south- sity and distribution in Peninsular Malaysia. Science of the central Trinidad, West Indies. Living World, Journal of the Total Environment , 571: 1069-1078. Trinidad and Tobago Field Naturalists’ Club , 2008:76-77. Pointier J. 2015 (ed.) Freshwater Molluscs of Venezuela and their Medical and Veterinary importance. Conch Ryan S. Mohammed Books Harxheim. 228p. University of the West Indies Zoology Museum, Smithsonian National Museum of Natural History, Department of Life Sciences (DLS), Faculty of Science Department of invertibrate Zoology . 1988, Catalogue and Technology (FST). Numbers 853930 and 853931. [email protected] Therriault, T.W., Docker, M.F., Orlova, M.I., Heath, D.D. and MacIsaac, H.J. 2004. Molecular resolution Kerresha Khan of the family Dreissenidae (Mollusca: Bivalvia) with DLS, FST, The University of the West Indies, St. HPSKDVLV RQ 3RQWR&DVSLDQ VSHFLHV LQFOXGLQJ ¿UVW Augustine. report of Mytilopsis leucophaeata in the Black Sea basin. Molecular phylogenetics and evolution , 30:479-489. Dennis C. Nieweg Williams, J.D., Warren Jr, M.L., Cummings, K.S., Natural History Museum Rotterdam (Het Natuurhistorisch), Harris, J.L. and Neves, R.J., 1993. Conservation status Archaeoconchology, Rotterdam, The Netherlands.

Field observations of potential cannibalism among tadpoles of the Trinidad stream frog Mannophryne trinitatis (Anura: Aromobatidae)

Cannibalistic behaviour has been reported among DQGEULHÀ\LQWKH¿HOG:HUHSRUWRQD¿HOGREVHUYDWLRQ WDGSROHVDQGLVSRWHQWLDOO\LQÀXHQFHGE\IRRGDYDLODELOLW\ of potential cannibalistic behaviour in tadpoles of M. DQGFRQVSHFL¿FGHQVLW\ -HIIHUVRQ et al . 2014). Cannibalism trinitatis in Trinidad and suggest further research. involves killing and feeding on an individual of the same 2Q$XJXVWDWK5-$DQG1)$REVHUYHG species (Fox 1975), not to be confused with scavenging, seven to eight tadpoles of M. trinitatis consuming a single which involves feeding on already dead individuals. FRQVSHFL¿F LQGLYLGXDO :H ZHUH XQDEOH WR GHWHUPLQH &DQQLEDOLVPRIIHUVGLUHFWEHQH¿WVWRWDGSROHVLQWKHIRUPRI if the individuals were scavenging on an already dead DPHDODQGDOVRSURYLGHVRWKHUGLUHFWDQGLQGLUHFWEHQH¿WV FRQVSHFL¿F RU ZKHWKHU WKLV ZDV D FDVH RI LQWUDVSHFL¿F such as shortened development time (Crump 1990) and SUHGDWLRQRUFDQQLELOLVPDVGH¿QHGE\)R[  0RVW UHGXFHG FRPSHWLWLRQ LQ VRPH SRSXODWLRQV -HIIHUVRQ et tadpoles detached from the dead tadpole when we moved al . 2014). Conversely cannibalism may pose selective leaves and shone our LED torches to gain a better view. disadvantages. For example, it may increase the risk of But one of the larger tadpoles appeared more aggressive contracting chytridomycosis (Altig et al. 2007), or there WKDQ WKH RWKHUV DQG VWD\HG IHHGLQJ ¿QDOO\ PRYLQJ WKH PD\ EH LQFOXVLYH ¿WQHVV FRVWV RI NLOOLQJ FORVH UHODWLYHV dead tadpole into an overhanging rock crevice apart from Observation and investigation across more species is the others (Figure 1). The site was along the Lopinot required to further understand how common cannibalism Rd., Trinidad just south of Lopinot Village (UTM 20N is among tadpoles (Fox 1975, Altig et al. 2007). 682925, 1181345). The microhabitat consisted of an Mannophryne trinitatis (Garman 1887) of the family isolated pool of water, within a rock gully, less than 10m Aromobatidae (formerly Dendrobatidae) are small frogs from the roadside. The ephemeral pool was on a slightly (males 29mm, females 30mm snout to vent length) which inclined rock surface surrounded by dense canopy cover are distributed across the Northern Range and Central ZLWKDQDOPRVWGULHGXSVWUHDPÀRZLQJQHDUE\7KHSRRO Range of Trinidad (Murphy et al . 2018). Cannibalism had leaves in it covering more than 50% of the surface of among large M. trinitatis tadpoles has been reported by the water, and macrofauna consisted solely of M. trinitatis Downie et al . (2001) mainly under laboratory conditions WDGSROHV7KHWDGSROHVZHUHYLVXDOO\LGHQWL¿HGXVLQJWKH Nature Notes 105

ORFDO¿HOGJXLGH 0XUSK\ et al . 2018) and we also observed ACKNOWLEDGEMENTS M. trinitatis adults near the pool with no other amphibian The authors would like to thank the organisers of the Latin species observed nearby. America and Caribbean Society of Conservation Biology Similar observations have been made before. Downie Congress for Conservation Biology and The University of et al . (2001) reported on larger M. trinitatis tadpoles the West Indies for facilitating new collaborations. DWWDFNLQJ DQG VRPHWLPHV HDWLQJ VPDOOHU FRQVSHFL¿F tadpoles which were introduced to their pool. This REFERENCES behaviour was observed in the laboratory, and under Altig, R., Whiles, M.R. and Taylor, C.L. 2007. What natural conditions at a location similar to that of the present do tadpoles really eat? Assessing the trophic status of REVHUYDWLRQ-RZHUVDQG'RZQLH  DOVRUHSRUWHG M. an understudied and imperiled group of consumers in trinitatis WDGSROHVNLOOLQJDQGIHHGLQJRQWKHLUFRQVSHFL¿FV freshwater habitats. Freshwater Biology 52, 386-395. under laboratory settings. Our additional observation of Crump, M.L. 1990. Possible enhancement of growth in M. trinitatis  WDGSROHV HDWLQJ FRQVSHFL¿FV XQGHU QDWXUDO tadpoles through cannibalism. Copeia , 1990: 560-564. conditions is therefore not surprising. However it does Downie, J.R., Livingstone, S.R., Cormack, J.R. 2001. highlight an area for further study. Selection of tadpole deposition sites by male Trinidadian Do tadpoles of M. trinitatis regularly kill and eat stream frogs, Mannophryne trinitatis (Dendrobatidae): other tadpoles of their own species or are they usually An example of anti-predator behaviour. Herpetological only scavenging on individuals who have died for other Journal , 11: 91-100. reasons? If they are just feeding on dead individuals, Fox, L.R. 1975. Cannibalism in natural populations. then does this help keep the water clean? Alternatively, Annual Review of Ecology and Systematics , 6: 87-106. LI LQGLYLGXDOV GR UHJXODUO\ NLOO FRQVSHFL¿FV ZKDW DUH Jefferson, D.M., Hobson, K.A., Chivers, D.P. 2014. Time WKH IDFWRUV WKDW LQÀXHQFH WKLV EHKDYLRXU"  &DQQLEDOLVP WRIHHG+RZGLHWFRPSHWLWLRQDQGH[SHULHQFHPD\LQÀXHQFH may provide a source of nutrition or reduce competition feeding behaviour and cannibalism in wood frog tadpoles in resource-poor environments (Crump 1990), and the Lithobates sylvaticus . Current Zoology , 60: 571-580. small isolated pools where the M. trinitatis tadpoles were Jowers, M.J. and Downie, J.R. 2005. Tadpole deposition observed were heavily shaded and likely to have low behaviour in male stream frogs Mannophryne trinitatis primary productivity. Alternatively do plentiful food (Anura: Dendrobatidae). Journal of Natural History , 39: sources modulate individual aggressiveness and boldness 3013-3028. and lead to reduced cannibalism? Ultimately does Murphy, J.C., Downie, J.R., Smith, J.M., Livingstone, FDQQLEDOLVPLQWKHVSHFLHVRIIHUDQ\¿WQHVVEHQH¿WVDQGGR S.R., Mohammed, R.S., Auguste, R.J., Lehtinen, R.M., they cannibalise related individuals? Eyre, M., Sewlal, J.N., Noriega, N., Casper, G.S., Anton, T., Thomas, R.A., Rutherford, M.G., Braswell, A.L., and Jowers, M.J. $¿HOGJXLGHWRWKHDPSKLELDQV and reptiles of Trinidad and Tobago. Trinidad and Tobago Field Naturalists’ Club. 336 p.

Renoir J. Auguste The University of the West Indies Zoology Museum, Department of Life Sciences, St. Augustine Campus, Trinidad and Tobago. [email protected]

Nicole F. Angeli Division of Amphibians and Reptiles, National Fig. 1. Mannophryne trinitatis WDGSROHIHHGLQJRQDFRQVSHFL¿F Museum of Natural History, Smithsonian Institution. individual. [email protected] 106 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

Possible predation on the frog Leptodactylus validus (Anura: Leptodactylidae) by the spider Ancylometes bogotensis (Ctenidae) in Trinidad, W.I.

Spiders are known to feed on a variety of species, ZLOO UHDGLO\ GLVWLQJXLVK LW IURP RWKHU Leptodactylidae LQFOXGLQJ UHSWLOHV &RUUH\   ¿VK 'HDFRQ et al . species on Trinidad, including Adenomera and L.  FUXVWDFHDQV %KXNDO et al  DQGDPSKLELDQV nesiotus 0XUSK\ et al    :H QRWHG WKH VLPLODULW\ :KLWH ,QGHHGSUHGDWLRQRQDPSKLELDQVE\VSLGHUV of A. bogotensis E\FRPSDULQJRXUSKRWRJUDSKVWRWKRVH KDV EHHQ UHFRUGHG TXLWH IUHTXHQWO\ DQG HQFRPSDVVHV LQ RWKHU UHFHQW SXEOLFDWLRQV RI WKH VSLGHU SUHGDWLQJ RQ D YDULHW\ RI VSHFLHV RI DPSKLELDQV 0HQLQ et al . 2005; D YDULHW\ RI VSHFLHV LQ 7ULQLGDG %KXNDO et al . 2015; 7ROHGR ,QWKH1HRWURSLFVWKHIDPLO\&WHQLGDHKDV 'HDFRQ et al :KLWH 7KLVLGHQWL¿FDWLRQZDV EHHQ UHSRUWHG DV RQH RI WKH GRPLQDQW JURXSV RI VSLGHUV FRQ¿UPHGE\'U+|IHURIWKH6WDWH0XVHXPRI1DWXUDO SUHGDWLQJRQDPSKLELDQV 0HQLQ et al %DUER et al . +LVWRU\.DUOVUXKHLQ*HUPDQ\ZKRDGGHGWKDWWKHVSLGHU  3LQWR DQG &RVWD&DPSRV   $OWKRXJK WKHVH was a female Ancylometes . Ancylometes bogotensis is SUHGDWLRQUHSRUWVKLJKOLJKWVSHFLHVRIIURJVEHORQJLQJWR IRXQGWKURXJKRXWWKH1HRWURSLFDOUHJLRQIURP%ROLYLDWR a variety of families, including Leptodactylidae, none, as 1LFDUDJXDDQGLVWKHRQO\VSHFLHVIURPWKLVJHQXVNQRZQ IDUDVZHDUHDZDUHRILQFOXGHWKHVSHFLHV Leptodactylus IURP 7ULQLGDG +|IHU DQG %UHVFRYLW   :H GLG QRW validus (Garman 1888). FROOHFWDQ\VSHFLPHQVDVDQDGGLWLRQDOSHUPLWLVUHTXLUHG 2Q  1RYHPEHU  DW K ZKLOH FRQGXFWLQJ WRFROOHFWZLWKLQWKH:LOGOLIH6DQFWXDU\$VVXFKZHGLG D KHUSHWRIDXQD ¿HOG WULS LQ WKH %XVK %XVK :LOGOLIH 6DQFWXDU\1DULYD6ZDPS7ULQLGDG 8701  DWDQHOHYDWLRQDSSUR[LPDWHO\±PDERYH VHDOHYHO ZHZLWQHVVHGSRVVLEOHSUHGDWLRQRQWKHIURJ L. validus E\WKHVSLGHU Ancylometes bogotensis (Keyserling   )LJXUH :HGLGQRWREVHUYHWKHLQLWLDOFDSWXUH RI WKH IURJ E\ WKH VSLGHU EXW ZH VWD\HG WR REVHUYH WKH EHKDYLRXUIRUDERXWWHQPLQXWHV7KHVSLGHUDSSHDUHGWR KDYHKDGLWVIDQJVJULSSHGRQWRWKHUHDUHQGRIWKHIURJ EHWZHHQLWVKLQGOHJV7KHIURJZDVDFWLYHDQGDSSHDUHGWR EHWU\LQJWRHVFDSHWKXVZHEHOLHYHWKHFDSWXUHRIWKHIURJ SUREDEO\RFFXUUHGMXVWSULRUWRRXULQLWLDOREVHUYDWLRQ7KH IURJZDVDEOHWRPRYHDIHZFHQWLPHWUHVEXWLWZDVXQDEOH WREUHDNIUHHRIWKHVSLGHU7KHDSSDUHQWSUHGDWLRQWRRN SODFHDORQJDWUDLOWKDWKDGOHDIOLWWHURQWKHIRUHVWÀRRU ZLWKPRVWO\FORVHGFDQRS\DERYH7KHUHZDVDWUHHVWXPS RQ WKH WUDLO DQG D WZRIRRWZLGH GLWFK RI ZDWHU QHDUE\ ZKHUHWKHDWWDFNRFFXUUHG7KHWUDLOZDVDSSUR[LPDWHO\ PDFURVV7KHIRUHVWW\SHLQ%XVK%XVK:LOGOLIH6DQFWXDU\ FRQVLVWV RI VHDVRQDO HYHUJUHHQ IRUHVW %HDUG   :H REVHUYHGPDQ\RWKHU L. validus LQWKHOHDIOLWWHUQHDUE\DQG DORQJWKHWUDLODQGZHVSHFXODWHWKDW A. bogotensis PD\EH FRPPRQO\SUHGDWLQJRQWKLVVSHFLHVLQWKHDUHD :H LGHQWL¿HG L. validus  EDVHG RQ WKH GHVFULSWLRQ LQ 0XUSK\   DQG 0XUSK\ et al    DQG WKH LGHQWL¿FDWLRQ ZDV VXEVHTXHQWO\ FRQ¿UPHG E\ -RKQ 0XUSK\IURPWKHSKRWRJUDSK Leptodactylus validus is a PHGLXPVL]HGWHUUHVWULDOIURJDWWDLQLQJDPD[LPXPVQRXW WRYHQWOHQJWKRIPPZKLFKLVZLGHVSUHDGWKURXJKRXW 7ULQLGDG DQG KDV EHHQ UHFRUGHG LQ %XVK %XVK :LOGOLIH 6DQFWXDU\ SULRU WR RXU REVHUYDWLRQ 0XUSK\   7KH tympanic fold of L. validus H[WHQGLQJWRWKHVKRXOGHUODFN RIDZKLWHODWHUDOEORWFKERUGHUHGE\DPRUHGRUVDOGDUN Fig. 1. Ancylometes bogotensis with captured Leptodactylus VWULSH DQG D ODFN RI D PLGGRUVDO VWULSH RQ WKH GRUVXP validus . Photo Renoir J. Auguste Nature Notes 107

QRWJDWKHUPHDVXUHPHQWVRIWKHIURJDQGWKHVSLGHUQRUGLG 1HRWURSLFDOVSLGHUJHQXV Ancylometes Bertkau (Araneae: ZHVWD\ORQJHQRXJKWRZLWQHVV A. bogotensis consuming Pisauridae). Insect Systematics and Evolution ± WKH L. validus :HSRVWXODWHWKDWRXUUHSRUWIXUWKHUDGGV Menin, M., de J. Rodrigues, D. and de Azevedo, C.S. 2005. WRWKHOLWHUDWXUHWKDWVSLGHUVDUHLPSRUWDQWSUHGDWRUVWRD 3UHGDWLRQRQDPSKLELDQVE\VSLGHUV $UDFKQLGD$UDQHDH  YDULHW\RIDPSKLELDQVSHFLHV LQWKH1HRWURSLFDOUHJLRQ Phyllomedusa   ± Murphy J.C. 1997 $PSKLELDQVDQG5HSWLOHVRI7ULQLGDG ACKNOWLEDGMENTS DQG7REDJR0DODEDU)ORULGD.ULHJHU3XEOLVKLQJS :HDUHJUDWHIXOWRWKH:LOGOLIH6HFWLRQRIWKH)RUHVWU\ Murphy, J.C., Downie, J.R., Smith, J.M., Livingstone, 'LYLVLRQ RI 7ULQLGDG DQG 7REDJR IRU LVVXLQJ HQWU\ SHU - S.R., Mohammed, R.S., Auguste, R.J., Lehtinen, R.M., PLWVWRWKH%XVK%XVK:LOGOLIH6DQFWXDU\ 3HUPLWQXPEHU Eyre, M., Sewlal, J.N., Noriega, N., Casper, G.S., RI DQGWRDOOWKHYROXQWHHUVIURPWKH7ULQLGDG Anton, T., Thomas, R.A., Rutherford, M.G., Braswell, DQG 7REDJR )LHOG 1DWXUDOLVWV¶ &OXE WKDW FDPH RXW ZLWK A.L. and Jowers, M.J. $¿HOGJXLGHWRWKHDPSKLE - XV GXULQJ WKH KHUSHWRORJ\ WULS RQ 1RYHPEHU  WK , 2017, LDQV DQG UHSWLOHV RI7ULQLGDG DQG7REDJR7ULQLGDG DQG HVSHFLDOO\0U3HWHU'LFNVRQZKR¿UVWVSRWWHGWKHSUHGD - 7REDJR)LHOG1DWXUDOLVWV¶&OXES WLRQ:HDUHDOVRWKDQNIXOWR'U+XEHUW+|IHUDQG-RKQ Pinto, R.O. and Costa-Campos, C.E. 2017. Predation on &0XUSK\IRUFRQ¿UPLQJLGHQWL¿FDWLRQRIWKHVSLGHUDQG Dendropsophus brevifrons  'XHOOPDQ  &UXPS   frog respectively. $QXUD+\OLGDH E\WKHJLDQW¿VKLQJVSLGHU Ancylometes rufus  :DOFNHQDHU   $UDQHDH &WHQLGDH  Alytes, REFERENCES International Journal of Batrachology ± Barbo, F.E., Rodrigues, M.G., Couto, F.M. and Sawaya, Toledo, L.F. 3UHGDWLRQRIMXYHQLOHDQGDGXOWDQX - R.J. 2009. Predation on Leptodactylus marmoratus UDQVE\LQYHUWHEUDWHVFXUUHQWNQRZOHGJHDQGSHUVSHFWLYHV $QXUD /HSWRGDFW\OLGDH  E\ WKH VSLGHU Ctenus medius Herpetological Review ± $UDQHDH &WHQLGDH  LQ WKH $WODQWLF )RUHVW VRXWKHDVW White, G.   2EVHUYDWLRQ RI D VSLGHU Ancylometes Brazil. Herpetology Notes ± bogotensis  &WHQLGDH SUH\LQJRQWKHIURJ Rhinella bee- Beard , J.S.   7KH QDWXUDO YHJHWDWLRQ RI 7ULQLGDG bei (Bufonidae) in Trinidad. Living World, Journal of the 2[IRUG&ODUHGRQ3UHVV Trinidad and Tobago Field Naturalists’ Club ± Bhukal, R., Rutherford, M.G. and Mohammed, R.S. 3UHGDWLRQRQDIUHVKZDWHUFUDE Dilocarcinus den- tatus  7ULFRGDFW\OLGDH E\ VHYHUDOWURSLFDOZROI VSLGHUV Renoir J. Auguste Ancylometes bogotensis  &WHQLGDH  LQ 7ULQLGDG :, 'HSDUWPHQWRI/LIH6FLHQFHV7KH8QLYHUVLW\RIWKH:HVW Living World, Journal of the Trinidad and Tobago Field ,QGLHV6W$XJXVWLQH7ULQLGDGDQG7REDJR Naturalists’ Club , 2015: 59± [email protected] Corey, D.T. 1988. Comments on a wolf spider feeding on a green anole lizard. Journal of Arachnology ± Sachin Maraj Deacon, A.E., Farrell, A.D. and Fraser, D.F. 2015. 'HSDUWPHQWRI/LIH6FLHQFHV7KH8QLYHUVLW\RIWKH:HVW 2EVHUYDWLRQV RI D VHPL±DTXDWLF VSLGHU DWWDFN$Q RYHU - ,QGLHV6W$XJXVWLQH7ULQLGDGDQG7REDJR ORRNHG ¿VK SUHGDWRU LQ D ZHOO±VWXGLHG HFRV\VWHP" Living World, Journal of the Trinidad and Tobago Field Rainer Deo Naturalists’ Club ± 7KH(O6RFRUUR&HQWUHIRU:LOGOLIH&RQVHUYDWLRQ)UHHSRUW Höfer, H. and Brescovit, A.D.  $ UHYLVLRQ RI WKH 7ULQLGDGDQG7REDJR Fifteenth report of the Trinidad and Tobago Birds Status and Distribution Committee Records submitted during 2017

Martyn Kenefick 1HZDOORYLOOH$YH6DQ-XDQ7ULQLGDG E-Mail: PDUW\QNHQH¿FN#KRWPDLOFRP

INTRODUCTION The Trinidad and Tobago Birds Status and Distribution DQG6FRWW:HLVHQVDXO 86$ DQG5RELQ5HVWDOO 8. GXULQJ Committee (TTBSDC) was established in 1995 to assess, $OOUHFRUGHGVLJKWLQJVVXPPDULVHGEHORZRFFXUUHG document and archive the occurrence of rare or unusual birds LQXQOHVVRWKHUZLVHVWDWHG in Trinidad and Tobago and thus provide reliable long-term monitoring of our rarer species. Records accepted 7KHRI¿FLDOOLVWRIWKHELUGVRI7ULQLGDGDQG7REDJR $ SUHVXPHG SDLU RI White-faced Whistling-Duck, together with the list of species considered by the committee Dendrocygna viduata found along an irrigation channel and details of all records accepted by the Committee can be ZLWKLQIDUPODQGDW2UDQJH*URYHRQ0D\LQFUHDVHGWR accessed on-line at http://rbc.ttfnc.org. Previous reports of IRXUELUGVRQWKH0D\ (JF et al  $OPRVWDOOVLJKWLQJV the committee were prepared by Hayes and White, (2000); of this austral wanderer occur between mid-May and mid- :KLWHDQG+D\HV  DQG.HQH¿FN DQQXDOO\ October.  $UFKLYHGUHFRUGVLQFOXGLQJSKRWRJUDSKLFVXEPLVVLRQV $ MXYHQLOH RU DGXOW IHPDOH Muscovy Duck, Cairina QXPEHUDWWKHHQGRI moschata ZDV SKRWRJUDSKHG À\LQJ RYHU 6XPDULD WUDFH The Committee has assessed all records submitted during &KDUOLHYLOOHRQ 2FWREHU 5$ 0% :KLOVWZHUHPDLQ ,QDOO records were adjudged, representing 56 different concerned over provenance of this species in the south-west species. This represents the highest number of submissions peninsula of Trinidad, all indications are that this bird could and the highest number of “reportable species” since the well be of wild origin. formation of the Committee. This illustrates both the continued $SDLURI American Wigeon, Mareca americana was growth and popularity of birdwatching nationally. We also IRXQGLQ&DURQL5LFH3URMHFWRQ0DUFK '1-5 7KLVLV wish to commend the quality of photographic submissions WKH¿UVWGRFXPHQWHGUHFRUGRIWKLVVSHFLHVIRUWKHLVODQGRI by so many observers and urge observers to continue to Trinidad in the last 22 years. On 8 March an adult female document and report their sightings to us. ZDVSKRWRJUDSKHGDW%RQ$FFRUGVHZDJHODJRRQV7REDJR Of the submissions assessed, in only nine cases did the (MKe). These constitute the latest dates for this migrant &RPPLWWHH¿QGWKHLGHQWL¿FDWLRQLQFRQFOXVLYH+RZHYHUWKH GXFNSUHYLRXVODWHVWEHLQJ)HEUXDU\ \HDUFRLQFLGHGZLWKDQXPEHURILGHQWL¿FDWLRQFKDOOHQJHV Two Northern Shoveler, Spatula clypeata OLNHO\ERWK for the Committee, principally with submissions of smaller IHPDOHVZHUHIRXQGDW%RQ$FFRUGVHZDJHODJRRQV7REDJR Elaenias and immature Ardea KHURQV&XUUHQWO\¿YH Elaenia RQ1RYHPEHUUHPDLQLQJXQWLOWKH1RYHPEHUDWOHDVW VXEPLVVLRQVUHPDLQRXWVWDQGLQJDQGDZRUNLQJSDUW\ZLWKLQ (LK). This is just the third documented account of this species the Committee aims to create a template to both scrutinise in the last 22 years. same and create a template to simplify the process for the Two pairs of Lesser Scaup, $\WK\D DI¿QLV were IXWXUH&RQFXUUHQWWRWKDWDVLPLODUWHPSODWHLVEHLQJZRUNHG SKRWRJUDSKHGVZLPPLQJRQ$UHQD5HVHUYRLURQ-DQXDU\ upon to clarify proven differences between immature plumage (MKe, FO). This species is one of our more regular migrant of the three Ardea KHURQVSHFLHVNQRZQWRRFFXU GXFNVKDYLQJEHHQIRXQGLQVHYHQRIWKHODVW\HDUV Records presented below follow the revised nomenclature $QDGXOW American Flamingo, Phoenicopterus ruber DQGWD[RQRPLFRUGHURIWKH6RXWK$PHULFDQ&ODVVL¿FDWLRQ wDV VHHQ À\LQJ RYHU &DURQL 6ZDPS RQ  0DUFK 0. Committee as at June 2018 (Van Remsen et al. 2018). et al .). Over the course of the next two months, numbers The Committee comprises the following members: LQFUHDVHGWRDPD[LPXPRIELUGVRQ0D\ZLWKRQH 0DUW\Q .HQH¿FN 6HFUHWDU\  *HRIIUH\ *RPHV 1LJHO UHPDLQLQJXQWLODWOHDVW-XQH /DOOVLQJK%LOO0XUSK\.ULV6RRNGHRDQG*UDKDP:KLWH $QLPPDWXUH Scaly-naped Pigeon, Patagioenas squamosa $JDLQWKHUHDUHLQVWDQFHVZKHUHZHKDYHEHQH¿WHGIURP was photographed perched in roadside mangrove south of VXSSRUWLQJLQWHUQDWLRQDOH[SHUWNQRZOHGJHWRDVVLVWXVZLWK Manzanilla on 25 October (BR). Whilst this species has FHUWDLQLGHQWL¿FDWLRQLVVXHV:HZLVKWRDFNQRZOHGJHWKH now colonised Little Tobago island and the adjacent Tobago YDOXDEOHDVVLVWDQFHSURYLGHGE\%LOO&ODUN-RQ'XQQ'U KHDGODQGVWKLVLVWKH¿UVWGRFXPHQWHGVLJKWLQJIRU7ULQLGDG .DPDO,VODP0LFKDHO2¶%ULDQ'DYLG6LEOH\-DPHV6PLWK $SDLURI Blue Ground Dove, Claravis pretiosa were 108 Trinidad and Tobago Birds Status and Distribution Committee Report 109 found on the Chaguaramas peninsula on 4 March (JF,FO) $EDVLFSOXPDJHGDGXOW Curlew Sandpiper, Calidris and subsequently seen and heard by many observers. Whilst ferruginea ZDVSKRWRJUDSKHGDW%ULFN¿HOGRQ'HFHPEHU this remains an uncommon and localised resident of dry  1/ 7KHUHKDYHQRZEHHQWKUHHGRFXPHQWHGUHFRUGV 7ULQLGDGVFUXEDQGIRUHVWLW¶VZDU\DQGVHFUHWLYHQDWXUH of this Eurasian shorebird in Trinidad, others being in May may belie its true abundance 2002 and September 2016. $ Dark-billed Cuckoo, Coccyzus melacoryphus was Two Buff-breasted Sandpiper, &DOLGULVVXEUX¿FROOLV IRXQGLQVFUXEERUGHULQJ&DURQL5LFH3URMHFWRQ$XJXVW ZHUHIRXQGLQDJULFXOWXUDO¿HOGVQRUWKRI:DWHUORR0DLQ5G 1/ 2QVXEVHTXHQWGD\VDWOHDVWWKUHHELUGVZHUHVHHQE\ on 30 September, one remaining until 15 October at least PDQ\REVHUYHUVZLWKWZRVWLOOSUHVHQWRQ$XJXVW$OO 1/ )LJ $OOGRFXPHQWHGUHFRUGVRIWKLVVRXWKERXQG documented sightings of this austral wanderer in the last migrant shorebird in the last 22 years, have been between \HDUVKDYHRFFXUUHGEHWZHHQ-XO\DQG$XJXVW 2Q-XQHDPRXOWLQJDGXOWPDOH Rufous-shafted Woodstar, Chaetocercus jourdanii was photographed at $VD:ULJKW1DWXUH&HQWUH 6* :KLOVWWKHLGHQWL¿FDWLRQ of Woodstars sometimes represents a challenge for the &RPPLWWHHWKLVLVWKH¿UVWGRFXPHQWHGLQGLYLGXDOWRVKRZ LWVXQLTXHWDLOVKDSHDQGSDWWHUQ7KHUHKDYHQRZEHHQ¿YH FRQ¿UPHGVLJKWLQJVRIWKLVHUUDWLFDXVWUDOZDQGHUHUZLWKDOO UHFRUGVIURPHDUO\0D\WRODWH$XJXVW Single Amethyst Woodstars, Calliphlox amethystina were documented as follows :- an immature male at Surry YLOODJH/RSLQRWRQ$SULO *: DQLPPDWXUHDW$VD Wright on 2 May (DP); an adult male at Yerette on 24 May 7) DQGDQRWKHULPPDWXUHDW$VD:ULJKWRQ-XQH 0.  6LQFH¿UVWIRXQGEDFNLQWKHUHKDYHQRZEHHQHLJKW documented sightings of this wandering hummingbird from PDLQODQG6RXWK$PHULFDDOORFFXUULQJEHWZHHQ$SULO and 26 July. $Q American Coot, Fulica americana was found in a ÀRRGHG¿HOGZLWKLQ&DURQL5LFH3URMHFWRQ-XO\ 5$  The extent of white on the blaze, reaching the top of the head, together with the date show this to be a wandering Fig. 1. Buff-breasted Sandpiper, &DOLGULVVXEUX¿FROOLV , Waterloo, )DFDULEDHD SUREDEO\IURPWKH/HVVHU$QWLOOHVIRUPHUO\ October 2017. Photo Nigel Lallsingh. NQRZQDV&DULEEHDQ&RRWDQGQRZFRQVLGHUHGFRQVSHFL¿F ZLWK$PHULFDQ&RRW 6HSWHPEHUDQG2FWREHU $UHFRUGEUHDNLQJWRWDORIIRXULQGLYLGXDO Double-striped Up to three Franklin’s Gulls, Leucophaeus pipixcan ¿UVW Thick-Knees, Burhinus bistriatus were found in widely IRXQGRQ'HFHPEHU VHH.HQH¿FN UHPDLQHG separate localities during the year as follows :- at Waller WKURXJKWR)HEUXDU\DWOHDVW$QDGXOWIRXQGDW3LJHRQ3W Field on 6 July (KF,DR); alongside a golf course at Trincity 7REDJRRQ$SULO 5/ LVWKH¿UVWGRFXPHQWHGUHFRUGIRUWKH on 10 July (MK,RJ); in Caroni Rice Project on 30 July island. Elsewhere a late northbound adult was photographed 1/ DQG¿QDOO\LQWKHXQXVXDOORFDWLRQRI4XHHQV3DUN DW%ULFN¿HOGRQ0D\ 1/ DQGDQHDUO\DUULYLQJELUGLQ¿UVW 6DYDQQD3RUWRI6SDLQRQ$XJXVW )2 2ILQWHUHVWWKLV ZLQWHUSOXPDJHIRXQGDWWKHVDPHVLWHRQ$XJXVW 1/  ODVWORFDOLW\LVWKHVLWHRIWKHYHU\¿UVWUHFRUGRIWKLVVSHFLHV The Audouin’s Gull, Ichthyaetus audouini ¿UVWIRXQG LQ7ULQLGDGEDFNLQ1RZGRFXPHQWHGLQIRXURIWKH LQ'HFHPEHU VHH.HQH¿FN ZDVDJDLQSUHVHQWDW last six years, these sightings have all occurred within the %ULFN¿HOGRQ-DQXDU\ 1/ DQGIURP0DUFKWR$SULO expected date range of early July to early September. PDQ\REVHUYHUV DQG¿QDOO\IXUWKHUQRUWKDW,QYDGHUV%D\RQ $Q Upland Sandpiper, Bartramia longicauda was $XJXVW :5 7KLVUHFRUGKDVQRZEHHQDFFHSWHGE\WKH SKRWRJUDSKHGDOVRRQ4XHHQV3DUN6DYDQQD3RUWRI6SDLQ 6RXWK$PHULFDQ&ODVVL¿FDWLRQ&RPPLWWHHRIWKH$PHULFDQ RQ$XJXVW -) 7KLVEULQJVWRWKHWRWDOQXPEHURI 2UQLWKRORJLVWV8QLRQDVWKH¿UVWFRQ¿UPHGVLJKWLQJIRU6RXWK sightings of this rare southbound migrant from continental $PHULFD 1RUWK$PHULFDLQWKHODVW\HDUVZLWKDOOEXWRQHELUGV $QDGXOW White-tailed Tropicbird, Phaethon lepturus RFFXUULQJEHWZHHQ$XJXVWDQG2FWREHU ZDVVHHQIURPWKHORRNRXWRQ/LWWOH7REDJRLVODQGRQ 110 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

-DQXDU\ .%'- )LJ 6XEVHTXHQWUHSRUWVFRQ¿UPHG The Gray Heron , Ardea cinerea ¿UVW IRXQG DW %RQ DEUHHGLQJSDLUZKLFKVXFFHVVIXOO\UDLVHGRQHFKLFN PDQ\ $FFRUG7REDJRRQ 'HFHPEHU VHH.HQH¿FN  REVHUYHUV :KLOVWORQJDQWLFLSDWHGWKLVLVWKH¿UVWFRQ¿UPHG ZDVUHSRUWHGXQWLO$SULO6LPLODUO\WKHLQGLYLGXDO¿UVW breeding of this species in Trinidad and Tobago. IRXQGDW%ULFN¿HOGRQ'HFHPEHU VHH.HQH¿FN  UHPDLQHGWKURXJKRXWWKH\HDU$GGLWLRQDOO\DELUG LQ¿UVWZLQWHUSOXPDJHZDVSKRWRJUDSKHGDW:DWHUORRRQ -XO\ 1/  $WOHDVWRQHRIWKH Glossy Ibis, Plegadis falcinellus ¿UVW found RQ1RYHPEHUDW%RQ$FFRUG7REDJR VHH .HQH¿FN UHPDLQHGXQWLO)HEUXDU\DWOHDVW $GGLWLRQDOO\VLQJOHELUGVZHUHSKRWRJUDSKHGDW&DURQL5LFH 3URMHFWRQ-XO\ 1/ DQG6DWQXULQHWUDFH3HQDORQ $XJXVW 5* 2QFHDUDUHZDQGHUHUIURPWKHPDLQODQGWKH\ now occur annually and have been found in most months of the year $QDGXOWPDOH Snail Kite, Rostrhamus sociabilis was photographed on Caroni Rice Project on 21 March. Later joined by two birds in immature/female plumage they remained XQWLO$SULODWOHDVW -)-*.0 2Q$SULODPDOHZDV SKRWRJUDSKHGDWIDUPODQGLQ$UDQJXH] -) DQGDIXUWKHU PDOHZDVDW2UDQJH*URYHRQ0D\,WLVSRVVLEOHWKDW only three birds in total were involved. Formerly a very Fig. 2. White-tailed Tropicbird, 3KDHWKRQOHSWXUXV Little Tobago rare wanderer from the mainland, they have now occurred island on 12 January 2017 photo Nigel Hacking. during the period March-May in four of the last seven years. $ Cory’s Shearwater, Calonectris diomedea was seen $ELUGRISUH\ZKLFKSX]]OHGPDQ\REVHUYHUVIRUDORQJ À\LQJVRXWKDORQJWKHFRDVWDW0DQ]DQLOODRQ-DQXDU\ ZKLOHZDVHYHQWXDOO\LGHQWL¿HGDVDK\EULG White-tailed Hawk, (TJ). This remains a rare migrant seabird to local waters, Geranoaetus albicaudatus x Savanna Hawk, Buteogallus with almost all sightings coming from the winward coast meridionalis . ,WZDV¿UVWIRXQGRQ-XO\ 5$et al. )(Fig. RI7REDJRDQG7ULQLGDG¶VHDVWHUQFRDVWOLQHIURP1RYHPEHU  DQGUHPDLQHGSUHVHQWXQWLOWKH\HDUHQG7KLVLVWKH¿UVW to May. ever documented report of such hybridisation in Trinidad. $QLPPDWXUH Agami Heron, Agamia agami was found LQÀRRGHGORZODQGIRUHVWHDVWRI%UDVVR6HFRGXULQJHDUO\ 0DUFK &) )LJ 7KLVLVWKH¿UVWGRFXPHQWHGUHFRUGRI WKLVVK\DQGHOXVLYHZDQGHUHUIURPPDLQODQG6RXWK$PHULFD since 1961

Fig. 4. Hybrid White-tailed Hawk, *HUDQRDHWXV DOELFDXGDWXV x Savanna Hawk, %XWHRJDOOXV PHULGLRQDOLV Caroni Rice Project 2017. Photo Jerome Foster. ,QHDFKRIWKHODVW¿YH\HDUVDSDLURI Brown-throated Parakeet, Eupsittula pertinax has been observed inspecting/ H[FDYDWLQJKROHVLQWHUPLWHQHVWVLQWKH$ULSRVDYDQQDDUHD On 5 December, three birds were found together, indicative of successful breeding (EC, ES). Fig. 3. Agami Heron, $JDPLDDJDPL Brasso Seco, March 2017. Two White-eyed Parakeet, Psittacara leucophthalmus Photo Carl Fitzjames. ZHUHSKRWRJUDSKHGFORVHWR7DOSDURRQ)HEUXDU\ )2  Trinidad and Tobago Birds Status and Distribution Committee Report 111

,QVRXWKZHVW7ULQLGDGDÀRFNRIDSSUR[LPDWHO\ZHUH $SDLURI Lesson’s Seedeaters, Sporophila bouvronides VHHQÀ\LQJLQWRURRVWLQ0DQJRWUHHVDW3OHDVDQWYLOOH6DQ were found in St. Joseph on 21 October (CC, JF). On the same Fernando on 9 June (RL) with varying numbers being seen in date, a male was photographed in farmland at El Socorro the general area on a number of dates subsequently (many (RJ, SR). The date pattern suggests natural wanderers from REVHUYHUV 7KHIHUDOÀRFNEDVHGLQWKHQRUWKHUQVXEXUEV the mainland, however escapees from the pet trade cannot of Port of Spain has been reported as far east as San Juan. EHHOLPLQDWHG1RUHSRUWVZHUHUHFHLYHGWKLV\HDUIURPWKH $ Small-billed Elaenia, Elaenia parvirostris was found historic breeding site in south Trinidad. DW2UDQJH*URYHRQ0D\ -) DQGWZRGLIIHUHQWELUGV Single male Yellow-bellied Seedeater, Sporophila were photographed at Carlsen Fields on 4 June and 10 June nigricollis ZHUHIRXQGDW2UDQJH*URYHRQ0D\ -) 0RUQH UHVSHFWLYHO\ 1/ 7KHUHDIWHUXSWRIRXUELUGVZHUHSUHVHQW Catherine on 24 September (FO) and Mt. St. Benedict on 9 in a small agricultural area alongside Caroni Rice Project on October (MC, JR). This follows the now anticipated trend of $XJXVWZLWKDWOHDVWRQHUHPDLQLQJXQWLOWKH$XJXVW sightings from the north-west peninsula and farmlands and 1/ et al .). These sightings fall within the newly established JUDVVODQGVDWWKHVRXWKHUQEDVHRIWKH1RUWKHUQ5DQJH2IWKH SDWWHUQRIVLJKWLQJVLQ7ULQLGDG,QDOOVLQFHELUGVKDYH 22 documented records archived since the formation of this been found; all but one between 11 May and 5 September. Committee, all but two were from these geographical areas. $ Lesser Elaenia, Elaenia chiriquensis was photographed $QDGXOWPDOH Scarlet Tanager, Piranga olivacea was DW%DERRQLDWUDFH&KDUOLHYLOOHRQ$XJXVW -) &XUUHQWO\ SKRWRJUDSKHGDW$VD:ULJKW1DWXUH&HQWUHRQ$SULO 3' et ¿YHRWKHUVXEPLVVLRQVDUHVWLOOXQGHUUHYLHZE\WKH&RPPLWWHH al  DQGDJDLQVHHQRQ$SULO %5 2IWKHGRFXPHQWHG This remains a scarce resident of Trinidad lowland scrub. northbound migration records of this species, all but one The complexity of Elaenia À\FDWFKHULGHQWL¿FDWLRQPD\ KDYHRFFXUUHGIURPWR$SULO PDVNLWVWUXHDEXQGDQFH $UHWXUQLQJDGXOWIHPDOH Black and White Warbler, 'XULQJWKH\HDU¿YHVLJKWLQJVRI Black-whiskered Vireo, Mniotilta varia was found once again in a patch of mangrove Vireo altiloquus ZHUHDVIROORZVDW*UDQ&RXYDRQ FORVHWR&DUOL%D\RQ2FWREHU 1/ IRULWVIRXUWKVXFFHVVLYH February (JF); Poole on 24 March (FO); Carlsen Field on winter. 16 June (JF); Caroni swamp on 16 September (WR) and For the second year running, what we assume to be 0RUQH%OHXRQ2FWREHU )$ 7KLVVSHFLHVLVDSDUWLDORU the same Tennessee Warbler, Leiothlypis peregrina was short-distance migrant, breeding both on many of the Lesser SKRWRJUDSKHGIHHGLQJLQDOLQHRIWUHHVDGMDFHQWWR%ULFN¿HOG $QWLOOHVDQGDOVRLVODQGVRIIRIWKH9HQH]XHODQFRDVWOLQH )LVKLQJ'HSRWRQ'HFHPEHU 1/ )LUVWGRFXPHQWHGLQ 2IWKHGRFXPHQWHGUHFRUGVLQWKHODVW\HDUVDOOEXW 7ULQLGDG DV UHFHQWO\ DV  WKHUH KDYH QRZ EHHQ ¿YH RQHUHÀHFWQRQEUHHGLQJGLVSHUVDO+RZHYHUZHQRZKDYH sightings of this southbound migrant, in three years. a June sighting. The question remains as to whether this is $IHPDOH Cerulean Warbler, Setophaga cerulea found merely a delayed migrant, or whether the species may now RQ'HFHPEHUDW*UDQ&RXYDUHPDLQHGXQWLOWKH\HDUHQG become resident. at least (JF et al .), although its preference for feeding in the Southern Rough-winged Swallow, Stelgidopteryx tops of an extremely large Samaan tree precluded positive UX¿FROOLV is a common and widespread Trinidad resident LGHQWL¿FDWLRQIRUVRPHZKLOH&RQVLGHUHG³YXOQHUDEOH´E\ VSHFLHVEXWLQ7REDJRLWLVUDUH2QHIRXQGRYHU%RQ$FFRUG WKH,QWHUQDWLRQDO8QLRQIRUWKH&RQVHUYDWLRQRI1DWXUHDQG sewage lagoons on 11 January (DJ) is just the fourth documented NQRZQWRKDYHVXIIHUHGDGHFUHDVHLQSRSXODWLRQE\PRUH record for the island. WKDQLQWKHODVW\HDUVWKLVLVWKHIRXUWKGRFXPHQWHG $ Bank Swallow, Riparia riparia was found at Bon sighting for Trinidad and Tobago and the only individual $FFRUGVHZDJHODJRRQVRQ1RYHPEHU &+5+ 6WLOO NQRZQWREHSUHVHQWIRUPRUHWKDQRQHGD\ a distinctly rare migrant to Tobago, all four records this The Bay-breasted Warbler, Setophaga castanea ¿UVW century have been from this location. IRXQGDW&DUOL%D\RQ'HFHPEHU VHH.HQH¿FN  1RUWKERXQG PLJUDQW Cliff Swallows, Petrochelidon UHPDLQHGXQWLODWOHDVW-DQXDU\$QRWKHUZDVIRXQG at pyrrhonota were observed on Caroni Rice Project on 4 7DOSDURRQ-DQXDU\ .0)2 6LQFH¿UVWIRXQGLQ 0DUFK 1/ 0DUFK -) $SULO -) DQG$SULO they have been documented every year since; but not, as (JF). Returning southbound birds were found at Trincity on yet, from Tobago. 6HSWHPEHU 0.1/ DQGDW%ULFN¿HOGRQ2FWREHU $ Chestnut-sided Warbler, Setophaga pensylvanica 1/ 7KLVLVE\IDUWKHODUJHVWHYHUQXPEHURIVLJKWLQJV LQEDVLFSOXPDJHZDVIRXQGDW%RQ$FFRUG7REDJRRQ LQ7ULQLGDG,WLVKRZHYHUXQFHUWDLQZKHWKHUWKLVUHÀHFWV )HEUXDU\ 0.H 7KLVLVWKH¿UVWGRFXPHQWHGUHFRUGIRU increased observer coverage, increased observer awareness Tobago in the last 22 years. Whilst still considered a very of KLUXQGLQH LGHQWL¿FDWLRQRUDWUXHFKDQJHLQPLJUDWLRQ UDUHVRXWKERXQGPLJUDQW¿YHKDYHEHHQIRXQGLQWKHODVW distribution. seven years. 112 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018

$IHPDOH Canada Warbler, Cardellina canadensis was IRXQGLQPDQJURYHDGMDFHQWWR%RQ$FFRUGVHZDJHODJRRQV RQ)HEUXDU\ 0.H 7KLVFRQVWLWXWHWKH¿UVWGRFXPHQWHG sighting for Tobago and just the second nationally of this migrant warbler which has a wide wintering distribution WKURXJKRXWQRUWKHUQ6RXWK$PHULFD $IHPDOH Baltimore Oriole, Icterus galbula was found DW$VD:ULJKW1DWXUH&HQWUHRQ 0D\ 5'$*/1 )LJ  This is just the third documented sighting of this northbound migrant in the last 23 years.

Fig. 6. Crested Caracara, &DUDFDUDFKHULZD\adult with nestling. 1HVWLQJ RI WKLV VSHFLHV ZDV ¿UVW GRFXPHQWHG LQ 0DUFK$SULO 2017, location withheld. Photo Rishi Goordial. Escaped cage and aviary species We are aware of a reintroduction project involving Muscovy Ducks &DLULQDPRVFKDWDfrom Point a Pierre Wildfowl Trust. Sightings of this species from the south-west peninsula of Trinidad may involve birds from this scheme. Village Weavers, 3ORFHXVFXOFXOODWXV remain in small numbers on Caroni Rice Project and Tricolored Munias, /RQFKXUD PDODFFD are spreading thoughout western and central Trinidad. Elsewhere, exotic parrot species continue to be reported. Painted Parakeet3\UUKXUDSLFWD has been photographed in both Princes Town and Port of Spain; Red-shouldered Macaw, 'LRSVLWWDFDQRELOLV from Waterloo; Red and Green Macaws, $UDFKORURSWHUXV from La Brea, Palo Seco, Vessigny, Fig. 5. Baltimore Oriole, ,FWHUXV JDOEXOD Asa Wright Nature 4XLQDP6DQWD)ORUD/DV&XHYDV%ODQFKLVVHXVH&XPXWR Centre, May 13 2017. Photo Ashley Grove DQG0RNDDQGDMilitary Macaw, Ara militaris from south 9DOVD\Q$Blue Bunting, &\DQRFRPSVDSDUHOOLQD at Orange $ÀRFNRI Bobolink, Dolichonyx oryzivorus was *URYHDQGDVenezuelan Troupial, ,FWHUXVLFWHUXV in Carlsen found inside Caroni Rice Project on 23 October (MC, JR). Fields are also presumed to be a loss to cage-bird owners . 7KLVVRXWKERXQGPLJUDQWIURPFRQWLQHQWDO1RUWK$PHULFD The provenance of VHHGHDWHU DQG VHHG¿QFK VSHFLHV is now found almost annually with 18 out of the last 20 FRQWLQXHVWREHDSUREOHP7KH&RPPLWWHHKDVWDNHQDGHFLVLRQ GRFXPHQWHGVLJKWLQJVEHLQJIURP2FWREHUWR1RYHPEHU that, unless there is supporting evidence to the contrary, all sightings will be considered under this category and that Additional records DVVHVVPHQWZLOOEHRQLGHQWL¿FDWLRQDORQHChestnut-bellied $FFHSWDEOHUHFRUGVZHUHDOVRUHFHLYHGIRUDIXUWKHU Seed-Finch, 2U\]RERUXVDQJROHQVLV was reported from South sightings of the following species whose status has been Oropouche and Carli Bay; an adult male Lesson’s Seedfinch, HVWDEOLVKHGEXWZKR¶VGLVWULEXWLRQFRQWLQXHVWREHPRQLWRUHG 6SRURSKLODERXYURQLGHVIURP1DULYDDQGDChestnut Munia, by the Committee. Scaled Dove&ROXPELQDVTXDPPDWD /RQFKXUDDWULFDSLOOD was photographed at Caroni Rice Project. Rufous-necked Wood-Rail, $UDPLGHVD[LOODULVRufescent Tiger-Heron, 7LJULVRPD OLQHDWXP Hook-billed Kite, Inconclusive records &KRQGURKLHUD[XQFLQDWXVBlack Hawk-Eagle, 6SL]DHWXV Submissions of the following species were deemed W\UDQQXV Rufous Crab-Hawk, %XWHRJDOOXVDHTXLQRFWLDOLV inconclusive :- Green Heron, %XWRULGHVYLUHVFHQV(Trinidad); Crane Hawk, *HUDQRVSL]DFDHUXOHVFHQVLittle Egret, Egretta Gray Heron, $UGHD FLQHUHD Lesser Elaenia, Elaenia garzetta; Crested Caracara, &DUDFDUDFKHULZD\(Fig. 6); FKLULTXHQVLV Cliff Swallow, 3HWURFKHOLGRQ S\UUKRQRWD Aplomado Falcon, )DOFRIHPRUDOLVVariegated Flycatcher, Louisiana Waterthrush6HLXUXVPRWDFLOODBlackburnian (PSLGRQRPXVYDULXVSummer Tanager, 3LUDQJDUXEUD Warbler, 6HWRSKDJDIXVFD Trinidad and Tobago Birds Status and Distribution Committee Report 113

ACKNOWLEDGEMENTS Records and contributions were received from the Tobago 2009. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG IROORZLQJ)DUDD]$EGRRO%ULDQG¶$EUHDX5DVKLG$OL 7REDJR)LHOG1DWXUDOLVWV¶&OXE  9LVDO $O5DQJHUVDPP\ $QQH %HOWRQ .DUHQ %OLVVDUG Kenefick, M. 2011. Eighth report of the Trinidad and Michael Boswell, Maydiel Canizares, Ernest Carman, Cyril Tobago rare birds committee: rare birds in Trinidad and Coomansingh, Randell Dewberry, Pete Dunn, Kris Falco, Tobago 2010. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG Theo Ferguson, Carl Fitzjames, Laurie Foss, Jerome Foster, 7REDJR)LHOG1DWXUDOLVWV¶&OXE 2011: 80-85. -RKQ*DJOLRQH6WHSKHQ*DVW5LVKL*RRUGLDO/DUU\*ULGOH\ Kenefick, M.  1LQWK UHSRUW RI WKH 7ULQLGDG DQG $VKOH\*URYH'DYLV*XQQ.QXW+DQVHQ&KULV+RZDUG Tobago rare birds committee: rare birds in Trinidad and Rosie Howard, David Huggins, Rodney Jagai, Doug Jagger, Tobago 2011. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG /DZUHQFH-DPHV7RP-RKQVRQ/DXUD.HHQH0DUW\Q.HQH¿FN 7REDJR)LHOG1DWXUDOLVWV¶&OXE 2012: 96-100. 0DWW.HOO\ 0.H 5LFKDUG/DNKDQ1LJHO/DOOVLQJK.DPDO Kenefick, M. 2013. Tenth report of the Trinidad and 0DKDELU7DUUDQ0DKDUDM/L]1DLSDXO'DUVKDQ1DUDQJ Tobago rare birds committee: rare birds in Trinidad and )HUR]H2PDUGHHQ'DYH3XOODQ%DUU\5DPGDVV0XNHVK Tobago 2012. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG Ramdass, Dave Ramlal, Darryl Ramlochansingh (DRa), 7REDJR)LHOG1DWXUDOLVWV¶&OXE  6DWDLVK5DPSHUVDG:HQGHOO5H\HV-HVVLFD5R]HN(OOHQ Kenefick, M. 2014. Eleventh report of the Trinidad and 6PLWK.ULV6RRNGHR*UDKDP:KLWH0LFKDHO:LOOLVRQDQG Tobago rare birds committee: rare birds in Trinidad and Barry Zimmer. Tobago 2013. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG 7REDJR)LHOG1DWXUDOLVWV¶&OXE 2014: 59-63. REFERENCES Kenefick, M. 2015. Twelfth report of the Trinidad and Hayes, F.E. and White, G. 2000. First report of the Tobago rare birds committee: rare birds in Trinidad and Trinidad and Tobago rare birds committee. /LYLQJ:RUOG Tobago 2014. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG -RXUQDO RI WKH 7ULQLGDG DQG 7REDJR )LHOG 1DWXUDOLVWV¶ 7REDJR)LHOG1DWXUDOLVWV¶&OXE 2015: 56-61. &OXE 2000: 39-45. Kenefick, M. 2016. Thirteenth report of the Trinidad and Kenefick, M. 2005. Third report of the Trinidad and Tobago rare birds committee: rare birds in Trinidad and Tobago rare birds committee: rare birds in Trinidad Tobago 2015. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG and Tobago in 2001-2003. /LYLQJ:RUOG-RXUQDORIWKH 7REDJR)LHOG1DWXUDOLVWV¶&OXE  7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE 2005: 56-61. Kenefick, M. )RXUWHHQWKUHSRUWRIWKH7ULQLGDGDQG Kenefick, M.  )RXUWK UHSRUW RI WKH 7ULQLGDG DQG Tobago rare birds committee: rare birds in Trinidad and Tobago rare birds committee: rare birds in Trinidad and Tobago 2016. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG Tobago in 2004 and 2005. /LYLQJ:RUOG-RXUQDORIWKH 7REDJR)LHOG1DWXUDOLVWV¶&OXE  7ULQLGDGDQG7REDJR)LHOG1DWXUDOLVWV¶&OXE  Remsen, J.V., Jr., Areta, J.I., Cadena, C.D., Jaramillo, Kenefick, M. 2008. Fifth report of the Trinidad and Tobago A., Nores, M., Pacheco, J.F., Perez-Emen, J., Robbins, rare birds committee: rare birds in Trinidad and Tobago M.B., Stiles, F.G., Stotz, D.F. and Zimmer, K.J. Version DQG /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG 22 February 2016 $FODVVL¿FDWLRQRIWKHELUGVSHFLHVRI 7REDJR)LHOG1DWXUDOLVWV¶&OXE  6RXWK$PHULFD$PHULFDQ2UQLWKRORJLVWV¶8QLRQ>2QOLQH@ Kenefick, M. 2009. Report of the Trinidad and Tobago $YDLODEOHDWKWWSZZZPXVHXPOVXHGXa5HPVHQ6$&& rare birds committee: rare birds in Trinidad and Tobago %DVHOLQHKWP $FFHVVHG0DUFK 2008. /LYLQJ:RUOG-RXUQDORIWKH7ULQLGDGDQG7REDJR White, G. and Hayes, F. 2002. Second report of the )LHOG1DWXUDOLVWV¶&OXE 2009: 46-49. Trinidad and Tobago rare birds committee. /LYLQJ:RUOG Kenefick, M. 2010. Seventh report of the Trinidad and -RXUQDO RI WKH 7ULQLGDG DQG 7REDJR )LHOG 1DWXUDOLVWV¶ Tobago rare birds committee: rare birds in Trinidad and &OXE 2002: 51-56. 114 Living World, J. Trinidad and Tobago Field Naturalists’ Club, 2018 Book Review

A Field Guide to the Amphibians & Reptiles of Trinidad & Tobago. Trinidad & Tobago Field Naturalists’ Club

by J.C. Murphy, J.R. Downie, J.M. Smith, S. M. Livingstone, R.S. Mohammed, R.M. Lehtinen, M. Eyre, J.N. Sewlal, N. Noriega, G.S. Casper, T. Anton, M.G. Rutherford, A.L. Braswell and M.J. Jowers. (R.J. August, Managing Editor; Foreword by R.A. Thomas) 2018

Twenty years ago, I bought ‘Amphibians and Reptiles in those two books. That sort of circle, is closed today as of Trinidad and Tobago’, published by the North American I write this modest review. Since then I have been always Krieger becoming, with ‘Turtles of Venezuela’ by Peter interested in the herpetofauna of Trinidad and Tobago, and 3ULWFKDUGDQG3HGUR7UHEEDXRQHRIWKH¿UVWKHUSHWRIDXQD of course of that of the Paria Peninsula. A few years ago, I books in my personal library. Then, I was amazed by do not remember who contacted who, John and I started an the similitude of the herpetofauna of these two island online friendship, so thatI was looking forward to A Field and adjacent islets with coastal Venezuela, separated by Guide to the Amphibians & Reptiles of Trinidad & Tobago barely 15km from the Venezuelan Paria Peninsula. When published by the renowned Trinidad & Tobago Field I received the book, my impression was that everything 1DWXUDOLVWV¶&OXEZKLFK¿OOVWKHJDSDIWHU0XUSK\   one needed to know was already there, there were only and surely as it happened with me, it would motivate a new few topics to investigate; but on the other hand, it also generation of researchers in herpetology. This new Field motivated me to investigate the Venezuelan area that is the Guide will certainly be a valuable source of information to most similar to those islands: the isolated Paria Peninsula. understand and preserve this unique biota, shared in part I was able to publish a note on the reptiles of Sucre State with northeastern Venezuela. one year late (Rivas and Oliveros 1997 “1998”). Coming The guide contains information provided by 20 back to Pritchard and Trebbau (1984), I noted a picture of investigators, directed by John Murphy, and edited by a mata mata turtle that arrived to the coast of Trinidad with Renoir Auguste, a promising young herpetologist from some barnacles in its carapace, due to its permanence in Trinidad. With 336 pages, A Field Guide to the Amphibians blackish water. What I read surprised me, how the waters & Reptiles of Trinidad & Tobago starts with a list of all IURPWKH2ULQRFRFDQFDUU\PDVVHVRIÀRDWLQJYHJHWDWLRQ DXWKRUVDQGWKHLUFXUUHQWDI¿OLDWLRQVDIRUHZRUGE\5REHUW with reptiles to the coast of Trinidad. That picture also Thomas, notable herpetologist from the US, who has called my attention because the photographer, Hans Boos, worked in South America. These are followed by a preface, had the same surname as a farm ubiquitous in northeastern acknowledgments, how to use this book, introduction Venezuela (decades later I noticed this farm have belonged the environment conservation, folklore, measurements to an uncle of H. Boos). & identifying features, waifs & questionable species, Years later, in March 2014, travelling with my colleague amphibian species accounts, reptile species accounts, six Tito Barros to Florida, USA, to visit Peter Pritchard to appendixes, glossary of terms, list of references cited in work on the book ‘Venezuela y sus Tortugas’, we made WKHWH[WDQG¿QDOO\DQLQGH[RIFRPPRQQDPHV a brief stop in Trinidad and took the opportunity to meet We can read in the preface about Anne, a pregnant Mr Boos. Once in the airport, Boos invited us to visit the green anaconda, found in Trinidad and shipped to the Emperor Valley Zoo and later visit his home to see his Bronx Zoo and how her babies were distributed to many personal library. Then, he showed us his slide collection other zoos, or the discovery of a luminous lizard found a and I was surprised to see that mata mata picture I had seen cave by Ivan Sanderson, among other histories, being a 20 year earlier in Pritchard and Trebbau’s and Murphy’s source of motivation to a next generation of individuals books. That is how my life history has become what it is, interested in herpetology. it was written and visualised by those photos that appeared 7KH ¿UVW SDUW RI WKH ,QWURGXFWLRQ GHVFULEHG EULHÀ\ Book Review 115 but precisely the rate of discoveries of new species of those people familiar or not with biodiversity issues. How amphibians and reptiles, lack of information, how the LWUHDGVRQSDJH³WKHGHVLJQRIWKLV¿HOGJXLGHDOORZV geographers consider Trinidad & Tobago a biodiversity ready access to the Trinidad and Tobago herpetofauna. The hub, the relationships of these islands with the surrounding vocabulary may be most familiar to naturalists, scientists land masses and features and diversity of the herpetofauna and amateur herpetologists, but for the novice we have that can be found. supplied a glossary and some detailed explanations”. The section of conservation is really useful for all Finally, a wider list of bibliographical references and an researchers, tourists, as well as hunters that need to know LQGH[RIFRPPRQQDPHV¿QLVKWKHPDVWHUSLHFH the legislation of the country as well as the laws that The team of authors of A Field Guide to the Amphibians protect the wildlife and organisation to grant the permits. & Reptiles of Trinidad & Tobago have provided us with a In the folklore chapter, the authors mention that Trinidad useful tool to know and preserve the life of the amphibians and Tobago is biologically and culturally very diverse, and reptiles from these islands. This guide complements ZLWKDYDULHW\RILQÀXHQFHVLQFOXGLQJ$IULFDQ&KLQHVH those of Kenny (1969), Murphy (1997), and Boos (2001) Indian and European immigrants and colonisers. This rich and those books edited by Hailey et al. (2011a,b). I will FXOWXUDOPRVDLFLVUHÀHFWHGLQIRONORUHDQGVXSHUVWLWLRQV recommend it to everyone interested in knowing the including folk medicine. Measurement and characteristics biodiversity of amphibians and reptiles of Trinidad and of the different groups, shows the main characteristics to Tobago, the Antilles and northern South America. identify the species. Profusely illustrated with drawings, albeit some of the drawings taken from other publications; REFERENCES however, the objective is met. In waif and questionable Boos, H.E.A. 2001. The snakes of Trinidad and Tobago. species, the authors have reviewed all available information W.L. Moody Jr. Natural History Series. Texas A & M and with provide new evidence about the species wrongly University Press. 328 p. cited for the Republic of Trinidad and Tobago. Hailey, A., Wilson, B.S. and Horrocks, J.A. (eds.) 2011. The species accounts make up the bulk of this treatise, Conservation of Caribbean island Herpetofaunas. Vol. I. including information on 129 species of amphibians and Conservation biology and the wider Caribbean. Brill, Leiden. reptiles (35 frogs, 30 lizards, 11 turtles, two species of 227 p. crocodiles, and 51 snakes). Ten of them had an exotic Hailey, A., Wilson, B.S. and Horrocks, J.A. (eds.) 2011. origin: the frogs Eleutherodactylus jonhtonei and Scarthyla Conservation of Caribbean island Herpetofaunas. Vol. II. vigilans , seven lizards ( Hemidactylus mabouia , Anolis Regional accounts of the West Indies. Brill, Leiden. 439 p. aeneus , A. extremus , A. richardii , A. sagrei , A. trinitatis Kenny, J.S. 1969. The Amphibia of Trinidad. Studies on the and A. wattsi ) and a fresh water turtle ( Trachemys scripta Fauna of Curacao and other Caribbean islands , 8: 1–78+15 elegans ). Each species account contains its previous plates. NQRZQQDPHFRPPRQQDPHVL]HLGHQWL¿FDWLRQVLPLODU Murphy, J.C. 1997. Amphibians and Reptiles of Trinidad species, distribution, habitat and biology, a dot map with and Tobago. Krieger Publishing, Malabar, FL. different colours dots according to the time of the record, Pritchard, P. and Trebbau, P. 1984. The Turtles of DQG ¿QDOO\ SURIXVHO\ LOOXVWUDWHG ZLWK PDQ\ SLFWXUHV DQG Venezuela. Society for the study of amphibians and drawing of the species and their habitat, most of them reptiles, contribution to herpetology , 2: 1-414. taken directly from the country, except few taken on Rivas, G. and Oliveros, O. 1997 “1998”. Herpetofauna outside such as Anolis extremus , an exotic species. del estado Sucre, Venezuela: lista preliminar de reptiles. 6L[DSSHQGLFHVDUHSURYLGHGWKH¿UVWLVRQ7ULQLGDG Memoria de la Sociedad de Ciencias Naturales La Salle , frog reproductive modes: spawn, hatching stages, and 147: 67–80. tadpoles (appendix I), followed by Handling Amphibians and Reptiles (appendix II), Amphibian chytridiomycosis, Gilson A. Rivas (appendix III) Snakebite (appendix IV), Herpetological Museo de Biología, Facultad Experimental de Ciencias Collections in Trinidad & Tobago (appendix V) and Universidad del Zulia, apartado postal 526, Maracaibo Research Stations and Lodging for Ecotourists (appendix 4011, Venezuela VI). The glossary of biological terms is really useful for all [email protected] Notes to Contributors Living World , the journal of The Trinidad and Tobago Field Naturalists’ Club, publishes articles on studies and observations of natural history carried out in Trinidad and Tobago, and in the Caribbean Basin. Contributors to Living World are not limited to members of the Club. Articles submitted for publication are sent to two referees for review. Articles are accepted on the condition that they are submitted only to Living World . Regarding a coauthored article, the senior author PXVWDI¿UPWKDWDOODXWKRUVKDYHEHHQRIIHUHGDQRSSRUWXQLW\ WRSHUXVHWKHVXEPLWWHG YHUVLRQDQGKDYH approved of its publication.

Articles may be emailed to: [email protected] or submitted online via our open access jounal platform (See https://ttfnc.org/livingworld/index.php/lwj/about/submissions#onlineSubmissions). In general, we follow the Council of Science Editors Style Manual (https://writing.wisc.edu/Handbook/ DocCSE.html). All articles, except for Nature Notes, should be accompanied by an abstract and a list of key words. Nature Notes LV D VHFWLRQ DOORZLQJ FRQWULEXWRUVWR GHVFULEH XQXVXDO REVHUYDWLRQVRQ RXU ÀRUD DQG IDXQD 7KH WLWOH RI each Nature Note should include key words and the note should not exceed three journal pages in length, including tables and photographs. Only a few key references should be included. References should follow the Name and Year system. Some examples:

1. Journals: The full title of a journal should be given. 0HPEHUVKLS LVRSHQWRDOOSHUVRQVRIDWOHDVW¿IWHHQ\HDUVRIDJHZKRVXEVFULEH WRWKHREMHFWVRIWKH Larsen, N.J. and Levesque, A. 2008. Range expansion of White-winged Dove ( Zenaida asiatica ) in the Lesser Antilles. Journal of Caribbean Ornithology , 21: 61-65. Quesnel, V., Farrel, T. F., Hilton, A., Hilton, J. and Zuniaga, L. 1996. The revegetation of the McClean Monument. Living World, Journal of The Trinidad and Tobago Field Naturalists’ Club, 1995-1996: 9-12.

2. Books and Monographs: Kenny, J. 2008. The Biological Diversity of Trinidad and Tobago. Maraval, Trinidad and Tobago: Prospect Press. 265 p.

3. Citation from Books and Monographs with Editors: Collins, C.T. 2002. Notes on the biology of the Band-rumped Swift in Trinidad. p. 138-143. In F.E. Hayes and S.A. Temple, eds. Studies in Trinidad and Tobago’s Ornithology Honouring Richard ffrench. St. Augustine, Trinidad and Tobago: Department of Life sciences, University of the West Indies, Occasional Paper No. 11.

4. Online References: Rutherford, M.G. 2012. Tucker Valley BioBlitz 2012 Summary. Field Naturalist 2012(4), p. 6-17 [Online]. Available at http://ttfnc.org/photojournals/2012-4.pdf (Accessed 02 February 2013)

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