A new snake from Trinidad, and its significance to the big picture

There is no doubt that there are many, many more species of amphibians and reptiles than previously thought. Peter Uetz, maintains the Reptile Database website, and he recently announced that in 2014, the number of known reptile species passed the 10,000 mark – and the year has not yet ended. In an email Uetz wrote the number of reptile species is, “10,038 species (including 79 described this year), up from 9,952 in April”. While some of these species are obviously different, many of them are cryptic, and it takes some detailed study of morphology or DNA to sort them out.

Trinidad is a relatively small island – 4800 square kilometers that has had its herpetofauna relatively well studied. The first list of herps was published in 1858, and the work by Mole and Urich at the turn of the 20th century examined the fauna extensively. William Beebe spent the last years of his life at Simla in the Arima Valley and published a fauna list as well as a paper on the ecology of the valley. Garth Underwood and Michael Emsley also studied and wrote about the island’s herpetofauna. Julian Kenny and Hans Boos were also actively working on the Trinidad herpetofauna for decades. And I made five- or six trips to the islands and examined museum specimens in the 1980-90’s for the 1997 book. During those trips, I frequently stayed at Simla and worked extensively in the Arima Valley. In 2010, I decided to take a second look at the fauna and since that date have made eight trips to investigate the herpetofauna of both Trinidad and Tobago. Additionally, I have been working with colleagues to compare museum material from the islands to those from the mainland – the results are startling.

It has become quite obvious to me that the diversity of reptiles on that 4800 sq km, well studied island (as well as the island of Tobago) is much greater than what I (or anybody else) thought it was in 1990.

Within the last few years, we have described Plica caribeana and Leptophis haileyi. The Plica is a Caribbean Coastal Range species, and the Leptophis appears to be a Tobago endemic. There are more species forthcoming – for the most part it is a matter of time and money to get the work done. But, preliminarily we have identified at least ten more species of squamate reptiles currently unrecognized from the islands or masquerading under the name of a widespread species.

In a forthcoming paper in the Journal of South American Herpetology, Teddy Angarita-Sierra describes a new, cryptic species of coffee snake, in the genus Ninia – from Trinidad. It would normally not surprise me, that a small, leaf litter dwelling snake would go un-noticed in the tropics. But, this snake is surprising because the of the type locality – Simla. A location that has to be the most closely examined piece of real estate on the island – in terms of its fauna.

Ninia atrata left, Nina franciscoi sp. n. right. T. Angarita-Sierra

Angarita-Sierra describes Ninia franciscoi, from a single specimen collected at Simla 6 March 1988 by William B. Montgomery and David Resnick. This specimen looks very much like the widespread Ninia atrata (also found at Simla) except for the number of upper labials contacting the primary temporal and most importantly, an unusually shaped hemipensis (see right).

Nina franciscoi sp n.,top, Nina atrata bottom.

After reviewing this paper – I went back and looked at about forty specimens of Ninia atrata from Trinidad and Tobago as well as all of my photographs, looking to see if any of the others had the primary temporal in contact with three upper labials. Some of these specimens were collected at Simla – none showed this trait.

So, what is the significance of this? Below is a power point slide I used in a recent presentation comparing the number of reptile species per 1000 km² in Trinidad & Tobago to Venezuela. The islands are much better studied than the mainland. Trinidad & Tobago have 4.2x more known species of reptiles per 1000 km² than does Venezuela (this is using the numbers from the 1997 book). In other words mainland South America (specifically Venezuela) most likely has a vast number of undescribed reptiles.

As for those undescribed Trinidad and Tobago squamates - one of them is a third species of Ninia. Below is typical Ninia atrata. 

Gymnophthalmus underwoodi on Tobago

The shiny lizard, Gymnophthalmus underwoodi, is one of those reptiles that is widespread and probably common,  but not particularly easy to find. It is small, lives in burrows or under objects, and is active only when the sun is shining - when it can significantly raise its body temperature. Its distribution ranges from Venezuela, the Guyanas, and Brazil northward into the Lesser Antilles. It has been reported as far north as Antigua. Humans may, at least in part, be responsible of its island distribution.The shiny lizard is frequently associated with lawns and gardens and as people transport their favorite plants from palace to place these tiny lizards may move with them. Additionally it is a unisexual species which provides the advantage of not needing a mate. In fact, G. underwoodi is one of four species in a complex of cryptic species (the others are: G. cryptus, G. leucomystax, and G. specious). Kiziran & Cole (1999) found G. cryptus is the ancestor of G. underwoodi.

Hardy (1982) commented on the lack of specimens of Gymnopthalmus underwoodi from Tobago, Cole et al did find find specimens from Tobago, nor did Murphy (1997). Yet some publications and websites have included Tobago in the distribution- the IUCN red list is an excellent example. It presence on Tobago would seem reasonable given its presence in Trinidad and Venezuela and the Antilles. However, a HerpNet search did not show any specimens from Tobago.

On 6 June 2013 Rick Lehtinen collected a single specimen of G. underwoodi in from of the Vegetable King in Castara, Tobago. Thus confirming its presence on the island. The specimen is deposited in the UWIMZ. Since that date the species has become quite abundant on the northeastern end of the island.

Literature

Cole, CJ, et al. Gymnophthalmus (Reptilia Teiidae) in the neotropics: genetics origin and systematics. American Museum Novitates (2994): 1-29

Hardy, J. D. 1982. Biogeography of Tobago, West Indies, with special reference to amphibians and reptiles, a review. Bulletin of the Maryland Herpetological Society 18(2):37–142.

Kizirian DA. Cole CJ. 1999. Origin of the Unisexual Lizard Gymnophthalmus underwoodi (Gymnophthalmidae) Inferred from Mitochondrial DNA Nucleotide Sequences. Molecular Phylogenetics and Evolution 11:394–400.

Murphy, J. C. 1997. Amphibans and reptiles of Trinidad ad Tobago. Krieger Publishing, Malabar, FL.

Is Chironius scurrulus part of the Trinidad herpetofauna?

In 1993 Dixon et al. reported on a specimen of Chironius scurrulus from Trinidad. The specimen, UMMZ 43965 is a juvenile that was collected by E. B. Williamson, an insect collector at the turn of the 20^th century (winter 1912). Wagler’s Sipo as it is sometimes called inhabits Brazil, SE Colombia, N Bolivia, Ecuador, Venezuela, E Peru, Trinidad, Guyana, Surinam, French Guiana according to the Reptile Database website, and probably has a type locality of Rio Japura, Brazil (based on comments in Kornacker 1999). Boos (2001) also reported this species as part of the Trinidad herpetofauna, following the previous authors.

The UMMZ specimen of Chironius scurrulus said to be from Trinidad

Recently I visited the UMMZ and examined the specimen of Chironius scurrulus, it is undoubtedly that species, but there is evidence to suggest it did not come from Trinidad.  Also collected by Williamson that winter on Trinidad were some Rhinella marina, Hypsiboas crepitans, Mannophryne trinitatis, and several specimens of the stream toad, Atelopus hoogmoedi. The last species is of course unknown from Trinidad – but it is known from Guyana, the other country visited by Williamson during the winter of 1912. Given the low probability of picking up the only known specimen of Chironius scurrulus and the only known specimens of Atelopus hoogmoedi from Trinidad on one trip, it seems likely that the specimens were instead collected in Guyana and at some point mislabeled as to their origins.

I would like to thank Greg Schneider at the UMMZ for searching the archives for information on these specimens, and to César Luis Barrio-Amorós, J. Roger Downie, and Joanna Smith for their perspectives into these records.

Citations

Boos, H.E.A. 2001. The snakes of Trinidad and Tobago. Texas A&M University Press, 270 pp.

Dixon, J. R., Wiest Jr, J. A., & Cei, J. M. 1993. Revision of the neotropical snake genus Chironius Fitzinger Serpentes, Colubridae. Torino: Museo regionale di scienze naturali.

Photos needed for the next Trinidad & Tobago Field Guide

We are making progress on the Trinidad and Tobago Field Guide and the goal to to have it ready for copy editing by the end of 2014. The extended dry season of 2014 has made it difficult to find animals and we still are looking for photos of  species we don't have represented. While it is improbable we will find live specimens of species known from a single specimen or just a few specimens we are looking for the best photography we can find. We are also looking for the best photography of Trinidad and Tobago specimens - that is we would rather have photos of specimens from the islands than from the mainland or other islands if at all possible.

So the following is a list of species we still need, or we need better photos than the ones we have. If you have photos that you are willing to let us use - send me (JCM) an email at: serpentresearch@gmail.com

Leptodactylus nesiotus, Pristimantis turpinorum, Mesoclemmys gibba, Anolis extremus, Anolis trinitatis, Anolis wattsi, Kentropyx striata, Drymarchon corais, Leptophis stimsoni,, Tantilla melanocephala Erythrolamprus cobella, Erythrolamprus ocellatus, Hydrops triangularis, Thamnodynastes ramonriveroi, Micrurus circinalis, Micrurus lemniscatus diutius, Lachesis muta. 

If you have great photos of other species we are interested. 

Bush Maco Trinidad & Tobago and a new Wildlife Policy for T&T

The following announcement  comes from the Bush Maco website. A new effort to document, track and share your Trinidad &  Tobago nature sightings. Bush Maco is just starting out, so they need your support and feedback. Many people can document much more than just one or two can. This is the power of citizen science. Your observations will be used to track the natural cycles of animal and plant species.

Fantastic news for wildlife in Trinidad and Tobago....the Wildlife Policy has been approved by Cabinet! Well done to the hardworking team who wrote it and to all the members of public who contributed to improving this policy through the public consultations!The National Wildlife Policy was formally approved by Cabinet in December, 2013. A Draft Wildlife Policy was developed for Trinidad and Tobago by a Cabinet-appointed Technical Committee and was made available to the general public for comment and public consultations were held during the period January to March, 2013 throughout Trinidad and in Tobago. Additionally, The Ministry of the Environment and Water Resources facilitated special meetings for major stakeholders to elicit views and recommendations on the Draft Policy.

These comments were then reviewed and used to revise the Draft by the Technical Advisory Committee. Once final adjustments were made, the Draft Policy was sent to Cabinet for consideration and on December 19, 2013 Cabinet adopted the National Wildlife Policy for Trinidad and Tobago.

The Ministry of the Environment and Water Resources, the Forest Division and the Tobago House of Assembly express gratitude to all those who participated and contributed to the process of finalising the Policy and as such, enhancing the policy framework for the sustainable management of wildlife resources of the country.  Development of an appropriate legislation and administrative framework for the implementation of the Wildlife Policy is currently underway.

Summary of the National Wildlife Policy

The Wildlife Policy provides guidance on the sustainable management of undomesticated animals and plants, whether introduced, resident or migratory, their parts or derivatives, and their habitats. It addresses issues related to endangerment and provides guidance on the management of threatened species. It also addresses key policy issues on the management of game species, wildlife habitat and the engagement of civil society in the management of wildlife.

Bush Maco is also on Facebook

The bird snakes (Pseustes & Spilotes) rearranged

The genus Pseustes Fitzinger, 1843 is composed of three known species, Pseustes poecilonotus, P. shropshirei and P. sulphureus. Pseuestes sulphureus may be the largest sized colubrid snake in the New World, although Drymarchon corias has a similar or possible greater size. But, both species may exceed 3 meters in total length. Pseustes has been classified as belonging to numerous other genera, over the years, including: Ahaetulla, Chironius, Coluber, Dipsas, Herpetodryas, Natrix, Phrynonax, Spilotes, Synchalinus, Thamnobius and Tropidodipsas. Pyron et al. (2013) found support that Pseustes sulphureus is the sister taxon to Spilotes pullatus, both members of the Colubrinae. However, the estimated phylogenetic position of this species was only based on a single 12S gene fragment as part of a large study. The study, used genes from 4161 species of squamates.  

In a forthcoming paper, Jaden et al. used multiple individuals of Pseustes across Central and South America and they analyzed the genus to infer the phylogenetic position of Pseustes within the Colubrinae, assess the relationship between Spilotes and Pseustes, and determine whether species of the genus Pseustes form a monophyletic group, infer phylogenetic relationships within Pseustes, and species-level diversity to resolve historical taxonomic debates. 

The authors examined four species from multiple specimens across their distribution and analysed one nuclear and two mitochondrial genes to determine the phylogenetic placement of the genus and infer relationships among Pseustes lineages. They found strong support for the paraphyly of Pseustes with respect to the monotypic genus Spilotes, both of which are nested within a clade of at least 23 other New World Colubrinae genera. 

The results produced a new view of the genera Psuestes and Spillotes. The authors resurrected the taxon P. polylepis for populations of P. poecilonotus from South America and moved P. sulphureus to the genus Spilotes which renders both genera monophyletic. Psuestes sulphureus is the type species of the genus Pseustes, and moving it to the genus Spilotes requires the allocation of the senior synonym Phrynonax be considered for the remaining Pseustes taxa.

Citation
Jadin RC., Burbrink FT, Rivas GA, Vitt LJ, Barrio‐Amorós CL, and; Guralnick RP. (2013). Finding arboreal snakes in an evolutionary tree: phylogenetic placement and systematic revision of the Neotropical birdsnakes. Journal of Zoological Systematics and Evolutionary Research. DOI: 10.1111/jzs.12055

New Version of the Herpetology of Trinidad & Tobago Web Site Now On-line

Today the next generation website for the Herpetofauna of T&T was put on-line. I am hoping the new web site and this blog will act to generate interest in the Field Guide. Please look it over and let me know if you encounter problems - with content or web access. Any account lacking accompanying photos means that we need photos for that species. JCM

The Trinidad Plica is now Plica caribeana

The lowland tropics were once though filled with widespread species, while moderate and higher elevations were thought to contain species with more restricted distributions. That idea is turning out to be partially incorrect. Widespread species now appear to be the exception, instead of the rule. A new study describes four species once considered to be the collared treerunner, a lizard known to the scientific community as Plica plica. The study was published in the open access journal ZooKeys.

The Trinidad Plica, is now Plica caribeana

The collared treerunner was originally described in 1758 and has been the subject of many biological, ecological, and behavioral studies in recent years. A new ZooKeys paper by John C. Murphy, Field Museum (Chicago) and Michael J. Jowers, Estación Biológica de Doñana (Sevilla, Spain) describe four new species formerly thought to be one.

“The collard treerunner was considered a single species ranging from Trinidad and Tobago and northern Venezuela southward into the Amazon Basin, south of the Amazon River.” Murphy said.” The Treerunners ancestor diverged about 25-30 million years ago, and throughout this time the South American continent has undergone dramatic remodeling, including the rise of the Andes, rising and falling sea levels, and changing climates that isolated populations for long periods of time, allowing them to become new species. Treerunners live on vertical surfaces, such as tree trunks, rock walls, and even buildings and they eat a variety of insects.

The new paper focuses on populations of this lizard in northern South America, but in an overall survey the authors examined specimens from across the Amazon basin and suspect there may be at least another five to seven undescribed species in what is currently considered the collared treerunner. The treerunners from Trinidad and northern Venezuela were 4.5% genetically different from those in southern Venezuela, and more than 5% different from those in Brazil. For comparison purposes humans and chimpanzees are less than 2% genetically different.

While some species may form by genetic divergence without showing any morphological differences from their ancestor, other often show subtle or obvious morphological differences that may be quite easy to detect. The latter is the case with the collard treerunners.

Some had as few as 92 scales around the body while others had 202 scales around the body. Some adult males have yellow heads while other have red heads, some have distinctive patterns of spots while others have transverse bands.

Unraveling cryptic species is important for a more complete understanding of biodiversity, evolution, and for long term conservation efforts.

The take home message here is that there are many more species of squamate reptiles (lizards and snakes) in the world than previously thought, and it is likely many species have and will disappeared before science is even aware of them. Cutting forests and draining swamps undoubtedly causes extinctions of the species depending upon those habitats. While none of the treerunners described in this paper are likely to be threatened with extinction this discovery and many other similar recent discoveries suggest our knowledge of biodiversity is lacking.

Citation
Murphy JC, Jowers MJ (2013) Treerunners, cryptic lizards of the Plica plica group (Squamata, Sauria, Tropiduridae) of northern South America. ZooKeys 355: 49–77. doi:10.3897/zookeys.355.5868

A new snake species from Tobago

Until now two snake species of the genus Leptophis (parrot snakes) have been known to  inhabit Trinidad and Tobago. One, L. stimsoni, is endemic to Trinidad’s Northern Range and known from relatively few specimens. The second is the diurnal, arboreal, brightly colored parrot snake Leptophis coeruleodorsus.

The genus Leptophis was first reported in Trinidad and Tobago by Reinhardt & Lütken (1862), and while it appeared on lists, no natural history information was reported until Mole (1924) described its diet. Oliver described the Trinidad & Tobago populations of Leptophis as  L. coeruleodorsus in 1942, and in 1948 re-classified it as a subspecies of L. ahaetulla. Oliver viewed Leptophis ahaetulla as a species with 147–183 ventrals, 135–185 subcaudals, 18–28 maxillary teeth, and a distribution extending from about 22º N latitude to 35ºS latitude (a straight line north-south distance of about 7500 km). Oliver’s 13 subspecies were based primarily on coloration, as well as ventral and subcaudal counts and he viewed much of the morphological variation as clinal.

The second known species of Leptophis on Trinidad is Leptophis stimsoni Harding, a Northern Range endemic and a member of the L. riveti Group. The first specimen was collected by Sanderson in 1937 from the summit of Mt. Aripo (Cerro del Aripo), and was considered to be L. riveti  by Emsley (1977). No other specimens were known until 1987 when two specimens, one from the Arima Valley and another from Cumaca Cave were collected. Harding (1995) described L. stimsoni as distinct from L. riveti.

Murphy et al. (2013) compared 11 specimens of the L. ahaetulla Group using DNA sequences from two mitochondrial genes (cytochrome b and 16S, 1,383 bp total) from island and mainland populations, reported on the variation in the morphology of 54 museum specimens of Leptophis a. coeruleodorsus; restrict the type locality of L. coeruleodorsus to Mt. St. Benedict; and they describe a new species of Leptophis from the island of Tobago that can be distinguished from L. coeruleodorsus on the basis of snout shape, upper labial architecture, elongated prefrontal scales, and ventral scale counts.

Leptophis haileyi is the only Tobago Leptophis to have a subacuminate snout in profile; the rostral is barely visible from above; the primary temporal is in contact with three or four upper labials, including the last; prefrontal scale is long, it has a high ventral count (173) and it is a male, with a tail/SVL ratio that falls with in the range of female L. coeruelodorsus.   All L. coeruleodorsus examined have two upper labials contacting primary temporal with last upper labial excluded from contact with primary temporal.

(a) Leptophis coeruleodorsus (b) L. haileyi

The new species is currently known from a single specimen from the northeast portion of Tobago in the vicinity of Roxborough and is named in honor of Adrian Hailey for his work on the Trinidad and Tobago herpetofauna. This description raises the number of endemic Tobago amphibians and reptiles to 11 taxa.

Citation
MURPHY, J. C., CHARLES, S. P., LEHTINEN, R. M., & KOELLER, K. L. (2013). A molecular and morphological characterization of Oliver’s parrot snake, Leptophis coeruleodorsus (Squamata: Serpentes: Colubridae) with the description of a new species from Tobago. Zootaxa, 3718(6), 561-574.

Trinidad Sphaerodactylus, I am looking for a banded one!

The dwarf day geckos of the genus Sphaerodactylus are primarily lizards of the Caribbean. The Reptile Database lists 101 currently recognized. Relatively few species are known from mainland Central and South America. Harris (1982 Occas. Pap Mus Zool. Univ Michigam  no. 704) reported five South American  spercies (S. heliconiae, S. lineolatus, S. molei, S. scapularis, and S. notatus). Species described since 1982 have been either Central American or West Indian (mostly Cuba and Hispanolia) and the only species present on Trinidad and Tobago has been thought to be S. molei, a gecko also in coastal Venezuela and Guyana. However, two musuem specimens with the locality data of "Trinidad" have been found that are clearly not S. molei.  I am writing this with the idea that readers will find this lizard. It is distinctive in that the body is transversely banded; and molei has longitudinal stripes. It is possible the locality data is wrong, it is possible the lizards were accidentally introduced from another island, but the specimens are not in great shape and getting scale count data and  other scale characteristics from them is at best problematic. So, should you find a tiny banded lizard, a Sphaerodactylus on Trinidad, send me an email or please take it to the Zoology Museum at UWI so we can identify it. Photos of a juvenile S. molei and one of the banded species are shown below.

The Second Trinidad Bioblitz

The second annual Trinidad Bioblitz has come and gone.  This year it was held in the very biodiverese Arima Valley, with the base camp at the Asa Wright Nature Center. The preliminary tally of species was - Vertebrates 139, Invertebrates 247, Fungus 30, Diatoms 7, Plants 317 for a grand total of 740 species!

Teams include: Mammals, Freshwater (Fish, Aquatic Insects, Crustaceans), Amphibia and Reptilia, Birds, Molluscs, Spiders and other Arachnids, Butterflies and Moths, Social Insects, Myriapods millipedes/ centipedes) and Worms, Flowering Plants, Ferns, Lichens, Orchids and Fungi.

The Arima Valley has been the crown jewel for Trinidad's biodiversity for centuries, the site has been the geographical location for much of the scientific research that has occurred on Trinidad including classic studies of bats, guppies, and the herpetofauna as well as many other higher taxa. However, active quarrying operations threaten the habitat, and the activity of bush meat hunters threaten larger animal populations. But, perhaps the most significant sign that the valley is under development - the installation of sidewalks along the road.

But, the Bioblitz results suggest the flora and fauna are doing well at least for the moment. Teams of experts and volunteers  searched the valley for 24 hours attempting to identify as many species as possible. But, it can be more than just a list, because the opportunity of natural history observations is under every log and in every fruiting tree.

 Informational displays by various groups including the Trinidad and Tobago Field Naturalist Club, UWI Zoology Museum, Asa Wright Nature Center and Wildlife Division were present to  inform the public. So what about the herpetofauana. Here are some photos of what was found.

An Arima Valley Bothrops cf asper. was sitting in the vegetation close to a
 chorus of Hypsiboans crepitans.

Top: a juvenile Chironius carinatus, bottom Chironius (Macrops)
septentrionalis. Both juveniles and very difficult to distinguish
from each other, but look at the relative size of the eyes

A Hypisboans crepitans hiding in the vegetation during the day.

Epictia tenella was found climbing a tree trunk, it was about a meter
off the ground and was probably headed for a rotten liana filled with
termites.

A juvenile Oxyrhopus petolarius

Anolis planiceps sleeping on a leaf.

A male Gonatodoes vittatus perhaps the most common seen
reptile during the Bioblitz

The leaf-nesting frog, Phyllomedusa trinitatus, 

Generic changes likely for some Trinidad Chironius

Convergent evolution sometimes covers the tracks of genetic lineages, so species can appear to be very similar but have different ancestors. Such appears to be the case with some of the snakes currently in the genus Chironius.  In Trinidad & Tobago, there are three species of Chironius, Machete savane, C. chironius chironius, the long-tailed machete, C. septentrionalis, and the smooth machete C. surrulus (this species is known from only a few specimens on Trinidad).  The sixteen species in the genus tend to have triangular shaped bodies in cross-section, large eyes, 14 or few scale rows at mid body,  they to to actively forage during the day and feed on frogs and lizards. They range from Central America to southern South America and inhabit some of the Lesser Antilles.

In a new study of squamate relationships, Pyron et al. (2013) found Chironius to be polyphyletic, with C, carinatus and C. quadricarinatus forming a clade and with nine other species that were included in the study, forming a second clade. The second clade contains C. septentrionalis and  C. scurrulus.
 Chironius carinatus is the type species of the genus, thus it and quadricarinatus (and whatever other species not included in the study are related to it) will retain the name Chironius. The second, however, will need a new name, and the oldest name available for the clade will likely be Macrops (= large eyes), a genus established by Wagler in 1830 for Linnaeus' species Coluber saturnius, which is now known as Chironius fuscus.

This result demonstrates how little we actually know about the Neotropical Herpetofauna, the prevalence of species that look similar but are genetically distinct, and the problems caused by the polytypic species concept. Sorting out the details of this promises to be challenging because several of the currently recognized species of Chironius have subspecies, including C. carinatus.

Citation
Pyron, RA, Burbrink FT, Wiens JJ. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13:93 doi:10.1186/1471-2148-13-93