The booid snakes (superfamily Booidae) are a near–circumglobally distributed group of macrostoman alethinophidian squamates, and several lineages are of significant conservation concern. A number of taxonomic changes have occurred among the superfamily Booidae over the last decade, including the resurrection and description of new families, elevation of a genus, elevation of 13 species, and the discovery of a new species. Here, we aim to synthesize existing knowledge of booid diversity, systematics, and conservation status. We provide a comprehensive checklist of all 66 species and 33 subspecies of booid snakes recognized herein, distributed among 14 genera and six families. For each species and subspecies, we evaluate taxonomy, distribution, type specimens, and conservation status.
INTRODUCTION
The first checklist of the boid (family Boidae Gray, 1825) snakes by Stull (1935) recognized 66 taxa (39 species, 27 subspecies) among 15 genera of what was then considered Boinae Gray 1825, a subfamily of Boidae (which also included Pythoninae and Loxocemus). Since that time, our understanding of the diversity and systematics of this group has changed a great deal (Table 1). For example, Stull's (1935) list of Boinae included the genera Tropidophis, Bolyeria, and Casarea; they are now considered representatives of distinct families (Tropidophiidae and Bolyeriidae) of henophidian snakes (e.g., Reynolds et al., 2014). Removing the taxa presently excluded from the booids, Stull's (1935) treatment included 30 species among 12 genera. In the decades following the publication of Stull's list, boid systematics were frequently revised and rearranged. Stimson (1969) published an updated checklist, recognizing 14 boid genera, 39 species, and 58 subspecies, including Xenoboa (=Corallus) cropanii, a genus no longer recognized. Kluge (1991) provided a de facto checklist of boids; he recognized 25 species among 8 genera (including Xenoboa). Since Kluge (1991), major reorganizations of boid genera have occurred over the last 10 years (e.g., Passos and Fernandes, 2008; Rivera et al., 2011; Reynolds et al., 2013a). Recent larger scale, species-level, molecular phylogenies (Pyron et al., 2013; Reynolds et al., 2014) have further suggested a historically incomplete representation of booid lineages and some discordance between taxonomy and phylogenetic relationships. These authors made a number of taxonomic rearrangements and suggestions, and subsequent work has accepted, expanded, or rejected these recommendations (Reeder et al., 2015; Figueroa et al., 2016; Streicher and Wiens, 2016; Zheng and Wiens, 2016; Uetz et al., 2017). Beginning with higher level systematics, these molecular phylogenies demonstrated inconsistencies in the placement of the booid family Calabariidae with respect to other alethinophidian lineages (Pyron et al., 2013; Reynolds et al., 2014; Harrington and Reeder, 2017). To resolve this, Pyron et al. (2014) erected additional booid families to accommodate distinct monophyletic lineages, simultaneously alleviating taxonomic issues related to phylogenetic uncertainty deeper in the booid phylogeny (the paraphyly of Boidae Gray 1825 given inconsistent placement of Calabariidae). This led to a narrowed definition of the Boidae to include only the New World genera Boa, Chilabothrus, Corallus, Epicrates, and Eunectes, all derived from an ancestral lineage in the Paleogene (Hsiang et al., 2015). Newly recognized families include Sanziniidae (Acrantophis and Sanzinia), Erycidae (Eryx), Charinidae (Charina, Exiliboa, Lichanura, Ungaliophis), and Candoiidae (Candoia). Not all workers embrace these changes, instead opting to remain agnostic regarding potential paraphyly until further phylogenetic relationships are worked out. Nevertheless, the recognition of families representing geographically, morphologically, and evolutionarily distinct lineages provides stability in the systematics and taxonomy of the group now and into the future, despite ongoing uncertainty in some phylogenetic relationships (Pyron et al., 2014). Continued efforts are ongoing to resolve some of the more challenging nodes in the booid phylogeny, including the application of genomic-scale data (e.g., Ruane and Austin, 2017) that suggests Calabariidae might be sister to the rest of the extant booids (Fig. 1).
Table 1.
The number of genera, species, and subspecies treated in checklists or systematic accounts of the Booidae since 1935.
![i0027-4100-162-1-1-t01.tif](ContentImages/Journals/harv/162/1/MCZ48.1/graphic/WebImages/i0027-4100-162-1-1-t01.jpg)
Figure 1.
Time-calibrated phylogeny of booid snakes based on an 11-gene supermatrix (after Reynolds et al., 2014). Calibration of the age of crown snakes was set to 145 mya (close to the mean inferred in Zheng and Weins, 2016), and the tree was inferred using the Bayesian algorithm implemented in the program BEAST v.1.8 (Drummond et al., 2012). This figure is therefore merely intended to show an approximation of divergence times and evolutionary relationships among booid genera to illustrate concepts discussed in the text.
![i0027-4100-162-1-1-f01.tif](ContentImages/Journals/harv/162/1/MCZ48.1/graphic/WebImages/i0027-4100-162-1-1-f01.jpg)
Additional recent taxonomic changes within the Booidae include the resurrection of the genus Chilabothrus for some West Indian boids. Thirteen new species of booids have also recently been recognized, largely resulting from phylogenetic studies of molecular data (Passos and Fernandes, 2008; Wood et al., 2008; Reynolds et al., 2013a, 2014; Card et al., 2016; Reynolds et al., 2018) or newly discovered species (Reynolds et al., 2016a).
Members of the superfamily Booidae are of Gondwanan origins (Noonan and Chippindale, 2006) and are distributed nearly circumglobally (Fig. 2). Major lineages are present in the Western hemisphere (Boidae + Charinidae; 43 species), Africa (Calabariidae + Erycidae; four species), Eurasia (Erycidae; 10 species), Madagascar (Sanziniidae; four species), and Oceania exclusive of Australia (Candoiidae; five species). Fossil booids are known from regions where they do not presently have extant representatives, such as Eastern North America (Holman, 1998; Mead and Schubert, 2013) and Western Europe outside the distal Balkan Peninsula (Szyndlar, 1991, 2009).
Booid taxonomy and systematics have experienced a considerable amount of flux over the last several decades, motivating a re-evaluation of the current state of taxonomy and diversity within the superfamily (sensu Pyron et al., 2014). This is especially relevant given the largely unknown conservation status of many of the world's booid snakes (Böhm et al., 2013; IUCN 2017) and the worrying prospects of some of those that have been evaluated (Tzika et al., 2008; Reynolds, 2011; Reynolds et al., 2016a; this work). Below, we provide generic and species accounts for all recognized species and subspecies of booids. For each account, we provide the taxonomic authority, a brief taxonomic history, type specimens, distribution, and conservation information for the species and subspecies. Unless we saw a specific need to do so, our checklist does not repeat lengthy synonymies that are available elsewhere (e.g., the excellent McDiarmid et al., 1999). All but one species of boa (superfamily Booidae, formerly family Boidae) are protected under the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). Six species are listed under CITES Appendix I and are noted below; the rest are listed under CITES Appendix II. Additional conservation designations have been assigned to some taxa, which are also noted. Our taxonomic presentation largely follows from McDiarmid et al., 1999, Wallach et al. (2014), Pyron et al. (2014), and Uetz et al. (2017), and these references contain full taxonomic histories for these species. We re-evaluate standing taxonomy from these sources and make several suggestions for taxonomic revision, recognizing 14 genera (Fig. 1), 66 species, and 33 subspecies of booid snakes. We have opted not to assign common names to the various species. We were uncomfortable imposing English language names because the majority of the species in this checklist occur in countries in which the native language is not English. A representative photograph is provided for each genus.
SPECIES ACCOUNTS FAMILY BOIDAE GRAY, 1825
Boa Linnaeus, 1758
Apart from a brief visit by the Malagasy boids (Acrantophis and Sanzinia), Boa or Constrictor has been considered a monotypic genus for over 100 years (1906–2009). Five species of Boa are now recognized based on morphological data, molecular data, or both. The genus has a vast mainland distribution, from northern Mexico to southern South America in Argentina, Paraguay, and Brazil, as well as continental and oceanic (St. Lucia and Dominica in the Lesser Antilles) islands. Species of Boa occur in a wide array of habitats, ranging from near-desert circumstances to tropical rainforests, and from sea level to about 2,000 m. They range in size from dwarfed island populations (of B. imperator) to >4.0 m in some South American B. constrictor. Although largely ground dwelling, they are capable of arboreal activity and will hunt in trees. Species of Boa take a wide taxonomic array of prey, including lizards, birds, and mammals (including marsupials, rodents, carnivores, bats, and primates). In recent years, species of Boa have been accidentally or purposely introduced to geographic areas where they formerly did not occur, with the potential of negatively affecting wildlife native to those areas (e.g., Quick et al., 2005; Romero-Nájera et al., 2007). Boa is sister to a clade comprising Chilabothrus + Corallus + Epicrates + Eunectes.
Table 2.
List of museum abbreviations used in the text with corresponding institution and location.
![i0027-4100-162-1-1-t02.tif](ContentImages/Journals/harv/162/1/MCZ48.1/graphic/WebImages/i0027-4100-162-1-1-t02.jpg)
Boa constrictor Linnaeus, 1758
Taxonomy. Originally described as Boa constrictor, this species has had a long taxonomic history placed in either Boa or Constrictor. Many names have been assigned to those two genera, and until recently, all have been placed into synonymy or have been described originally as subspecies or have been relegated to subspecific rank. Only recently has B. constrictor been partitioned into multiple species (Henderson and Powell, 2009; Hynková et al., 2009; Reynolds et al., 2014; Suárez-Atilano et al., 2014, 2017; Card et al., 2016). See McDiarmid et al., 1999 for a more complete synonymy. A number of subspecific epithets continue to be used by some sources (e.g., Uetz et al., 2017) but are not recognized by others (this work). This is partly owing to the preliminary molecular phylogenetic surveys of the genus, the unknown provenance of some samples used in these analyses (e.g., Hynková et al., 2009), and lack of a comprehensive morphological and molecular study of the genus. For example, B. c. amarali Stull 1932 is an epithet used to refer to populations from southern Brazil, possessing lower numbers of scale rows, ventrals, and caudals (Stull, 1932), but is not supported as distinct in other analyses (Hynková et al., 2009). We recognize four subspecies.
Type Specimens. Two syntypes, NRM 10 and NRM 20001, a third syntype is presumed lost (McDiarmid et al., 1999). Type locality “Indiis” (in error, fide Peters and Orejas-Miranda, 1970).
Boa constrictor constrictor Linnaeus, 1758
Taxonomy. Originally described as a full species, but subsequently relegated to subspecies rank with the description of multiple taxa that were either described as full species of Boa or Constrictor and then relegated to a subspecies of B. or C. constrictor, or originally described as subspecies of B. constrictor or C. constrictor. See McDiarmid et al., 1999 for a more thorough synonymy.
Boa constrictor longicauda Price & Russo, 1991
Taxonomy. Described as a subspecies of Boa constrictor based on tail length, color pattern, and scale characters.
Boa constrictor occidentalis Philippi, 1873
Taxonomy. Originally described as Boa occidentalis; Ihering (1910) subsumed it to a subspecies within Constrictor constrictor; Forcart used the trinomial Boa c. occidentalis. Bezerra de Lima (2016) considered B. occidentalis a distinct lineage within the B. constrictor complex.
Type Specimen. The type is unlocated (McDiarmid et al., 1999); type locality (fide Stimson, 1969) “Provinces Mendoza and San Juan, Argentina.”
Boa constrictor ortonii Cope, 1878
Taxonomy. Originally described as Boa ortonii; relegated to a subspecies of Constrictor constrictor by Schmidt and Walker (1943); Stimson (1969) was first to use the trinomial B. c. ortonii.
Type Specimen. ANSP 11402, “from Chilete, near Pacasmayo, 3000 feet [915 m] above the sea,” Peru (Cope, 1878:35).
Boa imperator Daudin, 1803
Taxonomy. Originally described as a full species, it was subsequently subsumed to a subspecies of Constrictor constrictor by Ihering (1910); Forcart (1951) recognized it as B. c. imperator; based on molecular data, a number of authors (Hynková et al., 2009; Reynolds et al., 2014; Suárez-Atilano et al., 2014, 2017; Card et al., 2016) suggested elevating it to full species. Bezerra de Lima (2016) considered B. imperator a distinct lineage within the B. constrictor complex, and this species is being recognized in contemporary treatments (e.g., Johnson et al., 2015; García-Padilla et al., 2016). Two subspecies are recognized.
Type Specimen. In the MNHN, but not definitely identifiable (J. Guibé in Stuart, 1963). Type locality “l'Amerique meridionale principalement au Mexique” was subsequently restricted to Córdoba, Veracruz, Mexico, by Smith and Taylor (1950); however, Dunn and Saxe (1950) favored the Colombian Chocó as the type locality.
Distribution. Southeastern Mexico, Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, and northwestern Colombia; includes many islands off the Caribbean/Atlantic and Pacific coasts of several of those countries. Introduced to Cozumel Island (Vázquez-Domínguez et al., 2012) and St. Croix (Golden, 2017).
Conservation Status. This species has not received an IUCN Red List assessment. Using IUCN Red List criteria, Acevedo et al. (2010) categorized this species as of Least Concern in Guatemala, as did Greenbaum and Komar (2010) for El Salvador, Townsend and Wilson (2010) for Honduras, and Sosa et al. (2010) for Costa Rica; Jaramillo et al. (2010) gave B. imperator an assessment of Vulnerable for Panama. Stafford et al. (2010) gave it a low Environmental Vulnerability Score for Belize.
Boa imperator imperator Daudin, 1803
Distribution. Southeastern Mexico, Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, and northwestern Colombia; includes many islands off the Caribbean/Atlantic and Pacific coasts of several of those countries (excluding the Islas de las Perlas in the Gulf of Panama). Introduced to Cozumel Island (Vázquez-Domínguez et al., 2012) and St. Croix (Golden, 2017).
Conservation Status. This taxon has not received an IUCN Red List assessment. Certain populations of B. i. imperator on the Bay Islands of Honduras have been severely affected by poaching for the pet trade, though some populations have recovered after protection in the Cayos Cochinos Archipelago Natural Marine Monument (Wilson and Cruz Diaz, 1993; Reed et al., 2007; Montgomery et al., 2015).
Boa imperator sabogae (Barbour, 1906)
Taxonomy. Originally described as Epicrates sabogae. Barbour and Loveridge (1929) considered it a subspecies of Constrictor constrictor; Forcart (1951) used Boa c. sabogae; Reynolds et al. (2014), recognizing B. imperator as a full species, used the trinomial B. i. sabogae.
Type Specimens. Description based on two syntypes (MCZ R6986) from Saboga Island, one of the Islas de las Perlas off the Pacific coast of Panama.
Boa nebulosa (Lazell, 1964)
Taxonomy. Originally described as a subspecies of Constrictor constrictor, it was elevated to species status by Henderson and Powell (2009) based on scale and pattern characters provided by Lazell (1964), as well as geographic isolation. Before Lazell's description, the boa population on Dominica was referred to as B. diviniloqua Günther (1888) and then fell under the taxonomic umbrella of the St. Lucia population (Constrictor orophias; e.g., Barbour, 1930, 1937). Bezerra de Lima (2016) considered B. nebulosa a distinct lineage within the B. constrictor complex. No subspecies are recognized.
Conservation Status. This species has been assessed as Least Concern based on IUCN Red List criteria, though that assessment is in review. Its distribution is limited to a small (790 km2) island and should certainly be assessed as Vulnerable. These boas are killed for the “medicinal” oil rendered from their fat and because they prey on domestic chickens (Henderson and Powell, 2009).
Boa orophias Linnaeus, 1758
Taxonomy. Originally described as a full species by Linnaeus, it was placed in the synonymy of Constrictor diviniloquus (or diviniloqua or diviniloquax) by Laurenti (1768) and subsequently by Duméril and Bibron (1844). Barbour (1914) regarded it as full species (C. orophias), but it was subsequently downgraded to subspecies by Amaral (1929). It was again elevated to species status by Stull (1935) as C. orophias, but Lazell (1964) considered it a subspecies of C. constrictor, as did Peters and Orejas-Miranda (1970; as Boa c. orophias). It was then given species rank by Henderson and Powell (2009) based on scale and pattern characters in Lazell (1964), as well as geography. Bezerra de Lima (2016) considered B. orophias a distinct lineage within the B. constrictor complex. No subspecies are recognized.
Boa sigma (Smith, 1943)
Taxonomy. Originally described as Constrictor constrictor sigma (mistakenly attributed to M. A. Smith [1943] by McDiarmid et al. [1999]); Zweifel (1960) relegated it to the synonymy of Boa constrictor imperator. Card et al. (2016) recommended elevation to species level based largely on molecular data, while Suárez-Atilano et al. (2017) suggest that the species is further defined ecologically and geographically. No subspecies are recognized.
Type Specimen. An adult female (USNM 46484) from María Madre Island in the Tres Marías Islands, Nayarit, Mexico (H. M. Smith, 1943).
Chilabothrus Duméril & Bibron, 1844
The genus Chilabothrus encompasses 13 recognized species restricted to the Greater Antillean Islands of Puerto Rico, Jamaica, Hispaniola, and the Puerto Rico Bank; as well as the Lucayan Archipelago (Bahamas and Turks and Caicos). These actively foraging nocturnal booids range in body size from <1 m to ∼4 m, with larger species being habitat and dietary generalists and smaller species frequently specializing (Rodriguez-Robles and Greene, 1996; Reynolds et al., 2016c). Members of Chilabothrus were considered to be congeneric with the genus Epicrates (e.g., Tolson, 1987), which is restricted to mainland Central and South America (and some continental islands). Phylogenetic analyses revealed that mainland Epicrates are phylogenetically sister to Eunectes (anacondas) and that West Indian representatives formed a monophyletic clade dating to the Miocene (Reynolds et al., 2013a). In addition to the recent elevation of the available name Chilabothrus for the West Indian clade, several taxonomic changes have occurred recently within the genus, including the elevation of three lineages to species (Reynolds et al., 2013a, 2018; Rodríguez-Robles et al., 2015), as well as the discovery of a previously unknown species (Reynolds et al., 2016a).
Chilabothrus angulifer (Bibron, 1840)
Taxonomy. Original name was Epicrates angulifer, generally attributed to Cocteau and Bibron (1838), though thought to be the description of Bibron (Smith and Grant, 1958) and occasionally given as Bibron 1843 (e.g., Henderson and Arias, 2001; Frynta et al., 2016). Genus was changed to Chilabothrus (Reynolds et al., 2013a). No subspecies are recognized.
Distribution. A species endemic to the island of Cuba and associated satellite islands (Sheplan and Schwartz, 1974; Schwartz and Henderson, 1991; Henderson and Arias, 2001). No subspecies are recognized, though a large degree of phenotypic variation is present across the island (Schwartz and Henderson, 1991; Henderson and Arias, 2001; Rodríguez-Cabrera et al., 2016).
Conservation Status. IUCN Red List Near Threatened (Day and Tolson, 1996). Although having a wide distribution and being locally common, this species is frequently persecuted when human encounters occur (Gundlach, 1880; Day and Tolson, 1996) and habitat loss has likely contributed to local declines (Tolson and Henderson, 1993). The species is also likely subject to mortality owing to vehicle strikes and invasive vertebrate predators (Rodríguez-Cabrera et al., 2016).
Chilabothrus argentum (Reynolds et al., 2016a)
Taxonomy. Discovered in situ in 2015, the species was named Chilabothrus argentum, as a lineage distinct from other members of the genus. No subspecies are recognized.
Distribution. A species endemic to the Conception Island Bank, located in the central Bahamas Archipelago. No additional populations or subspecies are known. Previous species lists for the Conception Island Bank included C. striatus (now C. strigilatus Reynolds et al., 2013a) based on anecdotal suggestion of a boa present on the bank (Schwartz et al., 1978; Franz and Buckner, 1998; Buckner et al., 2012).
Conservation Status. IUCN Red List Critically Endangered (Reynolds, 2017) based on extremely small extent of occurrence (EOO, a polygon containing all known populations) and area of occupancy (AOO, the actual area occupied within the EOO polygon), likely declining population size, and single known population (Reynolds et al., 2016a).
Chilabothrus chrysogaster (Cope, 1871)
Taxonomy. Originally described as Homalochilus chrysogaster (Cope, 1871), Stejneger (1904) moved it to Epicrates. It was subsequently moved into Chilabothrus with other West Indian Epicrates (Reynolds et al., 2013a). Two subspecies are recognized (Buden, 1975; see the C. schwartzi account).
Type Specimen. Holotype ANSP 10322, an adult of unknown sex. The holotype has since been lost, though presumably from “Turks Island,” a locality name that might have referred to either Grand Turk Island or South Caicos Island in the 19th century.
Chilabothrus chrysogaster chrysogaster (Cope, 1871)
Taxonomy. Considered to be a subspecies of Epicrates striatus by Stull (1935), and later as a subspecies of E. chrysogaster by Sheplan and Schwartz (1974).
Chilabothrus chrysogaster relicquus (Barbour & Shreve, 1935)
Taxonomy. Originally described as Epicrates relicquus by Barbour and Shreve (1935; a misspelling of relicqus, meaning relict); it was considered a subspecies by Sheplan and Schwartz (1974).
Chilabothrus exsul (Netting & Goin, 1944)
Taxonomy. Described from a specimen collected on Abaco Island (Netting and Goin, 1944); subsequently placed in the genus Chilabothrus (Reynolds et al., 2013a). No subspecies are recognized.
Distribution. Little Bahamas Bank, though questionable records exist for Grand Bahama (Schwartz and Henderson, 1991; Reynolds et al., 2016b). Most records are from the Abaco islands.
Conservation Status. IUCN Red List Vulnerable (Reynolds and Buckner, 2016). Threats include invasive vertebrate predators, habitat loss, persecution, and significant road mortality (Reynolds et al., 2016b). A possible extirpation from Strangers Cay, Bahamas (Netting and Goin, 1944; Henderson and Powell, 2009) likely represents an extirpation of Cubophis vudii and not C. exsul (Netting and Goin, 1944).
Chilabothrus fordii (Günther, 1861)
Taxonomy. Original name was Pelophilus fordii. Taxonomic changes included Chilabothrus maculatus Fischer 1888, Epicrates fordi Boulenger 1893 (subsequently referred to as Epicrates fordii), and Chilabothrus fordii Reynolds et al. 2013. Sheplan and Schwartz (1974) noted that the correct specific name is fordii, but that fordi should have been the proper name because the species is named for the individual Ford. According to Article 32.5 of the International Code for Zoological Nomenclature (ICZN), the original spelling does not constitute an “inadvertent error”; thus, the original specific epithet fordii stands. Subsequent authors have used both spellings, occasionally using the spellings interchangeably in separate treatments. Earlier works frequently used fordi (e.g., Schwartz, 1979; Henderson et al., 1987; Tolson, 1992; Tzika et al., 2008; Tolson and Henderson, 2011), while a mixture of older and most recent works recognize ICZN authority and use the spelling fordii (e.g., Tolson, 1987; Kluge, 1989; Reynolds et al., 2013a, 2014, 2015, 2016b,c). Three subspecies are recognized (Schwartz and Henderson, 1991; Tolson and Henderson, 1993; Henderson and Powell, 2004).
Type Specimen. Holotype BMNH 1946.1.1.55 (previously BMNH 1862.3.10.4), an adult female from the Dominican Republic (Wetherbee, 1987).
Chilabothrus fordii fordii (Günther, 1861)
Taxonomy. Stull (1935) considered Epicrates fordii to be a subspecies of E. inornatus; that same year, Barbour considered it to be a subspecies of E. fordii, along with E. f. monensis. Sheplan and Schwartz (1974) eventually sorted out the taxonomy, recognizing the subspecies.
Chilabothrus fordii agametus (Sheplan & Schwartz, 1974)
Taxonomy. Subspecies was described from a single male collected in 1960 by A.S. Rand and J.D. Lazell (Sheplan and Schwartz, 1974).
Type Specimen. MCZ R 62656, an adult male collected near Mole Saint-Nicholas, Département du Nord-Ouest, Haiti (Sheplan and Schwartz, 1974).
Chilabothrus fordii manototus (Schwartz, 1979)
Taxonomy. The original description of Epicrates fordi manototus was based on two specimens obtained by R. Thomas in 1966 and D. A. Daniels in 1976 (Schwartz, 1979).
Type Specimens. Holotype CM 60519, an adult female from Île á Cabrit, Département de l'Ouest, Haiti (Schwartz, 1979).
Chilabothrus gracilis Fischer, 1888
Taxonomy. Originally described as Chilabothrus gracilis. Boulenger (1893) placed it in Epicrates, then back to Chilabothrus when resurrected by Reynolds et al. (2013a).
Type Specimens. Two syntypes in the ZMH, now destroyed (Sheplan and Schwartz, 1974); type locality “Cap Hayti” (=Cap-Haïtien), Département du Nord, Haiti.
Chilabothrus gracilis gracilis Fischer, 1888
Taxonomy. First use of the trinomial was by Stull (1935) when she considered the taxon monensis to be a subspecies of E. gracilis; Sheplan and Schwartz (1974) provided a convincing argument that monensis was not a subspecies of E. gracilis (see C. monensis account).
Chilabothrus gracilis hapalus (Sheplan & Schwartz, 1974)
Chilabothrus granti (Stull, 1933)
Taxonomy. Originally described as a subspecies of Epicrates inornatus, Sheplan and Schwartz (1974) provided a new combination, identifying granti as a subspecies of Epicrates monensis. It had been informally referred to as a full species (e.g., Harvey and Platenberg, 2009; Platenberg and Harvey, 2010) owing to perceived uniqueness of this lineage as well as for conservation purposes. Rodríguez-Robles et al. (2015) provided a more thorough assessment for the recognition of Chilabothrus granti as a species distinct from C. monensis, further supported by Reynolds et al. (2015). No subspecies are recognized.
Distribution. The Puerto Rico Bank: northeastern Puerto Rico, Cayo Diablo, Culebra, St. Thomas, Jost Van Dyke, Tortola, Great Camanoe, and perhaps Guana (Rodríguez-Robles et al., 2015).
Conservation Status. The IUCN Red List has designated this species Endangered, largely because of continuing habitat destruction (Tolson, 1996a; Platenberg and Boulon, 2011); see also Reynolds et al. (2015). Although not currently CITES listed, it presumably would be considered a CITES Appendix I species.
Chilabothrus inornatus (Reinhardt, 1843)
Taxonomy. Originally described as Boa inornata (Reinhardt, 1843), it was subsequently placed in the genera Chilabothrus (Duméril & Bibron, 1844) and Epicrates (Boulenger, 1893). Reynolds et al. (2013a) resurrected the genus Chilabothrus to encompass the Greater Antillean boids. No subspecies are recognized.
Distribution. Currently restricted to the main island of Puerto Rico (Rivero, 1998), though a few individuals likely exist on Culebra Island and could represent a recent introduction (R.G.R., personal observation).
Conservation Status. IUCN Red List Least Concern (Mayer and Tolson, 2010) and CITES Appendix I. Threats include invasive vertebrate predators, habitat destruction, persecution, road mortality, and historical collection for liver oil extraction (Reagan, 1984; Wiley, 2003; Mayer and Tolson, 2010; USFWS, 2011). This species is widely considered to have recovered from the near-complete deforestation of the island of Puerto Rico in the early 20th century. The island is now reforested in many areas, and boas are common in more remote situations (Puente-Rolón, 2012). This species is also now readily found near human habitation and can persist in small habitat patches (Puente-Rolón et al., 2013). The species was likely extirpated from Vieques Island.
Chilabothrus monensis (Zenneck, 1898)
Taxonomy. Originally described as Epicrates monensis. Stull (1935) considered it a subspecies of E. gracilis, but Sheplan and Schwartz (1974) provided sound evidence for its recognition as distinct from E. gracilis (or E. inornatus). See also the Chilabothrus granti account. No subspecies are recognized.
Type Specimens. Five syntypes from Isla Mona in the ZMH (now destroyed; Sheplan and Schwartz, 1974).
Conservation Status. The IUCN Red List has designated this species Endangered (Tolson, 1996b); it is listed under CITES Appendix I (i.e., the most endangered and threatened with extinction). Threats include invasive vertebrate predators, especially cats, as well as habitat destruction owing to invasive pigs and rodents. As many as 70% of boas have scars or injuries caused by feral cats (Tolson, 1996b).
Chilabothrus schwartzi (Buden, 1975)
Taxonomy. Previously described as a subspecies of the Southern Bahamas boa (Epicrates chrysogaster schwartzi) by Buden (1975). This description was based on deceased animals and only one intact specimen (the holotype). The species was elevated based on the discovery of four wild individuals in 2018 and subsequent morphological and molecular phylogenetic analyses (Reynolds et al., 2018). No subspecies are recognized.
Distribution. A newly-recognized boa species endemic to the Crooked-Acklins Bank, southern Bahamas. Known from four museum specimens (KUH 260082-84; LSUMZ 27500) and four wild specimens (MCZ HO 28-31; Schwartz and Henderson, 1991; Reynolds, 2012; Buckner et al., 2012; Reynolds et al., 2018).
Conservation Status. This species has not been assessed based on IUCN Red List criteria. The species likely occurs across two large islands, though it is apparently rare or restricted to specific areas of these islands (Reynolds et al., 2018). Threats to the species are unknown, but likely include persecution, road mortality, and introduced vertebrate predators.
Chilabothrus striatus (Fischer, 1856)
Taxonomy. Originally described as Homalochilus striatus by Fischer (1856); Boulenger (1893) provided the first use of Epicrates striatus. A number of subspecies have been described, but not all are currently recognized—three are recognized here. See McDiarmid et al., 1999 for a more complete synonymy.
Chilabothrus striatus striatus (Fischer, 1856)
Taxonomy. Originally described as Homalochilus striatus by Fischer (1856); Stull (1935) was the first use of the trinomial. The epithet H. multisectus (Cope, 1862) was subsumed into C. s. striatus (Sheplan and Schwartz, 1974).
Chilabothrus striatus exagistus (Sheplan & Schwartz, 1974)
Chilabothrus striatus warreni (Sheplan & Schwartz, 1974)
Chilabothrus strigilatus (Cope, 1862)
Taxonomy. Originally described as Homalochilus strigilatus (Cope, 1862), it was subsequently placed in the genus Epicrates (Barbour, 1904). It was subsequently relegated to a subspecies of the Hispaniolan boa (C. striatus) by Stull (1935) and Sheplan and Schwartz (1974) until the species was elevated by Reynolds et al. (2013a) based on molecular data and placed into the genus Chilabothrus. Five subspecies are recognized.
Chilabothrus strigilatus strigilatus (Cope, 1862)
Chilabothrus strigilatus ailurus (Sheplan & Schwartz, 1974)
Taxonomy. This subspecies was first described by Sheplan and Schwartz (1974) based on material collected by G. Rabb in 1953 from Cat Island, Bahamas.
Chilabothrus strigilatus fosteri (Barbour, 1941)
Taxonomy. This subspecies was first described by Barbour (1941) based on material collected from the Bimini Islands by R. Foster and J. Huntington.
Chilabothrus strigilatus fowleri (Sheplan & Schwartz, 1974)
Taxonomy. This subspecies was first described by Sheplan and Schwartz (1974) based on material collected from the Andros Islands by Danny Fowler.
Chilabothrus strigilatus mccraniei (Sheplan & Schwartz, 1974)
Taxonomy. This subspecies was first described by Sheplan and Schwartz (1974) based on material collected from the Ragged Islands.
Chilabothrus subflavus (Stejneger, 1901)
Conservation Status. Considered Vulnerable based on a 1996 IUCN Red List assessment (Gibson, 1996), though this is likely to be revised to Endangered given negative population trends (S. Koenig, personal communication). It is listed in CITES Appendix I (i.e., the most endangered and threatened with extinction) and is listed on the Jamaica Wildlife Protection Act. The species is vulnerable to widespread habitat destruction, invasive predators, severe human persecution, road morality, and the introduced and potentially toxic Rhinella marina (Wilson et al., 2011; Newman et al., 2016).
Corallus Daudin, 1803
Nine species comprise Corallus, a genus of arboreal boas. Members of the genus occur from southeastern Guatemala, through much of Central America (although the distribution can be disjunct), into South America, with a limited distribution west of the Andes in Colombia and Ecuador. East of the Andes the genus is widespread in the Guianas, Amazonia, and the Atlantic Forests of southeastern Brazil. Species also occur on continental islands (e.g., Isla Margarita, Trinidad, and Tobago), as well as oceanic islands (the St. Vincent and Grenada banks in the southern Lesser Antilles). The various species occur in a wide range of habitats; as they are arboreal, however, they are precluded from nonforested areas. Elevational distribution is from sea level to about 1,000 m. Species of Corallus range in size from ∼1.2–2.1 m SVL. Diets of the various species include frogs, lizards, and a wide taxonomic range of birds and mammals (e.g., rodents, marsupials, and bats; Henderson, 2015). Corallus is sister to the Epicrates-Eunectes clade (e.g., Reynolds et al., 2014).
Corallus annulatus (Cope, 1875)
Taxonomy. Originally described as Xiphosoma annulatum; Boulenger (1893) provided the first combination of Corallus annulatus. Boa annulata was used by Rendahl and Vestergren (1940, 1941); Peters (1957) used the current combination after Forcart (1951) resurrected Corallus. No subspecies are recognized.
Distribution. This species has a disjunct range that extends from extreme southeastern Guatemala, into northern Honduras; southeastern Nicaragua (in the Caribbean lowlands at elevations of 70–185 m, Sunyer and Köhler, 2010); northeastern Costa Rica where it reaches elevations to at least 745 m (Sosa et al., 2010) and perhaps as high as 1,000 m, and Panama, where it occurs on both Atlantic and Pacific versants in the central and southern portions of the country, and from sea level to about 400 m; (Jaramillo et al., 2010); to northern Colombia west of the Andes (Henderson, 2015).
Corallus batesii (Gray, 1860)
Taxonomy. Originally described as Chrysensis batesii, but relegated to the synonymy of Corallus caninus by Boulenger (1893). Based on molecular (Vidal et al., 2005) and morphological data, Henderson et al. (2009) resurrected the species. No subspecies are recognized.
Distribution. Widely distributed in the Amazon basin. In Brazil north and south of the Rio Amazonas west of the Rio Negro, also in Amazonian Colombia, Ecuador, Peru, and Bolivia. It also occurs in northwestern Colombia north of the Andes, including the Río Magdalena valley. Elevational distribution is sea level to 1,100–1,200 m.
Conservation Status. According to the IUCN Red List this is a species of Least Concern because of its wide geographic distribution; it has no major threats, and it occurs in multiple protected areas (Rivas et al., 2016). It is a species that is popular in the pet trade, although ostensibly protected throughout most of its range.
Corallus blombergii (Rendahl & Vestergren, 1941)
Taxonomy. Originally described as a subspecies of Boa annulata; it continued to be recognized as a subspecies by Peters (1957) as Corallus annulata blombergi and by Peters and Orejas-Miranda (1970) as C. annulatus blombergi. Based on morphological characters, it was elevated to species rank by Henderson et al. (2001). No subspecies are recognized.
Type Specimen. An adult specimen in the NRM (no. 3141), from “Eastern Ecuador, Rio Zamora” (Rendahl and Vestergren, 1941).
Distribution. Known from Ecuador in the western lowlands of the Andes. Its distribution extends from Esmeraldas to Azuay provinces but is very disjunct (Valencia et al., 2008; Henderson, 2015); it occurs at elevations below 200 m. Two specimens from extreme southwestern Colombia (Tumaco, Nariño) have recently been identified as C. blombergii (Pinto-Erazo and Medina-Rangel, 2018).
Corallus caninus (Linnaeus, 1758)
Taxonomy. Originally described as Boa canina; it was briefly placed in the genus Xiphosoma (Fitzinger, 1843; Duméril and Bibron, 1844); Boulenger (1893) was first to use the combination Corallus caninus. Henderson et al. (2009) partitioned C. caninus into two species (C. batesii and C. caninus). No subspecies are recognized.
Distribution. Guyana, Suriname, French Guiana, eastern and southern Venezuela (states of Bolívar and Amazonas), and northeastern Brazil north of the Rio Amazonas and north and east of the Rio Negro (in the states of Amapá, Pará, Roraima, and Amazonas); elevational distribution is from sea level to about 200 m.
Conservation Status. Considered a species of Least Concern on the IUCN Red List because of its large extent of occurrence and because there are no immediate threats to its habitat (Oubotar et al., 2016). It is a species that is popular in the pet trade, although ostensibly protected throughout much of its range.
Corallus cookii Gray, 1842
Taxonomy. Originally described as Corallus cookii, it was relegated to subspecies rank by Stull (1935) as Boa enydris cookii; when Forcart (1951) resurrected Corallus, it became C. e. cookii; Roze (1966) recognized it as C. hortulanus cookii. Henderson (1997) elevated it to its former full species status as C. cookii. Recent molecular evidence (Colston et al., 2013; Reynolds et al., 2014) shows C. cookii nested within C. hortulanus; based on morphological characters and geography, Henderson (2015) continued to recognize it as a valid species. No subspecies are recognized.
Type Specimen. An 861-mm SVL male in the BMNH (1946.1.1.50); type locality “America” was amended to “West Indies” by Gray (1849); Henderson (1997) restricted it to St. Vincent in the Lesser Antilles.
Distribution. Endemic to the Lesser Antillean island of St. Vincent, where it is ecologically widespread. It has been encountered to at least 425 m above sea level (Henderson, 2015).
Conservation Status. The species has been assessed as Least Concern, though the listing is currently in review. We believe that the species should likely be considered Near Threatened to Vulnerable based on Red List criteria. Recently, illegally collected individuals have become available on the Internet for the pet trade. The total area of its range is less than 350 km2.
Corallus cropanii (Hoge, 1953)
Taxonomy. Described as the type species (X. cropanii) of a new genus (Xenoboa) by Hoge (1953). Based on its sister group relationship with Corallus caninus, and to avoid a paraphyletic taxon, Kluge (1991) placed X. cropanii in the genus Corallus. No subspecies are recognized.
Type Specimen. An adult male from Miracatu, São Paulo, Brazil; specimen in the IB, number 15200 (now presumably lost in the 2010 Instituto Butantan fire).
Conservation Status. It is listed as Endangered on the IUCN Red List (Marques, 2010). In our estimation, it should be considered Critically Endangered, based on diminishing habitat, proximity of urban areas, and apparent low population density. A boa was recently found alive based on a successful citizen science initiative and extensively studied. Nevertheless, very little is known about this rare species.
Corallus grenadensis (Barbour, 1914)
Taxonomy. Originally described as Boa grenadensis, Barbour (1935) subsequently relegated it to a subspecies of B. cookii; meanwhile, Stull (1935) synonymized it with B. enydris cookii; Barbour (1937) continued to recognize it as a subspecies of B. cookii. After Forcart (1951) untangled Boa, Constrictor, and Corallus and McDiarmid et al. (1996) did the same for C. enydris/hortulanus, Henderson (1997) resurrected Corallus grenadensis to full species status. Recent molecular evidence (Colston et al., 2013; Reynolds et al., 2014) shows C. grenadensis nested within C. hortulanus, though based on morphological characters and geographic isolation, Henderson (2015) continues to recognize it as a valid species. No subspecies are recognized.
Distribution. Islands on the Grenada Bank. It has been collected or observed on Bequia, Ile Quatre, Baliceaux, Mustique, Canouan, Mayreau, Union, Petit Martinique, Petit St. Vincent, Carriacou and Grenada. On Grenada, altitudinal distribution is from sea level to at least 530 m. Elevation is not likely to preclude C. grenadensis from occurring anywhere on the Grenadine Islands.
Conservation Status. This species has been given an IUCN Red List assessment of Least Concern, though that assessment is presently in review. Although it has a multi-island distribution, the total area of all the islands is ∼400 km2. We believe that it could be considered Near Threatened to Vulnerable by Red List criteria. Recently, illegally collected individuals have become available on the Internet for the pet trade.
Corallus hortulanus (Linnaeus, 1758)
Taxonomy. Originally described as Boa hortulana. The species has had a long, convoluted taxonomic history as, among others, a species of Boa, Corallus, or Xiphosoma; often referred to as Boa enydris or Corallus enydris, and with many other synonyms. Its taxonomy was finally disentangled by McDiarmid et al. (1996); McDiarmid et al., 1999 provide an updated synonymy. No subspecies are recognized.
Type Specimen. NRM no. Lin. 7; although considered missing (Andersson, 1899), it apparently resides in the Swedish Museum of Natural History (McDiarmid et al., 1999); the type locality is “America.”
Distribution. The Guianas and Amazonia (southern Colombia, southern Venezuela, Ecuador, Peru, Bolivia, Brazil. The distribution in Brazil also includes Cerrado, mesic enclaves in Caatinga, sand dune areas in Caatinga (Rodrigues, 1996), Pantanal (Marques et al., 2005; Guedes et al., 2014), Atlantic rainforest to about 26°08′S, and Ilha Grande and Ilha Santo Amaro off southeastern Brazil. Altitudinal distribution is from sea level to about 915 m (Henderson, 2015).
Conservation Status. Considered a species of Least Concern on the IUCN Red List based on its broad geographic distribution, “relatively large population, lack of widespread threats, and occurrence in numerous protected areas” (Calderón et al., 2016:1). This is a popular species in the pet trade, although it is protected over most of its range.
Corallus ruschenbergerii (Cope, 1875)
Taxonomy. Originally described as Xiphosoma ruschenbergerii by Cope (1875). It was relegated to a subspecies of Corallus cookii (with an incorrect spelling, ruschenbergi, which has often been duplicated) by Boettger (1898); finally placed in the synonymy of C. hortulanus cookii by Roze (1966). Based on morphological evidence, Henderson (1997) resurrected it from the synonymy of C. hortulanus at species rank; molecular data have supported that resurrection (Colston et al., 2013; Reynolds et al., 2014). No subspecies are recognized.
Distribution. Corallus ruschenbergerii occurs from southern Costa Rica (sea level to 300 m; Sosa et al., 2010), through Panama from sea level to 525 m (Jaramillo et al., 2010), including the offshore islands of Isla del Rey, Isla Contadora, Isla de Cébaco, and Isla Suscantupu); in Colombia east of the Andes in the llanos and adjacent foothills, and more or less north of the cordilleras Central and Oriental; and in Venezuela north of the Cordillera de Mérida and the Río Orinoco (and on Isla Margarita), and north and west of the Guiana Shield; also known from an intra-Andean locality (the Lake Maracaibo versant of the Cordillera de Mérida, Venezuela; Esqueda and La Marca, 2004); also on Trinidad and Tobago.
Conservation Status. Considered a species of Least Concern on the IUCN Red List on the basis of its large geographic range, “apparently large population size, occurrence in several protected areas, and apparently stable trend” (Acosta Chaves et al., 2016:1). Using IUCN Red List criteria, Sosa et al. (2010) gave this species an assessment of Endangered for Costa Rica.
Epicrates Wagler, 1830
For more than 120 years, the genus Epicrates basked in relative stability, with a single widespread species (E. cenchria) on the Neotropical mainland and eight species inhabiting island banks in the Bahamas and on the Greater Antilles. Based on molecular phylogeny and historical biogeography, Reynolds et al. (2013a) determined that Epicrates was restricted to the mainland clade and they resurrected Chilabothrus for the West Indian clade. In a review of the Epicrates cenchria complex (based on meristic, morphological, and color pattern data), Passos and Fernandes (2008) elevated four taxa that had long been considered subspecies of E. cenchria to species rank. Rivera et al. (2011) provided molecular support for the revision of Passos and Fernandes (2008). Currently, five species compose the genus, and all are restricted to the Neotropical mainland (Nicaragua to Argentina) and continental islands. The genus is ecologically widespread, and species occur in Amazonian and Atlantic forests, as well as xerophic Caatinga and grasslands. Species of Epicrates are largely ground dwelling and range in size from about 1,280 to 1,850 mm SVL (Passos and Fernandes, 2008). Their diets comprise a wide taxonomic range of ectothermic and endothermic vertebrates. Species of Epicrates are phylogenetically sister to Eunectes (e.g., Reynolds et al., 2013a).
Epicrates alvarezi Abalos, Baez, & Nader, 1964
Taxonomy. Originally described as a subspecies of Epicrates cenchria by Abalos et al. (1964), it was elevated to species status by Passos and Fernandes (2008). No subspecies are recognized.
Type Specimen. A presumably unnumbered adult female in the “Instituto de Animales Venenosos Jorge Washington Abaloz” from Forres, Department of Robles, Santiago Del Estero, Argentina (Passos and Fernandes, 2008).
Epicrates assisi Machado, 1945
Taxonomy. Originally described as a subspecies of Epicrates cenchria by Machado (1945) but elevated to species status by Passos and Fernandes (2008). No subspecies are recognized.
Type Specimen. An unnumbered adult male (presumably lost) in the Instituto Vital Brazil, from Campina Grande, Paraíba, Brazil (Passos and Fernandes, 2008).
Epicrates cenchria (Linnaeus, 1758)
Taxonomy. Originally described by Linnaeus (1758) as Boa cenchria. Wagler (1830) described the new genus Epicrates and E. cenchria became the type species of the genus. Subsequently, many names have been associated with E. cenchria; McDiarmid et al., 1999 and Passos and Fernandes (2008) provide extensive synonymies. No subspecies are recognized.
Epicrates crassus Cope, 1862
Taxonomy. Originally described as a full species by Cope (1862), it was relegated to a subspecies of Epicrates cenchria by Amaral (1929), then “considered” a full species by Lema (2002). No subspecies are recognized.
Distribution. Open formations on the Andean slopes of Bolivia, grasslands of Argentina and Paraguay, and Brazilian Cerrado (in the states of Rondônia, Pará, Mato Grosso, Tocantins, Goiás, Minas Gerais, São Paulo, and grassland of Mato Grosso do Sul, Paraná, and Rio Grandes do Sul) (Passos and Fernandes, 2008).
Epicrates maurus Gray, 1849
Taxonomy. Originally described as a full species, it was placed in the synonymy of E. cenchria by Boulenger (1893). Stull (1935) eventually recognized it as a subspecies of Epicrates cenchria; Chippaux (1986), however, considered it a full species, as did Gorzula and Señaris (1998). Nevertheless, McDiarmid et al., 1999 still considered it a subspecies of E. cenchria. Matz (2004) treated it as a full species and described two subspecies of E. maurus (both subspecies were subsequently placed in the synonymy of E. maurus by Passos and Fernandes, 2008). Passos and Fernandes (2008), in their review of the E. cenchria complex, considered it a full species. No subspecies are recognized.
Type Specimen. An adult male from Venezuela, BMNH 1946.1.10.40 (formerly BMNH 46.7.23. 2a + vi.6.3a).
Distribution. This species occurs in seasonally dry forest in Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, and northern Brazil, as well as on Trinidad, Tobago, and Isla Margarita; elevational distribution is 0–500 m. It may occur syntopically with E. cenchria where the savannah meets the forest (Passos and Fernandes, 2008).
Conservation Status. This species has received an IUCN Red List assessment of Least Concern, though the assessment is currently in review. Sosa et al. (2010), using IUCN criteria, gave it an assessment of Endangered in Costa Rica. Also using IUCN criteria, Jaramillo et al. (2010) gave an assessment of Least Concern for Panama.
Eunectes Wagler, 1830
This genus is composed of four species, and all are restricted to the South American mainland from Colombia and Venezuela to Argentina and the continental island of Trinidad. One species, Eunectes murinus, is one of the longest snakes in the world (to about 8.0 m, likely second in length to Malayopython reticulatus or Python sebae) and is certainly the most massive. All species are closely associated with water (e.g., rivers, swamps, “borrow” pits). Diets include a wide range of vertebrates, many of which are associated with aquatic habitats (turtles, crocodilians, wading birds, capybara; e.g., Rivas, 2000). All species are likely exploited for their hides. Species of Eunectes are phylogenetically sister to Epicrates (e.g., Reynolds et al., 2013a).
Eunectes beniensis Dirksen, 2002
Taxonomy. First described by Dirksen in 2002 and redescribed in Dirksen and Böhme (2005) based on color pattern differences from E. notaeus and E. deschauenseei. No subspecies are recognized.
Distribution. Known from the northeastern departments of Beni, Santa Cruz, and Pando in Bolivia; it may also occur in nearby Brazil. Elevational range is ∼115–350 m (Muñoz et al., 2016).
Conservation Status. The IUCN Red List has assessed the species as being of Least Concern based on its large estimated extent of occurrence (>45,000 km2; Muñoz et al., 2016). This species is collected for its skin and for its use as cooking fat; it is also killed because it feeds on chickens, dogs, and cats (Cortez et al., 2009). The Bolivian Government is presently assessing the feasibility of a sustainable harvest of anacondas within indigenous territories, and such activity would potentially affect the species.
Eunectes deschauenseei Dunn & Conant, 1936
Taxonomy. Described as a full species by Dunn and Conant (1936) based on a live individual. No subspecies are recognized.
Type Specimen. A female, ANSP 20891, “very probably” from Ilha de Marajó, Pará, Brazil (Dunn and Conant, 1936).
Eunectes murinus (Linnaeus, 1758)
Taxonomy. Originally described as Boa murina by Linnaeus. When Wagler (1830) described the genus Eunectes, B. murina became the type species. Aside from Stull (1935) resurrecting the old Linnaean name scytale to replace murinus, and the epithet E. barbouri being subsumed into E. murinus (Strimple et al., 1997; Dirksen, 2000), the species has had a remarkably stable taxonomic history. See McDiarmid et al., 1999, Dirksen (2002), and Dirksen and Böhme (2005) for a more complete synonymy. No subspecies are recognized.
Eunectes notaeus Cope, 1862
Taxonomy. Described by Cope in 1862, the species has had a stable taxonomic history. No subspecies are recognized.
FAMILY CALABARIIDAE GRAY, 1858
Calabaria Gray, 1858
Calabariidae is a monotypic family represented by a single extant species found in western Africa. Calabaria reinhardtii is a small oviparous species (<1.0 m SVL) that occurs in a variety of habitats, including primary and secondary swamp forest, cultivated fields, secondary dryland forest, and suburban environments; it is nocturnal, a specialized burrower, and often encountered underground, in leaf litter, or under cover objects. Its diet includes shrews and rodents (Luiselli and Akani, 1998; Luiselli et al., 2002). Calabariidae has been variously considered a pythonid (W. C. H. Peters, 1858), a boid, and a unique lineage, likely owing to a combination of unique and convergent morphological traits (Kluge, 1993). Recent molecular phylogenies suggest that Calabariidae is either nested within (Pyron et al., 2013; Reynolds et al., 2014) or sister to (Harrington and Reeder, 2017; Ruane and Austin, 2017) the booids; hence, we include it here as a booid family (fide Pyron et al., 2014), though we note that not all workers embrace this arrangement. The family Calabariidae was recognized by Pyron et al. (2014) to stabilize booid taxonomy in the face of inconsistent molecular and morphological phylogenetic hypotheses.
Calabaria reinhardtii (Schlegel, 1848)
Taxonomy. Described by Schlegel (1848) as Eryx reinhardtii, the type of the genus was given as Calabaria fusca Gray 1858. The genus was referred to as both Rhoptrura and Eryx in the 19th century until Boulenger (1893) stabilized the monotypic genus as C. reinhardtii. Kluge (1993) placed the species into Charina owing to shared morphological synapomorphies with North American Lichanura and Charina. No subspecies are recognized.
Type Specimen. The holotype is ZMB 1471, a specimen of unknown sex, likely from southeastern Ghana (Hughes and Barry, 1969).
FAMILY CANDOIIDAE PYRON ET AL., 2014
Candoia Gray, 1842
The Pacific boas are a widely distributed group recognized here as constituting five species and 12 subspecies (though O'Shea, 2007, recognized 14 subspecies). They occur from the Moluccas (=Maluku) in the west, to the Palau Archipelago in the North, the Loyalty Islands in the south, and American Samoa in the east. Species of Candoia occur in a variety of habitats, including rainforests and coffee and coconut plantations (in discarded husk piles; O'Shea, 1996). They are of moderate size (0.5–1.5 m SVL), stout and ground-dwelling to more slender and arboreal; viviparous; and the diet comprises mainly of lizards (primarily skinks) and murid rodents (Harlow and Shine, 1992). The genus Enygrus Wagler 1830 was originally used for these snakes, though a python (Python regius) was designated as the type specimen for the genus by Fitzinger (1843). This was resolved when Forcart (1951) recognized that Candoia Gray 1842 was the appropriate name for the South Pacific boas. It is worth noting that Gray (1842) initially provided the genus name Candoia, though the same author used Enygrus in subsequent work (Gray, 1849). Systematic revisions of the genus Candoia have been conducted at intervals. McDowell (1979) provided a detailed analysis of the biology, biogeography, and systematics of the group, recognizing the species C. bibroni, C. carinata, and C. aspera, though noting substantial regional variation in meristic characters. Smith et al. (2001) used morphological and squamation characters to further revise the carinata complex, recognizing two additional species (C. paulsoni and C. superciliosa) and 10 subspecies. Molecular phylogenetics have largely corroborated the proposed systematic revisions, based on both mitochondrial (Austin, 2000) and multilocus (Reynolds et al., 2014) datasets. Historical biogeographic analyses (Noonan and Chippindale, 2006; Noonan and Sites, 2010) have suggested an origin of the Candoia in the early Paleogene.
Candoia aspera (Günther, 1877)
Taxonomy. The species was originally described as Erebophis asper (Günther, 1877), despite frequent incorrect assertions that the original name given by Günther (1877) was Erebophis aspera. The name was changed to Enygrus asper by Boulenger (1893) and to Candoia aspera by Forcart (1951). Thus the specific epithet is now aspera, which is the correct declension of the Latin feminine Candoia. See McDowell (1979) and McDiarmid et al., 1999 for more detailed synonymies. Two subspecies are recognized.
Type Specimen. Holotype BMNH 1946.1.10.33 of unknown sex. The type locality is New Ireland Island (=Duke of York Island) in the Bismarck Archipelago.
Distribution. A species found below 1,300 m on the island of New Guinea and several nearby islands (e.g., Batanta, Misool, Waigeu); Biak and Japen islands; Seleo Island; Valise Island; Karkar Island; Umboi Island; islands in the Manus Group (Admiralty Islands); New Britain, Duke of York, New Ireland, and New Hanover in the Bismarck Archipelago (McDowell, 1979; Harlow and Shine, 1992; Austin, 2000; McCoy, 2015). Records from Bougainville Island in the Solomon Islands (Kinghorn, 1928) and Tokelau (Stull, 1935) are likely erroneous (McDowell, 1979).
Candoia aspera aspera (Günther, 1877)
Taxonomy. See the Candoia aspera account. It became the nominate subspecies of Candoia aspera with the description of E. a. schmidti.
Type Specimen. Holotype BMNH 1946.1.10.33, unknown sex, from New Ireland Island in the Bismarck Archipelago.
Candoia aspera schmidti (Stull, 1932)
Taxonomy. Described by Stull as Enygrus asper schmidti and subsequently recognized by Loveridge (1948). Later included in Candoia as C. aspera schmidti (Forcart, 1951).
Type Specimen. Holotype MCZ R29778, an adult female from near the Sepik River (=Kaiseriu Augusta River).
Candoia bibroni (Duméril & Bibron, 1844)
Taxonomy. Originally described as Tropidoboa de bibron (Hombron and Jacquinot, 1842), formalized to Enygrus bibroni by Duméril and Bibron (1844), and occasionally referred to as Enygrus bibronii thereafter (e.g., Boulenger, 1886). Genus Candoia was resurrected by Forcart (1951), though the specific epithet is incorrectly given therein as C. bibronii. See McDowell (1979) and McDiarmid et al., 1999 for more detailed synonymies.
Specimens of this species were described as Boa australis by Montrousier (1860). Boulenger (1893) split E. bibroni into Enygrus australis and Enygrus bibronii, and Roux (1913) used ventral counts to diagnose these as subspecies of C. bibroni. Forcart (1951) recognized these subspecies as C. b. bibronii and C. b. australis. Based on data in McDowell (1979), the subspecific status as currently recognized might be unwarranted by meristic characters. The subspecies were described based on ventral scale counts (Roux, 1913), which are variable across the species' range and do not correspond to geographic regions (McDowell, 1979). Although some geographic correlates exist for at least one skeletal characteristic (McDowell, 1979), this character does not form the basis of the subspecific description. Thus, we find insufficient reason to recognize the subspecies C. b. bibroni and C. b. australis.
Type Specimens. Syntypes MNHN 1313, 3276–77, 61, and 61A, likely from the Fijian Archipelago, possibly from the island of Viti Levu (Stimson, 1969; McDowell, 1979).
Distribution. A species found below 1,200 m elevation across a number of island archipelagos in the South Pacific (Melanesia and Polynesia), from the Solomon Islands east to American Samoa (McDowell, 1979; Allison et al., 2012a; Zug, 2013). Considered to range into the Solomon Islands, then west to Makira, Rennell, and Bellona islands, but not found on Guadalcanal, Malaita, or further northwest in the archipelago (McCoy, 2015). Known from the Loyalty Islands (possibly introduced), but not mainland New Caledonia (Bauer and Sadlier, 2000), Vanuatu, the Fijian Archipelago (including Rotuma Island; M. O'Shea, in litteris, 4.iv.2018) the Wallis and Fortuna islands, Samoa, and American Samoa (McDowell, 1979; Allison et al., 2012a). Possible records from Tokelau (Stull, 1935) are likely erroneous, and records from Tonga are suspect (McDowell, 1979).
Conservation Status. This species has been assessed as Least Concern based on IUCN Red List criteria. It is widespread and does not appear to show evidence of population decline, though individuals are often killed, and some might be poached for the pet trade (Allison et al., 2012a). These boas are somewhat common in the Loyalty Islands, New Caledonia, and are occasionally eaten there (Bauer and Sadlier, 2000).
Candoia carinata (Schneider, 1801)
Taxonomy. Described as Boa carinata (Schneider, 1801). It became the type species for the genus Candoia Gray (1842). The name Boa variegata (Thunberg, 1807) was synonymized with Candoia carinata (Bauer and Wahlgren, 2001). The genus was changed to Enygrus (Duméril and Bibron, 1844) but restored to Candoia by Forcart (1951). See McDowell (1979) and McDiarmid et al., 1999 for more thorough synonymies. Two subspecies are recognized.
Type Specimens. Lectotype ZFMK 35503 from an unknown locality, though restricted to Amboina, Indonesia (McDowell, 1979; Böhme et al., 1998). Lectotype of Boa variegata (= Candoia carinata fide Bauer and Wahlgren, 2001) ZIUU 313, likely from the Moluccas (Bauer and Wahlgren, 2001; Smith et al., 2001).
Candoia carinata carinata (Schneider, 1801)
Distribution. Distributed from the Sangihe Islands off Sulawesi (North Sulawesi Province) through the Maluku Islands (Maluku Province), including Tanimbar in the south and Seram in the north, through mainland New Guinea to the Owen Stanley Range (McDowell, 1979; O'Shea, 1996; Smith et al., 2001). Absent from Halmahera (fide Smith et al., 2001). Found at elevations below 1,525 m on New Guinea (O'Shea, 1996).
Candoia carinata tepedeleni Smith & Chiszar, 2001 (in Smith et al., 2001)
Taxonomy. First identified as Enygrus carinatus (Werner, 1899), later changed to Candoia carinata carinata (Stimson, 1969; affirmed by McDowell, 1979). This subspecies was described based on morphological and meristic characters (Smith et al. 2001).
Type Specimens. Holotype MCZ R72155, an adult male from Rabaul, New Britain, Bismarck Archipelago (Smith et al., 2001).
Candoia paulsoni (Stull, 1956)
Taxonomy. Apparently ignoring (or unaware of) Forcart's (1951) resurrection of Candoia, Stull (1956) described this taxon as a subspecies of Enygrus carinata (E. carinatus paulsoni). Candoia carinata paulsoni was elevated to a full species in the C. carinata complex by Smith et al. (2001). Six subspecies are recognized.
Candoia paulsoni paulsoni (Stull, 1956)
Taxonomy. Initially described as a subspecies, Enygrus carinatus paulsoni, of the E. carinatus complex (Stull, 1956) and later recognized as the nominotypical subspecies C. p. paulsoni (Smith et al., 2001).
Candoia paulsoni mcdowelli Smith & Chiszar, 2001 (in Smith et al., 2001)
Taxonomy. First described from New Guinea as part of Schlegel's (1872) Boa carinata; McDowell (1979) was first to recognize it as not C. carinata. The subspecies was recognized owing to meristic (squamation) characters (Smith et al., 2001).
Candoia paulsoni rosadoi Smith & Chiszar, 2001 (in Smith et al., 2001)
Taxonomy. Based on specimens from Misima Island and originally referred to as C. carinata by McDowell (1979; part) and O'Shea (1996, as the “paulsoni” phase). The subspecies was recognized based on meristic (squamation) characters (Smith et al. 2001).
Candoia paulsoni sadlieri Smith & Chiszar, 2001 (in Smith et al., 2001)
Taxonomy. Specimens of this subspecies were originally identified as Candoia carinata (e.g., McDowell, 1979), or C. c. paulsoni (O'Shea, 1996). The subspecies was recognized based on meristic (squamation) characters (Smith et al., 2001).
Candoia paulsoni tasmai Smith & Tepedelen, 2001 (in Smith et al., 2001)
Taxonomy. Like other subspecies in the C. paulsoni subcomplex, specimens of C. p. tasmai were considered to be C. carinata by Peters and Doria (1878), Stimson (1969), and McDowell (1979). The subspecies was recognized based on meristic (squamation) characters (Smith et al., 2001).
Distribution. Endemic to Indonesia, where it occurs from the eastern arm of North Sulawesi (the Talaud Islands) through the northern Maluku Islands. This subspecies is isolated from other members of the C. paulsoni complex by ∼800 km, though it is considerably closer to populations of C. carinata occurring in eastern Indonesia.
Candoia paulsoni vindumi Smith & Chiszar, 2001 (in Smith et al., 2001)
Taxonomy. Specimens of this subspecies were originally considered to be C. carinata by Sternfeld (1913) and McDowell (1979). The subspecies was recognized owing to meristic (squamation) characters (Smith et al., 2001).
Distribution. A subspecies endemic to Bougainville and Buka islands, central Solomon Islands. The subspecies possibly exists on satellites of Bougainville (Shortland, Ovau, and Fauro islands; Smith et al., 2001), but more likely they are inhabited by the nominate subspecies (M. O'Shea, in litteris, 4.iv.2018).
Candoia superciliosa (Günther, 1863)
Taxonomy. Originally described as Enygrus superciliosus by Günther but was relegated to synonymy with E. carinatus by Boulenger (1893). Smith et al. (2001) resurrected C. superciliosa as part of their C. carinata complex. Two subspecies are recognized, and multilocus phylogenetic analysis suggests that these subspecies are minimally diverged (Reynolds et al., 2014).
Candoia superciliosa superciliosa (Günther, 1863)
Candoia superciliosa crombiei Smith & Chiszar, 2001 (in Smith et al., 2001)
Taxonomy. Specimens of this subspecies were previously referred to as Enygrus carinatus (Sternfeld, 1913; Dryden and Taylor, 1969). Given the subspecific epithet C. s. crombiei based on number of ventral scales (Smith et al., 2001) and, presumably, on allopatry from the range of the subspecies C. s. superciliosa.
FAMILY CHARINIDAE GRAY, 1849
Subfamily Charininae Gray, 1849
Charina Gray, 1849
Two species comprise the genus and they are restricted to southwestern Canada and the western United States. These are short (<1.0 m total length), stout-bodied boas that frequent grassland, woodland, and forest; they are good burrowers and often are encountered under cover objects (rocks, logs, bark). The diet of smaller (younger) individuals include squamate eggs and lizards (Elgaria, Sceloporus); larger (older) boas take birds and mammals (insectivores and rodents; Rodríguez-Robles et al., 1999). Charina and Lichanura are sister taxa to Exiliboa and Ungaliophis (Reynolds et al., 2014).
Charina bottae (Blaineville, 1835)
Taxonomy. Originally described as Tortrix bottae; Gray (1849) described the genus Charina with T. bottae as the type species. It has had a fairly stable taxonomic history, with the only real deviation coming from Jan (1862), who placed C. bottae in the genus Pseudoeryx. See McDiarmid et al., 1999 for a complete synonymy. No subspecies are recognized.
Type Specimen. MNHN 730, from “Californie”; restricted to the “Coast Range, opposite Monterey, California” (Schmidt, 1953).
Charina umbratica Klauber, 1943
Taxonomy. Originally described as a subspecies of Charina bottae; Erwin (1974) first suggested elevating it to full species; Rodríguez-Robles et al. (2001) did elevate it to species rank based on morphological and molecular data. No subspecies are recognized.
Type Specimen. A (possibly immature) male, SDSNH 12101, from Fern Valley, near Idyllwild, Riverside Co., California.
Lichanura (Cope, 1861)
A genus of small (<1 m SVL) species restricted to the extreme southwestern United States and around the Gulf of California in Mexico. Individuals are largely fossorial throughout most of the year, occupying rodent burrows. When surface active, largely in the Spring, individuals are frequently crepuscular, though they transition to nocturnal surface activity as the weather warms. Diet largely comprises mammals (mostly rodents; Rodríguez-Robles et al., 1999). Original descriptions of the genus recognized two species (Cope, 1861; Stejneger, 1889a; Klauber, 1931, 1933), though these were later collapsed into a single species and multiple subspecies (Ottley, 1978; Yingling, 1982; Gorman, 1985; Spiteri, 1988, 1992). The genus shows a wide range of morphological variation (Limburg et al., 2011), on which historical taxonomic divisions were based (Yingling, 1982), and molecular phylogenetic data suggest at least three (mitochondrial) lineages, two of which are presently recognized at the specific level (Wood et al., 2008). Lichanura has previously been synonymized with Charina (e.g., Kluge, 1993; Murphy and Aguirre-Léon, 2002), though we follow most modern interpretations in recognizing both genera.
Lichanura orcutti Stejneger, 1889b
Taxonomy. Originally recognized as Lichanura orcutti (Stejneger, 1889b) and re-described along with L. simplex by Stejneger (1889a). Lichanura simplex is now considered a junior synonym (Wood et al., 2008). The subspecies L. trivirgata roseofusca (Cope, 1868) has occasionally been considered a synonym of L. orcutti (Yingling, 1982). This species likely comprises at least two lineages (Wood et al., 2008). No subspecies are recognized.
Distribution. North of the Tijuana River watershed in San Diego County, California, and east to the Colorado River and Gila River drainages in Arizona (Wood et al., 2008). Closely contacts L. trivirgata near Chula Vista, California.
Conservation Status. This species is recently recognized based on mitochondrial DNA and has not been assessed by IUCN criteria. It is widespread and locally common, though coastal populations have likely been reduced or extirpated in areas of heavy development. Nevertheless, the species is capable of surviving in close proximity to development (Diffendorfer et al., 2005; Lemm, 2006). Some populations have likely been affected by collection for the pet trade (Parizek et al., 1996) or possibly by road mortality (Rosen and Lowe, 1994).
Lichanura trivirgata Cope, 1861
Taxonomy. Originally described as Lichanura trivirgata (Cope, 1861), the species has previously been recognized as L. roseofusca (Cope, 1868) and Charina trivirgata. The subspecies L. trivirgata roseofusca (Cope, 1868) and L. trivirgata myriolepis (Cope, 1868) are considered synonyms of L. trivirgata (Ottley et al., 1980; Wood et al., 2008). Lichanura trivirgata gracia was described based on presumed regional coloration (Klauber, 1931). No subspecies are recognized.
Distribution. Wide ranging and locally abundant from south of the Tijuana and Otay watersheds to the tip of the Baja Peninsula and around the Sea of Cortés to coastal Sonora, Mexico. Occurs on continental islands off the Pacific coast of Mexico (Isla Cedros; Ottley, 1978) as well as islands in the Sea of Cortés (Murphy and Aguirre-Léon, 2002).
Subfamily Ungaliophiinae McDowell, 1987
Exiliboa Bogert, 1968b
Exiliboa is a monotypic genus restricted to southern Mexico. Exiliboa placata appears to be restricted to cloud forest, where it is frequently encountered under flat rocks. It is a small charinid (<0.5 m total length), and its diet might be confined to amphibian prey (frogs and salamanders; Campbell and Camarillo, 1992). Exiliboa and Ungaliophis are sister taxa to North American Charina and Lichanura (e.g., Reynolds et al., 2014).
Exiliboa placata Bogert, 1968b
Taxonomy. Described as the type species (E. placata) in a new genus (Exiliboa) by Bogert (1968b). No subspecies are recognized.
Type Specimen. A female (“possibly immature”) in the AMNH 100000 from the headwaters of the Río Valle Nacional on the northern slopes of the Sierra de Juárez, Oaxaca, Mexico (elevation ∼2,300 m; Bogert, 1968b).
Ungaliophis Müller, 1880
Two species comprise the genus Ungaliophis, and combined, they range from southern Mexico, through Central America, and into northwestern Colombia. They occur in tropical deciduous forest and lowland moist and wet forest to lower montane wet forest from near sea level to ∼2,300 m. Both species are small (<675 mm total length) and largely arboreal (often associated with epiphytic vegetation) but may be encountered on the ground as well (Corn, 1974; Köhler, 2003; McCranie, 2011). Diet in nature includes birds and bats (Dwyer, 2017; Solórzano and Carillo, 2017) and possibly frogs and lizards. Ungaliophis and Exiliboa are sister taxa to North American Charina and Lichanura (e.g., Reynolds et al., 2014).
Ungaliophis continentalis Müller, 1880
Taxonomy. Originally described as Ungaliophis continentalis. The name Peropodum guatemalensis Bocourt has caused some confusion regarding the genus Ungaliophis and this species, but it has been clarified by Stuart (1954) and McDiarmid et al., 1999. No subspecies are recognized.
Distribution. Pacific coastal plain and highlands of southern Mexico (southeastern Chiapas), Pacific coastal plain of southwestern Guatemala, Honduras, and Nicaragua; elevational distribution from 100 m to about 2,300 m.
Conservation Status. This species is being considered for listing as Near Threatened under IUCN Red List criteria, although the account is currently in review. Acevedo et al. (2010) provided an IUCN category of Vulnerable for this species in Guatemala, as did Townsend and Wilson (2010) for Honduras and Sunyer and Köhler (2010) for Nicaragua.
Ungaliophis panamensis Schmidt, 1933
Taxonomy. Originally described as Ungaliophis panamensis. Bogert (1968a) provided evidence for placing Ungaliophis danieli from Colombia (Prado, 1940) in the synonymy of U. panamensis. No subspecies are recognized.
Type Specimen. An adult female in the USNM 54029 from Cerro Brujo, Colon Province, Panama (elevation 2,000 feet [610 m]; Schmidt, 1933).
Distribution. Southern Nicaragua, Costa Rica, Panama to northwestern Colombia from near sea level to about 2,100 m (Villa and Wilson, 1990).
Conservation Status. This species was assessed as an IUCN Red List species of Least Concern because of its wide distribution, stable population, and lack of major threats (Sunyer and Ibáñez, 2015). Sunyer and Köhler (2010), using IUCN methodology, gave this species an assessment of Vulnerable for Nicaragua. Similarly, Sosa et al. (2010), also using IUCN Red List criteria, provided an assessment of Endangered for this species in Costa Rica.
FAMILY ERYCIDAE BONAPARTE, 1831
Eryx Daudin, 1803
The family Erycidae is a monogeneric group of relatively small (<1.0 m SVL) fossorial snakes. They occur in desert, near-desert, and dry woodland habitats; diet includes lizards, birds, and mammals (Rodríguez-Robles et al., 1999). They generally possess relatively small eyes, which can be oriented more dorsally rather than laterally in some species. A second genus (Gongylophis Wagler 1830) had been resurrected (McDowell, 1979; Tokar, 1995, 1996) and used to distinguish members lacking a mental groove (Tokar, 1995), but this arrangement rendered Eryx paraphyletic (Noonan and Chippindale, 2006; Reynolds et al., 2014). Eryx as a group is understudied from a systematics perspective, and there is little agreement on the number of species and subspecies that should be recognized. For example, Pyron et al. (2014) recognize 13 species, while Uetz et al. (2017) recognize 12 species. These differences are owing to the lack of consensus regarding whether E. vittatus is a separate species. Here we recognize 13 species, including E. vittatus owing to slight distinctiveness in recent molecular phylogenies (Lynch and Wagner, 2010; Reynolds et al., 2014), and no subspecies.
Eryx borrii Lanza & Nistri, 2005
Taxonomy. Described from a single specimen based on a relatively higher ventral scale count than other Somali Eryx (Lanza and Nistri, 2005). No subspecies are recognized.
Eryx colubrinus (Linnaeus, 1758)
Taxonomy. Originally described as Anguis colubrina (Linnaeus, 1758), the species was variously considered Eryx thebaicus (Reuss, 1834), E. jaculus (Jan and Sordelli, 1860–1866), and E. rufescens (Ahl, 1933). Flower (1933) considered E. thebaicus Stull 1932 conspecific with E. colubrinus. The subspecies E. c. loveridgei was recognized by Stull (1932) as a distinct lineage based on meristic and coloration differences from populations in Kenya (Stull, 1932), though Tokar's (1996) assessment suggests that these characters are clinal and that no subspecies are warranted. Lynch and Wagner (2010) suggest some phylogenetic divergence in the putative subspecies, though that analysis was based on a supermatrix approach; hence, additional information would be needed to warrant phylogenetic support for these subspecies.
Eryx conicus (Schneider, 1801)
Taxonomy. Originally described as Boa conica, this taxon later formed the basis for the description of the genus Gongylophis (Wagler, 1830). Gongylophis is no longer recognized following molecular phylogenetic reconstruction of the Erycidae (Reynolds et al., 2014), and the species is considered to be within the genus Eryx. No subspecies are recognized.
Type Specimen. Syntype ZMB 1470, likely from southeast India (Stimson, 1969); a second syntype in “Museo Barbyensi” was unlocated (McDiarmid et al., 1999).
Eryx elegans (Gray, 1849)
Taxonomy. Originally described as Cusoria elegans by Gray (1849), the species was placed in the genus Eryx by Blanford (1876). Analyses of morphology (Eskandarzadeh et al., 2013) and mitochondrial DNA (Rastegar-Pouyani et al., 2008) indicated that some E. jaculus had been misidentified as E. elegans, and that the two species might be conspecific in northern Iran. Further examination suggests that E. elegans is a distinct species in northern Iran and that the confusion likely is owing to an incomplete dichotomous key for these species (Zarrintab et al., 2017). No subspecies are recognized.
Eryx jaculus (Linnaeus, 1758)
Taxonomy. Originally described as Anguis jaculus by Linnaeus (1758), this species was subsequently assigned to the genus Eryx by Daudin (1803). Some sources recognize multiple subspecies (e.g., Tokar and Obst, 1993; Safaei-Mahroo et al., 2015; Uetz et al., 2017), though others have pointed out that the species is generally treated as monotypic (Sindaco et al., 2000). Morphological analyses of E. jaculus and E. elegans in Iran have demonstrated some confusion regarding species boundaries (Eskandarzadeh et al., 2013), though molecular phylogenies show they are likely distinct lineages (Lynch and Wagner, 2010). See McDiarmid et al., 1999 for a more complete synonymy. No subspecies are recognized.
Type Specimen. Lectotype NRM Lin-12, an adult of unknown sex from Egypt. The holotype is likely lost (Kluge, 1993).
Distribution. Southeastern Europe from Romania through the Balkan Peninsula; Mediterranean Islands including Sicily (Insacco et al., 2015) and Aegean islands (Sindaco et al., 2000); Iran (Gholamifard et al., 2012; Kazemi et al., 2015; Safaei-Mahroo et al., 2015) to western Turkey and the eastern Mediterranean Levant (Albaba, 2016); northern Africa, including Egypt (Marx, 1968) and west to Morocco.
Eryx jayakari Boulenger, 1888
Taxonomy. Recognized as a distinct species largely owing to squamation (Boulenger, 1888), and easily distinguishable from E. johnii (Zarrintab et al., 2017); oviparity is also a distinguishing trait. Eryx fodiens is a synonym (Stull, 1935; Stimson, 1969). No subspecies are recognized.
Eryx johnii (Russell, 1801)
Taxonomy. Originally described as Boa johnii. Synonyms include Boa anguiformis (Schneider, 1801), Eryx indicus (Schlegel, 1837), and E. maculatus (Hallowell, 1848). Occasionally misspelled as E. johni (e.g., Rastegar-Pouyani et al., 2008). No subspecies are recognized.
Type Specimen. Lectotype (illustration) designated by M. A. Smith (1943) from Tamil Nadu State, India (fide Wallach et al., 2014); subsequently lost (Stimson, 1969). Syntypes are unlocated (McDiarmid et al., 1999).
Eryx miliaris (Pallas, 1773)
Taxonomy. Originally described as Anguis miliaris, the species was placed into the genus Eryx by Eichwald (1831). Based on molecular phylogenetic analysis, Reynolds et al. (2014) found that this species might be conspecific with E. tataricus, or at least that additional work remains to designate species boundaries and diagnostic morphological characters for identification. Following Wallach et al. (2014), we continue to recognize it here. Two subspecies have been described, the nominate E. m. miliaris and E. m. nogaiorum (Nikolsky, 1910). Both Eskandarzadeh et al. (2013) and Zarrintab et al. (2017) suggest that E. miliaris does not occur in Iran. Given the uncertainty regarding the epithet E. miliaris vis-à-vis E. tataricus, we suggest not recognizing these subspecies pending much needed focal study of this potentially wide-ranging species. See McDiarmid et al., 1999 for a more complete synonymy.
Type Specimen. Holotype is presumed lost (Stimson, 1969; Kluge, 1993). A lectotype, MNKNU 27350 from the Dagestan region of Russia, was designated by Vedmederya et al. (2009).
Eryx muelleri (Boulenger, 1892)
Taxonomy. Originally described as Gongylophis muelleri to distinguish it from E. jaculus (Boulenger, 1892), the species was referred to Eryx by the same author the subsequent year (Boulenger, 1893). Tokar (1995) continued to recognize Gongylophis, though molecular phylogenetic analysis of Erycidae suggests that the name renders Eryx paraphyletic and current studies no longer recognize Gongylophis. Eryx muelleri is an oviparous species, likely a derived condition (Lynch and Wagner, 2010). No subspecies are recognized.
Eryx somalicus Scortecci, 1939
Taxonomy. Originally described as Eryx somalicus (Scortecci, 1939), the species was briefly moved to the genus Gongylophis (Lanza, 1990). Early representatives were likely misclassified as E. thebaicus (Boulenger, 1901; Parker, 1949). No subspecies are recognized.
Eryx tataricus (Lichtenstein, 1823)
Taxonomy. Originally described as Boa tatarica, it has variously been recognized as a separate species or as a subspecies of E. miliaris (e.g., Lambert, 2002). Based on molecular phylogenetic analysis, Reynolds et al. (2014) found that this species might be conspecific with E. miliaris, or at least that additional work remains to designate species boundaries and diagnostic morphological characters for identification. Following Wallach et al. (2014), we continue to recognize it here. Two subspecies have been described, the nominate E. t. tataricus (Lichtenstein, 1823) and E. t. speciosus (Tzarevsky, 1916), the latter of which is considered to occur in a small portion of the central part of the range in Pakistan (Khan, 2004). Eryx speciosus was recently recognized (Ananjeva et al., 2006), though without clear motivation or justification. Given the uncertainty regarding the epithet E. tataricus vis-à-vis E. miliaris, we suggest not recognizing these species or subspecies pending much needed focal study of this wide-ranging species.
Type Specimen. Lectotype ZMB 1461 (Bauer et al., 2002), an adult of unknown sex, likely from the Aral Sea region of Uzbekistan and Kazakhstan (Khan, 2002).
Eryx vittatus Chernov, 1959
Taxonomy. This species was long recognized as a subspecies of E. tataricus. It was elevated to species rank by Tokar (1989) based on comparative osteological data, though it continues to be recognized as a subspecies by some authors (e.g., Safaei-Mahroo et al., 2015). Recent molecular phylogenies have suggested that the lineage is evolutionarily distinct from both E. tataricus and E. miliaris; hence, it is recognized as a species by Pyron et al. (2014) and Wallach et al. (2014), and we recognize it here. No subspecies are recognized.
Eryx whitakeri Das, 1991
Taxonomy. Specimens of Eryx from the Western Ghats were considered E. conicus until Eryx whitakeri was described by Das (1991) owing to squamation and coloration differences. No subspecies are recognized.
Type Specimen. Holotype ZSI 24810, an adult female from Mangalore, Karnataka State, India (Das, 1991).
FAMILY SANZINIIDAE ROMER, 1956
Acrantophis Jan, 1860
The genus Acrantophis consists of two recognized species endemic to the island of Madagascar and satellites. These boas of moderate size (to ∼3.0 m total length for A. madagascariensis) occur in forested habitats. They are largely ground dwelling, nocturnal, and prey primarily on mammals, including lemurs (Glaw and Vences, 2007; Gardner et al., 2013). Acrantophis madagascariensis is largely restricted to northern Madagascar, while A. dumerili is restricted to southern Madagascar. Nevertheless, a broad zone of overlap occurs across the west-central portion of the island (Vences and Glaw, 2003). While these species may be distinguishable based on head squamation characteristics (Guibé, 1949, 1958; Vences and Glaw, 2003), a phylogenetic analysis of the mitochondrial locus 16S (Vences and Glaw, 2003) suggested that the two species are paraphyletic. Additional work using multilocus genetic data further supports the paraphyly of these two species, with some individuals identified as A. cf. dumerili belonging either to a southern population of A. madagascariensis or to a separate lineage of Acrantophis in southern Madagascar (Orozco-Terwengel et al., 2008; Reynolds et al., 2014). Both species are listed on CITES Appendix I, which prevents international trade in live individuals or parts obtained from these species.
Acrantophis dumerili Jan, 1860
Taxonomy. Originally described as Acrantophis dumerili. Boulenger (1893) assigned it to Boa (and Barbour, 1918, to Constrictor), but Stull (1935) resurrected Acrantophis. It was again briefly placed in the genus Boa by Kluge (1991), based on cladistic analysis of morphological characters and historical biogeographic inference. The genus Acrantophis was again resurrected after molecular phylogenetic analysis of Neotropical and Malagasy lineages (Vences et al., 2001). The species epithet is occasionally given as A. dumerilii (e.g., Vences and Glaw, 2003), though this would be an incorrect representation of the epithet honoring the individual A. M. Duméril. No subspecies are recognized.
Type Specimen. Holotype was in the MSNM but was destroyed in 1943. Type locality was questionable, but likely Amboasary, Madagascar (Henkel and Schmidt, 2000).
Distribution. A species endemic to southern Madagascar south of Antananarivo (Vences and Glaw, 2003; Glaw and Vences, 2007). No subspecies are recognized, though phylogenetic analysis suggests that species boundaries and phylogeographic variation are not well characterized (Orozco-Terwengel et al., 2008). This species has been reported from Reunion Island (Guibé, 1958; Kluge, 1991), but these records (e.g., MNHN RA 0.8161) are in error (Vences and Glaw, 2003; Wallach and Glaw, 2009).
Conservation Status. This species is listed on CITES Appendix I but is assessed as Least Concern based on IUCN Red List criteria. The species is widespread and does not appear to show evidence of population decline, despite persecution and habitat loss, and is tolerant of disturbed habitats (Raxworthy et al., 2011a).
Acrantophis madagascariensis (Duméril & Bibron 1844)
Taxonomy. Described as Pelophilus madagascariensis. Boulenger (1893) assigned it to Boa, but Stull (1935) resurrected Acrantophis. It was again briefly placed in the genus Boa by Kluge (1991) based on cladistic analysis of morphological characters and historical biogeographic inference. Acrantophis was resurrected after molecular phylogenetic analysis of Neotropical and Malagasy lineages (Vences et al., 2001). No subspecies are recognized, though phylogenetic analysis suggests that species boundaries and phylogeographic variation in Acrantophis are not well characterized (Orozco-Terwengel et al., 2008).
Type Specimens. Syntypes MNHN RA 0.3133, MNHN RA 0.7275, MNHN RA 0.8636, all presumably from Mahajanga, Madagascar (Henkel and Schmidt, 2000).
Distribution. A species endemic to northern Madagascar and satellite islands (e.g., Nosy Be) north of Antananarivo, with some records along the west coast north of Morondava (Vences and Glaw, 2003; Glaw and Vences, 2007).
Sanzinia Gray, 1849
The genus Sanzinia consists of two species, recognized herein, endemic to the island of Madagascar and satellites. These are moderate-sized boas (to ∼2.5 m total length) that occur in primary and secondary forests, as well as heavily disturbed areas. They occur at ground level as well as in trees, are nocturnal, and their diet consists largely of mammals, including lemurs (Glaw and Vences, 2007; Eberle and Kappeler, 2008). Sanzinia madagascariensis is largely restricted to eastern Madagascar, while S. volontany is restricted to western Madagascar. Both species occur at elevations up to 1,300–1,600 m (Henkel and Schmidt, 2000; Glaw and Vences, 2007; Vences et al., 2011). A small contact zone is potentially present southwest of Toalagnaro in extreme southeastern Madagascar (Vences and Glaw, 2003). The two species are distinguishable based on geographic location, as well as coloration, with S. volontany being largely brown and lacking the green-colored scales characteristic of S. madagascariensis. A number of studies have recognized the evolutionary distinctness of S. madagascariensis and S. volontany based on mitochondrial and multilocus phylogenetic analyses (Vences and Glaw, 2003; Glaw and Vences, 2007; Orozco-Terwengel et al., 2008; Reynolds et al., 2014). The genus is listed on CITES Appendix I, which prevents international trade in live individuals or parts obtained from these species.
Sanzinia madagascariensis (Duméril & Bibron, 1844)
Taxonomy. Described originally as Xiphosoma madagascariensis (Duméril & Bibron 1844), the species was assigned to Sanzinia (Gray, 1849). Boulenger (1893) placed it in Corallus, but Stull (1935) brought it back to Sanzinia. It was subsequently placed in the genus Boa by Kluge (1991) based on cladistic analysis of morphological characters and historical biogeographic inference. The genus Sanzinia was resurrected following molecular phylogenetic analysis of Neotropical and Malagasy lineages (Vences et al., 2001). No subspecies are recognized.
Type Specimens. Syntypes MNHN RA 43 and MNHN RA 7329, likely from Nosy Be Island, Madagascar (Duméril and Duméril, 1851), though also given as from Ankarana, Madagascar (Henkel and Schmidt 2000). A third possible syntype is ZMB 6400 (Bauer et al., 2002; Wallach et al., 2014).
Distribution. A species endemic to eastern Madagascar (Henkel and Schmidt, 2000; Glaw and Vences, 2007) and some satellites (e.g., Nosy Boraha = Île Sainte-Marie).
Conservation Status. This species is listed on CITES Appendix I and is assessed as Least Concern based on IUCN Red List criteria. The species is widespread across eastern Madagascar and occurs in a variety of natural and modified habitats (Henkel and Schmidt, 2000; Vences and Glaw, 2003; Glaw and Vences, 2007). It is occasionally collected for the pet trade (Vences et al., 2011).
Sanzinia volontany Vences & Glaw, 2003
Taxonomy. Initially recognized as the subspecies Sanzinia madagascariensis volontany (Vences and Glaw, 2003). The lineage has been shown to be genetically distinct (Orozco-Terwengel et al., 2008; Reynolds et al., 2014) and somewhat phenotypically diagnosable (Vences and Glaw, 2003) from eastern Madagascar S. madagascariensis. No subspecies are recognized.
Type Specimens. Holotype ZSM 804/2001, an adult male from Ankarafantsika Reserve (Vences and Glaw, 2003).
Distribution. A species endemic to western Madagascar and some satellites (e.g., Nosy Komba) (Henkel and Schmidt, 2000; Glaw and Vences, 2007; Bora et al., 2010).
Conservation Status. The genus is protected under CITES Appendix I, though the species is not currently assessed under IUCN Red List criteria. The species is widespread across western Madagascar and occurs in a variety of natural and modified habitats (Henkel and Schmidt, 2000; Vences and Glaw, 2003; Glaw and Vences, 2007). It is occasionally collected for the pet trade (Vences et al., 2011).
CONSERVATION
Despite some species of booids being among the most iconic of reptiles in general and perhaps including some of the most commercially sought-after species of snakes (e.g., Boa constrictor, Corallus caninus), only 59% of the 66 species have received IUCN Red List assessments (Table 3). Of those that have received assessments, 25 (37.9%) were designated Least Concern (i.e., species that are widespread and abundant). Only three species (Chilabothrus angulifer, C. chrysogaster, and Ungaliophis continentalis) are considered Near Threatened (i.e., close to qualifying for one of the threatened categories), and four (Chilabothrus exsul, C. subflavus, Epicrates alvarezi, and Exiliboa placata) are considered Vulnerable (i.e., considered to be facing a high risk of extinction in the wild). Five species (Boa orophias, Chilabothrus granti, C. monensis, Corallus blombergii, and C. cropanii) have been assessed as Endangered (i.e., considered to be facing a very high risk of extinction in the wild). The most endangered boa species globally (Chilabothrus argentum) is listed as Critically Endangered (i.e., considered to be facing an extremely high risk of extinction in the wild).
Table 3.
Summary of IUCN Red List assessmentsa for each family and genus (number of species). Asterisks indicate accounts that are still in review.
![i0027-4100-162-1-1-t03.tif](ContentImages/Journals/harv/162/1/MCZ48.1/graphic/WebImages/i0027-4100-162-1-1-t03.jpg)
With two exceptions (E. placata and U. continentalis, both family Charinidae, subfamily Ungaliophiinae), all of the species that have received IUCN assessments that are not Least Concern or Data Deficient belong to the Boidae, a family of 36 species in the Neotropics (the Neotropical mainland and associated islands plus the West Indies). Boa orophias (St. Lucia) and all the Chilabothrus have island distributions, and the two species of Corallus have very restricted distributions on the South American mainland. Many islands and island archipelagos are among the world's biodiversity hotspots (e.g., the Caribbean, Madagascar, Polynesia-Micronesia; Mittermeier et al., 2011), and islands harbor a high percentage of the Booidae (37.9% of the species and 81.8% of the subspecies). Perhaps not surprisingly, then, it is the island-restricted species that have most often received IUCN and CITES assessments (CITES Appendix I; IUCN Near Threatened to Critically Endangered) suggesting or indicating strong concerns for those species. Crucially, very little is apparently known about the conservation status of the Erycidae, with only 1 of the 13 species receiving an IUCN listing of Least Concern (E. jayakari). Given our occasionally problematic understanding of species boundaries and distributions in this group, further systematic and conservation study is greatly needed.
There are occasional discrepancies between the IUCN assessments and those of CITES. Several species that received IUCN assessments of Least Concern are listed under CITES Appendix I (both species of Acrantophis, Chilabothrus inornatus, and Sanzinia madagascariensis). Similarly, the range-wide IUCN assessments are sometimes at odds with an assessment (for which IUCN criteria were employed) for a particular country. We have noted those discrepancies in the species accounts.
It is worth noting that, aside from distributions (many in biodiversity hotspots), general habitat, fortuitous field observations on diet, predation, reproduction, or laboratory-based analyses of diet or reproduction, we know remarkably little about the ecology (or natural history) of most species of booids. Fewer than 20% have been the focus of prolonged, dedicated field studies and nearly all the species that have been the focus of such work are members of the Neotropical Boidae (e.g., Boa imperator, Chilabothrus angulifer, C. chrysogaster, C. granti, C. monensis, Corallus grenadensis, C. cropanii, Eunectes murinus, E. notaeus) and North American Charinidae (Charina bottae, Lichanura trivirgata); a lone exception is the African calabariid Calabaria reinhardtii. The world's boas face multiple challenges to their survival and persistence. These include habitat loss and alteration, introduced predators (e.g., cats, dogs, mongooses), accidental killing (vehicular traffic on roads), intentional killing, restricted population sizes and ranges, climate change, natural disasters (hurricanes, typhoons, volcanic eruptions), environmental pollution, depleted or shifting prey bases, and commercial and cultural exploitation (with thousands exported from their countries of origin for the pet or skin trade; Montgomery et al., 2015).
ACKNOWLEDGMENTS
We are appreciative of support from the University of North Carolina Asheville and the Museum of Comparative Zoology, Harvard University. Milwaukee Public Museum librarian Ruth King was instrumental in procuring literature, and Addison Wynn (USNM) provided critical information on Chilabothrus subflavus and Candoia aspera. We thank Mark O'Shea for updates on the distribution of several species of Candoia and Phil Bowles for providing information on IUCN Red List assessments. We are grateful to two anonymous reviewers for helpful comments on previous versions of this manuscript.
LITERATURE CITED
Appendices
INDEX TO SCIENTIFIC NAMES
Bold indicates illustrations.
Acrantophis 2, 4, 37–38, 40
dumerili 37
madagascariensis 37–38, 40
Anguis 33–35
colubrina 33
jaculus 34
miliaris 35
Boa 2, 4–5, 18–19, 37–38, 40
anguiformes 35
annulata 16, 17
australis 25
canina 17
carinata 26
cenchria 20
conica 33
constrictor 4–6, 8, 39
constrictor 5
diviniloquus 17
imperator 6, 8
longicauda 6
occidentalis 6
orophias 8
ortonii 6
sabogae 7
diviniloqua 7
enydris 19
cookii 17
grenadensis 18
hortulana 18
imperator 6–7, 41
imperator 7
sabogae 7
inornata 13
johnii 35
murina 22
nebulosa 7
occidentalis 6
orophias 8, 39–40
ortonii 6
sigma 8
tatarica 36
variegata 26
Boidae 1– 2, 4, 40–41
Boinae 1
Bolyeria 1
Bolyeriidae 1
Booidae 1–2, 4, 40
Bothrops caribbaeus 8
Calabaria 23, 40
fusca 23
reinhardtii 23, 41
Calabariidae 2, 23, 40
Candoia 2, 23–26, 40–41
aspera 24, 41
aspera 24
schmidti 25
bibroni 24, 25
australis 25
bibroni 25
carinata 25–27
carinata 26
paulsoni 26–27
tepedeleni 26
paulsoni 24, 26, 28
mcdowelli 27
paulsoni 27
rosadoi 27
sadlieri 27
tasmai 28
vindumi 27–28
superciliosa 28
crombiei 29
superciliosa 28–29
Candoiidae 2–3, 23, 40
Casarea 1
Charina 2, 23, 29–32, 40
bottae 29–30, 41
trivirgata 31
umbratica 29
umbratica x bottae 30
Charinidae 2, 29, 40–41
Charininae 29
Chilabothrus 2, 4, 8–10, 12–14, 19, 40
angulifer 9, 39, 41
argentum 9, 40
chrysogaster 10, 39, 41
chrysogaster 10
relicquus 10
exsul 10, 39
fordii 11
agametus 11
fordii 11
manototus 11
gracilis 12
gracilis 12
hapalus 12
granti 12–13, 39, 41
inornatus 13, 40
maculatus 11
monensis 12–13, 39, 41
schwartzi 13
striatus 9, 14
exagistus 14
striatus 14
warreni 14
strigilatus 9, 14–15
ailurus 15
fosteri 15
fowleri 15
mccraniei 15
strigilatus 15
subflavus 15, 39, 41
Chrysensis batesii 16
Constrictor 4–5, 18, 37
constrictor 6–8
orophias 7
Corallus 2, 4, 16–19, 38, 40
annulata blombergi 17
annulatus 16–17
blombergi 17
batesii 16–17
blombergii 17, 39
caninus 16–18, 39
cookii 17, 19
cropanii 18, 39, 41
enydris 19
cookii 17
grenadensis 18, 41
hortulanus 18
cookii 17, 19
ruschenbergerii 19
Cubophis vudii 11
Cusoria elegans 34
Elgaria 29
Enygrus 24, 26
asper 24
schmidti 25
australis 25
bibroni 25
bibronii 25
carinatus 26, 29
paulsoni 26–27
superciliosus 28
Epicrates 2, 4, 9–10, 12–14, 16, 19–21, 40
alvarezi 20, 39
angulifer 9
assisi 20
cenchria 19–21
chrysogaster 10
schwartzi 13
crassus 20–21
fordi 11
fordii 11
manototus 11
gracilis 12–13
inornatus 11–13
maurus 21
monensis 12–13
relicquus 10
sabogae 7
striatus 10, 14
subflavus 15
Erebophis asper 24
aspera 24
Erycidae 2–3, 32–33, 35, 40–41
Eryx 2, 23, 32–36
borrii 33
colubrinus 33
loveridgei 33
conicus 33, 36
elegans 34
fodiens 34
indicus 35
jaculus 33, 34, 35
jayakari 34, 40
johnii 34–35
maculatus 35
miliaris 35–36
miliaris 35
nogaiorum 35
muelleri 35
reinhardtii 23
rufescens 33
somalicus 33, 35
speciosus 36
tataricus 35–36
speciosus 36
tataricus 36
thebaicus 33, 35
vittatus 33, 36
whitakeri 36
Eunectes 2, 4, 9, 16, 20–22, 40, 41
barbouri 22
beniensis 21
deschauenseei 22
murinus 21–22
notaeus 22, 41
scytale 22
Exiliboa 2, 29, 31–32, 40
placata 31, 39
Gongylophis 32–33, 35
muelleri 35
Homalochilus
chrysogaster 10
multisectus 14
striatus 14
strigilatus 14
Lichanura 2, 23, 29–32, 40
orcutti 30
roseofusca 31
simplex 30
trivirgata 30–31, 41
gracia 31
myriolepis 31
roseofusca 30–31
Loxocemus 1
Malayopython reticulatus 21
Pelophilus fordii 11
madagascariensis 37
Peropodum guatemalensis 32
Pseudoeryx 29
Python regius 24
Python sebae 21
Pythoninae 1
Rhoptrura 23
Sanzinia 2, 4, 38, 40
madagascariensis 38–40
volontany 39
volontany 38–39
Sanziniidae 2–3, 37, 40
Sceloporus 29
Tortrix bottae 29
Tropidoboa de bibron 25
Tropidophiidae 1
Tropidophis 1
Ungaliophiinae 31, 40
Ungaliophis 2, 29, 31–32, 39–40
continentalis 32, 39, 40
danieli 32
panamensis 32
Xenoboa 2, 18
cropanii 2
Xiphosoma 17, 19
annulatum 16
madagascariensis 38
ruschenbergerii 19