Mabuyinae Mittleman, 1952

Hedges, S. Blair & Conn, Caitlin E., 2012, A new skink fauna from Caribbean islands (Squamata, Mabuyidae, Mabuyinae) 3288, Zootaxa 3288 (1), pp. 1-244 : 23-29

Subfamily Mabuyinae Mittleman, 1952

Neotropical Skinks

Type genus. Mabuya Fitzinger, 1826

Diagnosis. Mabuyid lizards with cylindrical bodies, four limbs, digits present (pentadactyl), smooth dorsal scales, no distinct boundary between gulars and ventrals, lower eyelid with semitransparent disc, 28–31 presacral vertebrae, no pterygoid teeth, no auricular lobules, smooth (not keeled) dorsal scales, and 24–36 scale rows around midbody (Mausfeld et al. 2002; Miralles et al. 2009a; data herein). Although placentotrophic viviparity has been mentioned as a diagnostic character for this group (Mausfeld et al. 2002; Miralles et al. 2010), it also occurs in some African species ( Flemming & Blackburn 2003).

Content. Sixty-one species placed in 16 genera: Alinea gen. nov., Aspronema gen. nov., Brasiliscincus gen. nov., Capitellum gen. nov., Copeoglossum , Exila gen. nov., Mabuya , Manciola gen. nov., Maracaiba gen. nov., Marisora gen. nov., Notomabuya gen. nov., Orosaura gen. nov., Panopa gen. nov., Psychosaura gen. nov., Spondylurus , and Varzea gen. nov. ( Table 1).

Species Distribution

Capitellum mariagalantae sp. nov.; Marie-Galante Skink Marie-Galante, Guadeloupe

Capitellum metallicum ( Bocourt 1879) ; Lesser Martinique Skink Martinique

Capitellum parvicruzae sp. nov.; Lesser Saint Croix Skink St. Croix, U.S. Virgin Islands

Copeoglossum arajara ( Reboucas-Spieker 1981) ; Caatinga Brazil

Bronze Skink

Copeoglossum aurae sp. nov.; Greater Windward Skink St. Vincent, the Grenadines, Grenada, Trinidad, & Tobago.

Copeoglossum margaritae sp. nov.; Margarita Skink Isla de Margarita, Venezuela

Copeoglossum nigropunctatum ( Spix 1825) ; South American Eastern Bolivia, Brazil, Colombia, Ecuador, French Spotted Skink Guiana, Guyana, Peru, Suriname, & Venezuela

Copeoglossum redondae sp. nov.; Redonda Skink Redonda

Exila nigropalmata ( Andersson 1918) ; Amazonian Gracile Skink Bolivia, southwestern Brazil, & southern Peru

Mabuya cochonae sp. nov.; Cochons Skink Îlet à Cochons, Guadeloupe

Mabuya desiradae sp. nov.; Désirade Skink La Désirade & Terre de Bas (Îles de la Petite Terre), Guadeloupe

Mabuya dominicana Garman, 1887 ; Dominica Skink Dominica

Mabuya grandisterrae sp. nov.; Grande-Terre Skink Grande Terre, Guadeloupe

Mabuya guadeloupae sp. nov.; Guadeloupe Skink Basse-Terre, Guadeloupe

Mabuya hispaniolae sp. nov.; Hispaniolan Two-lined Skink Hispaniola

Mabuya mabouya ( Bonnaterre 1789) ; Greater Martinique Skink Martinique

Mabuya montserratae sp. nov.; Montserrat Skink Montserrat

Manciola guaporicola (Dunn 1935) ; Neotropical Small-handed Bolivia & Brazil

Skink

Maracaiba meridensis ( Miralles et al. 2005b) ; Merida Skink Venezuela

Maracaiba zuliae (Miralles et al. 2009) ; Maracaibo Skink Venezuela

Marisora alliacea ( Cope 1876) ; Costa Rican Four-lined Skink Eastern Costa Rica & southeastern Nicaragua

Marisora aurulae sp. nov.; Lesser Windward Skink St. Vincent, the Grenadines, Grenada, Trinidad, & Tobago.

Marisora brachypoda ( Taylor 1956) ; Middle American Short- Mexico, Belize, Guatemala, Honduras (including Utila limbed Skink & Guanaja islands), Nicaragua, & western Costa Rica

Marisora falconensis ( Mijares-Urrutia & Arends 1997) ; Venezuela & Colombia

Venezuelan Coastal Skink

Marisora magnacornae sp. nov.; Corn Island Skink Great Corn Island, Nicaragua

Marisora roatanae sp. nov.; Roatán Skink Isla de Roatán, Honduras

Marisora unimarginata (Cope 1862) ; Middle American Long- Western Costa Rica & Panama limbed Skink

Notomabuya frenata (Cope 1862) ; Southern Neotropical Skink Argentina, Bolivia, Brazil, & Paraguay

Orosaura nebulosylvestris (Miralles et al. 2009) ; Venezuelan Venezuela

Cloud Forest Skink

Panopa carvalhoi (Reboucas-Spieker & Vanzolini 1990) ; Brazil & Venezuela

Amazonian Blue-tailed Skink

Species Distribution

Panopa croizati ( Horton 1973) ; Turimiquire Blue-tailed Skink Venezuela

Psychosaura agmosticha ( Rodrigues 2000) ; Caatinga Sharp- Brazil nosed Skink

Psychosaura macrorhyncha ( Hoge 1946) ; Atlantic Forest Sharp- Brazil nosed Skink

Spondylurus anegadae sp. nov.; Anegada Skink Anegada, British Virgin Islands

Spondylurus caicosae sp. nov.; Caicos Islands Skink Caicos Islands, Turks & Caicos

Spondylurus culebrae sp. nov.; Culebra Skink Culebra & Culebrita, Puerto Rico

Spondylurus fulgidus (Cope 1862) ; Jamaican Skink Jamaica

Spondylurus haitiae sp. nov.; Hispaniolan Four-lined Skink Hispaniola

Spondylurus lineolatus ( Noble & Hassler 1933) ; Hispaniolan Ten- Hispaniola lined Skink

Spondylurus macleani ( Mayer & Lazell 2000) ; Carrot Rock Skink Carrot Rock, British Virgin Islands

Spondylurus magnacruzae sp. nov.; Greater Saint Croix Skink St. Croix & Green Cay, U.S. Virgin Islands

Spondylurus martinae sp. nov.; Saint Martin Skink St. Martin

Spondylurus monae sp. nov.; Mona Skink Mona, Puerto Rico

Spondylurus monitae sp. nov.; Monito Skink Monito, Puerto Rico

Spondylurus nitidus ( Garman 1887) ; Puerto Rican Skink Cayo Luis Peña, Cayo Norte, Culebra, Desecheo, Icacos, Puerto Rico, & Vieques

Spondylurus powelli sp. nov.; Anguilla Bank Skink Anguilla, Dog Island, & St. Barts

Spondylurus semitaeniatus ( Wiegmann 1837) ; Lesser Virgin U.S. Virgin Islands, including Capella Island, Little Islands Skink Buck Island, and St. Thomas; British Virgin

Islands including Fallen Jerusalem, Ginger Island,

Great Camanoe Island, Guana Island, Little

Thatch Island, Mosquito Island, Necker Island,

Round Rock, Salt Island, Tortola Island, and

Virgin Gorda

Spondylurus sloanii ( Daudin 1803) ; Virgin Islands Bronze Skink U.S. Virgin Islands , including Capella Island , Little Buck Island , Little Saba Island, Water Island, and

St. Thomas; British Virgin Islands including

Little Tobago Island, Norman Island, Peter

Island, and Salt Island.

Spondylurus spilonotus ( Wiegmann 1837) ; Greater Virgin Islands St. Thomas & St. John, U. S. Virgin Islands

Skink

Spondylurus turksae sp. nov.; Turks Islands Skink Grand Turk Island & Gibbs Cay, Turks & Caicos

Varzea altamazonica (Miralles et al. 2006) ; Upper Amazon Bolivia, Peru

Floodplain Skink

Varzea bistriata ( Spix 1825) ; Lower Amazon Floodplain Skink Brazil, & French Guiana

Distribution. Mabuyinae is restricted to the Western Hemisphere (Americas) and is distributed from central Mexico (Colima in the west and Veracruz in the east) throughout Middle America and South America (primarily east of the Andes) as far south as central Argentina and Uruguay ( Fig. 8 ). Most (38 species) of the currently known species of Mabuyinae are distributed only on islands in the Caribbean: the Turks and Caicos Islands, Jamaica,

Hispaniola, the Puerto Rico Bank and region (including Mona, Monito, and St. Croix), the Lesser Antilles, Trinidad, Tobago, Isla de Margarita, Great Corn Island, Isla de San Andrés, and Isla de Providencia ( Figs. 1 , 9–11 ; no localities on the latter three islands are known with precision and therefore they are not individually mapped).

Most species are lowland, occurring below 1000 m, but four species (unrelated) occur primarily above 1000 m, with one of those reaching nearly 4000 m ( Miralles et al. 2009b).

Etymology. The subfamily name Mabuyinae is derived from the Genus Mabuya Fitzinger (1826) , which is in turn the name used by native peoples of the Americas, especially the Antilles, for various types of lizards.

Remarks. Skinks have been placed in one of the earliest-branching families of squamates, Scincidae Oppel (1811 ; see comment below), splitting from other scinciformatan families at 170 Ma (Hedges et al. 2009). That is twice as old as the mean age (84 million years) and nearly three times the modal age (67 million years) of any squamate family lineage ( Hedges & Kumar 2009). The divergence times of the informal groups of lygosomine skinks (the Egernia , Eugongylus , Lygosoma , Mabuya , and Sphenomorphus groups; Skinner et al. 2011) are comparable to divergences among squamate families. Scincidae (sensu lato) is also the largest of the 26 families of lizards, containing more than one-quarter (27%; 1,503 species) of the 5,537 known lizard species ( Uetz et al. 2011). As a result, skink genera and higher taxa have become too large to be manageable, inhibiting taxonomic revision and description of new species (e.g., species comparisons are simpler for a small genus than for a large genus). At the same time, phylogenetic studies have shown significant support for informal groups and higher-level taxa (Honda et al. 2000; Reeder 2003; Honda et al. 2003; Austin & Arnold 2006; Skinner et al. 2011). Considering all of this, it is appropriate at this time to make taxonomic adjustments to facilitate skink systematics. We recognize seven families of skinks, with content corresponding to the currently recognized subfamilies Acontinae and Scincinae , and those five informal groups of Lygosominae (type genera in parentheses): Acontidae Gray, 1839 ( Acontias Cuvier 1817 ); Egerniidae Welch, 1982 ( Egernia Gray 1838b ); Eugongylidae Welch, 1982 ( Eugongylus Fitzinger 1843 ); Lygosomidae Mittleman, 1952 ( Lygosoma Hardwicke & Gray 1827 ); Mabuyidae Mittleman, 1952 ( Mabuya Fitzinger 1826 ); Scincidae Oppel, 1811 ( Scincus Laurenti 1768 ); and Sphenomorphidae Welch, 1982

The authorship of the Family Scincidae has been ambiguous ( Speybroeck et al. 2010), being attributed to either Oppel (1811) or Gray (1825). Both authors were explicit in proposing a family group name for Scincus but used spellings different from that which is currently used for the family. Regardless of those spellings, and following either ICZN Article 29.5 or the Law of Priority, we consider Oppel (1811) to be the author. We agree with Skinner et al. (2011) in placing Feyliniidae Camp (1923) in the synonymy of Scincidae (sensu stricto). Under this arrangement, and considering the relationships of the families ( Skinner et al. 2011), we erect the Superfamily Lygosomoidea for the families Egerniidae, Eugongylidae, Lygosomidae, Mabuyidae , and Sphenomorphidae. We consider the Infraorder Scincomorpha Camp (1923) to contain only the seven families of skinks. At a higher level ( Vidal & Hedges 2009), the Suborder Scinciformata Vidal & Hedges (2005) includes the Scincomorpha + Cordylomorpha; the latter taxon includes Cordylidae Mertens (1937) , Gerrhosauridae Boulenger (1884) , and Xantusiidae Baird (1859) . This proposed rearrangement now provides more taxonomic "room" for partitioning, organizing, and better managing the great diversity of skink species.

Within the Family Mabuyidae , formal and informal suprageneric groups have been recognized based on morphological and molecular data over the past four decades ( Greer 1970a, b; Greer 1979; Honda et al. 2000; Mausfeld et al. 2002; Mausfeld & Vrcibradic 2002; Carranza & Arnold 2003; Honda et al. 2003; Jesus et al. 2005; Austin & Arnold 2006; Miralles et al. 2011; Skinner et al. 2011). Knowledge of mabuyid phylogeny has increased sufficiently such that a more structured and formal taxonomy of this large family (formerly the Mabuya Group) is warranted. Family group names (subfamilies and tribes) have been used for some clades in the past, and we propose that the subfamily rank be used for them, in addition to the Subfamily Mabuyinae for the group under study here. Those subfamilies include, along with their former informal group name and type genus (for new taxa): Chioniniinae subfam. nov. ( Cape Verde Skinks, Chioninia Gray 1845 ), Dasiinae subfam. nov. (Asian Mabuya Group, Dasia Gray 1839 ), and Trachylepidinae subfam. nov. (African Mabuya Group, Trachylepis Fitzinger 1843 ). Taxonomic content, as well as morphological and molecular support for each of these taxa, are included in the studies cited above.

The Subfamily Mabuyinae is a monophyletic and well-defined group including all members of what was previously the Genus Mabuya , a clade of American skinks (Mausfeld et al. 2002; Miralles & Carranza 2010). Moreover, we have identified 16 clades within Mabuyinae that can be defined by molecular and morphological evidence. Most of these clades were already identified in earlier molecular phylogenies ( Vrcibradic et al. 2006; Whiting et al. 2006; Miralles et al. 2009b; Miralles & Carranza 2010). Some have been recognized informally as species groups and "complexes" in the past (and in two cases, genera), but the membership of these groups has come into focus. Giving them formal taxonomic recognition now is appropriate. Initially we considered the rank of subgenus but realized that the number of species (61) would place the genus among the largest genera of skinks; i.e., those likely to be partitioned into multiple genera in the near future. We also considered that dozens of additional species are likely to be described from Central and South America in coming years, based on molecular phylogenies indicating the presence of undescribed species ( Miralles et al. 2009b; Miralles & Carranza 2010; and analyses herein). Moreover, time estimates among the American clades (see below) are comparable to those among skink genera in other parts of the world ( Skinner et al. 2011). It is also normal taxonomic practice to create superspecific taxa in order to organize biodiversity, especially, as in this case, when many new species are added to a genus ( Simpson 1961). While it is true that new binomials necessitate some changes in databases and guides, partitioning of large genera is beneficial for systematic work because it creates manageable taxa for study. Thus, for all of these reasons we have chosen to recognize these clades at the generic level.

There is one species name within Mabuyinae that we must address here: Tiliqua albolabris ( Gray 1838b) . The name is based on a single specimen of unknown locality, presumably from the Americas. It was synonymized in " Mabouya agilis " only a few years later by the original author ( Gray 1845), and then placed under " Mabuia aurata " (= Trachylepis aurata ) by Boulenger (1887:189). Dunn (1936) placed it in the synonymy of his wideranging " Mabuya mabouya " distributed through much of the Americas. Otherwise the name has been conspicuously absent from the literature, including synonymies and the latest databases (Uetz et al. 2009). The original description is but a single sentence that says, "Golden-green with a brown streak on each side [of] the head and body (enclosing the eyes and ears) edged above and below with a pale streak; lips white; tail elongate" (Gray 1838:292). This short description is consistent with many species of mabuyines. However, the holotype (the only described by Gray (1838). Without a specimen, figure, locality, or useful (diagnostic) description, the name is a nomen nudum.

Figures

FIGURE 1. Map of Caribbean region. Neotropical skinks occur, or once occurred, on the islands indicated.

FIGURE 8. Distribution of the 16 genera of Neotropical skinks, Family Mabuyidae, Subfamily Mabuyinae. (A) Genera occurring in the greater Caribbean region. (B) Genera occurring exclusively in South America. (C) Additional genera occurring in South America, including one with representatives on Caribbean islands.

FIGURE 9. Map showing the distribution of mabuyine skinks occurring on islands in the central, southern, and western Caribbean region. See Fig. 1 for location of the islands. The islands and species are: (A) Jamaica (Spondylurus fulgidus; hollow circle = specimen not examined); (B) Hispaniola (Spondylurus haitiae sp. nov., circle; Mabuya hispaniolae sp. nov., triangle; Spondylurus lineolatus, squares); (C) Turks and Caicos Islands (Spondylurus caicosae sp. nov., circles; Spondylurus turksae sp. nov., triangles; hollow triangle is Cotton Cay record (see text); (D) Margarita (Copeoglossum margaritae sp. nov.); and (E) Roatán (Marisora roatanae sp. nov.).

FIGURE 10. Map showing the distribution of mabuyine skinks occurring on islands in the greater Puerto Rico region. See Figure 1 for location of the islands. The islands and species are: (A) Puerto Rico (Spondylurus nitidus); (B) Mona (Spondylurus monae sp. nov., squares) and Monito (Spondylurus monitae sp. nov., circle); (C) Desecheo (Spondylurus nitidus); (D) Vieques (Spondylurus nitidus, circle), Culebra (Spondylurus nitidus, circles; Spondylurus culebrae sp. nov., squares), Cayo Luis Peña and Cayo Norte (S. nitidus, circles); (E) most of the U.S. and British Virgin Islands (Spondylurus macleani, diamond; Spondylurus sloanii, squares; Spondylurus semitaeniatus, circles; Spondylurus spilonotus, triangles); (F) Anegada (Spondylurus anegadae sp. nov.); and (G) St. Croix and Green Cay (Capitellum parvicruzae sp. nov., circle; Spondylurus magnacruzae sp. nov.,

FIGURE 11. Map showing the distribution of mabuyine skinks occurring in the Lesser Antilles and Trinidad and Tobago. See Fig. 1 for location of the islands. The islands and species are: (A) Anguilla, Dog Island, and St. Barts (Spondylurus powelli sp. nov., squares), and St. Martin (S. martinae sp. nov., circle); (B) Guadeloupe: Basse-Terre (Mabuya guadeloupae sp. nov., square), Grande-Terre (Mabuya grandisterrae sp. nov., triangle), Îlet à Cochons (Mabuya cochonae sp. nov., star), La Désirade and Terre de Bas, Petite-Terre (Mabuya desiradae sp. nov., diamonds), and Marie-Galante (Capitellum mariagalantae sp. nov., circle); (C) Martinique (Mabuya mabouya, circles; Capitellum metallicum, triangle); (D) St. Vincent, the Grenadines, and Grenada (Copeoglossum aurae sp. nov., circles; Marisora aurulae sp. nov., triangles); (E) Redonda (Copeoglossum redondae sp. nov., circle) and Montserrat (Mabuya montserratae sp. nov., square); (F) Dominica (Mabuya dominicana); (G) St. Lucia (Alinea luciae), (H) Barbados (Alinea lanceolata); (I) Trinidad (Copeoglossum aurae sp. nov., circles; Marisora aurulae sp. nov., triangle), and (J) Tobago (Copeoglossum aurae sp. nov., circles; Marisora aurulae sp. nov., triangles). Hollow symbols in A and J are literature records (specimens not examined). Where no specific localities within an island are known for a species, a symbol is placed centrally on the island: Alinea luciae, Capitellum mariagalantae, Capitellum metallicum, and Copeoglossum redondae sp. nov. The scale in (A) applies to all islands.