Lesser Windward Skink, Marisora aurulae (Family Scincidae)

Marisora aurulae Hedges & Conn 2012: 122
Mabuia agilis — Boulenger 1887:191 (part)
Mabuia aenea — Garman 1887:53 (part)
Mabuya mabouia — Barbour 1935:129 (part)
Mabuya mabouya mabouya — Dunn 1936:544 (part)
Mabuya bistriata — Powell et al. 1996:82 (part)
Mabuya sloanii — Mayer & Lazell 2000:883 (part)
Mabuya falconensis — Miralles et al. 2009:609 (part)
Mabuya mabouya — Henderson & Powell 2009:292 (part) 

Note. Pinto-Sanchez et al. (2015) proposed to maintain the genus Mabuya as a diverse taxon widely distributed throughout Central America, South America, and several Caribbean islands. According to their appraisal of species delimitation and diversity, there are 58 species within Mabuya including four candidate species revealed in the present study plus one candidate species proposed by Miralles et al. (2010). Of these, 23 named species do not have molecular data and were not evaluated here, suggesting the total number of species could be higher still.

Pinto-Sánchez NR, Calderón-Espinosa ML, Miralles A, Crawford AJ, Ramírez-Pinilla MP. 2015. Molecular phylogenetics and biogeography of the Neotropical skink genus Mabuya Fitzinger (Squamata: Scincidae) with emphasis on Colombian populations. Molecular phylogenetics and evolution. 93:188-211.

Size. 80.9 mm SVL in males, 89 mm SVL females. Differs from all T&T lizards in having five digits on each hand and foot, overlapping head scales, 30–32 mid-body scale rows, the only exception is the Greater Windward skink. This species differs from Greater Windward skink, Copeoglossum aurea by having the parietals in contact, 2-4 chin scales in contact with the infralabials between then postmental and first sublabial. This species also has a more faded pattern, few if any dorsal spots and a dark lateral stripe that fades posteriorly. Distribution. The southern Lesser Antilles: Young's Island (a St. Vincent satellite), the Grenadines (Mayero Island, Carriacou, and Petit Bateau), Grenada, Trinidad, and Tobago.

The Rain Lizard, Kentropyx striata (Family Teiidae)

Lacerta striata Daudin 1802: 247
Centropyx striatus — Duméril & Bibron 1839: 151
Centropyx decodon Cope 1862: 495
Centropyx Renggerii Peters 1869: 63
Centropyx striatus — Boulenger 1885: 340
Kentropyx striatus — Peters & Donoso-Barros 1970: 152
Kentropyx striatus striatus - Hoogmoed 1973: 302
Kentropyx striatus viridicervis - Hoogmoed 1973: 312

Size: Males 124 mm, females 84 mm SVL; hatchlings 31−34 mm SVL; the tail is at least 2.2 times the SVL; hind legs about 63% SVL. Snout pointed from above, sloping in profile; rostral scale triangular, nasals in contact on the midline of the crown; scales on crown large plates; eye diameter three-fourths of eye-nostril distance; ear opening slightly less than the eye's diameter; tympanum visible. Anterior and lateral neck scales granular and keeled; mid-dorsum with 14 rows of large, overlapping, heavily keeled scales; laterally are 16 rows of small keeled scales; ventral scales large, heavily keeled, overlapping and arranged in 14 longitudinal rows, distinguishing it from Cnemidophorus (with eight longitudinal rows) and Ameiva(8−10 longitudinal rows); tail completely covered with large, keeled scales; upper surface of the hind limbs with small keeled scales, anterior surface of limbs with very large keeled scales; forelimbs with large scales on dorsal and anterior surfaces; fourth finger with 15−19 lamellae, the fourth toe with 24−28 lamellae; femoral pores number 6−7 in both sexes; males are larger than females and have a pair of preanal spurs on each side of central pre-anal scales; dorsum green with tan to white dorsolateral stripes, and blue-white spotting on the flanks; and brown lateral stripes; orange venter in males, white ventral surface in females; venter of tail orange in both sexes.

Habitat: Savanna wetlands and swamps.

A semi-aquatic lizard that climbs trees and shrubs; much of its distribution in Trinidad coincides with sugarcane and natural savanna, where it uses streamside (canals and ditches) trees and shrubs. It seeks refuge in water when approached while basking and during the rainy season it occupies small islands of vegetation until the water recedes; activity centered in the early morning but can be seen mid-day; sleeping on the tall grass. Diet: Insects and arachnids, also frogs and lizards. Reproduction: On Trinidad eggs laid in July-August and hatch in October-November. On the mainland, females reach sexual maturity at 74 mm SVL and produce 3−12 eggs per clutch, and produce more than one clutch per season; eggs laid at night in open, short, grassy fields; nest chamber about 3.5 cm deep with a 7 cm tunnel leading to a chamber at a 40° angle; eggs laid 10 October, hatched between 11−15 February, an incubation period of about 124 days.

Caribbean Treerunner, Plica caribeana (Family Tropiduridae)

Plica caribeana Murphy & Jowers 2013: 59
Hypsibatus agamoides – Court 1858: 440
Uraniscodon plica – Boulenger 1885, 2: 180 (in part)
Plica plica – Burt & Burt 1931: 282 (in part)
Tropidurus plica – Frost 1992: 1 (in part)


Size: 124 mm, tail twice SVL; smallest measured 57 mm SVL, probably about size at hatching. Identification: A brown-green lizard with heavily keeled, overlapping scales giving it a spiny appearance; snout rounded from above; rostral rectangular; canthus short and distinct; ear opening has anterior and ventral tufts of spiny scales, a character that will readily distinguish this lizard from all other Trinidad and Tobago lizards; mid-dorsal crest starts on posterior head with a few enlarged spines and extends onto tail; dorsal scales are keeled, overlapping, rhomboidal, and decrease in size laterally; ventral scales smooth, slightly larger than dorsal scales and overlap; well-developed lateral fold of spiny scales from axillary region to base of tail, a character that will separate it from all other Trinidad and Tobago lizards; front limbs well developed with long fingers; tail laterally compressed. Color green with red-brown transverse bands, each having black spots; limbs and tail banded; a red-brown band from eye to ear; venter orange-yellow in males and orange-brown in females; males with a white throat and black gular pouch, females with gray throat and brown gular pouch; has color change ability.

Forest and forest-edge using vertical surfaces of tree trunks, rock faces, and buildings, holes at the base of trees for escape.

Diurnal. Diet: Insectivorous but will feed on other invertebrates. Hatchlings are more terrestrial than adults; occur in colonies of 6−12 adults and juveniles, on trunks of large trees, stone walls, bridges, ruins, old houses, and caves; usually head down, but in whatever position they may be the head and forebody raised.

Is Plica caribeana still present on Tobago? In the early 1990's Plica was observed by JCM near Easterfield, Tobago. No specimens were collected. In 2011 I started searching for this lizard on Tobago and failed to find it. The locality where it was observed had been deforested and developed.

Murphy, J.C. and Jowers, M.J. 2013. Treerunners, cryptic lizards of the Plica plica group (Squamata, Sauria, Tropiduridae) of northern South America. ZooKeys, (355), p.49.

Multi-colored Tree Lizard, Polychrus marmoratus (Family Polychrotidae)

Polychrus marmoratus inhabits much of South America east of the Andes, including Guyana, French Guyana, Suriname, Venezuela, Colombia, Ecuador, Peru, Amazonian Brazil, and Brazil’s Atlantic Forest; Paraguay. It also occurs on the Isla de Margarita. Court (1858) first reported it from Trinidad, and it was first reported from Tobago by Barbour (1916). Its presence on Trinidad, Tobago, and the nearby Bocas Islands of Gaspar Grande, Monos, and Chacachacare was confirmed and detailed by Boos (1984, 1990). The multi-colored tree lizard is arboreal, spending much of its time in understory bushes, and the forest canopy. It is often found in pairs.  The diet includes a variety of invertebrates as well as fruits. It will drink droplets of water off of leaves. 

Mole's Day Gecko, Sphaerodactylus molei (Family Spaerodactylidae)

Sphaerodactylus molei Boettger 1894: 80.
Sphaerodactylus buergeri Werner 1900: 264.
Sphaerodactylus venezuelanus Roux 1927: 254.
Sphaerodactylus boettgeri: Donoso-Barros 1968.

Type locality: Caparo, Trinidad. Distribution. Antilles, Guyana, Venezuela, Margarita Island (Venezuela), Trinidad, Tobago.

Adult males and females measure 20-28.0 mm and 20.8-27.7 mm respectively; smallest specimen 16.6 mm. Dorsal body scales strongly keeled, minute, homogeneous, 63-83 from groin-axilla. Ventrals 31-39. Scales around mid-body 65-81. Supra-nasals two; larger anterior ones separated by 1-3 small inter-nasals. Rostral deeply notched behind, filled by 1-3 small scales; median cleft short. Fourth supralabial lies below anterior half of' eye; fourth infralabial below center of eye. Subcaudal scales to three times width of supra-caudals, alternately arranged. Dorsal surfaces light brown with a light dorsolateral stripe, variously faded or accentuated on trunk, but never broken. These are the smallest lizards on Trinidad and Tobago and habitat is difficult to define. They are often in crevices in buildings, but probably also occur under tree bark.

References

Boettger O. 1894. A preliminary list of the reptiles and batrachians of the Island of Trinidad. With descriptions of two new species. J. Field Natural. Club, Trinidad, Port of-Spain, 2: 77-90

Cole CJ, Townsend CR, Reynolds RP, MacCulloch RD, Lathrop A, 2013. Amphibians and reptiles of Guyana, South America: illustrated keys, annotated species accounts, and a biogeographic synopsis. Proceedings of the Biological Society of Washington 125 (4): 317-578; plates: 580-620.

Gorzula S & Senaris, JC. 1999. In: Contribution to the herpetofauna of the Venezuelan Guayana. I: a data base. Scientia Guaianae, Caracas, No. 8 [1998], 269.

Harris DM. The Sphaerodactylus (Sauria: Gekkonidae) of South America. University of Michigan Museum of Zoology, Occasional Papers (704):1-31.

Lynn WG. 1959. Some Reptiles and Amphibians from Trinidad. Herpetologica 15 (3): 113-117.

Mertens R. 1972. Herpetofauna tobagana. Stuttgarter Beitr. zur Naturkunde nr. 252 22 pp.

Rivas GA et al. 2012. Reptiles of Venezuela: an updated and commented checklist. Zootaxa 3211: 1–64

Roux J. 1927. Contribution à l’erpétologie du Vénézuéla. Verhandlungen dem Naturforschenden Gesellschaft in Basel 38: 252—261.

Werner F. 1900. Ueber Reptilien und Batrachier aus Columbien und Trinidad. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 50:262—272.

Luminous Lizard, Oreosaurus shrevei (Family Gymnophthalmidae)

Size: 42 mm SVL, but may exceed 50 mm. Tail is 1.7x SVL, hind legs 37–42% of SVL. Oreosaurus shrevei can be distinguished from all other Trinidad and Tobago microteiids by the separation of the nasal scales by the frontoparietal scale and the pattern of two lateral scale rows equaling the width of one dorsal row. Adult males have a single row of lateral ocelli. Fourth finger 9–13 lamellae; fourth toe 15–18 lamellae. Habitat: Lives along streams and is frequently found under rocks within a meter of the stream. Biology: Nocturnal, but may be active during the day; maybe semi-aquatic. Diet: small arthropods. Reproduction: deposits its eggs in arboreal situations. Females may use communal nesting sites. The lateral ocelli of males have been reported to be bioluminescent, but instead reflective. The species seems to be heat sensitive and uses cool microhabitats.

Common Green Iguana, Iguana iguana (Family Iguanidae)

Size. Males 580 mm SVL, 2010 mm TL; females 410 mm SVL, 1440 mm TL. The largest lizard in T&T and the western hemisphere. Snout blunt, head rectangular, body robust, a large pendulous dewlap, and a vertebral crest with the tallest spines on the nape will readily distinguish it from all other lizards. The body is covered with small, quadrangular, imbricate, keeled scales; ventral scales larger than dorsals. Adults usually green but may be gray, red, tan, or blue with obscure transverse bands; pale green beneath. Juveniles may be confused with adult Polychrus, which has a pointed snout and large plate like scales on the top of its head and moves slowly, iguanas are very fast.

This lizard is relatively common in urban areas with large trees. Juvenile cohorts may stay together, terrestrial during the day, arboreal at night; adults may be solitary or in colonies, along streams or ponds; may use urban parks or tree plantations; also in secondary and primary forests. Escape response is to dive into water from perch when disturbed. Iguanas are strict herbivores and they have special chambers in their digestive system for processing difficult to digest plant material. Reproduction. Males turn gold or red-orange during breeding season. A month of courtship precedes copulation, usually in early dry season; females construct a nest burrow, these may be communal shared with each female depositing eggs in a separate camber, clutch sizes average about 35 eggs; incubation is 65 to 115 days; hatchlings 70-80 mm; maturity reached in the second or third year at about 250 mm SVL. Predators. Snakes, raptors, cats, coatis, tyra, and humans are predators. If restrained they will bite and lash tail.

Spot-nosed Gecko, Gonatodes humeralis (Family Sphaerodactylidae)

A small, slender lizard, moderately long tail. Scales on snout larger than those on the back of the head. Males usually have a dark brown or red-brown dorsum; two rows of spots on flanks may be present; with some gray, black, yellow mottling; light spot of yellow, orange or gray on snout, ante-humeral bar or stripe present in both sexes. Female's dorsum brown-gray, indistinct vertebral stripe; snout pointed from above, round in profile; canthus rounded and indistinct; dorsum covered in small granular scales; ventrals larger than dorsal scales and imbricate; 16−21 lamellae on fourth toe wider than digit.

Coloration will separate this species from other Trinidad Gonatodes, but it may be most easily confused with G. ceciliae which has tiny scales in the canthal region, about 15 scales occur between the eye and the nostril, while G. humeralis has only seven or eight scales in this region.

Distributed in Venezuela, the Guianas, Trinidad, and Amazonia (Brazil, Bolivia, Ecuador, Peru).

Forest and forest edge habitats, usually low on tree trunks, rock walls, and in low vegetation as well as the canopy; male usually with several females. A diurnal, ambush predator. This lizard was studied at São Luís, Maranhão, Brazil. Its body temperature was strongly correlated with environmental temperatures (air and substrate), but air temperature had a greater effect on the males’ body temperatures after removing the effect of the substrate temperature; sexes differed in perch height, males perched higher, but the sexes did not differ in the tree trunk diameter used; both sexes use the largest tree trunks available in the environment; trunks with deeper leaf litter at the base were selected, deeper leaf litter is likely to provide a retreat from predators. It may also increase the humidity in the lizard’s microhabitat. Diet consists mostly of arthropods (isopods, spiders, pseudoscorpion, scorpions, millipedes, and insects) as well as land snails and annelid worms. Sexual maturity for both sexes is attained at 31 mm SVL. There is evidence of communal nesting in some populations with nest locations in ant nests, rotten palm trunks, rotten log, and a rotten liana; females produce a single egg per clutch and lay eggs in rapid succession, 52−110 day incubation period (longer for eggs laid in captivity).

House Gecko, Hemidactylus mabouia (Family Gekkonidae)

Gekko mabouia Moreau De Jonnès 1818: 138

Hemidactylus mabouia - Duméril & Bibron 1836: 362

Type locality: Restricted to St Vincent in the Lesser Antilles (Stejneger, 1904), but the type specimen apparently does not belong to the taxon to which this name is usually applied. Instead it is a H. angulatus, which is similar to those from the Greater Antilles and Colombia, particularly the latter (Kluge, 1969) [from Carranza & Arnold 2006].

Adults have a SVL up to 63 mm; hatchlings are about 20 mm SVL; tails are about 1.2x the body length. Diagnosis. Dorsum covered with small granular scales studded with trihedral; tubercles in 6–14 longitudinal rows. Ventral scales overlap and are large than dorsals. Upper labials 8–12, lower labials 7–9. Lamellae on fourth finger 7–9, lamellae on fourth toe 7–8. Total pores in males 29–33. Color change in this lizard can be dramatic. Similar species. It is most easily confused with Hemidactylus palaichthus which has a more spiny appearance and more lamellae on the base of toes four and five that extend to the base of the digit. Distribution. Probably occurs island wide and associated with human modified habitats, most often buildings and frequently near lights that attract insects. Other common names. House gecko, wall mabuia, African woodslave, twenty-four hours.

Duméril, A.M. C. and G. Bibron. 1836. Erpetologie Générale ou Histoire Naturelle Complete des Reptiles. Vol.3. Libr. Encyclopédique Roret, Paris, 528 pp.

Kluge, A. G. 1969. The evolution and geographical origin of the New World Hemidactylus mabouia-brooki complex (Gekkonidae, Sauria). Misc. PubI. Mus. Zool. Univ. Michigan 138:1-78.

Moreau de Jonnès, A. 1818. Monographie du mabouja des murailles, on Gecko Mabouja des Antilles. Bull. Scient. Soc. Philomath. Paris, Ser. 3,5:138-139.

Stejneger, L. 1904. The herpetology of Porto Rico. Rept. United States Natl. Mus. 1902: 549-724.

Trinidad Anole, Anolis trinitatis (Family Dactyloidae)

Anolis trinitatis Reinhardt & Lütken 1862: 269.
Anolis vincenti Garman 1887: 46.
Anolis trinitatis trinitatis — Underwood In Williams et al. 1959: 212.
Anolis trinitatis vincentii — Underwood In Williams et al. 1959: 213.
Anolis trinitatis — Schwartz & Henderson 1991: 349.

Holotype: ZMUC (UZM) R.37145. Type locality: “Trinidad”. Revised to “Kingstown, St. Vincent” by Lazell 1972.

Distribution: Anolis trinitatis inhabits the Lesser Antilles (St. Vincent, St. Lucia, Chateaubelair Island) and it has been introduced to Trinidad (Lazell 1972). May be present in Venezuela’s Peninsula de Paria (Donoso-Barros 1968).

Anolis trinitatis hybridizes with Anolis aeneus on Trinidad (Lazell 1972). However, hybrids show greatly reduced reproductive fitness (Gorman et al. 1971). Males, which reach 74 mm snout-to-vent, are green to green-blue, with blue stippling on the head and anterior trunk. They have yellow coloring on the throat and ventral surface, and the area around the eye is dark. Males have a large dewlap that extends into the abdominal region. Females are duller and have a smaller dewlap.

Synonymy: Boulenger (1885) listed Anolis trinitatis (and Anolis aeneus) as synonyms of Anolis alligator, which is now A. roquet.


Karyotype: Gorman et al. (1968) reported Anolis trinitatis and Anolis aeneus have 36 and 34 chromosomes, respectively. Hybrids have 35 chromosomes.

Literature
Donoso-Barros, R. 1968. The lizards of Venezuela (checklist and key). Carib. J. Sci. 8 (3-4): 105-122

Gorman GC. 1969. The zoogeography of Lesser Antillean Anolis lizards; an analysis based upon chromosomes and lactic dehydrogenases. Bull. Mus. Comp. Zool. Harvard 138 (3): 53-80.

Gorman GC, Atkins L. 1968. Natural hybridization between two sibling species of Anolis lizards: Chromosome cytology. Science 159 (March): 1358-1360

Gorman G, Licht P, Desshauser HC, Boos JO 1971. Reproductive failure among the hybridizing Anolis lizards of Trinidad. Syst. Zool. 20: 1–18

Lazell JD. 1972. The anoles (Sauria: Iguanidae) of the lesser Antilles. Bull. Mus. comp. Zool. Harvard 143 (1): 1-115

Poe S. 2004. Phylogeny of anoles. Herpetological Monographs 18: 37-89

Poe S. 2013. 1986 Redux: New genera of anoles (Squamata: Dactyloidae) are unwarranted. Zootaxa 3626 (2): 295–299

Reinhardt J, Lütken CH 1862. Bidrag tii det vestindiske Öriges og navnligen tii de dansk-vestindiske Oers Herpetologie. Vidensk. Meddel. Naturhist. For. Kjöbenhavn (10-18): 153-291 [1862]

Schwartz A, Henderson RW. 1991. Amphibians and Reptiles of the West Indies. University of Florida Press, Gainesville, 720 pp.

Williams EE, Quesnel VC, Kenny JS, Underwood G. 1959. The anoles of the Eastern Caribbean (Sauria, Iguanidae) Part. I. Preface; Part II. Two sibling species of anoles in Trinidad; Part. III. Revisionary notes. Bull. Mus. Comp. Zool. Harvard 121 (5): 185-226 + 1 plate

Anolis wattsi (Dactyloidae)

Anolis wattsi Boulenger 1894: 375. 

Type locality: Antigua. Restricted to St. John’s, St. John Parish, Antigua, by Lazell (1972). Syntypes: BMNH 1946.8.29.12-13. White and Hailey (2006) give a detailed account of this lizard from Trinidad. It is known from Anguilla, Antigua, Barbuda, St. Martin, St. Eustatius, St. Christopher, Nevis, St. Lucia, and was discovered in Trinidad in 1992. See this web site: http://www.ahailey.f9.co.uk/anolis.htm.

Giant Crown Anole, Anolis richardi Duméril & Bibron

Anolis Richardi Duméril & Bibron 1837: 141
Ptychonotus (Ctenodeira) richardii — Fitzinger 1843: 66
Anolis richardii — Boulenger 1885: 37
Anolis trossulus Garman 1887: 38
Anolis trossulus — Barbour 1916
Anolis richardii richardii — Underwood In Williams et al. 1959


Holotype: MNHN Ig. 53 (fide Underwood 1959); MCZ Type locality: “Tortola”. Revised to “Crown Point, Tobago” by Lazell 1972.

Distribution Lesser Antilles: Grenadines, Grenada, Tobago, St. Vincent; Surinam (introduced according to Hoogmoed 1981: 281). The Tobago population seems to have been introduced from the Grenadines or Grenada.

This is large, morphologically variable anole, with males reaching 140 mm snout-to-vent. It is often seen sitting on tree trunks with its head down. It has a dark green or brown dorsal surface, with a green-gray to yellow ventral surface. Its dewlap is orange, yellow, or gray-green. Females and juveniles often have a yellow or cream-colored lateral stripe. This species reaches relatively high population densities along streams.

Barbour T. 1916. Amphibians and reptiles of Tobago. Proc. Biol. Soc. Washington 29: 221-224

Boulenger G.A. 1885. Catalogue of the lizards in the British Museum (Natural History). Vol. 2, Second edition. London.

Creer DA, Kevin de Queiroz, Todd R. Jackman, Jonathan B. Losos and Allan Larson 2001. Systematics of the Anolis roquet series of the southern Lesser Antillies. Journal of Herpetology 35 (3): 428-441.

Duméril AMC, Bibron G. 1837. Erpétologie Générale ou Histoire Naturelle Complete des Reptiles. Vol. 4. Libr. Encyclopédique Roret, Paris, 570 pp.

Fitzinger L. 1843. Systema Reptilium, fasciculus primus, Amblyglossae. Braumüller et Seidel, Wien: 106 pp.

Garmam S. 1887. On West Indian reptiles. Iguanidae. Bull. Essex Inst. 19: 25-50.

Gorman GC, Atkins, L. 1968. The relationships of the Anolis of the roquet species group (Sauria: Iguanidae). III. Comparative study of display behavior. Breviora (284): 1-31

Hoogmoed MS. 1981. Introduced species of reptiles in Surinam. Notes on the herpetofauna of Surinam VIII. Amphibia-Reptilia 1: 277-285

Köhler G. 2014. Characters of external morphology used in Anolis taxonomy—Definition of terms, advice on usage, and illustrated examples. Zootaxa 3774 (2): 201–257

Lazell JD. 1972. The anoles (Sauria: Iguanidae) of the lesser Antilles. Bull. Mus. comp. Zool. Harvard 143 (1): 1-115

Malhotra A, Thorpe RS. 1999. Reptiles & Amphibians of the Eastern Caribbean. MacMillan, London & Oxford

Mertens R. 1972. Herpetofauna Tobagana. Stuttgarter Beitr. zur Naturkunde nr. 252 22 pp.

Poe S. 2004. Phylogeny of anoles. Herpetological Monographs 18: 37-89

Roughgarden J. 1995. Anolis Lizards of the Caribbean: Ecology, Evolution, and Plate Tectonics. Oxford University Press, 200 pp.

Schwartz A., Henderson RW. 1991. Amphibians and Reptiles of the West Indies. University of Florida Press, Gainesville, 720 pp.

Williams EE. Quesnel VC. Kenny JS, Underwood G. 1959. The anoles of the Eastern Caribbean (Sauria, Iguanidae) Part. I. Preface; Part II. Two sibling species of anoles in Trinidad; Part. III. Revisionary notes. Bull. Mus. Comp. Zool. Harvard 121 (5): 185-226 + 1 plate