Diapherodes Gray, 1835

Frank H. Hennemann, Oskar V. Conle & Daniel E. Perez-Gelabert, 2016, Studies on Neotropical Phasmatodea XVI: Revision of Haplopodini Günther, 1953 (rev. stat.), with notes on the subfamily Cladomorphinae Bradley & Galil, 1977 and the descriptions of a new tribe, four new genera and nine new species (Phasmatodea: “ Anareolatae ”: Phasmatidae: Cladomorphinae), Zootaxa 4128 (1), pp. 1-211 : 59-63

5.4. Genus Diapherodes Gray, 1835

Type-species: Mantis gigantea Gmélin, 1789: 2055 , by subsequent designation of Kirby, 1904a: 362.

Diapherodes Gray, 1835: 33 (in part). Burmeister, 1838: 560, 573 (in part). Westwood, 1859: 84 (in part). Saussure, 1872: 185 (in part). Stål, 1875: 31.

Bolivar, 1888: 140 (in part). Kirby, 1904a: 362 (in part). Redtenbacher, 1908: 434 (in part). Rehn, 1909: 200.

Wolcott, 1922: 23.

Wolcott, 1936: 35.

Rehn & Hebard, 1938: 53, pl. 4: 22 (♀). Wolcott, 1948: 50.

Wolcott, 1951: 50.

Moxey, 1972: 85 (in litt.; in part).

Langlois & Lelong, 1997: 43, fig. 20 (♂, ♀).

Brock, 1998c: 33.

Langlois, Lelong, Rastel, Polidori & Dorel, 2000: 46, fig. 19-g (♂, ♀) & photos 10–12 (egg).

Lelong, Langlois, Rastel & Dorel, 2003: 48, fig. 19a–g (♂, ♀) & photos 11–13 (egg).

Lelong & Langlois, 2005: 264.

Otte & Brock, 2005: 120 (in part).

Langlois, Lelong & Rastel, 2006: 42, figs. 19 a–g (♂, ♀, nymph), photos 5–6 (♀) & 27–29 (egg). Aplopus, Redtenbacher, 1908: 429 (in part).

Rehn, 1904: 68 (in part).

Rehn, 1909: 200.

Wetmore, 1916: 58.

Wolcott, 1923: 23.

Wolcott, 1936: 35.

Wolcott, 1948: 50.

Wolcott, 1951: 50.

Otte & Brock, 2003: 301 (in part).

Aplopus (?), Rehn & Hebard, 1938: 52.

Cyphocrana, St. Fargeau & Audinet-Serville, 1825: 445.

Audinet-Serville, 1831: 61.

Haplopus, Westwood, 1859: 85 (in part).

Kirby, 1904a: 363 (in part).

Mantis, Drury, 1773: 88 , 89, pl. 44: 1 & 50 (in part).

Fabricius, 1775: 274.

Fabricius, 1787: 227.

Gmélin, 1789: 2054-2055.

Olivier, 1792: 634.

Fabricius, 1793: 13.

Phasma, Fabricius, 1798: 187 , 188.

St. Fargeau & Audinet-Serville, 1825: 101.

Westwood, 1859: 34, pl. 22: 7, 7a, 7b (♀). Phasma (Diapherodes), de Haan, 1842: 109 . [Not: Diapherodes, Saussure, 1872: 183 . à Venupherodes n. gen.]

Description: ♀♀, ♂♂. Medium-sized to very large (body lengths: ♀♀ 92.0–197.0 mm, ♂♂ 60.0–126.0 mm) and moderately slender to very robust Haplopodini. ♀♀ in particular, often broad to very broad and massive insects with the body distinctly oval in cross-section and the thorax and abdomen conspicuously broadened; brachypterous. ♂♂ moderately slender, body cylindrical or sub-cylindrical in cross-section, with scale-like tegmina and well developed alae. Body surface of ♀♀ ± shiny, colouration usually plain bright green (rarely brown or with brown markings), ♂♂ brown or green. Head globose, indistinctly longer than wide. Vertex roundly convex, tuberculose and armed with a pair of ± prominent tubercles or blunt spines. Antennae with all antennomeres moderately thickened and longer than head and thorax combined. Scapus dorsoventrally flattened, longer than pedicellus. Pronotum tuberculose, with or without a ± prominent pair of blunt spines or tubercles in anterior half. Mesothorax at best slightly more than 2x the length of head and pronotum combined, in ♂♂ slender and roughly parallel sided, in ♀♀ ± strongly widened towards the posterior and constricted anteriorly. Mesonotum anteriorly armed with 2–6 distinct spiniform tubercles or spines; surface otherwise smooth, sparsely granulose or tuberculose. Meso- and metapleurae of ♀♀ ± strongly expanded and laterally projecting over mesonotum; armed with a longitudinal marginal row of ± prominent tubercles, teeth or spines. Meso- and metasternum either smooth, both granulose, or only the mesosternum with a few scattered granules. ♀♀ with scale-like tegmina which at best slightly project over posterior margin of metanotum. Alae at best slightly longer than the tegmina, but usually much shorter and may be vestigial. Tegmina of ♂♂ oval, longer than metanotum and with a ± distinct conical central hump. Alae at least reaching to posterior margin of abdominal tergum V. Anal region of alae transparent pink or violet (♀♀), plain transparent white, pink or orange (♂♂). Abdomen distinctly longer than head and complete thorax combined; unarmed in ♂♂, tergites sometimes finely multi-carinate in ♀♀. Abdomen of ♀♀ ± broadened and swollen basally or sub-basally, with segments II and/or III the broadest, then gradually narrowing towards the apex. Abdomen of ♂♂ slender and cylindrical with the terminal three segments slightly broader than previous. In ♀♀ segments II–IV transverse to quadrate, V to VII slightly longer than wide; in ♂♂ II–VII parallelsided and 1.5–2.5x longer than wide. Tergum VII of ♀♀ with lateral margins ± rounded and/or expanded, or with a triangular posterolateral lobe; in ♂♂ parallel-sided or with a small lobe posterolaterally. Praeopercular organ of ♀♀ indistinct; formed by a single posteromedian granule or wart-like tubercle on sternum VII. Cerci very small, oval in cross-section and tapered towards the apex; slightly in-curving in ♂♂, often hidden underneath anal segment in ♀♀. Epiproct very small and mostly hidden underneath anal segment. Vomer short and broad; roughly triangular. Subgenital plate of ♀♀ keeled, narrowed towards a ± pointed apex and extending considerably beyond apex of abdomen. Poculum of ♂♂ strongly convex and cup-like. Legs of moderate length and rather broad. Profemora at best about as long as mesothorax, hind legs not reaching to apex of abdomen. Medioventral carina of femora armed with a longitudinal row of spines (often indistinct on profemora). Both dorsal carinae of meso- and metafemora occasionally with a sub-apical lobe, the ventral carinae each with 1–2 distinct and pointed sub-apical spines. Some carinae of mid and hind legs may be ± elevated and ledge-like in ♀♀ of certain species. Basitarsi short, at best slightly longer than following two tarsomeres combined.

Eggs: Medium-sized to large (capsule length 3.7–5.6 mm), ovoid and cylindrical in cross-section; capsule 1.5–2.0x longer than wide. Polar-area ± impressed in lateral aspect. Capsule surface strongly coriaceous and covered with irregular raised tubercles and ridges to a variable degree. Micropylar plate generally drop or heartshaped, narrowed towards anterior end and broadened basally, at least 1/3 the length of capsule; central portion sculptured like capsule. Posterior end of micropylar plate with a wide triangular gap. Median line distinct. Operculum flat to slightly convex, circular and in the centre structured like capsule; no conspicuous central capitulum. Colouration irregularly grey or brown.

Differentiation: Closely related to the Cuban Venupherodes n. gen. and Haplopus Burmeister, 1838 . Slender representatives of the jamaicensis -group in particular resemble stockier representatives of Haplopus .

However, Diapherodes is absent in Cuba and easily distinguished from Venupherodes n. gen. by: the larger size (♀♀ in particular); unicolourous head of ♂♂; elevated and prominently armed mesopleurae which laterally project over the mesonotum; mesothorax which is gradually broadened towards the posterior; lack of a longitudinal median keel of the mesonotum; presence of tegmina and alae; basally or sub-basally broadened abdomen; simple praeopercular organ and longer subgenital plate of ♀♀. The eggs at once differ by the less strongly sculptures capsule surface and longer, elongate micropylar plate (→ Table 33 ).

From Haplopus it differs by: the broader and more robust body (♀♀ in particular); relatively shorter mesothorax (at best a little more than 2x the length of head and pronotum combined); shorter basitarsi and plain anal region of the alae of both sexes; oval body cross-section and more decidedly broadened meso- and metathorax of ♀♀, as well as the smaller and much more slender cerci ♂♂. Furthermore, ♀♀ are mostly plain bright green, whereas ♀♀ of Haplopus are brown or grey. The eggs clearly differ from those of Haplopus by having the capsule surface much more prominently sculptured and lacking a distinct hat- or knob-like central capitulum.

Comments: The classification of Diapherodes has varied through the years, but in the way treated by Redtenbacher (1908: 434) the genus was paraphyletic and as interpreted by Moxey (1972: 85 ff., in litt.) included five generic units. Several features of the insects and eggs indicate two species-groups within the genus, here recognized as the gigantea -group and jamaicensis -group. These two groups are geographically rather well separated, members of the gigantea -group being restricted to the Lesser Antilles, and the three known species of the jamaicensis -group occuring only on the Greater Antillean islands of Jamaica and Puerto Rico (→ 6.).

The systematic level of sub-genera was considered for these two groups of species in Diapherodes , but due to the lack of satisfying distinctive features in the eggs in particular, interpreting these solely as species-groups appears more appreciable. For a better understanding of the present arrangement a detailed comparison of these two species-groups here proposed appears warranted and is presented below (→ Table 10 ).

Distribution ( Fig. 375 ): Lesser Antilles ( Guadeloupe, Dominica, Martinique, Saint Lucia, Grenada, & Saint Vincent → gigantea -group, Fig. 381 ) and Greater Antilles ( Puerto Rico & Jamaica → jamaicensis -group).

Species & subspecies included:

I. Gigantea -group

1. Diapherodes angulata (Fabricius, 1793: 13) [ Mantis ]. rev. stat.

= Haplopus grayi Kaup, 1871: 36 . n. syn. [Distribution: Guadeloupe]

2. Diapherodes dominicae (Rehn & Hebard, 1938: 53) . n. comb. [Distribution: Dominica]

3. Diapherodes gigantea gigantea (Gmélin, 1789: 2055) [ Mantis ]. = Mantis gigas Drury, 1773: 89 , pl. 50 (♀). = Cyphocrana cornuta St. Fargeau & Audinet-Serville, 1825: 445 . [Distribution: Saint Vincent & Grenada]

4. Diapherodes gigantea saintluciae n. ssp.

[Distribution: Saint Lucia]

5. Diapherodes martinicensis Lelong & Langlois, 2005: 264 . [Distribution: Martinique]

II. Jamaicensis -group

6. Diapherodes achalus (Rehn, 1904: 68) [ Aplopus ]. n. comb. = Diapherodes longiscapha Redtenbacher, 1908: 435 . n. syn. [Distribution: Puerto Rico]

7. Diapherodes jamaicensis (Drury, 1773: 88, pl. 44: 1 (♂)) [ Mantis ]. = Mantis bispinosa Fabricius, 1775: 274 .

= Haplopus christopheri Westwood, 1859: 84 , pl. 33: 4, 4a (♀). n. syn. = Diapherodes glabricollis Gray, 1835: 33 . n. syn. = Haplopus murinus Redtenbacher, 1908: 429 . n. syn. = Dipaherodes pulverulentus Gray, 1835: 34 . n. syn. [Distribution: Jamaica]

8. Diapherodes laevicollis Redtenbacher, 1908: 434 . [Distribution: Jamaica]

Keys to the species and subspecies of Diapherodes

♀♀*

1. Meso- and metapleurae strongly expanded and with a prominent marginal row of teeth or spines; pronotum with a distinct pair of anterior spines; alae vestigial, <½ the length of tegmina; Lesser Antilles........................ 2. ( gigantea -group)

- Meso- and metapleurae tuberculose; no anterior spines on pronotum; alae ± as long as tegmina; Greater Antilles ( Jamaica & Puerto Rico)......................................................................... 6. ( jamaicensis -group)

2. Body length> 11 cm; mesonotum> 1.5x longer than head and pronotum combined;> 50 antennomeres................. 3

- Body length <10 cm; mesonotum only 1.3x longer than head and pronotum combined ( Fig. 144 ); 37 antennomeres; Martinique ( Fig. 381 ).............................................................................. martinicensis

3. Body length <15 cm; mesonotum with numerous spiniform tubercles or spines; carinae of mid and hind legs not conspicuously expanded and dorsal carinae not dentate............................................................... 4

- Very large, body length> 15 cm ( Figs. 120–122 ); mesonotum unarmed or at best with six enlarged spines in anterior portion; mid and hind legs with carinae ± strongly expanded and acutely dentate ( Fig. 132 ); Saint Vincent & Grenada ( Fig. 381 )........................................................................................... gigantea gigantea

4. Small, body length <14 cm; head bi-tuberculate............................................................. 5

- Large, body length> 14 cm, head prominently bi-cornute ( Fig. 107 ); Guadeloupe ( Fig. 381 ).................... angulata

5. Spines of meso- and metapleurae considerably longer than width of base, acute; tegmina large and overlapping ( Figs. 136–137 ); Saint Lucia ( Fig. 381 ).................................................... gigantea saintluciae n. ssp.

- Spines of meso- and metapleurae small and blunt ( Figs. 109–110 ); tegmina small and leaving a distinct gap between each other ( Figs. 109–110 ); Dominica ( Fig. 381 )............................................................... dominicae

6. Slender insects; mesothorax slightly widened towards the posterior,> 2.5x longer than head and pronotum combined; abdominal tergum VII gently broadened and indistinctly broader than VI............................................... 7

- Very massive insects ( Figs. 183–184 ); mesothorax strongly broadened towards the posterior, only about 2.1x longer than head and pronotum combined; abdominal tergum VII prominently laterally expanded; almost 2x broader than VI ( Fig. 186 ); Jamaica...................................................................................... laevicollis

7. Slender insects ( Figs. 167–170 ); head globose and distinctly bi-cornute ( Fig. 173–174 ); mesonotum granulose and with 2–4 anterior spines, not keeled; abdomen slender; Jamaica................................................. jamaicensis

- Broader insects ( Fig. 157 ); head longer than wide and faintly bi-tuberculate ( Fig. 159 ); mesonotum smooth and with a longitudinal median carina; abdomen distinctly broadened sub-basally; Puerto Rico................................. achalus

* body lengths given are including the subgenital plate ♂♂

1. Mesonotum irregularly granulose, tuberculose or spinose; metasternum and pleurae granulose; Lesser Antilles ( Fig. 381 ).......................................................................................... 2 ( gigantea -group) - Mesonotum unarmed (may have 2–8 anterior spines), metasternum and pleurae unarmed; Greater Antilles ( Jamaica & Puerto Rico).............................................................................. 6 ( jamaicensis -group) 2. Mesonotum tuberculose or spinose; colouration of body, tegmina and costal region of alae brown...................... 3 Mesonotum sparsely granulose; body, tegmina and costal region of alae green ( Figs. 101–102 ); Guadeloupe ( Fig. 381 ).................................................................................................... angulata 3. Head distinctly bi-cornute; anterior margin of tegmina white or pale yellowish..................................... 4 - Cephalad pair of tubercles obsolete ( Fig. 113 ); anterior margin of tegmina bright yellow and green with a dark green stripe interiorly ( Fig. 117 ); Dominica ( Fig. 381 )........................................................... dominicae 4. Mesothorax elongate> 1.7x longer than head and pronotum combined; alae at best reaching to posterior margin of abdominal tergum VI; ventral portion of tegmina without distinct colouration............................................... 5 - Mesothorax short, only 1.5x longer than head and pronotum combined; alae reaching to posterior margin of abdominal tergum VII ( Figs. 146–147 ); central portion of tegmina bright purple; Martinique ( Fig. 381 )...................... martinicensis 5. Body length> 9 cm; tubercles of pro- and mesonotum dull green ( Fig. 125 ); alae at best reaching half way along abdominal tergum VI; Saint Vincent & Grenada ( Fig. 381 )................................................ gigantea gigantea - Body length <9 cm; tubercles of pro- and mesonotum black; alae reaching to posterior margin of abdominal tergum VI; Saint Lucia ( Fig. 381 ).................................................................. gigantea saintluciae n. ssp. 6. Mesonotum with 2–8 distinct paired anterior spines; alae at best reaching to abdominal tergum VI; tegmina plain green; Jamaica............................................................................................. 7 - Mesonotum unarmed, at most with some minute granules; alae projecting over tergum VII; tegmina with a brown longitudinal median stripe ( Fig. 158 ); Puerto Rico................................................................ achalus 7. Slender; mesonotum 2x longer than head and pronotum combined; alae reaching to posterior of abdominal tergum VI ( Figs. 171–172 ); scapus and pedicellus greenish or brown ( Fig. 175 ).......................................... jamaicensis - More robust; mesonotum 1.8x longer than head and pronotum combined; alae reaching to posterior of abdominal tergum VII ( Fig. 185 ); scapus and pedicellus black............................................................. laevicollis

Figures

FIGURE 381. Maps. Distribution of the species of the gigantea - group of Diapherodes Gray, 1835 in the Lesser Antilles.

FIGURES 375 – 376. Maps. 375. Known distribution of Diapherodes Gray, 1835 and the two species-groups here defined: Gigantea - group [green] and Jamaicensis - group [red]; 376. Distribution of Haplopus Gray, 1835. Although there have not yet been definite records from the southern islands of the Bahamas Caicos and Turks Islands, Haplopus is most certainly also present on these islands. Hence, they are here included in the distribution range.

FIGURES 144 – 147. Diapherodes martinicensis Lelong & Langlois, 2005. 144. ♀ HT: Martinique, trace Duclos (dorsal view) [MNHN]; 145. ♀: captive reared from Martinique [coll. FH, No. 0750 - 1]; 146. ♂: captive reared from Martinique [coll. FH, No. 0750 - 8]; 147. ♂: captive reared from Martinique [coll. FH, No. 0750 - 5].

FIGURES 120 – 123. Diapherodes gigantea gigantea (Gmélin, 1789). 120. ♀: Saint Vincent [NHMW, No. 838]; 121. ♀: captive reared from Grenada [coll. FH, No. 0359 - 1]; 122. ♀: captive reared from Grenada [coll. FH, No. 0359 - 28]; 123. ♂, captive reared from Grenada [coll. FH, No. 0359 - 33].

FIGURES 124 – 135. Diapherodes gigantea gigantea (Gmélin, 1789). 124. ♀ head and pronotum [coll. FH, No. 0359 - 29]; 125. ♂ head and pronotum [coll. FH, No. 0359 - 33]; 126. ♀ mesothorax (dorsal view) [coll. FH, No. 0359 - 29]; 127. ♀ mesosternum [coll. FH, No. 0359 - 29]; 128. ♂ apex of abdomen (lateral view); 129. ♂ apex of abdomen (dorsal view); 130. ♀ apex of abdomen (lateral view); 131. ♀ apex of abdomen (dorsal view); 132. Right hind leg of ♀ from Grenada (posterolateral view) [coll. FH, No. 0359 - 1]; 133. Right hind leg of ♂ from Grenada (posterolateral view) [coll. FH, No. 0359 - 33]; 134. Egg (dorsal view) [coll. FH, No. 0359 - E] 135. Egg (lateral view) [coll. FH, No. 0359 - E].

FIGURES 103 – 108. Diapherodes angulata (Fabricius, 1793). 103. ♀ NT, apex of abdomen (lateral view) [MNCN]; 104. ♀ NT, apex of abdomen (dorsal view) [MNCN]; 105. Apex of ♂ abdomen (lateral view) [MNHN]; 106. Right hind leg of ♀ NT (posterolateral view) [MNHN]; 107. Head and pronotum of ♀ NT [MNHN]; 108. Head and pronotum of ♂ [MNHN].

FIGURES 136 – 139. Diapherodes gigantea saintluciae n. ssp .. 136. ♀ PT: Saint Lucia, Barre de l’Isle Trail [coll. ASPER]; 137. ♀ PT: Saint Lucia, Barre de l’Isle Trail [coll. ASPER]; 138. ♂ PT: Saint Lucia, Barre de l’Isle Trail [coll. ASPER]; 139. ♂ PT: Saint Lucia, Union Agricultural School [FSCA].

FIGURES 109 – 112. Diapherodes dominicae (Rehn & Hebard, 1938). 101. ♀ HT: Dominica [ANSP]; 102. ♀ Dominica, Croix [coll. ASPER]; 103. ♀ apex of abdomen (lateral view): Guadeloupe, Croix [coll. ASPER]; 104. ♀ apex of abdomen (dorsal view): Guadeloupe, Croix [coll. ASPER].

FIGURES 183 – 189. Diapherodes laevicollis (Redtenbacher, 1908). 183. ♀ HT: Jamaica (dorsolateral view) [NHMW, No. 839]; 184. ♀ HT: Jamaica (dorsal view) [NHMW, No. 839]; 185. ♂: Jamaica, Saint Andrew, Parrish [FSCA]; 186. ♀ HT: Apex of abdomen (dorsal view—apex of subgenital plate reconstructed based on ♀ in NHMJ) [NHMW, No. 839]; 187. ♀ HT: Apex of abdomen (lateral view—apex of subgenital plate reconstructed based on ♀ in NHMJ) [NHMW, No. 839]; 188. ♂ apex of abdomen (dorsal view) [FSCA]; 189. ♂ apex of abdomen (dorsal view) [FSCA].

FIGURES 167 – 170. Diapherodes jamaicensis (Drury, 1773). 167. ♀: captive reared from Jamaica (typical green specimen with a short subgenital plate) [coll. FH, No. 0428 - 20]; 168. ♀: captive reared from Jamaica (green specimen with a brownish wash and a very short subgenital plate) [coll. FH, No. 0428 - 19]; 169. ♀: captive reared from Jamaica (typical green specimen with a long subgenital plate and strongly developed leg-armature) [coll. FH, No. 0428 - 17]; 170. ♀, PLT of synonym Haplopus murinus Redtenbacher, 1908: Jamaica (brown colour-form) [NHMW, No. 834].

FIGURES 171 – 176. Diapherodes jamaicensis (Drury, 1773). 171. ♂: captive reared from Jamaica [coll. FH, No. 0428 - 26]; 172. ♂: captive reared from Jamaica [coll. FH, No. 0428 - 35]; 173. ♀ head and pronotum: captive reared from Jamaica (specimen with large and irregularly tuberculose horns) [coll. FH, No. 0428 - 31]; 174. ♀ head and pronotum: captive reared from Jamaica (specimen with moderately sized, conical horns) [coll. FH, No. 0428 - 32]; 175. ♂ head and pronotum: Jamaica: Portland, Parrish (specimen two large anterior mesonotal spines) [FSCA]; 176. ♂ head and pronotum: captive reared from Jamaica (specimen with a cluster of six anterior mesonotal spines) [coll. FH, No. 0428 - 37].

FIGURES 157 – 160. Diapherodes achalus (Rehn, 1904). 157. ♀: Puerto Rico, Mayaguez [ANSP]; 158. ♂: Puerto Rico, Parador Casa Grande [FSCA]; 159. ♀ head and pronotum: Puerto Rico, Mayaguez [ANSP]; 160. ♂ head and pronotum Puerto Rico, Parador Casa Grande [FSCA].

FIGURES 100 – 102. Diapherodes angulata (Fabricius, 1793). 100. ♀: Guadeloupe, Trois Rivères [MNHN]; 101. ♂: Guadeloupe [MNHN]; 102. ♂: Guadeloupe [MNHN].

FIGURES 113 – 119. Diapherodes dominicae (Rehn & Hebard, 1938). 113. ♂ head and pronotum: Guadeloupe, Picard River track [coll. ASPER]; 114. ♀ HT, head and pronotum [ANSP]; 115. ♂ apex of abdomen (lateral view) [coll. ASPER]; 116. ♂ apex of abdomen (dorsal view) [coll. ASPER]; 117. Tegmina of ♂ [coll. ASPER]; 118. Egg (lateral view), reproduced from REM-photograph by Lelong et al. (2003: appendix II, fig. 11); 119. Egg (dorsal view), reproduced from REM-photograph by Lelong et al. (2003: appendix II, fig. 12).

Tables

TABLE 10. Comparison of the gigantea- group and jamaicensis- group of Diapherodes Gray, 1835.

	gigantea -group	jamaicensis -group
Body length (♂♂)	64.0–126.0 mm	61.5–76.5 mm
Body length (♀♀)	92.0–197.0 mm	104.0–164.5 mm
Pronotum	With a pair of prominent anterior spines	No anterior spines
Mesonotum (♂♂)	Granulose or tuberculose	Smooth
Mesonotum (♀♀)	No median carina; surface armed with several blunt tubercles or spines	With an indistinct longitudinal median carina; surface ± granulose or tuberculose
Meso- and metapleurae (♂♂)	Granulose or tuberculose	Smooth
Meso- and metapleurae (♀♀)	Strongly deflexed and armed with a marginal row of prominent teeth or spines	Gently expanded and with a marginal row of granules or acute tubercles
Meso- and metasternum (♂♂)	Granulose	Smooth
Metasternum (♀♀)	Granulose or tuberculose	Smooth
Alae (♀♀)	Vestigial; <half the length of tegmina	± as long as tegmina
Abdominal tergum VII (♀♀)	Lateral margins gently rounded	With a small posterolateral lobe or distinctly deflexed laterally
Epiproct (♀♀)	Very small and mostly fully concealed by anal segment	Large triangular to shield-shaped and projecting over anal segment
Egg (capsule length)	4.1–5.6 mm	3.5–3.8 mm
Egg (micropylar plate)	Elongate; ± ½ as long as capsule	About as long as wide; ± 1/3 the length of capsule
Distribution	Lesser Antilles (Fig. 381)	Greater Antilles (Puerto Rico & Jamaica)