Nesocyrtosoma Marcuzzi

Hopp, Katie J. & Ivie, Michael A., 2009, A Revision Of The West Indian Genus Nesocyrtosoma Marcuzzi (Coleoptera: Tenebrionidae), The Coleopterists Bulletin (mo 8) 63, pp. 1-138 : 11-15

Taxonomy of Nesocyrtosoma Marcuzzi

As stated above, members of Nesocyrtosoma can be recognized by a combination of four characters — 1) clypeal membrane concealed; 2) epipleuron very broad with subparallel margins in the anterior third, then gradually narrowing and terminating at the base of the fifth abdominal ventrite; 3) femora slightly to moderately emarginate on the inner surface near the apex; and 4) female genital tract with an abruptly expanded apical spermatheca connected to a bursa-less vagina by an apical spermathecal accessory gland, and a cylindrical stiffened cuticular tube with squared-off ends at the base of the spermathecal accessory gland. The stiffened cuticular tube is not unique to Nesocyrtosoma as many tenebrionids seem capable of forming thicker cuticle in almost any location in the female genital tract (W. Tschinkel, pers. comm.), but it was discovered during this study that the particular structure of the stiffened cuticular tube in the species of Nesocyrtosoma is unique. It is thus termed the Nesocyrtosoma -stiffened cuticular tube (NSCT). There is a short section of either the vagina or accessory gland that is constricted at the beginning of the vagina (i.e., the point farthest from the vaginal opening), and the NSCT is located on the spermathecal accessory gland just above this constriction. The NSCT is unique in that it is a long, cylindrical tube that is weakly constricted near the apex. Both ends of the NSCT are squared-off, giving it a rectangular shape. The accessory gland can usually be seen within the NSCT and is weakly to strongly pigmented. This pigmentation becomes extremely weak at the point of constriction and then becomes very strong near the exiting point within the NSCT ( Figs. 77, 78 ).

The NSCT is different than the stiffened cuticular tube seen in Apsida chrysomelina Lacordaire and Hesiodus longitarsus Champion ( Fig. 81 ). The stiffened cuticular tube in these two species is located in the same place as in species of Nesocyrtosoma . However, the stiffened cuticular tube begins as a continuation of the vagina and is lacking the constriction seen in the NSCT. The shape of the stiffened cuticular tube is more robust and ovate in A. chrysomelina and H. longitarsus , and the apical end of the stiffened cuticular tube is tapered into the accessory gland ( Fig. 81 ). Finally, the accessory gland is lacking pigmentation within the stiffened cuticular tube except for a faint portion near the apex immediately before it exits the stiffened cuticular tube ( Fig. 81 ).

With the discovery of the NSCT, H. caraibus was eliminated from this study. Although the type series of this species was not examined, specimens determined by T. Spilman as H. caraibus , and the types of H. longitarsus , Hesiodus brasilensis Champion , Hesiodus conspurcatus Champion , Hesiodus debilis Champion , Hesiodus egaensis Champion , Hesiodus ellipticus Champion , Hesiodus jasoni Champion , and Hesiodus sordidus Champion were all examined. Two females of H. longitarsus , the type species of Hesiodus designated by Lucas (1920) and subsequently by Gebien (1938 – 1942 a), were dissected. It was determined that H. longitarsus has a distinctly different stiffened cuticular tube than Nesocyrtosoma (as discussed above). The specimens from Dominica and Guadeloupe determined as H. caraibus by T. Spilman have the same stiffened cuticular tube structure as in H. longitarsus . Thus, based on these dissections, H. caraibus should remain in Hesiodus and is not included any further in this revision.

Nesocyrtosoma is redefined based on Marcuzzi’s (1976, 1999) and Doyen’s (1989) work and current findings. Sixteen of the 18 species are redescribed, 27 new species are described, all but three species are illustrated, and a key to species is provided below.

Nesocyrtosoma Marcuzzi, 1976

Cyrtosoma (Nesocyrtosoma) Marcuzzi 1976: 137 [ Nesocyrtosoma Marcuzzi is a nomen nudum under ICZN Art. 13.3, because no type species was designated. We recognize the type species as Cyrtosoma (Nesocyrtosoma) inflatum Marcuzzi 1976 as per an application to the International Commission on Zoological Nomenclature by Hopp et al., submitted]; Marcuzzi 1984: 102; Marcuzzi 1991: 235, 246; Garrido and Gutiérrez 1996: 281–282; Marcuzzi 1999: 81.

Nesocyrtosoma ; Doyen 1989: 280–281, 299; Doyen and Poinar 1994: 45–50; Poinar et al. 2001: 205; Arillo and Ortuño 2005: 22; Perez-Gelabert 2008: 115; Vitali 2008: 11.

Hesiodobates Kaszab and Schawaller 1984: 1–6 , Fig. 1 ; Doyen 1989: 279; Poinar 1992: 154 [Type species { Hesiodobates antiquus Kaszab and Schawaller by original designation]. [Synonymy by Doyen and Poinar 1994: 45].

Cyrtosoma (Pachycyrtosoma) Marcuzzi 1999: 81–82 , Figs. 1–6 [Type species Cyrtosoma (Pachycyrtosoma) merkli Marcuzzi by original designation]; Perez-Gelabert 2008: 115. New Synonymy.

Nesocyrtoma Perez-Gelabert 1999: 31 [lapsus calami].

Serrania Garrido 2003: 49–52 , Fig. 1 [Type species Diaperis viridula Zayas by monotypy]. New Synonymy.

Diagnosis. The members of this genus are distinguished from all other genera of the Cnodalonini by the combination of the following external characters: clypeal membrane concealed beneath the epistoma ( Fig. 4 ); epipleuron terminating at 5th abdominal ventrite ( Fig. 33 ); femur with apical portion emarginate ( Fig. 37 ). The females are unique in the combination of an abruptly expanded apical spermatheca connected to a bursa-less vagina by an apical spermathecal accessory gland with a cylindrical stiffened cuticular tube at the base of the spermathecal accessory gland termed the Nesocyrtosoma -stiffened cuticular tube (NSCT) ( Fig. 77 ).

Redescription (modified from Marcuzzi 1976, 1999 and Doyen 1989). Male. 3.5– 12.0 mm long, 1.5–6.5 mm wide. Variable in color, usually black, sometimes purple, metallic greenish blue/purple or shining bronze. Body variable in shape, ranging from short and extremely convex to elongate and weakly convex ( Figs. 116, 117 , 157, 158 ). Glabrous, except with extremely short setae extending from punctures on head, legs and abdominal ventrites. Head small, round, punctate; punctures variable in size with short golden setae; frontoclypeal suture present; clypeal membrane concealed. Antenna 11-segmented; clavate to weakly clavate, nearly filiform; apical 4, 5 or 6 antennomeres apically widened forming a loose short to elongate club ( Figs. 15, 16, 17 ); apical antennomere either subcircular or longitudinally ovate; apical 4, 5, or 6 antennomeres with stellate sensoria ( Fig. 14 ). Mandibles barely projecting beyond labrum when closed; usually bifid at apex; prostheca present; molar lobe present, mola flat with small serrations on surface ( Figs. 5, 6, 7 ). Apical maxillary palpomere securiform or triangular ( Fig. 19 ); apical labial palpomere rectangular ( Fig. 21 ). Mentum with median keel raised anteriorly to a point ( Figs. 10, 12 ); ventral portion of eye sometimes reaching subgenal sulcus ( Fig. 9 ); postgena with or without distinct punctures ( Fig. 12 ). Dorsal surface of pronotum punctate, punctures variable in size and density. Pronotal marginal bead complete laterally, anterior and posterior margins usually with marginal bead effaced medially ( Fig. 24 ); anterior edge evenly emarginate; posterior edge bisinuate; anterior angles of pronotum acute ( Fig. 159 ) to obtuse ( Fig. 101 ), strongly ( Fig. 159 ) to weakly ( Fig. 249 ) produced and apically narrowly ( Fig. 159 ) to widely ( Fig. 101 ) rounded; lateral edge of pronotum usually evenly curved to base or sinuate at base ( Figs. 23, 24 ); pronotum usually evenly convex. Hypomeron with or without distinct punctures ( Fig. 27 ). Prosternal process usually apically rounded; marginal grooves opposite coxae either joined or not apically; setose to glabrous. Elytral striae variable, ranging from distinctly impressed striae to seemingly absent ( Figs. 213 , 116 , 221 , 108 ); scutellary striae variable, from 1–10 punctures long. Mesoventrite thin or broad antero-posteriorly, excavate, receiving prosternal process ( Figs. 123 , 135 ). Epipleuron extremely broad at base, gradually narrowing toward apex until terminating at the fifth abdominal ventrite ( Fig. 33 ). Metaventrite either 1/2 antero-postero length of mesocoxa or subequal to antero-postero length of mesocoxa; discrimen present; katepisternum present. Metathoracic wing vestigial or fully developed ( Fig. 34 ). Legs long and slender to short and broad; usually punctate with short, golden seta; trochanter off-set, highly reduced and setose ( Fig. 36 ); apical portion of femur emarginate ( Fig. 37 ), usually reaching beyond edge of elytron; tibiae straight to strongly curved; two apical tibial spurs on each tibia; pro- and mesotibia often with highly reduced apical ventral tooth, usually difficult to see; metatibia usually modified with apical ventral tooth varying in proximity to apex and variable in size, with a ventral longitudinal patch of setae, ventrally apically emarginate or rarely unmodified ( Figs. 39–73 ); tarsal formula 5-5-4; tarsi long, slender; first metatarsomere usually subequal in length to 2nd and 3rd tarsomeres; apical metatarsus usually subequal in length to first three tarsomeres combined; all but last tarsomere with dense pad of golden setae on venter; apical tarsomere with two distinct longitudinal rows of sparse setae. Abdomen with visible ventrites 1–3 connate, 4 and 5 movable with visible membrane; abdominal depressions on 4th and 5th ventrites usually reduced to indistinct slits, but sometimes present as strongly impressed pits ( Figs. 74, 75 ); intercoxal process of first ventrite usually rounded at apex, sometimes with nipple at apex ( Fig. 76 ). Ventral surface variably punctate; punctures variable in size, density and degree of impression; short golden seta emerging from each puncture. Aedeagus with basal piece and parameres simple; curved, canoe-like; parameres approximately 1/3 entire length of aedeagus. Ends of parameres range from blunt to sharply rounded ( Figs. 270–304 ).

Notes. Male genitalia do not provide useful characters for diagnosing species of Nesocyrtosoma . There is some variation in the degree of curvature, entire length (which correlates to species size), and the degree of sharpness of the apical portion of the parameres. However, these differences are difficult to quantify and are therefore not used to diagnose species.

Female. Identical to male, except tibia usually straight and always lacking apical tooth on metatibia. Ovipositor with pair of basal paraprocts, shorter than coxites; proximal coxite considerably longer than the three distal lobes combined ( Fig. 83 ). Spermathecal accessory gland extending apically from the bursa-less vagina; stiffened cuticular tube long, cylindrical, with squared-off ends; spermatheca greatly enlarged at the apex of the spermathecal accessory gland ( Fig. 77 ).

Larvae. Unknown.

Biology. Very little is known about the biology of Nesocyrtosoma , however, there is some general knowledge of the biology of the subfamily to which it belongs, Stenochiinae (5 Coelometopinae ) ( Bouchard et al. 2005). According to Doyen (1989), stenochiines primarily occupy forest and woodland habitats in the tropics and subtropics. Larvae inhabit rotten wood and adults are frequently found associated with various sorts of dead wood, and most are nocturnally active.

Nesocyrtosoma follows these generalizations — it has primarily been collected on dead wood, on trees, under bark and rocks, and by beating dead tree limbs, dead vines, thorn scrub and other vegetation. Specimens found under bark are often found during the day, while those on the surface are usually found at night, indicating they are nocturnally active. Specimens have also been taken at light and in flight intercept traps and Malaise traps, indicating some species may fly.

The males of most species of Nesocyrtosoma display sexual dimorphism in the form of a very small to large tooth on the ventral apical surface of at least the metatibia (there is usually a tooth on the pro- and mesotibia when one is present on the metatibia, but these are generally more difficult to detect than the tooth on the metatibia). In some species this modification is simply a patch of setae extending from the apex up to 1/3 of the length of the metatibia, but others have an apical tibial tooth, ranging from vestigial, which is generally difficult to detect, to quite sharp and large. These are likely used to help the male grasp the female during mating.

The larvae remain unknown, but are expected to be found in some type of rotten wood or debris.

Distribution. Species of Nesocyrtosoma are known from the West Indian islands of Cuba, Hispaniola, Puerto Rico, St. John and St. Thomas of the U. S. Virgin Islands, and Montserrat ( Figs. 262–269 ).

Etymology. Nesocyrtosoma is a Greek word in the neuter form. The International Commission on Zoological Nomenclature (1999) states in Art. 34.2 that the ending of a Latin or latinized adjectival or participial species-group name must agree in gender with the generic name with which it is at any time combined [Art. 31.2]; if the gender ending is incorrect, it must be changed accordingly (the author and date of the name remain unchanged [Art. 50.3.2]). Thus, the following species names are changed to agree with the neuter form of Nesocyrtosoma : Apsida cubanensis Kulzer to Nesocyrtosoma cubanense (Kulzer) , Cnodalon turquinensis Zayas to Nesocyrtosoma turquinense (Zayas) , Cnodalon parallelus Zayas to Nesocyrtosoma parallelum (Zayas) , Cnodalon cristalensis Zayas to Nesocyrtosoma cristalense (Zayas) , Cnodalon cuproso Zayas to Nesocyrtosoma cuprosum (Zayas) , and Cnodalon elongatus Zayas to Nesocyrtosoma elongatum (Zayas) .

Figures

Figs. 77–78. Female genitalia characters. 77) Nesocyrtosoma cuprosum female genital tract with Nesocyrtosoma-stiffened cuticular tube (NSCT); 78) N. cuprosum, closeup view of the NSCT.

Figs. 79–83. Female genitalia characters. 79) Cnodalon viride female genital tract lacking stiffened cuticular tube; 80) Nesocyrtosoma virens female genital tract with NSCT; 81) Hesiodus longitarsus female genitalia with a stiffened cuticular tube not consistent with NSCT; 82) N. virens defensive glands (stained with acid fuschsin); 83) Nesocyrtosoma cuprosum ovipositor.

Figs. 1–4. Dorsal view of head. 1) Nesocyrtosoma otus with punctures on the dorsal surface of head smaller than a single eye facet (sp 5 small puncture); 2) Nesocyrtosoma bankense with dorsal punctures on the surface of the head subequal or larger than a single eye facet (lp 5 large puncture; 3) Cyrtosoma picea with clypeal membrane exposed (CM 5 clypeal membrane); 4) Nesocyrtosoma inflatum with clypeal membrane concealed.

Figs. 29–34. Characters of the elytra and metathoracic wing. 29) Nesocyrtosoma skelleyi with elytral interstriae roundly convex; 30) Nesocyrtosoma cubanense with elytral interstriae flat; 31) Nesocyrtosoma otus with anterodorsal corners of elytron with a notch to receive prothorax; 32) Nesocyrtosoma hispaniolae with epipleuron terminating at base of 5th abdominal ventrite; 33) Strongylium sp. with epipleuron continuing to apex of 5th abdominal ventrite. 34) Nesocyrtosoma otus metathoracic wing.

Figs. 35–38. Characters of the legs. 35) Nesocyrtosoma inflatum with trochanters reduced and setose; 36) Nesocyrtosoma otus with metafemur emarginate at apex; 37) Nesocyrtosoma virens with protibia with dorsal longitudinal groove; 38) Nesocyrtosoma serratum with ventral metatibial serrations.

Figs. 112–119. 112–115: Nesocyrtosoma neibaense. 112) Habitus, dorsal; 113) Habitus, lateral; 114) Pronotum; 115) Thoracic sterna. 116–119: Nesocyrtosoma inflatum. 116) Habitus, dorsal; 117) Habitus, lateral; 118) Pronotum; 119) Thoracic sterna.

Figs. 153–160. 153–156: Nesocyrtosoma crenulatum. 153) Habitus, dorsal; 154) Habitus, lateral; 155) Pronotum; 156) Thoracic sterna. 157–160: Nesocyrtosoma otus. 157) Habitus, dorsal; 158) Habitus, lateral; 159 Pronotum; 160) Thoracic sterna.

Figs. 10–18. 10–13: Ventral portion of head. 10) Nesocyrtosoma guerreroi ocular depression; 11) Nesocyrtosoma bankense ocular depression; 12) Nesocyrtosoma virens distinct postgenal punctures; 13) Nesocyrtosoma otus median keel of mentum broad. 14–18: Antenna. 14) Nesocyrtosoma crenulatum with antennomeres 10 and 11 with stellate sensoria; 15) Nesocyrtosoma guerreroi with antenna clavate, apical six antennomeres enlarged to form club; 16) Nesocyrtosoma crenulatum with antenna moderately clavate, apical six antennomeres enlarged to form club; 17) Nesocyrtosoma inflatum with antenna weakly clavate, nearly filiform, apical six antennomeres enlarged to form club; 18) Strongylium atrum with filiform antenna.

Figs. 5–9. 5–7: Mandible of Nesocyrtosoma hispaniolae. 5) View from back; 6) View from side; 7) View from front. 8–9: Ventral portion of eye. 8) Nesocyrtosoma lacrima with ventral portion of eye not reaching the subgenal sulcus; 9) Nesocyrtosoma darlingtoni with ventral portion of eye reaching the subgenal sulcus.

Figs. 19–22. 19–20: Left maxilla; mp 5 maxillary palp. 19) Nesocyrtosoma hispaniolae with apical maxillary palpi securiform; 20) Tenebrio molitor (redrawn from Doyen 1966) with apical maxillary palpi rectangular. 21–22: Labium; lp 5 labial palp. 21) Nesocyrtosoma hispaniolae with apical labial palpi rectangular; 22) Strongylium atrum with apical labial palpi triangular.

Figs. 23–28. Characters of the pronotum, hypomeron and prosternal process. 23) Nesocyrtosoma bestiola pronotum with anterior marginal bead effaced, lateral marginal bead complete, and posterior margin absent; 24) Nesocyrtosoma inflatum pronotum with anterior and posterior marginal beads effaced medially and lateral marginal bead complete; 25) Nesocyrtosoma lacrima pronotum with anterior marginal bead absent, lateral marginal bead complete, and posterior marginal bead absent; 26) Strongylium atrum pronotum with anterior and posterior marginal bead complete and lateral marginal bead absent; 27) Nesocyrtosoma cubanense hypomeron with marginal bead and distinct punctures; 28) Cnodalon viride prosternal process apically narrowly rounded, nearly to a point.

Figs. 95–103. 95–98: Nesocyrtosoma gebieni. 95) Habitus, dorsal; 96) Habitus, lateral. 97) Pronotum; 98) Thoracic sterna. 99–103: Nesocyrtosoma tumefactum. 99) Habitus, dorsal; 100) Habitus, lateral; 101) Pronotum; 102) Thoracic sterna; 103) Lateral view of prosternal process showing apical, medial bump.

Figs. 243–250. 243–246: Nesocyrtosoma cubanense. 243) Habitus, dorsal; 244) Habitus, lateral; 245) Pronotum; 246) Thoracic sterna. 247–250: Nesocyrtosoma dentatum. 247) Habitus, dorsal; 248) Habitus, lateral; 249) Pronotum; 250) Thoracic sterna.

Figs. 209–216. 209–212: Nesocyrtosoma serratum. 209) Habitus, dorsal; 210) Habitus, lateral; 211) Pronotum; 212) Thoracic sterna. 213–216: Nesocyrtosoma skelleyi. 213) Habitus, dorsal; 214) Habitus, lateral; 215) Pronotum; 216) Thoracic sterna.

Figs. 217–224. 217–220: Nesocyrtosoma nearnsi. 217) Habitus, dorsal; 218) Habitus, lateral; 219) Pronotum; 220) Thoracic sterna. 221–224: Nesocyrtosoma productum. 221) Habitus, dorsal; 222) Habitus, lateral; 223) Pronotum; 224) Thoracic sterna.

Figs. 104–111. 104–107: Nesocyrtosoma turquinense. 104) Habitus, dorsal; 105) Habitus, lateral; 106) Pronotum; 107) Thoracic sterna. 108–111: Nesocyrtosoma hispaniolae. 108) Habitus, dorsal; 109) Habitus, lateral; 110) Pronotum; 111) Thoracic sterna.

Figs. 120–127. 120–123: Nesocyrtosoma merkli. 120) Habitus, dorsal; 121) Habitus, lateral; 122) Pronotum; 123) Thoracic sterna. 124–127: Nesocyrtosoma scabrosum. 124) Habitus, dorsal; 125) Habitus, lateral; 126) Pronotum; 127) Thoracic sterna.

Figs. 128–135. 128–131: Nesocyrtosoma lacrima. 128) Habitus, dorsal; 129) Habitus, lateral; 130) Pronotum; 131) Thoracic sterna. 132–135: Nesocyrtosoma simplex. 132) Habitus, dorsal; 133) Habitus, lateral; 134) Pronotum; 135) Thoracic sterna.

Figs. 39–49. Male metatibia of the Nesocyrtosoma s. s. species-group species. 39) Nesocyrtosoma bestiola; 40) N. ferruginea; 41) N. tumefactum; 42) N. turquinense; 43) N. hispaniolae; 44) N. neibaense; 45) N. inflatum; 46) N. merkli; 47) N. scabrosum; 48) N. lacrima; 49) N. simplex.

Figs. 50–61. Male metatibia of the Purple species-group species. 50) Nesocyrtosoma basilense; 51) N. dolosum; 52) N. purpureum; 53) N. crenulatum; 54) N. otus; 55) N. darlingtoni; 56) N. mutabile; 57) N. larseni; 58) N. parallelum; 59) N. cristalense; 60) N. cuprosum; 61) N. elongatum.

Figs. 62–73. Male metatibia of the Serrania species-group species. 62) N. bankense; 63) N. garridoi; 64) N. puertoricense; 65. N. serratum; 66) N. skelleyi; 67) N. nearnsi; 68) N. teresitae; 69) N. curvum; 70) N. guerreroi; 71) N. virens; 72) N. cubanense; 73) N. dentatum.

Figs. 74–76. Abdominal characters. 74) Nesocyrtosoma inflatum with abdominal depressions on 4th and 5th ventrites reduced to indistinct slits (AbS 5 abdominal slits); 75) Nesocyrtosoma otus with abdominal depressions on 4th and 5th ventrites present as impressed pits (AbP 5 abdominal pits); 76) Nesocyrtosoma skelleyi with first visible abdominal ventrite with a nipple at the apex.

Figs. 270–280. Illustrations of male genitalia in dorsal and lateral view of the Nesocyrtosoma s.s. species-group species. 270) Nesocyrtosoma bestiola; 271) N. ferruginea; 272) N. tumefactum; 273) N. turquinense; 274) N. hispaniolae; 275) N. neibaense; 276) N. inflatum; 277) N. merkli; 278) N. scabrosum; 279) N. lacrima; 280) N. simplex.

Figs. 281–292. Illustrations of male genitalia in dorsal and lateral view of the Purple species-group species. 281) Nesocyrtosoma basilense; 282) N. dolosum; 283) N. purpureum; 284) N. crenulatum; 285) N. otus; 286) N. darlingtoni; 287) N. mutabile; 288) N. larseni; 289) N. parallelum; 290) N. cristalense; 291) N. cuprosum; 292) N. elongatum.

Figs. 293–304. Illustrations of male genitalia in dorsal and lateral view of the Serrania species-group species. 293) Nesocyrtosoma bankense; 294) N. garridoi; 295) N. puertoricense; 296) N. serratum; 297) N. skelleyi; 298) N. nearnsi; 299) N. teresitae; 300) N. curvum; 301) N. guerreroi; 302) N. virens; 303) N. cubanense; 304) N. dentatum.

Figs. 262–264. Nesocyrtosoma species distribution maps. 262) Distributions of N. bestiola, N. ferruginea, N. merkli, N. altagracia, N. basilense, N. dolosum, N. darlingtoni, N. cristalense, and N. dentatum; 263) Distributions of N. virens, N. purpureum, N. crenulatum, N. otus, and N. elongatum; 264) Distributions of N. tumefactum, N. turquinense, N. scabrosum, N. simplex, N. parallelum, N. fernandoi, N. serratum, N. teresitae, and N. productum.

Figs. 265–267. Nesocyrtosoma species distribution maps. 265) Distributions of N. gebieni, N. hispaniolae, N. neibaense, N. inflatum, N. skelleyi, and N. nearnsi; 266) Distributions of N. mutabile, N. garridoi, N. guerreroi, and N. cubanense; 267) Distributions of N. larseni and N. cuprosum.

Figs. 268–269. Nesocyrtosoma species distribution maps. 268) Distributions of N. bankense, N. curvum, and N. puertoricense; 269) Distribution of N. lacrima on Montserrat.