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A new Cryptotermes (Blattodea (Isoptera): Kalotermitidae) from Honduras and known distribution of New World Cryptotermes species.

Cryptotermes Banks (Blattodea (Isoptera): Kalotermitidae), now with 29 endemic species (31 total) is the most speciose kalotermitid genus in the New World. The West Indies and Caribbean mainland, now with 22 species, has a disproportionately diverse number of Cryptotermes compared to the rest of the world. After the world revision by Bacchus (1987), the West Indian group was revised by Scheffrahn & Krecek (1999), adding 12 new species. Another new West Indian species, C. bracketti Scheffrahn & Krecek, was added by Scheffrahn et al. (2006). The latest mainland endemic Cryptotermes to be described include C. abruptus Scheffrahn & Krecek from Mexico (Scheffrahn et al. 1998) and C. colombianus Casalla, Scheffrahn & Korb from Colombia (Casalla et al. 2016).

Cryptotermes brevis (Walker), endemic to the coastal desert of Peru and Chile (Atacama), is the most economically important and widespread kalotermitid pest of dry wood worldwide (Scheffrahn etal. 2009). Both Cryptotermes dudleyi Banks, a pest species from the Indian subcontinent, and C. havllondi (Sjostedt), a pest from equatorial Africa, have been introduced by human commerce to the New World. I herein describe Cryptotermes garifunae sp. nov. from a single colony collected on the Caribbean coast of Honduras. I also update the distribution of all described Cryptotermes species in the New World from records in the University of Florida Termite Collection and from the literature.

Materials and Methods

Microphotographs (Figs. 1, 2) were taken as multi-layer montages using a Leica M205C stereomicroscope controlled by Leica Application Suite version 3 software (Leica Geosystems, Inc., Norcross, Georgia, USA). Preserved specimens were taken from 85% ethanol and suspended in a pool of Purell[R] Hand Sanitizer (GOJO Industries, Akron, Ohio, USA) to position the specimens on a transparent Petri dish background.

Cryptotermes distribution records were taken either from unpublished localities in the University of Florida Termite Collection in Davie, Florida, or from the literature (Table 1). Distribution maps (Figs. 3-5) were prepared using ArcMap 10.3 software (ESRI, Redlands, California, USA).

Results

Cryptotermes garifunae Scheffrahn sp. nov. 2018 (Figs. 1, 2)

DEALATED MALE IMAGO (Fig. 1, Table 2). Head and nota light yellowish brown. Postclypeus hyaline. Chevron pattern on fore wing scales slightly darker than mesonotum. Legs very light yellow concolorous with abdominal sternites. Eyes dark grey, occupying 2/5 distance between vertex and genal margin, the latter of which are closer; ellipsoid with rectate margin at antennal socket. Ocelli moderately large, hyaline, touching eyes; oval except for acute wisp at dorsal margin. Antennae article formula 2 > 3 < 4 < 5. Pronotum wider than long, distinctly narrower than head width at eyes; anterior and posterior margins nearly rectate, sides slightly convex. Arolia present.

SOLDIER (Fig. 2, Table 3). Head, in lateral view, grading from hyaline at the cervical margin to dark ferruginous brown at frontal flange; in dorsal view coloration is a patchwork of reddish brown to dark ferruginous brown corresponding with thickness of cuticular rugosity.

Mandibles concolorous with frons. Anterior pronotal margin yellowish brown, remainder yellowish. Head capsule cuboidal in dorsal view, sides nearly parallel until anterior protrusion of frontal flange; dorsal outline of head capsule, in lateral view, forming a weak "s" shape from flange to occiput.

Texture of dorsal rugosity moderate; more rugose in anterior half including frontal flange and frons. Frons deeply concave. Frontal flange robust, elevated, with median notch continuous with mid-vertex concavity; in dorsal view, flange forms circular 120[degrees] arch. Eye spots very faint.

Labrum short, apex upturned. Mandibles short, angular, bent about 120[degrees]; finely rugose. Dentition weakly developed.

In dorsal view, frontal horns large, globular, projecting beyond the frontal flange reaching the posterior margin of the postclypeus; genal horns small blunt, apex in line with anterior margin of antennal socket. In lateral view, frontal horns nearly semicircular, projecting beyond base of genal horns. Antenna with 11 articles; formula 2 > 3 = 4 < 5. Anterior margin of pronotum incised with weak irregular sinuosity; anterolateral corners square, lateral margins and posterior margin form an evenly rounded outline.

TYPE MATERIAL

HOLOTYPE soldier HONDURAS: Kerala, Laguna Guaimoreto (16.0132[degrees]N, 85.9184[degrees]W, elev. 6 m asl), 29-V-2007, J.A. Chase (UF no. HN273).

ETYMOLOGY

Named after the Garifuna people who live along the coastline of Honduras.

DIAGNOSIS

The dealated imago of C. garifunae, along with C. fatulus (Light) and C. undulans Scheffrahn & Krecek, are the smallest of the New World Cryptotermes. However, the C. garifunae imago has a lighter yellowish coloration than the other 2. The soldier of C. garifunae is the smallest of the New World Cryptotermes with the exception of C. fatulus (Light) which is of similar size but lacks head capsule rugosity.

In the key given by Scheffrahn and Krecek (1999), the new species will key out at couplet 10 leading to C. darlingtonae sp. nov. and modified as follows:
10. Frontal horns, in lateral view, about 3 times larger
than genal horns                                                   11
10'. Frontal horns, in lateral view either subequal or
about 5 times larger than genal horns                              12
11. Genal horns projecting forward and only slightly
recessed behind frontal horns, left mandible 0.68 to
0.75 mm long (Figs. 40-42)                                          C.
darlingtonae sp. nov.
11'. Genal horns blunt, apex in line with anterior
margin of antennal fossae, left mandible 0.57 mm long      C. garifunae
12. Frontal horns, in lateral view, about 5 times
larger than genal horns; genal horns projecting more
dorsally and greatly recessed behind frontal horns,
left mandible 0.63 to 0.72 mm long (Figs. 67-69)           C. pyrodomus
12'. Frontal horns, in lateral view, subequal;
smaller species, left mandible 0.56 to 0.62 mm long
(Figs. 19-21)                                            C. aequacornis
                                                          n. sp.


In addition to Cryptotermes localities reported in Casalla et al. (2016), Scheffrahn & Krecek (1999), Scheffrahn et al. (2003), and Scheffrahn et al. (2009); Figures 3 to 5 include 1,452 new Cryptotermes records and localities recorded in the University of Florida Termite Collection. Only endemic C. brevis localities are included in these figures because of its expansive non-endemic pest localities (Scheffrahn et al. 2009). Some localities yielded more than 1 sample of the same Cryptotermes species. New records are from Florida and Georgia, the West Indies (The Bahamas, Cayman Islands, Cuba, Dominica, Dominican Republic, Grenada, Guadeloupe, Haiti, Jamaica, Puerto Rico, Turks and Caicos Islands, Trinidad and Tobago, and The US Virgin Islands), Central America (Belize, Guatemala, Honduras, Mexico, Nicaragua, and Panama), and South America (Bolivia, Ecuador, French Guiana, Paraguay, Peru, and Venezuela).

Discussion

Although the imago morphology is quite conserved, the head capsules of Cryptotermes soldiers are variously adorned with protuberances and rugosities that facilitate their phragmotic defensive strategies. The most extreme case of phragmosis is exemplified by Cryptotermes cryptognathus from Jamaica, which has a wine cork-shaped head capsule and functionless mandibles (Scheffrahn et al. 1998). Cryptotermes chasei, on the other hand, has very long crushing mandibles and weak cephalic phragmosis (Scheffrahn 1993). Cryptotermes garifunae is intermediate between these 2 soldier forms.

Presently, C. garifunae and C cubioceps are the only Cryptotermes known from their type localities. Cryptotermes bracketti is known only from San Salvador Island, The Bahamas, but it is present throughout the island. All other New World Cryptotermes species show much greater distributions (Figs. 3-5). Cryptotermes cubioceps was described by Emerson (1925) from a single soldier collected in Guyana. It has not been collected again, even though considerable collecting efforts have been conducted in French Guiana (Bourguignon et al. 2011; Davies 2002). Whereas C. garifunae is the smallest New World Cryptotermes, C. cubioceps, with a head width of 2.12 mm, is the largest.

Figures 3 to 5 have reduced the Wallacean shortfall, defined as the state of incompleteness in understanding of geographical distributions of taxa (Lomolino 2004) for New World Cryptotermes. The maxium range extensions are increased as follows: C abruptus - 437 km; C aequicornis - 864 km; C. cavifrons - 681 km; C cylindroceps - 471 km; C. chacoensis - 379 km; and C veruculosus - 2,447 km.

Acknowledgments

Thanks to Jim Chase for persisting through thick brush and a heavy mosquito presence to collect C. garifunae.

References Cited

Bacchus S. 1987. A taxonomic and biometric study of the genus Cryptotermes (Isoptera: Kalotermitidae). Tropical Pest Bulletin 7: 1-91.

Bourguignon T, Leponce M, Roisin Y. 2011. Beta - diversity of termite assemblages among primary French Guiana rain forests. Biotropica 43: 473-479.

Casalla R, Scheffrahn R, Korb J. 2016. Cryptotermes colombianus a new drywood termite and distribution record of Cryptotermes in Colombia. ZooKeys 596: 39-52.

Constantino R. 2000. A new Cryptotermes from the Brazilian Atlantic forest (Isoptera: Kalotermitidae). Sociobiology 36: 525-530.

Constantino R, Cancello EM. 1992. Cupins (Insecta, Isoptera) da Amazonia Brasileira: distribuicao geografica e esforco de coleta. Revista Brasileira de Biologia 52: 401-413.

Davies RG. 2002. Feeding group responses of a Neotropical termite assemblage to rain forest fragmentation. Oecologia 133: 233-242.

Emerson AE. 1925. The termites of Kartabo, Bartica District, British Guiana. Zoologica (New York) 6: 291-459.

Fontes LR. 1998. Novos aditamentos ao "Catalogo dos Isoptera do Novo Mundo," e uma filogenia para os generos neotropicais de Nasutitermitinae, pp. 309-412 In Fontes LR, Filho EB [eds.], Cupins: o Desafio do Conhecimento. Fundacao de Estudos Agrarios Luiz de Queiroz, Sao Paulo, Brazil.

Fontes LR, Milano S. 2002. Termites as an urban problem in South America. Sociobiology 40: 103-151.

Light SF. 1933. Termites of western Mexico. University of California Publications in Entomology 6: 79-164.

Light SF. 1935. The Templeton Crocker Expedition of the California Academy of Sciences, 1932. No. 20. The termites. Proceedings of the California Academy of Sciences 21: 233-256.

Lomolino MV. 2004. Conservation biogeography, pp. 293-296 In Lomolino MV, Heaney LR [eds.], Frontiers of Biogeography: New Directions in the Geography of Nature. Sinauer Associates, Sunderland, Massachusetts, USA.

Nickle DA, Collins MS. 1990. The termite fauna (Isoptera) in the vicinity of Chamela, State of Jalisco, Mexico. Folia Entomologica Mexicana 77: 85-122.

Nickle DA, Collins MS. 1992. The termites of Panama (Isoptera), pp. 208-241 In Quintero D, Aiello A [eds.], Insects of Panama and Mesoamerica: Selected Studies. Oxford University Press, Oxford, United Kingdom.

Nutting WL. 1970. Composition and size of some termite colonies in Arizona and Mexico. Annals of the Entomological Society of America 63:1105-1110.

Roisin Y 2003. Cryptotermes chacoensis, a new species from native South American inland habitats (Isoptera: Kalotermitidae). Sociobiology 42: 319-327.

Scheffrahn RH. 1993. Cryptotermes chasei, a new drywood termite (Isoptera: Kalotermitidae) from the Dominican Republic. Florida Entomologist 76: 500-507.

Scheffrahn RH, Krecek J. 1999. Termites of the genus Cryptotermes Banks (Isoptera: Kalotermitidae) from the West Indies. Insecta Mundi 13: 111-171. High resolution images are available at: https://figshare.com/articles/West_lndies_Cryptotermes_lsoptera_Kalotermitidae_/6144137

Scheffrahn RH, Jones SC, Krecek J, Chase JA, Mangold JR, Su NY. 2003. Taxonomy, distribution, and notes on the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Puerto Rico and the US Virgin Islands. Annals of the Entomological Society of America 96: 181-201.

Scheffrahn RH, Krecek J, Chase JA, Su NY. 1998. Cryptotermes abruptus, a new drywood termite (Isoptera: Kalotermitidae) from southeastern Mexico. Florida Entomologist 81: 188-193.

Scheffrahn RH, Krecek J, Chase JA, Maharajh B, Mangold JR. 2006. Taxonomy, biogeography, and notes on termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of the Bahamas and Turks and Caicos Islands. Annals of the Entomological Society of America 99: 463-486.

Scheffrahn RH, Krecek J, Ripa R, Luppichini P. 2009. Endemic origin and vast anthropogenic dispersal of the West Indian drywood termite. Biological Invasions 11: 787-799.

Scheffrahn RH, Krecek J, Su NY, Roisin Y, Chase JA, Mangold JR. 1998. Extreme mandible alteration and cephalic phragmosis in a drywood termite soldier (Isoptera: Kalotermitidae: Cryptotermes) from Jamaica. Florida Entomologist 81: 238-240.

Snyder TE. 1934. Two new termites from Costa Rica. Proceedings of the Biological Society of Washington 47: 95-97.

Snyder TE. 1956. Termites of the West Indies, the Bahamas and Bermuda. Journal of Agriculture of the University of Puerto Rico 40: 189-202.

Rudolf H. Scheffrahn (1)

(1) University of Florida, Fort Lauderdale Research & Education Center, 3205 College Avenue, Davie, Florida 33314, USA; E-mail: rhsc@ufl.edu

Corresponding author; E-mail: rhsc@ufl.edu

Caption: Fig. 1. Dorsal (A) and lateral (B) views of the Cryptotermes garifunae female dealate (vertex of head capsule partially collapsed). Arrow of fore tarsus (C) points to arolium of dealate.

Caption: Fig. 2. Dorsal (A), anterodorsal (B), lateral (C), and ventral (D) views of Cryptotermes garifunae soldier head capsule. Arrows in lateral view of area near antennal fossa (E) point to frontal horn (FH) and genal horn (GH).

Caption: Fig. 3. Distribution of Cryptotermes species (Group 1) in Florida, the West Indies, and Caribbean Basin from the University of Florida termite collection and literature records. "UF collection ALL" represents other collection locations in the University of Florida termite collection.

Caption: Fig. 4. Distribution of Cryptotermes species (Group 2) in the West Indies, and Caribbean Basin from the University of Florida termite collection and literature records.

Caption: Fig. 5. Distribution of Cryptotermes species in South America from the University of Florida termite collection and literature records. Only endemic records of C. brevis are shown.

Please Note: Illustration(s) are not available due to copyright restrictions.
Table 1. Literature localities of New World Cryptotermes species not
encompassed in the University of Florida collection.

Species              Latitude           Longitude

C. cavifrons      32.3[degrees]N      64.76[degrees]W
C. chacoensis     20.695[degrees]S    61.929[degrees]W
C. chacoensis     25.046[degrees]S    58.059[degrees]W
C. colombianus    11.323[degrees]N    74.109[degrees]W
C. contognathus   21.3[degrees]S      40.96[degrees]W
C. cubicoceps      6.383[degrees]N    58.7[degrees]W
C. darwini         1.26[degrees]S     90.43[degrees]W
C. dudleyi         9.93[degrees]N     84.09[degrees]W
C. dudleyi         1.46[degrees]S     48.5[degrees]W
C. dudleyi         7.13[degrees]S     34.84[degrees]W
C. dudleyi        22.9[degrees]S      43.21[degrees]W
C. dudleyi        12.48[degrees]N     81.68[degrees]W
C. fatulus         2.025[degrees]S    80.735[degrees]W
C. fatulus        21.64[degrees]N    106.56[degrees]W
C. fatulus        19.21[degrees]N    104.68[degrees]W
C. fatulus         0.95[degrees]S     91.14[degrees]W
C. havilandi       1.46[degrees]S     81.68[degrees]W
C. havilandi       1.29[degrees]S     48.47[degrees]W
C. havilandi      23.96[degrees]S     46.33[degrees]W
C. havilandi      22.9[degrees]S      43.21[degrees]W
C. havilandi       3.72[degrees]S     38.54[degrees]W
C. longicollis    23.06[degrees]N    106.21[degrees]W
C. longicollis    19.527[degrees]N   105.075[degrees]W
C. longicollis     9.22[degrees]N     79.85[degrees]W
C. verruculosus    6.383[degrees]N    58.7[degrees]W

Species                    Location

C. cavifrons      Bermuda
C. chacoensis     Paraguay: Nueva Asuncion
C. chacoensis     Argentina: P. N. Rio Pilcomayo
C. colombianus    Colombia: P. N. Tayrona
C. contognathus   Brazil: Espfrito Santo, Praia das Neves
C. cubicoceps     Guyana: Kartabo
C. darwini        Galapagos Islands, Floriana Island
C. dudleyi        Costa Rica: San Jose
C. dudleyi        Brazil: Para, Belem
C. dudleyi        Brazil: Parafba, Joao Pessoa
C. dudleyi        Brazil: Rio de Janeiro
C. dudleyi        Colombia: San Andres Island
C. fatulus        Ecuador: Palmar
C. fatulus        Mexico: Maria Madre Island
C. fatulus        Mexico: Jalisco, Barra de Navidad
C. fatulus        Galapagos Islands, Isabela Island
C. havilandi      Brazil: Para, Belem
C. havilandi      Brazil: Para, Icoaraci
C. havilandi      Brazil: Sao Paulo, Santos
C. havilandi      Brazil: Rio de Janeiro
C. havilandi      Brazil: Ceara, Fortaleza
C. longicollis    Mexico: 30 km S. Matzatlan
C. longicollis    Mexico: Chamela
C. longicollis    Panama
C. verruculosus   Guyana: Kartabo

Species                   Reference

C. cavifrons      Snyder 1956
C. chacoensis     Roisin 2003
C. chacoensis     Roisin 2003
C. colombianus    Casalla et al. 2016
C. contognathus   Constantino 2000
C. cubicoceps     Emerson 1925
C. darwini        Light 1935
C. dudleyi        Snyder 1934
C. dudleyi        Constantino & Cancello 1992
C. dudleyi        Fontes & Milano 2002
C. dudleyi        Fontes & Milano 2002
C. dudleyi        Fontes & Milano 2002
C. fatulus        Bacchus 1987
C. fatulus        Light 1935
C. fatulus        Nutting 1970
C. fatulus        Light 1935
C. havilandi      Constantino & Cancello 1992
C. havilandi      Constantino & Cancello 1992
C. havilandi      Fontes 1998
C. havilandi      Fontes 1998
C. havilandi      Fontes 1998
C. longicollis    Light 1933
C. longicollis    Nickle & Collins 1990 (*)
C. longicollis    Nickle & Collins 1992
C. verruculosus   Emerson 1925

(*) Misidentified as C. fatulus

Table 2. Measurements (mm) of the Cryptotermes garifunae sp. nov. male
dealate (n = 1).

Measurement

Head length with labrum       1.05
Head length to postclypeus    0.96
Head width, maximum at eyes   0.84
Eye diameter, maximum         0.26
Eye to head base, minimum     0.12
Ocellus diameter, maximum     0.05
Pronotum, maximum length      0.61
Pronotum, maximum width       0.77
Total length without wings    5.63

Table 3. Measurements (mm) of the Cryptotermes garifunae sp. nov.
soldier (n = 3).

Measurement                                    maximum   minimum   mean

Head length to tip of mandibles                1.39      1.23      1.31
Head length to tip frontal horns               1.11      0.98      1.06
Frontal flange width                           1.02      1.00      1.01
Frontal horns, outside span                    0.74      0.74      0.74
Head width, maximum                            1.04      0.96      0.99
Head height, excluding postmentum              0.70      0.63      0.65
Pronotum, maximum length                       0.72      0.67      0.70
Pronotum, maximum width                        0.96      0.95      0.96
Left mandible length, tip to ventral condyle   0.44      0.39      0.41
Total length                                   5.13      3.81      4.25
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Title Annotation:Research
Author:Scheffrahn, Rudolf H.
Publication:Florida Entomologist
Date:Dec 1, 2018
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