Expanded New World distributions of genera in the termite family Kalotermitidae

The New World Kalotermitidae constitute about one-fourth of all termite species in this region. The geographic distributions of all fifteen kalotermitid genera are updated based on records in the University of Florida Termite Collection and in the literature.  Range-appropriate exotic records of four pest species are also given.  New distribution records are presented on maps, and representative soldier photographs are provided for each genus.

Expanded New World distributions of genera in the termite family Kalotermitidae

Introduction
The termite family Kalotermitidae Froggatt 1897 is composed of nearly 500 extant species worldwide (Casalla on water can vary drastically.Among New World Kalotermitidae, their home wood can be free of exposure to precipitation for most (Incisitermes, Marginitermes, Pterotermes, and Tauritermes) or all of the year (Cryptotermes brevis (Walker) in all cases and Incisitermes minor (Hagen) and Marginitermes hubbardi (Banks) infesting structural wood).Other species flourish in wood that is water-saturated constantly or for long periods (e.g., Comatermes perfectus (Hagen), and some Calcaritermes, Glyptotermes, Neotermes, and Rugitermes).Some New World kalotermitids, e.g., N. castaneus (Burmeister), inhabit galleries excavated within the xylem or heartwood of living trees.The soundness of host wood also varies remarkably from almost completely rotten (soft enough to pinch) to decay-free wood in structural or xeric habitats.The Kalotermitidae are generalist wood feeders and their range is dictated by habitat and climate, not by host wood preference (Scheffrahn pers. obs.).Few large-scale generic distribution maps have been published for termites.For the Kalotermitidae, Emerson (1969) produced world maps for Calcaritermes, Incisitermes, and Kalotermes.New World maps of Calcaritermes (Scheffrahn, 2011), Eucryptotermes Fort Lauderdale Research and Education Center, University of Florida, Institute of Food and Agricultural Sciences, Davie, Florida, U.S.A.
The purpose of this report is to update the distributional data for all genera of New World Kalotermitidae.Soldier photographs of a genus representative are also provided as an aid in identifying genera.

Materials and Methods
The distribution maps (Figs 1-8) are taken from 2,174 unique kalotermitid localities (Fig 1) obtained from expert sampling expeditions (Scheffrahn et al., 2018b) and acquisitions from other collectors housed with the author in the University of Florida Termite Collection (UFTC) located in Davie, Florida.Additional localities were taken from the literature (Table 1).
Generic identification of 15,911 kalotermitid samples was based mostly on Krishna's 1961revisionary key, Krishna et al.'s 2013updated key, and Constantino's 2002 key.Cryptotermes dudleyi Banks and Cr.havilandi Sjöstedt were identified using Scheffrahn and Křeček (1999).All UFTC localities were georeferenced either from hard copy maps before 1997, GPS receiver coordinates beginning in 1997, or by consulting Google Earth™.
with a Leica DFC 425 module run with Leica Application Suite version 3 software.Specimens were suspended and positioned in a transparent pool of Purell® hand sanitizer (70% EtOH) within a clear plastic Petri dish.

CALCARITERMES (Figs 2, 9E)
Calcaritermes ranges from Georgia to the vicinity of Rio de Janeiro.Soldiers are characterized by a large apical spur on the fore tibia.Ca. rioensis Krishna) and Bolivia (new species).The relict Ca. nearcticus (Snyder) forms an allopatric range in central Florida to the Georgia coastline (Scheffrahn et al., 2001) and does not occur in Neotropical (southeastern) Florida.With the exception of Ca. temnocephalus, pseudergates of this genus can be identified by their mesonotal rasp which is suggested to aid microbial cultivation in their foraging galleries (Scheffrahn, 2011).In addition to Ca. nearcticus, ten species are described from the Neotropics and single curious species from the Indomalayan Region (Maiti & Chakraborty, 1981).This genus contains no pest species.The UFTC contains three undescribed new species from Bolivia, Ecuador and Venezuela.1).

COMATERMES PERFECTUS (Figs 3, 10G)
Records from the UFTC (Fig 3) reveal that this monotypic genus occurs at elevations between 360-1370 m in mesic to wet forests from Honduras (new record) to Bolivia and the Lesser Antilles.Comatermes perfectus is found on all major islands of the Lesser Antilles except for the low-lying islands of Anguilla, Barbados, and Barbuda, and the volcanic peaks of Nevis and Antigua.Although the dark body coloration and wing venation of the imago is similar to Paraneotermes, Krishna (1961) concluded that the long wavy hairs on the head and pronotum justify generic status.Specimens in the UFTC collection show that Co. perfectus has a polymorphic soldier caste grading from very long headed individuals to much smaller, short-headed morphs.Constantino (2002) includes Co. perfectus as a pest of coffee plants.and literature (lit., Table 1).
CRYPTOTERMES (Figs 4,5,9F) The diversity of New World Cryptotermes, at 28 endemic species (30 total), exceeds that of all other kalotermitid genera of the region.The West Indies and Caribbean mainland have, at 21 species, the greatest diversity and abundance of this genus.Soldier head capsule phragmosity and rugosity vary greatly among New World Cryptotermes (Scheffrahn & Křeček, 1999).The UFTC contains three new undescribed species from Ecuador, Guadeloupe (Lesser Antilles), and Honduras.The most northern native distribution for the genus is for Cr.cavifrons on Jekyll Island, Georgia (31°N) and the southernmost is Valparaiso, Chile (33°S) for Cr.brevis (Scheffrahn et al., 2009).Three species have extended invasive New World distributions.Cryptotermes brevis, endemic to the Peruvian coastal desert (new inland record 13 km NE Huanuco, Peru) and Chilean Atacama desert, is the most destructive and widespread pest in the New World (Scheffrahn et al., 2009) especially from Florida to Brazil.I have excluded (Fig 4) isolated localities that either represent interceptions or static, non-invasive localities such as Ontario, Canada (Myles, 1995); Los Angeles, CA; Sussex DE; W. Simsbury, CT; Anchorage, AK; Long Island, NY; and Middletown, RI.  1).
Cryptotermes dudleyi, a pest endemic to the Indian subcontinent, has been introduced throughout the Caribbean Basin, especially Jamaica, and into South America (Fig 5).Cryptotermes havilandi, a pest species native to Africa, is common in the Lesser Antilles.Both of these species are listed as major Neotropical pests by Constantino (2002).The specimens in the UFTC were all collected from non-structural wood such as tree branches and fence posts.
of E. breviceps from French Guiana (J.Šobotník unpubl.)extends the range of this genus north of the equator.Constantino (2002) lists E. hagenii as a possible pest.I have never collected this remarkable genus.

GLYPTOTERMES (Figs 7, 10A)
Except for the phragmotic species, many Glyptotermes soldiers have rather unadorned head capsules and are difficult to identify.Glyptotermes soldiers tend to have cylindrical head capsules and steep frons.Many species are small, tend to nest in sound wood, and thus, are difficult to collect.The first records of this genus are provided from Bolivia, Peru, Paraguay, Ecuador, and most of the Lesser Antilles (Fig 7).Twenty-three species are recorded from the Neotropics and, surprisingly, six are described from Costa Rica.Poor illustrations of early descriptions lack resolution to aid in identification.This genus is in dire need of revision.1).

INCISITERMES (Figs 6, 10D)
With the exception of I. immigrans (Snyder) along the Pacific coast south to Peru, this genus is extremely rare in South America (Fig 6).A rather large portion of its range includes the southern Nearctic due, in large part, to the endemic range of I. minor in the West and I. snyderi (Light) in the East.Along with Cryptotermes and Neotermes, Incisitermes is a dominant genus in the West Indies and Caribbean coastline.The most diagnostic characters among Incisitermes soldiers are the sharply incised anterior margin of the pronota and the enlarged third antennal articles.Some species, such as I marginipennis (Latreille) and I. schwarzi (Banks) have large and small soldier morphs.Soldier size ranges widely, with I. marginipennis among the largest (head width > 2mm) and a new species, also from Mexico among the smallest (HW < 1mm).There are fifteen New World species of Incisitermes.Incisitermes tabogae (Snyder) is a synonym of I. schwarzi and I. nigritus (Snyder) is a synonym of I. platycephalus (Light).The UFTC contains seven undescribed species.This genus is in need of revision.Emerson (1969) includes records for I. galapagoensis (Banks), I. immigrans, and I. pacificus (Banks) from the Galápagos Islands and one or more undescribed species from Central America.He suggested that I. immigrans colonized Ecuador and Peru from Pacific island populations although the opposite seems more likely.and literature (lit., Table 1).
Incisitermes minor is endemic to the southwestern Nearctic where it is a major structural pest.The abundance of I. minor in Southern California, both in buildings and outdoor habitats such as woodpiles, fencing, and tree branches, is striking.This densely populated region is prone to earthquakes so many houses have wood-frame construction making I. minor the most destructive structural pest in southern California.As would be expected, I. minor has been transported by infested goods or by infested boats (Scheffrahn & Crowe, 2011) to other parts of the Nearctic, especially Florida (Scheffrahn, 2013).Although exotic structural records of I. minor in Florida are relatively common, it appears that I. minor cannot disperse from structure to structure or live in outdoor habitats.I have excluded isolated localities that either represent interceptions or static, non-invasive records such as Ontario, Canada (Grace et al., 1991); Tacoma, WA (Light, 1934a); Evanston, IL; Fergus Falls, MN; and Franklin, TN.

KALOTERMES (Figs 8, 10F)
Soldiers of Kalotermes are smaller and have a more quadrate pronotum than Neotermes.Only two species of Kalotermes are known in the New World, K. approximatus Snyder in the Southeastern U.S. and Bermuda and K. gracilignathus Emerson from Juan Fernández Island (Emerson, 1924, Fig 8).Kalotermes approximatus does not occur in Neotropical Florida.The genus lacks pest species.and literature (lit., Table 1).

MARGINITERMES (Figs 3, 9B)
The soldier of Marginitermes, with enormous third antennal articles, is the most distinctive of the New World Kalotermitidae.Two species occur in central and southwestern America and a third species occurs in Australia (Scheffrahn & Postle, 2013).Marginitermes hubbardi (Banks) is sympatric with I. minor and P. occidentis (Walker) in the Sonoran Desert of Arizona and Mexico, while M. cactiphagus Myles occupies the milder arid regions of Mexico and Central America (Fig 3).According to Light (1934b), M. hubbardi is the second most destructive kalotermitid of the region after I. minor.I have excluded one record of M. hubbardi from Odessa, Florida, as a static, non-invasive locality.

NEOTERMES (Figs 2, 10B)
Many Neotermes species are large or very large (e.g., N. mona (Banks); Křeček et al., 2000) and some species have dimorphic soldiers (e.g., N. jouteli (Banks); Scheffrahn et al., 2000).The distribution of Neotermes is the greatest and most continuous of any new world kalotermitid genus extending for 7,700 km from Bermuda [N.castaneus (Burmeister)] to coastal southern Chile [N.chilensis (Blanchard)] and elevations from sea level to 1831 m in Venezuela (Neotermes sp., Scheffrahn, 2015).Neotermes castaneus occurs in Bluefields, Nicaragua, while a new species of Neotermes inhabits the Peruvian coast in the proximity of Lima considered among the rainiest and driest habitats, respectively, in the New World.Interestingly, both of the latter species often reside beneath live cambium in standing trees.Some Neotermes species are minor pests of orchard trees (Constantino, 2002) including cacao and mango.
With 26 described species, Neotermes is the second most specious New World genus after Cryptotermes.Neotermes is in need of revision as synonyms and undescribed species abound for New World inhabitants.

PARANEOTERMES (Figs 8, 10E)
Paraneotermes simplicicornis (Banks) is unique among all kalotermitid species in that it has subterranean characteristics (Light, 1937).Paraneotermes, along with Comatermes and Pterotermes, is a monotypic genus that occurs in the Sonoran desert, but Light (1933) reports Pa.simplicicornis in the northern mesic Neotropics around San Blas, Mexico (26°N,Fig 8).Light (1937) reported that the imagos of Pa.simplicicornis readily bury themselves in the soil and avoid direct colonization of wood.Lack of literature since Light (1937), suggests this species is probably of little economic significance either as a structural wood or tree orchard pest.(Howell, 1984) as a static, non-invasive locality.

RUGITERMES (Figs 8, 10E)
Rugitermes soldiers are recognized by a subtle frontal ridge that extends medially from the dorsal antennal margin.Some images have a striking coloration contrast between the head and pronotum.Twelve Neotropical species are recorded, but soldiers are difficult to identify to species and genetic sequences suggest many more species remain to be described.The UFTC holdings now extend the northern limit of Rugitermes to Mexico and Belize (Fig 8).Honduras, Venezuela, Peru, Bolivia, Paraguay, Trinidad, Tobago are also added to the range of this genus.All species are Neotropical with the lone exception of Rugitermes athertoni (Light) from the Papuan Region (Light, 1932).Rugitermes laticollis Snyder holds the high elevation record for termites of 3600 m in Ecuador (Scheffrahn, 2015).

TAURITERMES (Figs 3, 9G)
Limited to the Neotropics, the frontal rugosity of the Tauritermes soldiers approaches that of some Cryptotermes, but Tauritermes has much longer mandibles.Three species of Tauritermes are described and herein is the first report from Bolivia and Paraguay (Fig 3).Tauritermes is recorded from rather more arid Neotropical habits with the exception of the surprising Manaus locality (ann.rainfall > 2m) of a new species (Dambros et al., 2013).Constantino (2002) has observed minor damage by Tauritermes in Brazil.

Fig 1 .
Fig 1. Map of New World Kalotermitidae localities from the University of Florida Termite Collection (UF) and literature (lit., Table1).
Fig 2 provides the first records of Calcaritermes from Peru (Ca.temnocephalus Silvestri and

Fig 2 .
Fig 2. Map of New World Calcaritermes and Neotermes localities from the University of Florida Termite Collection (UF) and literature (lit., Table1).

Fig 5 .
Fig 5. Map of New World exotic Cr. dudleyi and Cr.havilandi localities from the University of Florida Termite Collection (UF) and literature (lit., Table1).

Fig 6 .
Fig 6.Map of New World Eucryptotermes, Incisitermes, I. minor exotic, and I. minor endemic localities from the University of Florida Termite Collection (UF) and literature (lit., Table1).

Fig 7 .
Fig 7. Map of New World Glyptotermes and Proneotermes localities from the University of Florida Termite Collection (UF)and literature (lit., Table1).

Table 1 .
Collection localities from the literature used to supplement those of the University of Florida Termite Collection.

Table 1 .
Collection localities from the literature used to supplement those of the University of Florida Termite Collection.(Continuation)

Table 1 .
Collection localities from the literature used to supplement those of the University of Florida Termite Collection.(Continuation)

Table 1 .
Collection localities from the literature used to supplement those of the University of Florida Termite Collection.(Continuation)