Diorhabda sublineata

Diorhabda sublineata
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Coleoptera
Family:	Chrysomelidae
Genus:	Diorhabda
Species:	D. sublineata
Binomial name
	Diorhabda sublineata; (H. Lucas, 1849)
Synonyms
	Galeruca sublineata H. Lucas, 1849; Diorhabda elongata sublineata (H. Lucas, 1849);

Diorhabda sublineata is a leaf beetle known as the subtropical tamarisk beetle (STB). The species was first described by Hippolyte Lucas in 1849. It feeds on tamarisk trees from Portugal, Spain and France to Morocco, Senegal, Algeria, Tunisia, Egypt, Yemen, and Iraq.^[1] It is used in North America as a biological pest control agent against saltcedar or tamarisk (Tamarix spp.), an invasive species in arid and semi-arid ecosystems (where the STB and its closely related sibling species also may be less accurately referred to as the 'saltcedar beetle', 'saltcedar leaf beetle', 'salt cedar leaf beetle', or 'tamarisk leaf beetle') (Tracy and Robbins 2009).

Taxonomy

The STB was first described from Annaba, Algeria as Galeruca sublineata H. Lucas (1849). Reiche and Saulcy (1858) erroneously placed G. sublineata as a junior synonym to the sibling species G. elongata Brullé (the Mediterranean tamarisk beetle, Diorhabda elongata). Weise (1893) created the genus Diorhabda and proposed the variety Diorhaba elongata var. sublineata (H. Lucas). Gressitt and Kimoto (1963) proposed the subspecies D. e. sublineata. Tracy and Robbins (2009) restored D. sublineata (H. Lucas) as a valid species based on comparisons of the male and female genitalia, and provided illustrated taxonomic keys separating the STB from the four other sibling species of the D. elongata (Brullé) species group: Diorhabda elongata, Diorhabda carinata (Faldermann), Diorhabda carinulata (Desbrochers), and Diorhabda meridionalis Berti and Rapilly. In literature prior to 2009, D. sublineata was usually also referred to as D. elongata, or as a subspecies or color variant of D. elongata.

Host plants

Field collections in Spain and north Africa reveal that the STB feeds on at least five species of tamarisks, including Tamarix gallica which hybridizes with the widely invasive T. ramosisima in western North America. The STB can become numerous in Egypt and Senegal, but reports on defoliation of tamarisk are lacking (Tracy and Robbins 2009). Extensive laboratory host range studies verified that STB is a specialist feeder on tamarisks, feeding only on plants of the tamarisk family, Tamaricaceae. In laboratory and field cage studies, the STB will also feed and complete development on Frankenia shrubs, distant relatives of tamarisks in the same plant order Caryophyllales, but STB greatly prefer to lay eggs upon tamarisk (Milbrath and DeLoach 2006).

Life cycle

The STB overwinters as adults on the ground. Adults become active and begin feeding and mating in the early spring when tamarisk leaves are budding. Eggs are laid on tamarisk leaves and bark and hatch in about a week in warm weather. Three larval stages feed on tamarisk leaves for about two and a half weeks when they crawl to the ground and spend about 5 days as a C-shaped inactive prepupa before pupating about one week. Adults emerge from pupae to complete the life cycle in about 4–5 weeks in the summer. (For images of various life stages of a related species, see Diorhabda carinulata at Commons.) Five generations of STB occur through spring and fall in central Texas (Milbrath et al. 2007). Similar to the northern tamarisk beetle, adults begin to enter diapause in the late summer and early fall, ceasing reproduction and feeding to build fat bodies before seeking a protected place to overwinter (Lewis et al. 2003). Larvae and adults are sensitive to shorter daylengths as the summer progresses that signal the coming of winter and induce diapause (Bean et al. in prep.). Robert Bartelt and Allard Cossé (USDA-ARS, Peoria, Illinois) found that male STB emit a putative aggregation pheromone, similar to that found in Diorhabda carinulata (Cossé et al. 2005), that could serve to attract both males and females to certain tamarisk trees.

Biological control agent

Establishment of the STB as a biological control agent for tamarisk along the Rio Grande in West Texas was confirmed in 2010 (Knutson 2010). Populations of STB from around 35°N latitude near Sfax, Tunisia were initially released by the USDA Agricultural Research Service in south Texas in 2005, but failed to establish. During the spring and summer of 2009, populations from around 34° latitude near Marith, Tunisia were released in south and west Texas (Tracy and Robbins 2009). These populations were showing promising signs of establishment on the Rio Grande, such as near Presidio, Texas at Alamito Marsh, by the fall of 2009 (MacCormack 2009). By August 2010, the STB had defoliated about 23 miles of tamarisk along the Rio Grande near Presidio, but it was causing concern by also defoliating the related but non-target athel tamarisk (Tamarix aphylla) trees, a taller species of tamarisk used around Presidio and neighboring Mexican communities for shade (Haines 2010). The STB may be better adapted to subtropical interior desert habitats and subtropical Mediterranean habitats than other Old World tamarisk beetles that are being introduced, such as the Mediterranean tamarisk beetle, Diorhabda elongata. The northern tamarisk beetle, Diorhabda carinulata, is probably better adapted to northern cold deserts in North America where it is widely established, and the larger tamarisk beetle, Diorhabda carinata, is probably better adapted to warm temperate grasslands and deserts (Tracy and Robbins 2009).

Tamarisk does not usually die from a single defoliation from tamarisk beetles, and it can resprout within several weeks of defoliation. Repeated defoliation of individual tamarisk trees can lead to severe dieback the next season and death of the tree within several years (DeLoach and Carruthers 2004). Tamarisk beetle defoliation over the course of at least one to several years can severely reduce the nonstructural carbohydrate reserves in the root crowns of tamarisk (Hudgeons et al. 2007). Biological control of tamarisk by the STB will not eradicate tamarisk but it has the potential to suppress tamarisk populations by 75–85%, after which both STB and tamarisk populations should reach equilibrium at lower levels (DeLoach and Carruthers 2004, Tracy and DeLoach 1999).

A primary objective of tamarisk biological control with the STB is to reduce competition by exotic tamarisk with a variety of native riparian flora, including trees (willows, cottonwoods, and honey mesquite), shrubs (wolfberry, saltbush, and baccharis), and grasses (alkali sacaton, saltgrass, and vinemesquite). Unlike expensive chemical and mechanical controls of tamarisk that often must be repeated, tamarisk biological control does not harm native flora and is self-sustaining in the environment. Recovery of native riparian grasses can be quite rapid under the once closed canopy of repeatedly defoliated tamarisk. However, tamarisk beetle defoliation can locally reduce nesting habitat for riparian woodland birds until native woodland flora are able to return. In some areas, tamarisk may be replaced by grasslands or shrublands, resulting in losses of riparian forest habitats for birds (Tracy and DeLoach 1999). Releases of tamarisk beetles in southern California, Arizona, and along the Rio Grande in western New Mexico, are currently delayed until concerns can be resolved regarding safety of tamarisk biological control to nesting habitats of the federally endangered southwestern willow flycatcher, Empidonax traillii Audubon subspecies extimus Phillips, which will nest in tamarisk (see DeLoach et al. 2000, Dudley and DeLoach 2004).

References

Bean, D.W.; Keller, J.C. in prep.: Characteristics of diapause induction in populations of Diorhabda elongata collected from sites in Europe, Africa and Asia: Implications for tamarisk (Tamarix spp) biocontrol in North America. For publication in Biological Control.
Cossé, A.A.; Bartelt, R.J.; Zilkowski, B.W.; Bean, D.W.; Petroski, R.J. 2005: The aggregation pheromone of Diorhabda elongata, a biological control agent of saltcedar (Tamarix sp.): Identification of two behaviorally active components. Journal of Chemical Ecology, 31(3): 657–670. PDF
DeLoach, C.J.; Carruthers, R. 2004: Biological control programs for integrated invasive plant management. In: Proceedings of Weed Science Society of America Meeting, Kansas City, MO. Weed Science Society of America (CD-ROM). 17 pp. PDF
DeLoach, C.J.; Carruthers, R.I.; Lovich, J.E.; Dudley, T.L.; Smith, S.D. 2000: Ecological interactions in the biological control of saltcedar (Tamarix spp.) in the United States: toward a new understanding. In N. R. Spencer (ed.), Proceedings of the X International Symposium on Biological Control of Weeds, 4–14 July 1999, Montana State University. Bozeman, Montana, pp. 819–873. PDF
Knutson, A. 2010: Tunisian beetle establishes on the Rio Grande river. Beetle-Mania; Biological Control of Saltcedar, Spring, 2010: 2(1): 2. PDF
Dudley, T.L. DeLoach, C.J. 2004: Saltcedar (Tamarix spp.), endangered species, and biological weed control-can they mix? Weed Technology, 18(5): 1542–1551. PDF^{[permanent dead link‍]}
Gressitt, J.L.; Kimoto, S. 1963: The Chrysomelidae (Coleopt.) of China and Korea, Part 2. Pacific Insects Monograph, 1B: 301–1026.* Lucas, P.H. 1849: Exploration scientifique de l'Algeria. Zoologie (Paris), 2: 542–546. (In French)
Haines, T. 2010: Attack of the hungry salt cedar beetles. The International (Presidio, Presidio County, Texas), 26 August 2010: 24(3): 1,3. PDF
Hudgeons, J.L.; Knutson, A.E.; Heinz, K.M.; DeLoach, C.J.; Dudley, T.L.; Pattison, R.R.; Kiniry, J.R. 2007: Defoliation by introduced Diorhabda elongata leaf beetles (Coleoptera: Chrysomelidae) reduces carbohydrate reserves and regrowth of Tamarix (Tamaricaceae). Biological Control, 43: 213–221. PDF
Lewis, P.A.; DeLoach, C.J.; Knutson, A.E.; Tracy, J.L.; Robbins, T.O. 2003: Biology of Diorhabda elongata deserticola (Coleoptera: Chrysomelidae), an Asian leaf beetle for biological control of saltcedars (Tamarix spp.) in the United States. Biological Control, 27: 101–116. PDF
MacCormack, J. 2009: Tiny bugs sent to devour trees that are big pests. San Antonio Express-News, 3 November 2009: 1,6. [1]^{[permanent dead link‍]} PDF
Milbrath, L.; DeLoach, C.J. 2006: Host specificity of different populations of the leaf beetle Diorhabda elongata (Coleoptera: Chrysomelidae), a biological control agent of saltcedar (Tamarix spp.). Biological Control, 36: 32–48. PDF
Milbrath, L.; DeLoach, C.J.; Tracy, J.L; 2007: Overwintering survival, phenology, voltinism, and reproduction among different populations of the leaf beetle Diorhabda elongata (Coleoptera: Chrysomelidae). Environmental Entomology, 36(6): 1356–1364. PDF
Reiche, L.; Saulcy, F. 1858: Espèces nouvelles ou peu connues de Colèoptères, recueillies par M.F. de Saulcy, member de l’Institute, dans son voyage en Orient, et dècrites par M.M. L. Reiche et Fèlicien de Saulcy. Annales de la Société Entomologique de France, Sér., 3, 6: 5–60. (In French)
Tracy, J.L.; DeLoach, C.J. 1999: Biological control of saltcedar in the United States: Progress and projected ecological effects. In: Bell, C.E. (Ed.), Arundo and Saltcedar: The Deadly Duo, Proceedings of the Arundo and Saltcedar Workshop, 17 June 1998. Ontario, California, 111–154. PDF
Tracy, J.L.; Robbins, T.O. 2009: Taxonomic revision and biogeography of the Tamarix-feeding Diorhabda elongata (Brullé, 1832) species group (Coleoptera: Chrysomelidae: Galerucinae: Galerucini) and analysis of their potential in biological control of Tamarisk. Zootaxa, 2101: 1–152. PDF
Weise, J. 1893: Chrysomelidae. In: Erichson, W. (ED.), Naturgeschichte der Insecten Deutschlands, 61 (73): 961–1161. (in German)

Notes

^ Tracy and Robbins (2009) provide a detailed review of the distribution, biogeography, biology, and taxonomy of D. sublineata that is a general source for most of this article.

External links

Data related to Diorhabda sublineata at Wikispecies
Media related to Diorhabda sublineata at Wikimedia Commons
Texas Agri-Life Extension Leaflet; Biological Control of Saltcedar (Tunisia source population, not mentioned, is D. sublineata). PDF
Texas Agri-Life Extension Newsletter; Beetle-Mania; Biological Control of Saltcedar in Texas, Volume 1, No. 2, Summer 2009 PDF
Texas Agri-Life Research and Extension Program; Biological Control of Saltcedar: Using Natural Enemies to Combat an Invasive Weed Competing with Texas' Water Resources (Tunisia source population, not mentioned, is D. sublineata). PDF
USDA Agricultural Research Service and Texas Agri-Life Research and Extension Service Report of Information to the Public; Progress on Biological Control of Saltcedar in the Western U.S.: Emphasis -- Texas 2004-2009. PDF

[Tracy-1] Tracy and Robbins (2009) provide a detailed review of the distribution, biogeography, biology, and taxonomy of D. sublineata that is a general source for most of this article.

[1]