Family Cixiidae

Subfamily Cixiinae

Tribe Pentastirini Emeljanov, 1971

Genus Melanoliarus Fennah, 1945.

Note: Melanoliarus was originally described as a subgenus of Oliarus.  Oliarus as historically defined was of world distribution, but Emeljanov (2001) and later Holzinger et al. (2003) asserted that the New World Oliarus did not belong to the genus in the strict sense, and removed all New World species into Melanoliarus (new status, genus), except for a few species explicitly placed in Reptalus or Pentastiridius. The features of Melanoliarus (especially as compared with Oliarus and Reptalus) remain incompletely elaborated.  Melanoliarus is unlikely to be monophyletic as currently comprised, and additional changes in the composition of the genus are highly likely. 

Type species: Oliarus (Melanoliarus) maidis Fennah, 1945.

Synonyms:

= Oliarus Stål, 1862a in part (auctorum) (Type species Cixius walkeri Stål, 1859); Emeljanov 2001 (all Nearctic and Neotropical species of Oliarus auctt. moved to Melanoliarus by implication Emeljanov 2001: 122).

Distribution: Nearctic and Neotropical.

Recognized species

There are 48 species reported north of Mexico and 30 additional species from the Neotropics as follows:

1 Melanoliarus acicus (Caldwell, 1947a) - USA: AZ, CA, TX; Mexico (Baja California, Sinaloa,  Sonora)
    = Oliarus acicus Caldwell, 1947a: 76.
    = Melanoliarus acicus (Caldwell, 1947a); comb. by implication Emeljanov 2001: 122.
2 Melanoliarus altanatus (Caldwell, 1951) - USA: TX; Mexico (Chiapas, Jalapa, Federal District, Hidalgo, Morelos, Veracruz)
     = Oliarus altanatus Caldwell, 1951: 35.
     = Melanoliarus altanatus (Caldwell, 1951); comb. by implication Emeljanov 2001: 122.
3 Melanoliarus altanus (Ball, 1934b) - USA: AZ
    = Oliarus altanus Ball, 1934b: 276.
    = Melanoliarus altanus (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
4 Melanoliarus apache (Ball, 1934b) - USA: AZ, CA; Mexico (Sonora)
     = Oliarus apache Ball, 1934b: 275.
     = Melanoliarus apache (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
5 Melanoliarus aridus (Ball, 1902a) - USA: AL, AR, AZ, CA, CO, GA, ID, IL, IN, KS, IL, LA, MO, MS, MT, NC, ND, NE, NM, OH, OK, OR, SC, SD, TN, TX, UT, VA, WY; CAN: ON; Mexico (Durango)
     = Oliarus aridus Ball, 1902a: 151.
    = Melanoliarus aridus (Ball, 1902a); comb. by implication Emeljanov 2001: 122.
6 Melanoliarus arizonensis (Mead & Kramer, 1982) - USA: AZ
    = Oliarus arizonensis Mead & Kramer, 1982: 428-432.
    = Melanoliarus arizonensis (Mead & Kramer, 1982); comb. by implication Emeljanov 2001: 122.
7 Melanoliarus bispinus (Caldwell, 1947b) - USA: OK, TX
    = Oliarus bispinus Caldwell, 1947b: 150-151.
    = Melanoliarus bispinus (Caldwell, 1947b); comb. by implication Emeljanov 2001: 122.
8 Melanoliarus caldwelli (Mead & Kramer, 1982) - USA: NM, TX
    = Oliarus caldwelli Mead & Kramer, 1982: 523-527.
    = Melanoliarus caldwelli (Mead & Kramer, 1982); comb. by implication Emeljanov 2001: 122.
9 Melanoliarus californicus (Van Duzee, 1914) - USA: CA
    = Oliarus californicus Van Duzee, 1914: 36.
    = Melanoliarus californicus (Van Duzee, 1914); comb. by implication Emeljanov 2001: 122.
10 Melanoliarus canyonensis (Mead & Kramer, 1982) - USA: CA, NM
    = Oliarus canyonensis Mead & Kramer, 1982: 535-537.
    = Melanoliarus canyonensis (Mead & Kramer, 1982); comb. by implication Emeljanov 2001: 122.
11 Melanoliarus catus (Caldwell, 1947b) - USA: AZ
    = Oliarus catus Caldwell, 1947b: 147.
    = Melanoliarus catus (Caldwell, 1947b); comb. by implication Emeljanov 2001: 122.
12 Melanoliarus chuliotus (Ball, 1934b) - USA: AR, DC, DE, FL, GA, IL, KS, MS, NC, OH, SC, TN
    = Oliarus chuliotus Ball, 1934b: 271.
    = Melanoliarus chuliotus (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
    = Oliarus gladensis Caldwell, 1951: 35; syn. by Mead & Kramer 1982: 478-480.
13 Melanoliarus coconinus (Ball, 1934b) - USA: AZ, CO, NV, TX, UT; CAN: BC
    = Oliarus coconinus Ball, 1934b: 274.
    = Melanoliarus coconinus (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
14 Melanoliarus concinnulus (Fowler, 1904) - Mexico (Guerrero, Veracruz) (reported in error: USA: TX)
    = Oliarus concinnulus Fowler, 1904: 92.
    = Melanoliarus concinnulus (Fowler, 1904); comb. by implication Emeljanov 2001: 122.
    = Oliarus texanus Metcalf, 1923: 181; syn. by Ball 1934: 275 (error, see Mead & Kramer 1982: 389, 436).

15 Melanoliarus complectus (Ball, 1902a) - USA: FL; Cuba, Dominican Republic, Haiti, Jamaica, Puerto Rico (Inc. Caja de Muertos, Mona and Vieques Islands), Virgin Islands (Jost van Dyke, St. Croix, St. Thomas, St. Vincent) (reported in error: USA: AZ, Mexico)
    = Oliarus complectus Ball, 1902a: 152.
    = Melanoliarus complectus (Ball, 1902a); comb. by implication Emeljanov 2001: 122.
    = Oliarus (Melanoliarus) campestris Fennah, 1945b: 141; syn. by Caldwell 1951 (in Caldwell & Martorell 1951): 138-141.
    = Oliarus franciscanus (Stål, 1859: 273); senior syn. by Van Duzee 1917a: 309 (error).
16 Melanoliarus corvinus (Ball, 1934b) - USA: AZ, CA, TX
    = Oliarus corvinus Ball, 1934b: 273.
    = Melanoliarus corvinus (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
17 Melanoliarus difficilis (Van Duzee, 1912) - USA: FL, TX
    = Oliarus difficilis Van Duzee, 1912a: 494.
    = Melanoliarus difficilis (Van Duzee, 1912a); comb. by implication Emeljanov 2001: 122.
    = Oliarus vittatus Metcalf, 1923: 181; syn. by Mead & Kramer 1982: 408, 411.
18 Melanoliarus dondonius (Ball, 1934b) - USA: AZ, CA, CO, NM, NV, OR, TX, UT; CAN: AB, BC, SK; Mexico (Baja California, Mulge, San Francisco, San Lorenzo, and Santa Inez Islands, Gulf of California)
    = Oliarus dondonius Ball, 1934b: 276.
    = Melanoliarus dondonius (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
19 Melanoliarus ecologus (Caldwell, 1947a) - USA: AL, AR, CT, DC, DE, GA, IL, KS, KY, MD, MS, NC, NJ, OH, OK, PA, SC, TN, VA (reported in error: CAN: NS, ON)
    = Oliarus ecologus Caldwell, 1947a: 76.
    = Melanoliarus ecologus Caldwell, 1947a; comb. by implication Emeljanov 2001: 122.
20 Melanoliarus eximus (Caldwell, 1947b) - USA: CA, NM
    = Oliarus eximus Caldwell, 1947b: 145.
    = Melanoliarus eximus (Caldwell, 1947b); comb. by implication Emeljanov 2001: 122.
21 Melanoliarus forcipatus (Caldwell, 1947b) - USA: AZ, TX
    = Oliarus forcipatus Caldwell, 1947b: 146.
    = Melanoliarus forcipatus (Caldwell, 1947b); comb. by implication Emeljanov 2001: 122.
22 Melanoliarus franciscanus (Stål, 1859) - Puerto Rico, Haiti, Jamaica, Isle of Pines, Cuba, Mexico (Baja California)
(reported in error: USA: AZ, CA, CO, CT, DC, FL, IA, MA, MD, KS, NC, NJ, NY, OH, PA, UT; CAN: AB, BC)
    = Cixius franciscanus Stål, 1859: 273.
    = Oliarus franciscanus (Stål, 1859); comb. by Stål 1862a: 306.
    = Melanoliarus franciscanus (Stål, 1859); comb. by implication Emeljanov 2001: 122.
    = Oliarus fransiscanus (Stål, 1859); missp. by Strickland 1940: 87.

23 Melanoliarus hesperius (Van Duzee, 1917a) - USA: CA, NV, OR
    = Oliarus hesperius Van Duzee, 1917a: 307.
    = Melanoliarus hesperius (Van Duzee, 1917a); comb. by implication Emeljanov 2001: 122.
24 Melanoliarus humilis (Say, 1830) - USA: AZ, CT, DC, DE, IA, IL, IN, KS, MA, MD, MI, MN, MS, MT, NC, NH, NJ, NM, NV, NY, OH, OR, PA, UT, WI; CAN: AB, BC, ON, QC, SK
    = Flata humilis Say, 1830: 241.
    = Oliarus humilis (Say, 1830); comb. by Van Duzee 1890: 389.
    = Melanoliarus humilis (Say, 1830); comb. by implication Emeljanov 2001: 122.
25 Melanoliarus kieferi (Mead & Kramer, 1982) - USA: AZ, UT
    = Oliarus kieferi Mead & Kramer, 1982: 559-561.
    = Melanoliarus kieferi (Mead & Kramer, 1982); comb. by implication Emeljanov 2001: 122.
26 Melanoliarus knullorum (Mead & Kramer, 1982) - USA: AZ, TX
    = Oliarus knullorum Mead & Kramer, 1982: 529-531.
    = Melanoliarus knullorum (Mead & Kramer, 1982); comb. by implication Emeljanov 2001: 122.
27 Melanoliarus littoralis (Ball, 1934b) - USA: FL; Dry Tortugas (Garden Key)
    = Oliarus littoralis Ball, 1934b: 274.
    = Melanoliarus littoralis (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
28 Melanoliarus lobatus (Caldwell, 1938b) - USA: NM, TX
    = Oliarus lobatus Caldwell, 1938b: 305.
    = Melanoliarus lobatus (Caldwell, 1938b); comb. by implication Emeljanov 2001: 122.
29 Melanoliarus montanus (Metcalf, 1923) - USA: DE, GA, IL, KS, MD, MO, MS, NC, NJ, NY, OH, PA, TN, VA
    = Oliarus montanus Metcalf, 1923: 179.
    = Melanoliarus montanus (Metcalf, 1923); comb. by implication Emeljanov 2001: 122.
30 Melanoliarus papagonus (Ball, 1934b) - USA: AZ
    = Oliarus papagonus Ball, 1934b: 272.
    = Melanoliarus papagonus (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
31 Melanoliarus pima (Kirkaldy, 1907b) - USA: AZ, CA, NM, UT
    = Oliarus pima Kirkaldy, 1907b: 62.
    = Oliarus nogalanus Ball, 1934b: 272 (syn. Mead & Kramer 1982: 390, 550-551, 554; see also Ball 1937: 180).
    = Melanoliarus pima (Kirkaldy, 1907b); comb. by implication Emeljanov 2001: 122.
32 Melanoliarus placitus (Van Duzee, 1912a) - USA: AL, AR, DC, DE, FL, GA, IL, KS, MD, MO, MS, NC, NJ, NY, OH, OK, PA, SC, TN, VA; CAN: ON
    = Oliarus placitus Van Duzee, 1912a: 493.
    = Melanoliarus placitus (Van Duzee, 1912a); comb. by implication Emeljanov 2001: 122.
33 Melanoliarus pygmaeus (Ball, 1937) - USA: AZ
    = Oliarus pygmaeus Ball, 1937: 180.
    = Melanoliarus pygmaeus (Ball, 1937); comb. by implication Emeljanov 2001: 122.
34 Melanoliarus quinquelineatus (Say, 1830) - USA: AL, AR, CT, CO, DE, FL, GA, IA, IL, IN, LA, MA, MD, ME, MN, MS, NC, NJ, NY, OH, OK, PA, RI, SC, TN, VA, WI; CAN: NS, ON, QC
    = Flata quinquelineata Say, 1830: 241.
    = Oliarus quinquelineatus (Say, 1830); comb. by Provancher 1889: 223.
    = Oliarus quinquelineata (Say, 1830);unjustified emendation by Smith 1900: 89.
    = Melanoliarus quinquelineatus (Say, 1830); comb. by implication Emeljanov 2001: 122.
35 Melanoliarus retentus (Caldwell, 1947b) - USA: AZ, CO, UT
    = Oliarus retentus Caldwell, 1947b: 148.
    = Melanoliarus retentus (Caldwell, 1947b); comb. by implication Emeljanov 2001: 122.
36 Melanoliarus sablensis (Caldwell, 1951) - USA: CT, DC, DE, FL, GA, IA, IL, IN, KY, MA, MD, MN, MO, NC, NH, NJ, NY, OH, PA, SC, TN, WV; CAN: NB, NS, ON
    = Oliarus sablensis Caldwell, 1951: 34.
    = Melanoliarus sablensis (Caldwell, 1951); comb. by implication Emeljanov 2001: 122.
37 Melanoliarus sementinus (Ball, 1902a) - USA: AZ, CA, CO, NM, UT
    = Oliarus sementinus Ball, 1902a: 152.
    = Melanoliarus sementinus (Ball, 1902a); comb. by implication Emeljanov 2001: 122.
38 Melanoliarus slossonae (Van Duzee, 1912a) - USA: FL; Cuba; Puerto Rico; Virgin Islands (Great Camanoe, Guana, Jost Van Dyke, Little Thatch, St. John, St. Thomas, Tortola) (reported in error: USA: MS, TX)
    = Oliarus hyalinus Van Duzee, 1908: 487 (nec. Fieber) (preoccupied).
    = Oliarus slossoni Van Duzee, 1912a: 494; replacement name for preoccupied O. hyalinus Van Duzee, 1908.
    = Oliarus borinquensis Caldwell, 1951: 137 (in Caldwell & Martorell 1951); syn. by Mead & Kramer 1982: 424-426.
    = Oliarus slossonae Van Duzee, 1912a: 494; emendation by Mead & Kramer 1982: 424.
    = Melanoliarus slossonae (Van Duzee, 1912a); comb. by implication Emeljanov 2001: 122.
39 Melanoliarus sonoitus (Ball, 1937) - USA: AZ, CA, NM, TX; Mexico (Chihuahua, Tamaulipas)
    = Oliarus sonoitus Ball, 1937: 179.
    = Oliarus nigravittus Caldwell, 1938b: 304; syn. by Mead & Kramer 1982: 554-556, 558.
    = Melanoliarus sonoitus (Ball, 1937); comb. by implication Emeljanov 2001: 122.
40 Melanoliarus sylvaticus (Caldwell, 1947b) - USA: CA
    = Oliarus sylvaticus Caldwell, 1947b: 151.
    = Melanoliarus sylvaticus (Caldwell, 1947b); comb. by implication Emeljanov 2001: 122.
41 Melanoliarus texanus (Metcalf, 1923) - USA: OK, TX; Mexico (Chiapas, Michoacán, Veracruz)
    = Oliarus texanus Metcalf, 1923: 181.
    = Melanoliarus texanus (Metcalf, 1923); comb. by implication Emeljanov 2001: 122.
42 Melanoliarus teximus (Caldwell, 1947b) - USA: TX
    = Oliarus eximus teximus Caldwell, 1947b: 146.
    = Oliarus teximus Caldwell, 1947b; status by Mead & Kramer 1982: 517-518, 521.
    = Melanoliarus teximus (Caldwell, 1947b); comb. by implication Emeljanov 2001: 122.
43 Melanoliarus truncatus (Van Duzee, 1929b) - USA: CA; Mexico (Baja California)
    = Oliarus truncatus Van Duzee, 1929b: 72.
    = Melanoliarus truncatus (Van Duzee, 1929b); comb. by implication Emeljanov 2001: 122.
44 Melanoliarus uncatus (Caldwell, 1947b) - USA: AZ
    = Oliarus uncatus Caldwell, 1947b: 151.
    = Melanoliarus uncatus (Caldwell, 1947b); comb. by implication Emeljanov 2001: 122.
45 Melanoliarus vicarius (Walker, 1851) - USA: FL, GA, IL, MS, NC, SC (Reported in error: USA: CO, MA, MD, NJ, TX)
    = Cixius vicarius Walker, 1851: 343.
    = Oliarus vicarius (Walker, 1851); comb. by Distant 1907a: 280.
    = Oliarus vitreus Metcalf, 1923: 180; syn. by Mead & Kramer 1982: 506, 509.
    = Melanoliarus vicarius (Walker, 1851); comb. by implication Emeljanov 2001: 122.
46 Melanoliarus viequensis (Caldwell, 1951) (in Caldwell & Martorell 1951) - USA: FL; Belize, Haiti, Puerto Rico (Vieques Island); Virgin Islands (Anegada, Antigua, Guana, St. John)
    = Oliarus viequensis Caldwell, 1951: 141.
    = Melanoliarus viequensis (Caldwell, 1951); comb. by implication Emeljanov 2001: 122.
47 Melanoliarus yavapanus (Ball, 1934b) - USA: AZ, CA, TX, UT
    = Oliarus yavapanus Ball, 1934b: 274.
    = Melanoliarus yavapanus (Ball, 1934b); comb. by implication Emeljanov 2001: 122.
48 Melanoliarus zyxus (Caldwell, 1947a) - USA: AZ, CA, CO, ID, NM, NV, OR, TX, UT; CAN: BC; Mexico (Coahuila, Nayarit, Nuevo León)
    = Oliarus zyxus Caldwell, 1947a: 76-77.
    = Melanoliarus zyxus (Caldwell, 1947a); comb. by implication Emeljanov 2001: 122.

Neotropical species (combinations by implication Emeljanov 2001: 122). (based on unpublished list by Lois O'Brien, distributions may be incoomplete)

1 Melanoliarus abacus (Caldwell 1947) - Mexico
   Melanoliarus aridus (Ball 1902) - Mexico, US
2 Melanoliarus aztecus (Caldwell 1947) - Mexico
3 Melanoliarus atkinsae (Myers 1928) - Cuba
4 Melanoliarus beebei (Metcalf 1945) - British Guiana
    O. borinquensis see Melanoliarus slossonae
5 Melanoliarus chiriquiensis (Fowler 1904) - Panama
6 Melanoliarus cinereus (Wolcott 1921) - Puerto Rico
7 Melanoliarus complectus (Ball 1902) - Puerto Rico, Cuba, US
     = O. campestris Fennah 1945 Antigua
8 Melanoliarus concinnulus ( Fowler 1904) - Mexico
9 Melanoliarus dimidiatus (Berg 1879) - Argentina
10 Melanoliarus excelsus (Fowler 1904) - Mexico (Baja California)
    Melanoliarus franciscanus (Stål 1859) - South America
11 Melanoliarus fulvus (Walker 1858) - Brazil
13 Melanoliarus humeralis (Fowler 1904) - Guatemala
14 Melanoliarus kindli (Bourgoin & Wilson 1998) - French Guiana
15 Melanoliarus insignior (Fowler 1904) - Panama, Guatemala
16 Melanoliarus isolatus (Caldwell 1947) - Mexico
17 Melanoliarus lacteipennis (Fowler 1904) - Panama
18 Melanoliarus latifrons (Walker 1851) - location unknown
19 Melanoliarus lunata (Fabricius 1803) - West Indies, Isle of Pines, South America
20 Melanoliarus maidis (Fennah 1945) - Trinidad
21 Melanoliarus nanus (Caldwell 1947) - Mexico
22 Melanoliarus nigro-alutaceus (Fowler 1904) - Mexico

    Oliarus omani Metcalf 1938 - Cyclopoliarus
23 Melanoliarus opalinus (Fennah 1945) - Venezuela
24 Melanoliarus orizicola (Bourgoin & Wilson 1998) - Peru
25 Melanoliarus pinicolus (Osborn 1926) -  Cuba
26 Melanoliarus propior (Fowler 1904) - Mexico
27 Melanoliarus quadratus (Fennah 1945) - Venezuela
28 Melanoliarus rarus (Caldwell 1947) - Mexico (Baja California)
     Melanoliarus slossonae (Van Duzee 1912) - Puerto Rico, Virgin Islands, US
       = O. borinquensis Caldwell 1952; see Mead & Kramer 1982
29 Melanoliarus transitorius (Berg 1883) - Uruguay
     Melanoliarus viequensis (Caldwell 1951) - Puerto Rico, USA: FL
30 Melanoliarus villosa (Fabricius 1775) - South America

Economic Importance:

Apparently limited - not reported as pests.

Known host plants: 

Nymphs of cixiids are subterranean, feeding on roots and possibly fungi.  The significance of adult host records is unclear. Many cixiids are presumed to be polyphagous (as adults), most often on woody plants.

[Still to come]

Hosts from Wilson et al. 1994; plant names from USDA PLANTS or Tropicos.

Recognition:

Head angularly incised posteriorly, mesonotum with 5 carinae (intermediate carinae sometimes obsolete), 10 or fewer teeth at apex of hind tarsimere (not definitively separated from Reptalus).

Melanoliarus placitus (all photographs by Kimberley Shropshire, University of Delaware)

Collecting

Melanoliarus is most readily collected at lights (particularly near a new moon); specimens may also be found by beating or vacuum sampling woody vegetation, or sometimes by the inspection of tree trunks.

Molecular resources: As of this writing, Genbank has data for M. slossonae, placitus, humilis, vicarius, and 4 undet sp.; Barcode of Life has M. humilis and M. quinquelineatus (both as Oliarus).

Selected references:

Ball, E. D. 1934. The genus Oliarus and its allies in North America (Homoptera: Fulgoridae). Journal of the Washington Academy of Science 24: 268-276.

Bourgoin T. & Wilson M. R. 1998. Description of two new species of South American Oliarus Stål (Homoptera: Fulgoromorpha: Cixiidae), including a rice-associated species from Peru. Proceedings of the Entomological Society of Washington 100(1): 108-113.

Caldwell, J. S. 1947. New species of Oliarus Stål from southwestern United States and Mexico (Homoptera: Cixiidae). Pan-Pacific Entomologist 23: 145-151.

Caldwell, J. S. and L. F. Martorell. 1951 [dated 1950]. Review of the Auchenorynchous [sic] Homoptera of Puerto Rico. Part II. The Fulgoroidea except Kinnaridae. Journal of Agriculture of the University of Puerto Rico 34(2): 133-269.

Emeljanov, A. F. 2001. The generic position of some Nearctic Pentastirini (Homoptera: Fulgoroidea: Cixiidae). Zoosystematica Rossica 9: 122.

Fennah, R. G. 1945. The Fulgoroidea, or lanternflies, of Trinidad and adjacent parts of South America. Proceedings of the United States National Museum 95(3184): 411-520.

Holzinger, W. E., A. F. Emeljanov and I. Kammerlander. 2002. The family Cixiidae Spinola 1839 (Hemiptera: Fulgoromorpha) – a review. Pp. 113-138. In: Holzinger, W. (ed.). Zikaden: Leafhoppers, Planthoppers, and Cicadas (Insecta: Hemiptera: Auchenorrhyncha). Denisia, Volume 4. Oberosterreichisches Landesmuseum, Linz, Austria. 556 pp.

Kramer, J. P. 1983. Taxonomic study of the planthopper family Cixiidae in the United States (Homoptera: Fulgoroidea). Transactions of the American Entomological Society 109: 1-57.

Mead, F. W. and J. P. Kramer. 1982. Taxonomic study of the planthopper genus Oliarus in the United States (Homoptera: Fulgoroidea” Cixiidae). Transactions of the American Entomological Society 107: 381 569.

Van Duzee, E. P. 1929. A new Oliarus. Pan-Pacific Entomologist 6: 72.

Wilson, S. W., C. Mitter, R. F. Denno, and M. R. Wilson.1994. Evolutionary patterns of host plant use by delphacid planthoppers and their relatives. In: R. F. Denno and T. J. Perfect, (eds.). Planthoppers: Their Ecology and Management. Chapman and Hall, New York. Pp. 7-45 & Appendix