Metaphycus Mercet 1917,

Zuparko, Robert L., 2015, Annotated Checklist of California Encyrtidae (Hymenoptera), Zootaxa 4017 (1), pp. 1-126: 35-41

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Metaphycus Mercet 1917


Metaphycus Mercet 1917 

Hosts. Hemiptera  : Coccidae  , Diaspididae  , Eriococcidae  , Kermesidae  , Pseudococcidae  , Triozidae 

alberti ( Howard 1898 a: 247)  ( Aphycus  )

Type. USNM

Distribution. W (Los Angeles, Riverside, San Diego, Santa Barbara)

Host/habitat. Ceroplastes  sp., Coccus hesperidum  , Parthenolecanium persicae 

Remarks. Originally described from Australia, this species has also been recorded from Hawaii and South Africa. Stauffer & Rose (1997) reported this species from Riverside county on C. hesperidum  , while Bernal et al. (2001) reported it on citrus from the southern San Joaquin Valley (Fresno, Kern and Tulare counties) without specifying exactly which county it occurred in. They also reared this species from a batch of Coccus pseudomagnoliarum  , noting that the exact host still needs to be confirmed. Kapranas et al. (2007) failed to find this species in their survey of Coccus hesperidum  in southern California citrus. A single specimen from Solano County ( UCDC) may belong to this species as well.

angustifrons Compere 1957: 227 

Type. USNM

Distribution. E (Riverside, San Bernardino, San Diego)

Host/habitat. Coccus hesperidum 

Remarks. In the 1950 s, this species was repeatedly imported into California from the Orient against Coccus hesperidium  and Saissetia oleae  , but it was never considered established here ( Bartlett 1978 a; Lampson & Morse 1992) until Kapranas et al. (2007) found it 50 years later in a survey of C. hesperidum  parasitoids. Three scale species ( Coccus pseudomagnoliarum  , Pulvinaria psidii  and S. oleae  ) have been reported as hosts of M. angustipennis  ( Noyes & Hayat 1994; Noyes 2001), based upon citations from Bartlett (1978 a) and Cock (1985) but I consider these records dubious, at best. For C. pseudomagnoliarum, Barlett  (page 61) noted that M. angustipennis  (among other species) reportedly “showed some interest in this scale in the laboratory”, and for S. oleae, Bartlett  (page 69) simply reported that the parasitoid was imported in a biocontrol program, without any evidence that the scale was actually an acceptable host. The record of P. psidii  is traceable to its importation into Bermuda in the 1950 s, when Bennett & Hughes (1959) reported that many parasites from California of “unknown value” against the scale were imported, and among these was M. angustifrons  , which did not become permanently established there. This species is very close to M. stanleyi  , and Rugman-Jones et al. (2011), suggest that records of the former may be applicable to the latter.

anneckei Guerrieri & Noyes 2000 

Type. PPRI

Distribution. E (Alameda, Los Angeles, Riverside, San Diego, Santa Clara, Ventura, San Francisco Bay area, Central Valley)

Host/habitat. Coccus hesperidum  , Saissetia miranda  , S. oleae  , Waxiella mimosae  ; NEW: Parasaissetia nigra  ( UCRC)

Remarks. Early in the 20 th century, the South African species Metaphycus lounsburyi  was exported to several countries (including Australia) in biocontrol programs directed against Saissetia oleae  . However, Guerrieri & Noyes (2000) realized that two parasitoid species were involved, and described the second one as M. anneckei  , noting that much of the literature about M. lounsburyi  may refer to M. anneckei  . In 1916, “ M. lounsburyi  ” was imported from Australia and became established in southern California ( Smith & Compere 1928), but Noyes (in litt.) thinks this was likely M. anneckei  . Bartlett (1978 a) reported that M. lounsburyi  (presumably M. anneckei  ) may be the second most effective parasitoid of the black scale in the State, while Kennett (1986) and Lampson & Morse (1992) found it to be much less important. Specimens labelled as this species are recorded from “ Lecanium viridis  ” (=? Coccus viridis Green  ) and Parthenolecanium corni  (both UCRC), but I suspect these are misidentifications.

argyrocomus ( Compere 1947: 10)  ( Erythraphycus  ) Type. USNM

Distribution. C (Lassen, Marin, Monterey, Riverside) Host/habitat. Eriococcus  sp.

armitagei ( Compere 1926 a: 30)  ( Aenasioidea  )

Type. USNM

Distribution. C (Los Angeles, Marin)

Host/habitat. The original description was based on a specimen collected on a Quercus  sp. in Los Angeles, which led Compere to opine that the host was likely to be a Kermes  sp., but such an association has never been established. A single specimen ( RLZC) was collected on Quercus agrifolia  in Marin County, indicating that the species extends throughout California’s central coast area.

californicus ( Howard 1898 a: 245)  ( Aphycus  )

Type. USNM

Distribution. N (Butte, Calaveras, Contra Costa, El Dorado, Lassen, Los Angeles, Madera, Marin, Monterey, Nevada, Plumas, Riverside, San Benito, San Bernardino, San Joaquin, San Luis Obispo, Santa Barbara, Santa Clara, Sonoma, Stanislaus, Tulare, Yolo)

Host/habitat. Eulecanium  sp., Mesolecanium nigrofasciatum  , Parthenolecanium corni  , P. pruinosum 

Remarks. Timberlake (1916) thought that M. oregonensis  ( Howard 1898 a) was probably the male of M.

californicus  , and later suggested that M. pulvinariae ( Howard 1881)  might be synonymous as well ( Compere & Annecke 1961). This same paper noted that M. californicus  was indistinguishable from the imported Palearctic M. insidiosus ( Mercet 1921)  , although there is no evidence that the latter ever established in California. They also reported that M. californicus  is likely to be confused with M. stanleyi Compere  morphologically, although these two species can be separated based on host species. Two specimens from Santa Barbara County  ( USNM) that seems to be referable to this species were reared from Physokermis  insignicola  .

calvus ( Compere 1947: 11)  ( Erythraphycus  )

Type. USNM

Distribution. C (Contra Costa, Marin, Riverside, San Benito, San Bernardino, Santa Clara, Solano, Stanislaus)

Host/habitat. Collected on an Eriogonum  sp. ( Polygonaceae  )

Remarks. Compere described this species with only the first two funicular segments darkened, but here I include a series of specimens which have the 3 rd and 4 th segments darkened as well.

clauseni ( Timberlake 1918: 358)  ( Pseudococcobius  )

Type. USNM

Distribution. C (Alameda, Calaveras, Contra Costa, Imperial, Kern, Los Angeles, Marin, Napa, Riverside, San Benito, San Bernardino, San Diego, Santa Clara, Sonoma, Stanislaus)

Host/habitat. Amonostherium lichtensioides  , Eriococcus adenostomae  , E. palustris  , E. sp.; NEW: Eriococcus coccineus  ( UCRC)

Remarks. Timberlake (1918) recorded this species from an “ Erium  sp.” on cactus, which Peck (1951) considered possibly an Amonostherium  species. Essig (1926) reported the association with E. palustris  on Spartina foliosa  ( Poaceae  ), which is limited to the high tide zone in the San Francisco Bay area. I have collected specimens over a range of ecological habitats, including the Sierra foothills, inland chapparal, and the coastal plain.

coquilletti ( Howard 1898 a: 244)  ( Aphycus  ) Type. USNM

Distribution. N (Los Angeles) Host/habitat. Pulvinaria bigeloviae 

eriococci ( Timberlake 1916: 631)  ( Aphycus  )

Type. USNM

Distribution. N (Contra Costa)

Host/habitat. Coccus hesperidum  , Eriococcus quercus 

Remarks. Originally described from Utah, Essig (1926) reported this species from California, without providing a more specific location. I’ve found only one specimen from California, from Mt. Diablo State Park.

eruptor ( Howard 1881: 364)  [New state record] ( CSCA, RLZC) Type. USNM

Distribution. W (Marin, San Diego)

Host/habitat. Ceroplastes cirridpediformis  , C. floridensis  , C. sp., “ Lecanium  sp.”

flammeus Compere 1947: 13 

Type. USNM

Distribution. C (Alameda, Calaveras, Contra Costa, Kern, Los Angeles, Marin, Mendocino, Plumas, Santa Clara, Sierra, Solano, Tehama)

Host/habitat. Parthenolecanium quercitronis 

fumipennis ( Timberlake 1918: 356)  ( Pseudococcobius  ) Type. USNM

Distribution. N (Riverside, San Bernardino) Host/habitat. Eriococcus  sp., Phenacoccus solani  Remarks. Timberlake (1918) reported this species was reared from Pseudococcus solani (Cockerell)  in southern California (San Bernardino County). However, it is likely that Timberlake’s identification of the host was based on Essig’s (1909 or 1914) characterization, which was a misidentification of Phenacoccus solani  ( Ben-Dov 2006 b). Certainly, in a later work Essig (1926: 833) reported the parasitoid (as Pseudococcobius fumipennis  ) “Reared from Phenacoccus solani Ferris  in southern California” and did not include any mention of Pseudococcus solani  , suggesting that he recognized the initial record of this species to be a mistake. Further, while Phenacoccus solani  is known from throughout California ( McKenzie 1967), Pseudococcus solani  is known only from New Mexico ( Ben-Dov 2006 b).

funicularis Annecke 1965: 227 

Type. SANC

Distribution. E (Alameda, Marin, Monterey, Santa Cruz, Yolo)

Host/habitat. Pulvinaria delottoi  , Pulvinariella mesembryanthemi  , P. sp.

Remarks. This species was introduced from South Africa with M. stramineus  in a biocontrol program against P. delottoi  and P. mesembryanthemi  from 1978–1983, and along with Encyrtus saliens  , is credited with successfully controlling both scale species ( Tassan & Hagen 1995). The record of M. funicularis  being used in a biocontrol program against Saissetia oleae ( Lampson & Morse 1992)  appears to be in error.

fuscipennis ( Howard 1898 a: 241)  ( Aphycus  )

Type. USNM

Distribution. C (Alameda, Amador, Calaveras, Contra Costa, El Dorado, Kern, Marin, Monterey, San Benito, San Bernardino, Santa Barbara, Santa Clara, Sonoma, Stanislaus)

Host/habitat. Eulecanium  sp. on Arctostaphylos pungens  ( Ericaceae  )

Remarks. Both Howard (1898 a) and Compere (1947) reported this species from undetermined “ Lecanium  ” species, but this generic name has since been suppressed under Eulecanium Cockerell. Essig (1926)  ascribed the original host record from Arctostaphylos  in Sonoma County to Parthenolecanium corni  , but this conclusion is questionable since P. corni  has not otherwise been associated with any Arctostaphylos  species.

hageni Daane & Caltagirone 1999: 14 

Type. EMEC

Distribution. E (Tehama)

Host/habitat. Saissetia oleae 

Remarks. This is a European species, imported from Spain in 1985 during a biocontrol program against Saissetia oleae ( Daane & Caltagirone 1999)  and has been confused with both M. anneckei  and M. lounsburyi ( Guerrieri & Noyes 2000)  .

helvolus ( Compere 1926 a: 25)  ( Aphycus  )

Type. USNM

Distribution. E (Alameda, Contra Costa, Kern, Los Angeles, Marin, Merced, Modoc, Monterey, Orange, Placer, Riverside, San Bernardino, San Diego, San Francisco, San Mateo, Santa Clara, Stanislaus, Tulare, Ventura)

Host/habitat. Ceroplastes destructor  , C. helichrysi  , C. sp., Coccus hesperidum  , C. pseudomagnoliarum  , Coccus viridis  , Eucalymnatus tessellatus  , Marsipococcus proteae  , Parasaissetia litorea  , P  . nigra, P. sp., Parthenolecanium corni  , P. persicae  , Protopulvinaria pyriformis  , Pulvinaria aethiopica  , P. psidii  , P. urbicola  , Pulvinariella mesembryanthemi  , Saissetia coffeae  , S. nigrella  , S. oleae  , S. somereni  , S. sp.; NEW: Coccus africanus  , Parlatoria pergandii  (both UCRC)

Remarks. This is a south African species which was established in California in 1937 in a biocontrol program for Saissetia oleae  , and is one of its most effective imported natural enemies ( Bartlett 1978 a; Kennett 1986; Daane et al. 1991; Lampson & Morse 1992). Noyes (2001) listed M. helvolus  as a biocontrol agent of Aonidiella aurantii  , but this is misleading—the paper he cited ( Bellows & Morse 1988) merely noted the effect of pesticides (applied for A. aurantii  control) on parasitoids of other species. Guerrieri & Noyes (2000) note that Aspidiotus  sp. (Homoptera: Diaspididae  ) and Rastrococcus mangiferae (Green)  (Homoptera: Pseudococcidae  ), have been reported as hosts, but they consider these records questionable.

howardi ( Cockerell 1898: 276)  ( Aphycus  )

Type. USNM

Distribution. N (Lassen, Riverside)

Host/habitat. Eriococcus tinsleyi  , E. sp.

Remarks. Two specimens ( UCDC) from Imperial County may belong to this species as well.

immaculatus (Howard 1894: 235)  ( Aphycus  ) [New combination]

Type. USNM

Distribution. C (Los Angeles)

Host/habitat. Aonidiella aurantii 

Remarks. The species was described in Aphycus  , based on a single male. Timberlake (1916) was unable to locate the type, and opined that if it was a true Aphycus  , then the host record of A. aurantii  was “undoubtedly erroneous”. The holotype is slide type # 1474 at the USNM, and belongs in Metaphycus  , which means the original host record may be correct.

inviscus Compere 1940 a: 20 

Type. BMNH

Distribution. E (Contra Costa, Fresno, Marin, Santa Cruz, Sonoma, Tulare)

Host/habitat. Saissetia oleae  , S. sp.

Remarks. Native to South Africa, this species was imported into California at least twice. A specimen mentioned in the original description was reared from the Riverside Insectary in 1924 —probably imported in one of the shipments of parasitoids made by E.W. Rust in the 1920 s ( Compere 1940 b)—but there is no record of this species being released then. A second importation was made in 1979, and the species was recovered soon thereafter in the Central Valley ( Kennett 1986). Bartlett (1978 a) reported a third importation of this species in 1958, which apparently failed to establish. However, Annecke & Mynhardt (1972) considered this effort involved a species that is distinct from M. inviscus  and described it as M. bartletti  (now a junior synonym of M. lounsburyi  ).

kermicola ( Timberlake 1916: 583)  ( Aenasioidea  )

Type. USNM

Distribution. N (Los Angeles)

Host/habitat. Allokermes essigi  , A. galliformis 

Remarks. The host record of Allokermes  (= Kermes  ) nigropunctatus (Ehrhorn & Cockerell)  cited in Essig (1926) is a misidentification of A. essigi (Miller & Gimpel)  .

lecanii ( Howard 1898 a: 242)  ( Aphycus  )

Type. USNM

Distribution. C (Alameda, Los Angeles, Santa Barbara) Host/habitat. Eulecanium pubescens  , E. sp., Parthenolecanium corni  , P. quercifex  , Physokermes insignicola 

lounsburyi ( Howard 1898 a: 244)  ( Aphycus  )

Type. USNM

Distribution. E (Alameda, Marin, Riverside, San Diego, San Mateo, Santa Barbara, Santa Clara, Solano, Tulare, Ventura, Yolo)

Host/habitat. Ceroplastes floridensis  , Coccus capparidis  , C. hesperidum  , C. pseudomagnoliarum  , Lichtensia viburni  , Parthenolecanium corni  , Saissetia coffeae  , S  . oleae

Remarks. This species is native to South Africa, and has been exported as a parasitoid of Saissetia oleae  to several countries. However, Guerrieri and Noyes (2000) discovered that material identified as M. lounsburyi  included a second species, which they described as M. anneckei  , noting that previous literature references to M. lounsburyi  may be referrable to M. anneckei  . One such reference includes Smith and Compere’s (1928) report of material imported from Australia into California in 1916, which Noyes (in litt.) thinks was likely M. anneckei  . In 1958, a species identified as M. inviscus  was imported from South Africa into California ( Bartlett 1978 a), but Annecke & Mynhardt (1972) considered this material represented a new species, which they described as Metaphycus bartletti  . Based on a comparison of types, Guerrieri & Noyes (2000) synonymized M. bartletti  under M. lounsburyi  . This makes M. lounsburyi  represents one of the most widespread and important natural enemies of S. oleae  in California ( Smith & Compere 1928; Bartlett 1978 a; Kennett 1986, Daane et al. 1991; Lampson & Morse 1992). In his report on the parasitoids of S. oleae  in central and northern California, Kennett (1986) reported distinct biological differences between the taxa he identified as M. lounsburyi  and M. bartletti  : the latter was one of the most common species found (in both the interior valley as well as the coastal and subcoastal areas), while the former appeared to be largely limited to the more temperate coastal area. Noyes (in litt.) suspects Kennett’s M. lounsburyi  was really M. anneckei  , and his M. bartletti  was M. lounsburyi  . Two specimens that appear close to M. lounsburyi  were taken in Contra Costa and Kern counties ( RLZC).

luteolus ( Timberlake 1916: 636)  ( Aphycus  )

Type. USNM

Distribution. W (Butte, Fresno, Kern, Los Angeles, Orange, Riverside, San Bernardino, San Diego, Tulare, Ventura, Yolo, Yuba)

Host/habitat. Coccus hesperidum  , C. pseudomagnoliarum  , Parthenolecanium corni  , Pulvinaria psidii  , Pulvinariella mesembryanthemi  , Saissetia coffeae  , S. oleae 

Remarks. This species may prove to be a junior synonym of M. flavus (Howard) ( Guerrieri & Noyes 2000)  . These authors also note that the host record of Saissetia coffeae  was incorrect, based upon an error in Herting (1972). However, S. coffeae  is now considered the senior synonym of S. hemisphaerica  ( Ben-Dov 2006 a), and there are several records of that species serving as a host for M. luteolus  . Coccus viridis  has been noted as host of M. luteolus ( Noyes & Hayat 1994: 398)  , based on its use in biocontrol programs against the scale in Bermuda and Hawaii. However, M. luteolus  failed to establish in both cases, and there is no record of it successfully attacking this host ( Bennett & Hughes 1959; Bartlett 1978 a). Specimens ( UCRC) have been reportedly reared from Aonidiella aurantii  and Parlatoria pergandii  (both Hemiptera  : Diaspididae  ), but these records need to be confirmed.

matteolus ( Compere 1947: 11)  ( Erythraphycus  )

Type. USNM

Distribution. C (Contra Costa, Imperial, Kern, Los Angeles, Marin, Merced, Napa, Riverside, San Bernardino, Solano, Stanislaus)

Host/habitat. Host unknown, but this species is common in chaparral, collected off Adenostoma fasciculatum  , Chilopsis linearis arcuata  ( Bignoniaceae  ), Juniperus californicus  ( Cupressaceae  ) and Larrea tridentata  ( Zygophyllaceae  ).

Remarks. This species was described based on a single male specimen ( Compere, 1947). There are a series of females ( EMEC, RLZC) that match the original description exactly, except for the color of the fore- and hind tibiae. Given the propensity for sexual dimorphism in the Encyrtidae  , I have no hesitation in considering these females conspecific with M. matteolus  . The specimen from Merced County ( EMEC) was labeled “ex Desmia  on wild grape”, but I think it likely this simply indicates the specimen was collected on grape ( Vitaceae  ) infested with a Desmia  species, and does not reflect an actual rearing record.

physokermis ( Timberlake 1916: 606)  ( Aphycus  ) Type. USNM

Distribution. C (Alameda, Contra Costa, Humboldt, Plumas, Santa Barbara) Host/habitat. Physokermis  insignicola 

psyllidis Compere 1943: 72 

Type. USNM

Distribution. C (Los Angeles, Orange, Santa Barbara, Santa Clara, Ventura)

Host/habitat. Bactericera cockerelli 

Remarks. This is one of only three species in the genus recorded as psyllid parasitoids ( Guerrieri & Noyes 2000).

stanleyi Compere 1940 a: 20 

Type. Lost

Distribution. E (Los Angeles, Orange, Riverside, San Bernardino, San Diego, Santa Barbara, Tulare, Ventura  )

Host/habitat. Ceroplastes brevicauda  , C. sp., Coccus alpinus  , C. celatus  , C. hesperidum  , C. pseudomagnoliarum  , C. viridis  , Eucalymnatus tessellatus  , Lichtensia chilianthi  , Parasaissetia nigra  , P. sp., Protopulvinaria pyriformis  , Pulvinariella mesembryanthemi  , Pulvinaria psidii  , Saissetia coffeae  , S. nigrella  , S. oleae  , Saissetia somereni  , S. sp.; NEW: Pulvinarisca jacksoni  ( UCRC)

Remarks. Although the name is properly ascribable to Compere 1940 a, the full description is found in Compere 1940 b. Native to Africa, this species was imported into California in 1937 in a biocontrol program against Saissetia oleae  , and immediately became established ( Compere 1940 a, 1940 b; Bartlett 1978 a). This species was recovered in southern California over 55 years later at very low levels ( Lampson & Morse 1992), but was not found in central or northern California ( Kennett 1986; Daane et al. 1991). Bernal et al. (2001) reported this species on citrus from the southern San Joaquin Valley (Fresno, Kern and Tulare counties) without specifying exactly in which county this species occurred. This species is very similar to M. californicus  , separable only by differences in hosts ( Compere & Annecke 1961). A host record of Coccus subhemisphaericus (Newstead)  is doubtfully referred to M. stanleyi  ( Compere 1940 a).

stramineus Compere 1940 a: 28 

Type. BMNH

Distribution. E (Alameda, Monterey, San Mateo)

Host/habitat. Pulvinaria delottoi  , Pulvinariella mesembryanthemi 

Remarks. Imported from South Africa from 1978–1983 in a biocontrol program against iceplant scales, this species, along with Encyrtus saliens  and Metaphycus funicularis  , established immediately and are credited with the successful control of those species ( Tassan & Hagen 1995).

trimblei ( Dozier 1936: 183)  ( Aenasioidea  )

Type. USNM

Distribution. N (San Bernardino)

Host/habitat. Parthenolecanium quercifex 

Remarks. The only primary literature record of this species is the original description, based on specimens from Caledonia, Pennsylvania. Noyes (2001) noted the type was from California, but I suspect this was a misreading of “ Caledonia ”. However, there is a single specimen of this species collected from San Bernardino County ( UCRC).

zebratus ( Mercet 1917: 138)  ( Aphycus  )

Type. MNMS

Distribution. E

Host/habitat. Ceroplastes floridensis  , Drepanococcus cajani  , Eriopeltis festucae  , E. lichtensteini  , Lecanopsis formicarum  , Luzulaspis luzulae  , Parthenolecanium corni  , P. persicae  , P. pomeranicum  , P. rufulum  , Pulvinaria vitis  , Saissetia oleae 

Remarks. This is a Palearctic species, imported into California in 1986 in a biocontrol program against Saissetia oleae  . The species was released in northern and central California (Alameda, Contra Costa, Fresno, Glenn, Madera & Tehama counties), recovered and considered established, although the exact localities were not specified ( Lampson & Morse 1992). Questionable host records for this species include Aonidiella orientalis  , Planchonia arabidis  , Nipaecoccus  sp., Phenacoccus aceris  and Trionymus perrisi  .


Remarks. In addition to the described species listed above, I have seen specimens that appear to represent up to 31 undescribed morphospecies from the state ( LACM, RLZC, SBMN, UCDC, UCFC, UCRC).


Smithsonian Institution, National Museum of Natural History


R. M. Bohart Museum of Entomology


ARC-Plant Protection Research Institute, National Collection of Fungi: Culture Collection


University of Newcastle


University of California, Riverside


California State Collection of Arthropods


Agricultural Research Council-Plant Protection Research Institute


Essig Museum of Entomology


Natural History Museum of Los Angeles County


University of Central Florida












Metaphycus Mercet 1917

Zuparko, Robert L. 2015


Daane 1999: 14


Annecke 1965: 227


Compere 1957: 227

argyrocomus (

Compere 1947: 10

calvus (

Compere 1947: 11


Compere 1947: 13

matteolus (

Compere 1947: 11


Compere 1943: 72


Compere 1940: 20


Compere 1940: 20


Compere 1940: 28

trimblei (

Dozier 1936: 183

armitagei (

Compere 1926: 30

clauseni (

Timberlake 1918: 358

fumipennis (

Timberlake 1918: 356

zebratus (

Mercet 1917: 138

eriococci (

Timberlake 1916: 631

kermicola (

Timberlake 1916: 583

luteolus (

Timberlake 1916: 636

physokermis (

Timberlake 1916: 606

alberti (

Howard 1898: 247

californicus (

Howard 1898: 245

coquilletti (

Howard 1898: 244

fuscipennis (

Howard 1898: 241

howardi (

Cockerell 1898: 276

lecanii (

Howard 1898: 242

lounsburyi (

Howard 1898: 244

eruptor (

Howard 1881: 364