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PROCEEDINGS
OF THE
California Academy of Sciences
FOURTH SERIES
Vol. XL
SAN FRANCISCO PUBLISHED BY THE ACADEMY 1974-1975
COMMITTEE ON PUBLICATION GrorcE E. LinpsAy, Chairman Diana R. Younc, Editor PauLt H. ARNAUD, JR. WILLIAM N. ESCHMEYER
ALAN E. LEvITON
No.
CONTENTS OF VOLUME XL
WILLIAMS, STANLEY C. A new genus of North American scorpions with a key to the North American genera of Vaejovidae (Scorpionida: Vaejovidae). Published October SIC) 7) Laas eRe ein nee Ee en
CARTER, ANNETTA M. The genus Cercidium (Leguminosae: Caesalpinioideae) in the Sonoran Desert of Mexico and the United States. Published October 30, 1974 _..
HALLACHER, LEoN E. The comparative morphology of ex- trinsic gasbladder musculature in the scorpionfish genus Sebastes (Pisces: Scorpaenidae). Published October 30, LAr pees WRENS Bet he Pd ES Ds I 2 SS
Murpuy, Rosert W. A new genus and species of euble- pharine gecko (Sauria: Gekkonidae) from Baja California, Mexico: Published) October 30) 1907/4 2
Murpuy, Rospert W. Two new blind snakes (Serpentes: Leptotyphlopidae) from Baja California, Mexico with a contribution to the biogeography of peninsular and insular herpetofatna. Published February 18, 1975
CHEN, Lo-cHat. The rockfishes, genus Sebastes (Scor- paenidae), of the Gulf of California, including three new species, with a discussion of their origin. Published Feb- ELUTE WN 6 BLS Au LO) [is an Si) ee Se ee
Haves, ALAN H. The larger moths of the Galapagos Islands (Geometroidea: Sphingoidea & Noctuoidea). Published PAUSES LS AUN ()9) ees eee Me. SP Tee ee ee, ee
SmitH, ALAN Reip. New species and new combinations of ferns from Chiapas, Mexico. Published August 8, 1975
ZAVORTINK, THOMAS J. A new genus and species of eucerine bee from North America (Hymenoptera: Anthophoridae). ECD LIS ECE AU UIST. sO), 1,0 aera ee mers ek Beene Seer
GoLp, J. R., anp G. A. E. GALL. The taxonomic structure of six golden trout (Salmo aguabonita) populations from the Sierra Nevada, California (Pisces: Salmonidae). Pub- Mi Se dwAUIEUS ts Sal Ooms ek) ee Ree ees
Pages
17-57
59-86
87-92
93-107
109-141
145-2081
209-230
231-242
243-263
No.
No.
No.
No.
11g
WAS
1S.
» 14:
Se
EscHMEYER, WILLIAM E., AND JoHN E. RANDALL. The scorpaenid fishes of the Hawaiian Islands, including new species and new records (Pisces: Scorpaenidae). Published (CCC ae Neel BSA ee ee eee 1 ee
Howarp, L. D. Muscular anatomy of the hind limb of the sea otter (Enhydra lutris). Published October 3, 1975
McCoskErR, JOHN E., AND RicHarp H. RosENBLATT. The moray eels (Pisces: Muraenidae) of the Galapagos Islands, with new records and synonymies of extralimital species. Published-@ctober ee 10S eee
Davis, JOHN, AND W. Z. LipIcKER, JR. The taxonomic status of the southern sea otter. Published October 3, 1975 —
PAxTon, JOHN R. Heraldia nocturna, a new genus and species of pipefish (family Syngnathidae) from eastern Aus- tralia, with comments on Maroubra perserrata Whitley. Publisheds@ctoher 31075. - 2 ot ee ee
imGgexito: Volume ky Seaenrer er te Saar sss ee eA ee
265-334
335-416
417-427
429-437
439-447 449-460
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
/
FOURTH SERIES ’ Walt La haled ok 7%
Vol. XL, No. 1, pp. 1-16; 5 figs.; 1 table. October 30, 1974
A NEW GENUS OF NORTH AMERICAN SCORPIONS WITH A KEY TO THE NORTH AMERICAN GENERA OF VAEJOVIDAE (SCORPIONIDA: VAEJOVIDAE)
By
Stanley C. Williams
Research Associate in Entomology, California Academy of Sciences San Francisco 94118
Apstract: A new genus of North American vaejovid scorpions is described and named Nullibrotheas. Nullibrotheas is a monotypic genus, the only species being Nullibrotheas allenii (Wood). The systematic relationships of NV. allenii are analyzed and it is concluded that the closest known relatives are in the vaejovid genera Uroctonus and Vaejovis. A key to the North American genera of Vaejovidae is included.
INTRODUCTION
In 1863, Horatio Wood described a new species of North American scorpion which he named “Scorpius allenii” (Wood, 1863a). This new taxon was based on specimens collected by L. J. Xantus de Vesey from Cabo San Lucas, in Baja California, Mexico, presumably between 1859 and 1861. This new species had the distinction of being one of the first species of North American scorpions to be described and named, and it has remained one of the most misunderstood taxa in North America. The actual identity and phylogenetic relationships of this species remained unknown until just recently.
The main purposes of this study are to more clearly define the species ‘allenii, to determine its geographical distribution, to study its systematic affinities, and to erect a new genus to permit appropriate taxonomic placement of ‘allenii. Most of the specimens cited as new records are deposited in the collections of the California Academy of Sciences.
[1]
2 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
ACKNOWLEDGMENTS
Much appreciation is due the following individuals and their respective institutions for cooperation and loan of specimens which materially aided this study: M. A. Cazier, Arizona State University; J. A. Chemsack, California Insect Survey, University of California, Berkeley; W. J. Gertsch, American Museum of Natural History; C. F. Harbison, San Diego Museum of Natural History; H. W. Levi, Harvard Museum of Comparative Zoology; P. H. Arnaud, Jr., California Academy of Sciences. Thanks are due Richard and Mary Lou Adcock for providing transportation to the islands in the Gulf of California aboard their boat Marisla in 1968, to Thomas and Doris Hearne, who sponsored an expedition to study the Gulf Islands aboard their boat Muy Pronto in 1969, and to Richard and Elenore Dwyer who sponsored another research expedition to the Gulf Islands aboard their boat Sea Quest in 1970.
Appreciation is owed to Carolyn Mullinex for making technical drawings, to V. F. Lee for technical and field assistance, and to C. F. Williams for clerical assistance. This study was partially supported by the National Science Foundation through research grants GB 7679 and GB 23674 and the California Academy of Sciences through the use of the research facilities of the Department of Entomology.
Nullibrotheas Williams, new genus
TyYPE-SPECIES: Scorpius allenii Wood, 1863.
Description. Pedipalp palms greatly swollen, fingers shorter than palm or carapace; brachium with posterior border of inferior surface with six long trichobothria; fingers with supernumerary denticles flanking single principal row of denticles medially. Pectines with fulcra sub-triangular in shape; middle lamellae with elongate basal piece and one row of subcircular to ovate sclerites; males with longer comb, longer teeth, and greater number of teeth than females. Males with large distinct genital papillae on inner margins of genital operculum. Sternum pentagonal, with almost parallel sides; length approximates width or is slightly longer. Stigma of book lungs minute, circular in shape. Last tarsomere of walking legs with one row of short spines on ventral surface; two pedal spurs. Two median eyes on low ocular tubercule; median eyes entirely on anterior half of carapace; lateral eyes three per group, third eye reduced to degenerate, often giving appearance of two-eyed condition.
Nullibrotheas allenii (Wood). (Figures 1-4; table 1.) Scorpius allenti Woop, 1863a, p. 360 (original description) ; 1863b, p. 107.
Broteas allenii, MARx, 1887, p. 91. Uroctonus privus Karscu, 1879, p. 103 (original description) ; HyJELLE, 1972, p. 28-29.
Vor. XL] WILLIAMS: NEW SCORPION GENUS 3
TaBLE 1. Measurements (in millimeters) of Nullibrotheas allenii Wood, SCW #116 (1), topotypes from Cabo San Lucas, Baja California Sur, Mexico.
Male Female Total length 42 30 Carapace, length 5.0 4.5 width (at median eyes) 4.1 Saf) Metasoma, length segment I (length/width) DP ores 1.5/2.4 segment II (length/width) 2.6/2.7 1.9/2.2 segment III (length/width) 2.9/2.6 2.0/2.0 segment IV (length/width) 3.5/2.4 2.6/1.8 segment V (length/width) SW oe 4.3/1.8 Telson length 6.0 4.2 Vesicle (length/width) 4.4/2.4 2.8/1.8 depth 2.0 1.4 Aculeus, length 1.6 1.4 Pedipalp Humerus (length/width) 3.4/1.7 3.0/1.4 Brachium (length/width) 3.9/1.6 3.5/1.4 Chela (length/ width) oll Bell 6.7/2.5 depth 4.8 3.6 movable finger, length 4.3 3.5 fixed finger, length 2.4 D2 Pectines, teeth (left/right) 11/11 8/9 Sternum (length/ width) Taye elysle? Stigma (circular), diameter 0.2 0.2 Number middle lamellae 6 5 Genital operculum (length/ width) 0.8/1.6 0.6/1.7
Uroctonus mordax (part), KRAEPELIN, 1894, p. 194 (synonymy of U. privus).
Broteas formosus Marx, 1889, p. 211.
Broteas alleni, EWING, 1928, pp. 3, 6 (part) ; HOFFMANN, 1931, pp. 332-333; KRAEPELIN, 1899, p. 176 (lists as doubtful species of chactid); GerrtscH, 1958, pp. 2-5; GerTscH and SOLEGLAD, 1966, p. 1; Diaz Najera, 1970, pp. 113-114.
Broteochactas allenii (part), BANKs, 1910, p. 188.
Type DATA. Scorpius allenii Wood, two cotypes (one male, one female) ; collected at Cabo San Lucas, Baja California Sur, Mexico; male cotype deposited in U. S. National Museum (Type number S-5, jar 2), the location of the female cotype is unknown. The male cotype is intact, badly yellowed, and not well preserved. This specimen was designated as a lectotype by Gertsch (1958). It appears to be a subadult.
The type of Uroctonus privus Karsch is a holotype female with the following
4 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Sy
Ficure 1. Nullibrotheas allenti (Wood), dorsal and ventral views of male topotype from Cabo San Lucas, Baja California Sur, Mexico.
VoL. XL] WILLIAMS: NEW SCORPION GENUS
On
data attached ‘“Californien; leg. graber.” It is deposited in the Zoological Museum of Berlin, DDR; catalog number 3036. The holotype is a small juvenile that does not differ significantly from the juveniles of NV. allenii collected recently in the Cape region of Baja California, Mexico. The holotype is somewhat faded with age but the color pattern is still apparent; the metasoma is severed between segments 2 and 3.
REDESCRIPTION BASED ON TOPOTYPES FROM CABO SAN Lucas. Variable species throughout range in coloration, cuticular granulation, hirsuteness, and body size; apparently forming many more or less isolated regional races. Typically with base color of cuticle golden yellow with underlying dusky to dark markings on carapace and mesosomal dorsum; pedipalps with dark reddish brown fingers; metasoma with inferior keels with more or less distinct dusky stripes; other parts of cuticle may or may not show underlying dusky markings.
Carapace. Anterior margin usually with deep median emargination and rounded lateral ends; lateral eyes generally three per group (sometimes appar- ently two per group); median eyes two in number, located on elevated ocular tubercule; median diad completely on anterior half of carapace. Sternum large, pentagonal; lateral sides almost parallel; length approximates width or is slightly longer; with deep median longitudinal furrow along posterior half.
Mesosoma. Terga 1 to 6 usually lustrous; seventh tergum with one pair short, obsolescent lateral keels, these granular; terga 1 to 6 lacking distinctly developed dorsal keels; sterna smooth and lustrous; stigma small, circular in shape; all sterna lack keels.
Metasoma. Dorsal and dorsolateral keels present and granular on segments I to IV; inferior lateral keels smooth to crenulate on I, irregularly crenulate on II and III, serrate on IV and V. Inferior median keels smooth to obsolescent on I, crenulate to obsolescent on IT, crenular on III, serrate on IV and V.
Telson. Aculeus short, less than one-half length of vesicle; larger individuals normally with laterally swollen vesicle.
Pectines. Male with middle lamellae consisting of elongate basal piece and about five or six subcircular to ovate sclerites in single row; fulcra subtriangular; pectinal teeth six to nine in females, 10 to 14 in males; female comb much smaller than male, proximal one-fourth of female comb lacking teeth.
Genital operculum. In males completely divided longitudinally; in females divided only in posterior region of furrow; males with very large, conspicuous genital papillae, these lacking in females.
Chelicerae. Fixed finger with one simple tooth and one greatly elevated bicuspid tooth; movable finger with superior border armed with three or four simple teeth in addition to terminus; inferior border with three or four small denticles, these not conspicuous.
6 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 2. Nullibrotheas allenii (Wood) figured from topotype male from Cabo San Lucas, Baja California Sur, Mexico. Top. Ventral view of pedipalp brachium. Bottom. Mesosomal sternum showing openings to book lungs.
Pedipalps. Fingers very short, palm very deep and swollen; keels obsolete to smooth except for pronounced inferior keel; palm surface lustrous, smooth to finely granular. Movable finger distinctly shorter than carapace; fixed finger shorter than metasomal segment III. Each finger armed with one longitudinal row of small denticles; no distinct gap between fingers when chela closed. Fingers very hirsute. Brachium with six trichobothria on posterior border of inferior surface.
Walking legs. Two pedal spurs; one row of minute, longitudinally arranged hairs on ventral surface of last tarsomere.
VoL. XL] WILLIAMS: NEW SCORPION GENUS 7
Ficure 3. Nullibrotheas allenii (Wood), figured from topotypes from Cabo San Lucas, Baja California Sur, Mexico. Top left. Ventral view of male showing prosomal sternum, genital opercula, and pectines. Top right. Ventral view of female showing prosomal sternum, genital opercula, and pectines. Bottom left. Right anterolateral aspect of carapace showing three lateral eyes. Bottom right. Right anterolateral aspect of carapace showing two lateral
eyes.
GEOGRAPHICAL DISTRIBUTION. Known only from the southern region of the Baja California peninsula and associated islands. Specimens have been collected from Cabo San Lucas north to El Coyote on Bahia Concepcion. Reports of this species in the United States appear to be due to errors in locality records and to mistaken identification of this species.
New records. Known from the following localities in Baja California Sur, Mexico: 1 mi. SW. Rancho Canipole, elevation 800 ft., 16 May 1969, (S. C. Williams), 2 juveniles; 4 mi. SW. San Miguel de Comondu, elevation 900 it., 15 May 1969, (S. C. Williams), 2 juveniles; 5 mi. SW. San Miguel de Comondu,
CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
3 °
AA QC Q )
ou c ot
Ficure 4. Nullibrotheas allenii (Wood), figured from topotypes from Cabo San Lucas, Baja California Sur, Mexico. Top left. Ventral view of male metasoma. Top right. Movable pedipalp finger of male. Bottom left. Female telson. Bottom right. Male telson.
Vor. XL] WILLIAMS: NEW SCORPION GENUS 9
elevation 1000 ft., 2 July 1968, (S. C. Williams, M. A. Cazier), 1 male, 1 female, 1 juvenile; 5 to 10 mi. SW. San Miguel de Comondu, elevation 1000 ft., 3 July 1968, (S. C. Williams, M. A. Cazier), 1 female, 3 juveniles; San Jose de Comondu, 15 February 1966, (V. Roth), 1 male, 1 female; 4 mi. W. La Purisima, elevation 375 ft., 1 July 1968, (S. C. Williams, M. A. Cazier), 1 female; Rancho Las Parras, 26 May 1970, (S. C. Williams, V. F. Lee), 1 juvenile; 9.9 mi. N. Loreto, 27 May 1970, (S. C. Williams, V. F. Lee), 1 juvenile; Loreto city dump, just N. Loreto, 20 June 1964, (Chris Parrish), 1 female; 8 mi. S. Loreto, base of La Gigantia, 27 January 1965, (V. Roth), 1 male juvenile; Puerto Escondido, 17 mi. S. Loreto, 27 May 1970, (S. C. Williams, V. F. Lee), 1 juvenile; 22 mi. NE. Santo Domingo, 15 February 1966, (V. Roth), 1 female, 1 juvenile; 51 mi. N. El Crucero, elevation 400 ft., 14 May 1969, (S. C. Williams), 1 male, 3 females, 7 juveniles; 10 mi. N. El Crucero, 15 February 1966, (V. Roth), 1 male juvenile, 1 female juvenile; Bahia Concepcion near E] Coyote, 26.40 N., 111.50 W., 17 February 1966, (V. Roth), 3 males; 10.3 mi. SE. Santa Rita, 27 July 1968, (S. C. Williams, M. A. Cazier), 1 male, 1 female, 1 juvenile; 49.3 mi. SE. Santa Rita, 27 July 1968, (S. C. Williams, M. A. Cazier), 1 juvenile; 49.3 mi. SE. Santa Rita, 27 July 1968, (S. C. Williams, M. A. Cazier), 1 male, 4 juveniles; 31.0 mi. W. Los Aripes, elevation 800 ft., 25 July 1968, (S. C. Williams, M. A. Cazier), 3 juveniles; 1 mi. E. Los Aripes, 8 July 1968, (S. C. Williams, M. A. Cazier), 1 male; 1.4 mi. W. El Coyote, 30 December 1958, (H. B. Leech), 1 male, 3 juveniles; La Paz, 1 to 3 February 1965, (V. Roth), 1 female, 3 juveniles; 2 mi. E. La Paz, elevation 50 ft., 5 July 1968, (S. C. Williams, M. A. Cazier), 1 male; 12.4 mi. E. La Paz on road to Las Cruces, Arroyo Agua de los Pozos, 4 January 1959 (A. Leviton), 1 juvenile; 2 mi. NW. Los Pozos, 23 July 1968, (S. C. Williams, M. Bentzien, W. Fox), 8 juveniles; 14.5 mi. E. La Paz on road to Las Cruces, 4 January 1959, (H. B. Leech), 1 juvenile; .25 mi. N. La Paz airport, 12 July 1968, (S. C. Williams, M. A. Cazier), 1 juvenile; 6 mi. N. La Paz on road to Pichilingue, 6 January 1959, (H. B. Leech), 1 juvenile; 12.5 mi. N. La Paz on road to Pichilingue, 29 December 1958, (H. B. Leech), 1 female; Puerto Balandra, approximately 13 mi. N. La Paz, 3 September 1963, (P. R. & D. L. Craig), 1 juvenile female; 14 mi. NE. La Paz along A. Balandra road, 14 July 1968, (S. C. Williams, M. A. Cazier), 2 males, 1 female, 1 juvenile; 14 mi. NE. La Paz along A. Balandra road, 9 July 1968, (S. C. Williams, M. A. Cazier), 1 juvenile; 75 mi. NW. La Paz, elevation 200 ft., 4 July 1968, (S. C. Williams, M. A. Cazier), 1 female; 5 mi. SW. La Paz, 2 August 1968, (S. C. Williams, M. A. Cazier), 1 juvenile; Las Cruces, 5.0 mi. SW. Las Cruces, 30 July 1968, (S. C. Williams, M. A. Cazier), 2 females, 5 males, 4 juveniles; 5 to 6 mi. SW. La Paz, elevation 25 ft., 5 July 1968, (S. C. Williams, M. A. Cazier), 1 juvenile; 14.8 mi. N. Todos Santos, elevation 500 ft., 24 July 1968,
10 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
7
Guaymas 112° my 110°
a Ista Coronados
Loreto Isla Carmen e
Q° Isla Danzante
26°
v Q Isla Santa Catalina
Q_ ‘sla Santa Cruz ® Isla San Diego
Isla San Jose
Isla San Francisco
Isla Partida Isla Espiritu Santo \ Isla Cerralvo
Scale of Miles
Vor. XL] WILLIAMS: NEW SCORPION GENUS 11
(S. C. Williams, M. A. Cazier), 1 female, 6 juveniles; 5.5 mi. NW. Todos Santos, road to La Pastora, 13 January 1959, (H. B. Leech), 1 juvenile; 5.9 mi. N. Todos Santos, elevation 500 ft., 24 July 1968, (S. C. Williams, M. A. Cazier), 14 males, 18 females; 9 mi. N. Todos Santos, elevation 200 ft., 4 May 1969, (S. C. Williams), 1 female; 3.5 mi. S. El Pescadero, elevation 20 ft., 23 July 1968, (S. C. Williams, M. A. Cazier), 1 male, 1 female, 4 juveniles; 4 mi. N. Tinaja, 23 July 1968, (S. C. Williams, M. Bentzien, W. Fox), 1 female, 1 juvenile; Santiago, 19 August 1964, (H. W. Campbell), 1 male, 1 female; Boca de la Sierra, near Miraflores, 10 February 1966, (V. Roth), 1 female; Bahia de los Frailes, 9 March 1947, (I. La Rivers), 1 male; 1.5 mi. NE. Punta Palmilla, elevation 50 ft., 16 July 1968, (S. C. Williams, M. A. Cazier), 4 males, 13 females: 1.5 mi. NE. Punta Palmilla, elevation 50 ft., 17 July 1968, (S. C. Williams, M. A. Cazier), 1 juvenile; 2.2 mi. SW. Punta Palmilla, 17 July 1968, (S.C. Williams, M. A. Cazier), 3 juveniles; 3.5 mi. SW. Punta Palmilla, 17 July 1968, (S. C. Williams, M. A. Cazier), 1 juvenile; 3.9 mi. SW. Punta Palmilla, 17 July 1968, (S. C. Williams, M. A. Cazier), 4 males, 6 females; 16.5 mi. NW. Cabo San Lucas, 23 July 1968, (S. C. Williams, M. Bentzien, W. Fox), 5 juveniles; 5 mi. N. Cabo San Lucas, 21 July 1968, (S. C. Williams, M. A. Cazier), 1 juvenile; 4 mi. N. Cabo San Lucas, 21 July 1968, (S. C. Williams, M. A. Cazier), 2 females, 1 juvenile; 2 mi. N. Cabo San Lucas, 19 July 1968, (S. C. Williams, M. A. Cazier), 1 male, 1 female, 1 juvenile; 3 mi. E. Cabo San Lucas, 18 July 1968, (S. C. Williams, M. A. Cazier), 2 females; Cabo San Lucas, 20 July 1968, (S. C. Williams, M. A. Cazier), 3 males, 9 females, topotypes; Isla Carmen, Puerto Ballandra, 24 May 1970, (S. C. Williams, V. F. Lee), 2 juveniles; Isla Carmen, Puerto Ballandra, 23 March 1971, (V. F. Lee), 1 female, 1 juvenile; Isla Carmen, Marquer Bar, 22 June 1964, (R. C. Banks), 1 juvenile; Isla Santa Catalina, 27 April 1964, (C. H. Croulet), 3 females, 6 juveniles; Isla Santa Catalina, NE. side, 10 April 1962, (Members Belvedere Expedition), 1 female, 1 juvenile; Isla Santa Catalina, NW. end, Punta Arena, 24 June 1964, (C. Parrish), 1 male; Isla Santa Catalina, cave on S. end, 25 June 1964, (R. C. Banks), 1 juvenile; Isla Santa Catalina, SW. end, 26 April 1964, (A. J. Sloan), 1 male, 1 female, 4 juveniles; Isla Santa Catalina, W. side, 9 April 1962, (Members Belvedere Expedition), 5 juveniles; Isla Coyote, 19 April 1962, (George Lindsay), 1 female, 2 juveniles; Isla San Diego, WNW. side, 18 April 1962, (C. Parrish, M. Soule), 1 female; Isla San Diego, 19 April 1962, (George Lindsay), 1 juvenile; Isla San Jose, N. side, 26 March 1971, (V. F. Lee), 3 juveniles; Isla San Jose, NE. side of Arroyo de Aguada, 11 April 1962, (Members Belvedere Expedition), 1 female, 3 juveniles; Isla San Jose, S. end near lagoon mouth, 27 June 1964, (C. Parrish), 1 juvenile; Isla San Jose, SW. end, near cattle ranch, 24 April 1964, (A. J. Sloan), 2 females; Isla San Jose, Los Ostiones, 12 April 1962, (Members Belvedere Expedition), 2 females, 1 juvenile; Isla Cayo, 28 June 1964, (C. Parrish, G. Lindsay,
12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
I. Wiggins), 1 male, 2 females, 2 juveniles; Isla San Francisco, SW. side, 25 May 1969, (S. C. Williams), 1 juvenile; Isla San Francisco, S. end, 17 April 1962, (R. GC. Banks), 1 male; Isleta Rock, N. Partida Island, 20 April 1962, (M. Soule), 1 male, 1 juvenile; Isla Partida, central valley, elevation 25 ft., 9 July 1968, (S. C. Williams, M. Bentzien, W. Fox), 1 male, 2 juveniles; Isla Partida, dry lake, 29 June 1964, (I. Wiggins), 1 female; Isla Ballena, 21 April 1962, (C. Parrish), 3 juveniles; Isla Espiritu Santo, SW. shore, 7 July 1968, (S. C. Williams, M. A. Cazier), 1 male, 2 females, 10 juveniles, 1 female with 23 in litter; Isla Espiritu Santo, Bahia San Gabriel, 24 May 1969, (S. C. Williams), 5 juveniles; Isla Espiritu Santo, Bahia Canon, 29 June 1964, (I. Wiggins), 2 juveniles; Isla Espiritu Santo, Bahia Candelaro, 30 June 1964, (I. Wiggins), 1 juvenile; Isla Cerralvo, Piedras Gordas, 17 May 1970, (S. C. Williams, V. F. Lee), 5 juveniles; Isla Cerralvo, sand dune area on SW. side, 16 April 1962, (C. Parrish), 1 juvenile; Isla Cerralvo, Arroyo Aguaje, 15 April 1962, (G. Lindsay), 2 juveniles; Isla Cerralvo, Rancho Ruffo, 16 April 1962, (I. Wiggins, G. Lindsay), 1 juvenile; Isla Magdalena, Puerto Magdalena, 17 March 1957, (R. Zweifel), 1 male; Santa Margarita Island, June 1970, (USNM), 1 specimen.
Remarks. Nullibrotheas allenii is an extremely variable species in coloration, cuticular granulation, hirsuteness of metasoma and telson, and body size. Throughout its distribution it appears to form series of local races with some evidence of genetic continuity between many of the races. For this reason it appears best to treat this as one variable species rather than as a complex of many poorly defineable, separate ones.
In regard to size, adults in most populations range from 30 to 45 millimeters in length. A few local populations have been found in which adults have attained much larger sizes, in the range of 50 to 60 millimeters. Such giant races have been found in the area of La Paz and on some of the islands of the gulf, notably on Isla Catalina.
In regard to color, most populations have a golden yellow base color with underlying dark coloration on the carapace and mesosomal dorsum. Some populations have the dark markings reduced to faint dusky markings, notably on the Magdalena Plain, while others may have these markings very dense and dark, notably in the volcanic habitats, such as around Comondu. South of Todos Santos the dark coloration of the mesosomal dorsum is divided longitu- dinally by a thin unpigmented stripe thus creating the appearance of one pair of dark longitudinal stripes. North of the Todos Santos area the metasomal dorsum is usually completely pigmented giving no indication of stripes. Around the Todos Santos and Las Cruces areas individuals are variable and show all forms of intermediate conditions.
The granulation of the cuticle, especially on the carapace, vesicle, and
Vou. XL] WILLIAMS: NEW SCORPION GENUS 13
pedipalp palms is highly variable and difficult to interpret. It appears that the amount of granulation is determined by age, sex, length of time since last moult, and individual genetics. Most individuals with more distinctive granulation are larger than average.
The hirsuteness of the vesicle and metasoma is another variable character. In the Cabo San Lucas area the metasoma and telson are usually only moderately hirsute. The northern and island populations generally had both vesicle and metasomal segments IV and V considerably more hirsute than the southern populations. Samples taken from intermediate areas indicate the variation in hirsuteness is to some extent individual, and developmental, and over the range is somewhat clinal.
The collection records of this species indicate that individuals probably spend most of their life cycle in burrows in the ground. They do, however, appear to leave the burrow and seek shelter under suitable surface rocks during part of the year.
Of the hundreds of specimens collected during this study only two females were observed with litters on their backs. Both of these were discovered under the protection of moderate-size flat rocks during July. The females appear normally not to leave their protective shelters at time of birth or while carrying young on their backs. Both females had only small to moderate numbers of young in their litters. The first litter was found 4 miles N. of Tinaja on 23 July 1968 and consisted of the mother and 8 young. The second litter was found on Isla Espiritu Santo on 7 July 1968 and consisted of the mother and 23 young.
During the summer of 1970 the male cotype in the U. S. National Museum was examined and compared with topotypes recently collected at Cabo San Lucas. The type was almost identical to a sub-adult male topotype except for being more yellowish, less hirsute, and lacking the characteristic color pattern (three characteristics commonly altered in this way by poor preservation and age). It was interesting that the cotype was labeled “Broteas alleni Wood” by an old hand-written label included in the specimen jar. This was apparently a later addition or alteration of the original data since this species was originally named Scorpius allenii. It is notable that when Gertsch (1958) studied and redescribed the types two cotypes were present. In 1970 only one of the cotypes could be located, this presumably is the lectotype designated by Gertsch in 1958.
TAXONOMIC RELATIONSHIPS OF NULLIBROTHEAS. Nullibrotheas allenii was first placed within the genus Scorpius by Wood in 1863 where it remained until 1887 at which time George Marx placed it within the chactid genus Broteas. Since this species has been relatively unstudied it has essentially remained placed in the genus Broteas and family Chactidae until now. However, Banks (1910) apparently realized that ‘allenii? was not properly placed within the genus Broteas because he assigned this species to the genus Broteochactas thus
14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
retaining it within the family Chactidae. Banks’ generic assignment was, however, not followed by others working with North American scorpions.
It is interesting that European workers did not agree with the systematic placement of ‘allenii’ by American workers. In 1879, Karsch studied a juvenile specimen of NV. allenii and named it Uroctonus privus, thus placing it within the family Vaejovidae. Later Kraepelin (1899), considered U. privus (a junior synonym of JN. allenii) to be a junior synonym of Uroctonus mordax, thus also retaining NV. allenii within the family Vaejovidae. Kraepelin (1899) did not see the relationship between ‘“‘Broteas allenii” and Uroctonus privus, but he did indicate that he believed ‘‘B. allenii” to be a good species, and doubted that it should be placed in the Chactidae.
The family placement of the genus Nullibrotheas is made somewhat difficult by the obscure relationship between the families Chactidae and Vaejovidae. The primary characteristic that has been used to differentiate these two families is the presence of two lateral ocelli at each anterolateral corner of the carapace in the Chactidae. The family Vaejovidae is contrasted by having three or more lateral ocelli at each anterolateral corner of the carapace. Numbers of lateral ocelli are not satisfactory criteria alone for placement of scorpions within a family. Several species which clearly belong to families other than the Chactidae have secondarily lost lateral ocelli. For example, in the Diplocentridae several species belonging to the Didymocentrus group show only two lateral ocelli at each anterolateral corner of the carapace. These were even considered by Stahnke (1968) to represent a new diplocentrid genus which he called Bioculus. Gertsch (1972) has found several species of Vaejovis and Uroctonus which either have external signs of the third lateral ocellus completely absent or in an obsolescent state. Actually, most species of Vaejovis and Uroctonus have the third lateral ocellus showing signs of becoming reduced in size or approaching obsolescence.
Nullibrotheas allenii clearly does not belong in the genus Broteas because it has the openings to the book lungs completely circular rather than linear, and the last tarsomere of the walking legs has one row of ventral hairs (not two parallel rows of spine-like hairs). Nullibrotheas allenii also clearly does not belong in the genus Broteochactas because it has six trichobothria in a posterior row on the ventral surface of the pedipalp brachium (not seven such trichobothria), and also because the walking legs have the last tarsomere with one row of ventral hairs (not with abundant irregular hairs).
Study of the lateral ocelli of Nudlibrotheas allenii reveals two distinct eyes at each anterolateral corner of the carapace in some specimens. However, individuals usually show evidence of a third reduced or degenerate ocellus. This line of evidence indicates that this genus should be placed within the family Vaejovidae. Furthermore, in other characteristics it appears somewhat related to the genera Uroctonus and Vaejovis (both members of the family Vaejovidae).
Vor. XL] WILLIAMS: NEW SCORPION GENUS 115)
Nullibrotheas can be clearly distinguished from Vaejovis and Uroctonus in that it has the ventral surface of the pedipalp tibia (brachium) with six trichobothria in a posterior row (not with two or three such trichobothria) and has minute, round openings to the book lungs (not slit-like or oval).
Considering all the morphological evidence available it appears that Nullibrotheas has no known close living relatives, but that it is more closely related to the vaejovid genera Uroctonus and Vaejovis than to any other living ones known. Nullibrotheas is therefore here considered to be a member of the family Vaejovidae.
KEY TO THE NORTH AMERICAN GENERA OF THE FAMILY VAEJOVIDAE
ies oOkslings Openings rOUnd. =e eee ee ee Nullibrotheas, new genus Book lung openings not round; but oval or elongate 2 2. Metasoma with single, unpaired, inferior median keel on segments I to IV _
NS Ne ee SS BB ns Mee A et ee genus Syntropis Metasoma with inferior median keels of segments I to IV paired or obsolescent —.-
oes. 5 es Bg a Oe a a Fe ae en aS Be EO re Be) 8) 3. Lateral ocelli four per group; pectines with middle lamellae angular and not Giubie == ee _.. genus Anuroctonus Lateral eyes three or “fewer s per group; eee ay Paddle lamellae subcircular —_ ir ee A a a we Se oe NAN a OA eno te 4
4. Inferior margin of movable cheliceral finger with one long, dark, conspicuous tooth; pedipalps with ventral surface of brachium with over 20 trichobothria _. genus Hadrurus
Inferior margin of movable cheliceral finger with small teeth, minor denticles,
denticle-like crenulation, or completely lacking denticles; pedipalps with ventral surface of brachium with two or three trichobothria along posterior border
5. Pectines with middle lamellae usually composed of 9 or fewer circular sclerites;
openings to book lungs usually oval or long oval in shape and generally with length
less than three times width; pedipalp with ratio of palm width to palm depth
generally greater than 1.05; pedipalps with ventral surface of brachium with (HLOVREYEY «(Hs ClO oY Oe oa yp ein rs ee ee genus Uroctonus
Pectines with middle lamellae usually composed of 10 or more circular sclerites;
openings to book lungs usually elongate or slit-like and generally with length greater
than three times width; pedipalp with ratio of palm width to palm depth generally
less than 1.05; pedipalps with ventral surface of brachium always with 2 trichobothria
(Meverswithethtee)) hp een eens ee osetia ie Sosa Bae eae ee he ee bee 6 6. Chelicerae with inferior margin of avable. Gee an one or several small denticles Ormcrenula tions 5 en a ek ee ee ee es bie a) genus Paruroctonus Chelicerae with inferior margin i movable finger smooth and completely lacking CLETULALILONS we temas ear er oe see NE a ee Rae es edo _.. genus Vaejovis
REFERENCES Banks, N. 1909-1910. The scorpions of California. Pomona College Journal of Entomology, vol. 2, no. 2, pp. 185-190. D1az-NaAjera, A. 1970. Contribucion al conocimiento de los alacranes de Mexico (Scorpionida). Revista de Investigation en Salud Publica, vol. 30, no. 2, pp. 111-122.
16 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
EwIne, H. 1928. The scorpions of the western part of the United States with notes on those occurring in northern Mexico. Proceedings of the United States National Museum, vol. 73, no. 9, pp. 1-24. GertscH, W. J. 1958. Results of the Puritan-American Museum Expedition to western Mexico. 4. The scorpions. American Museum Novitates, no. 1903, pp. 1-20. GertscH, W. J., AND M. SOLEGLAD 1966. The scorpions of the Vejovis boreus group (sub-genus Paruroctonus) in North America (Scorpionida: Vejovidae). American Museum Novitates, no. 2278, pp. 1-54. 1972. Studies of North American scorpions of the genera Uroctonus and Vejovis (Scorpionida, Vejovidae). Bulletin of the American Museum of Natural History, vol. 148, pp. 547-608. lBlpowinon if, ab 1972. Scorpions of the Northern California Coast ranges (Arachnida: Scorpionida). California Academy of Sciences, Occasional Paper, no. 92, pp. 1-59. HOFFMANN, C. C. 1931. Monografias para la entomologia medica de Mexico. Monografia num. 2, los scorpiones de Mexico. Primera parte: Diplocentridae, Chactidae, Vejovidae. Anales del Instituto de Biologica, Universidad de Mexico, vol. 2, no. 4, pp. 291-408. KarscuH, R. 1879. Scorpionologische Beitrage. Ueber scorpione. Mittheilungen des Munchener Entomologischen Vereins, vol. 3, pp. 97-136. KRAEPELIN, K. 1894. Scorpiones und Pedipalpi. Das Tierreich, vol. 8, p. 182. Marx, G. 1887. Remarks on the types of Scorpionidae described by Wood. Proceedings of the Entomological Society of Washington, vol. 1, pp. 90-94. 1889. Arachnida in the scientific results of explorations by the U.S. Fish Commission steamer Albatross. Proceedings of the United States National Museum, vol. 12, pp. 207-211. STAHNKE, H. L. 1968. Some diplocentrid scorpions from Baja California del Sur, Mexico. Proceedings of the California Academy of Sciences, fourth series, vol. 35, no. 14, pp. 273-320. Woop, H. C. 1863a. Descriptions of new species of north American pedipalpi. Proceedings of the Academy of Natural Sciences of Philadelphia, April, pp. 107-112. 1863b. On the pedipalpi of North America. Journal of the Philadelphia Academy of Natural Sciences, series 2, vol. 5, pp. 357-376.
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 2, pp. 17-57; 15 figs.; 1 table. October 30, 1974
THE GENUS CERCIDIUM (LEGUMINOSAE: CAESALPINIOIDEAE) IN THE SONORAN DESERT OF MEXICO AND THE UNITED STATES
By Annetta M. Carter
Herbarium, Department of Botany, 2017 LSB, University of California, Berkeley, California 94720
Cercidium comprises nine taxa, five of which occur in the Sonoran Desert of Mexico and southwestern United States (figs. 1, 2). Of these five, only one, C. praecox (Ruiz and Pavon) Harms, has a widespread distribution, occurring from western and southern Mexico south to Peru. Of the remaining taxa, two occur in Texas and northern Mexico and two in Argentina. Thus, it can be seen that the greatest concentration of species occurs in the Sonoran Desert (Shreve, 1951, 1964).
Cercidiums, with their green branches which at a distance appear leafless much of the year, and are laden from March into June with a profusion of yellow, caesalpinoid flowers, are among the most conspicuous and characteristic trees of the Sonoran Desert. The newcomer to this region tends to refer to all these green-barked trees as palo verde, but if he goes into the field with a paisano, he soon learns that there is not only palo verde, but there are also dipua (dipuga), palo brea, and palo estribo (fig. 3). One other green-branched tree, closely related to Cercidium, also occurs in the Sonoran Desert. It is Parkinsonia aculeata Linnaeus, known locally in Baja California as junco. Continued experience with palos verdes in the field, especially in the Sierra de la Giganta of Baja California Sur, made it evident that they merited special attention in order to understand the relationships within the group.
[17]
18 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
As with so many Sonoran Desert plants, most species of Cercidium present an entirely different aspect in the dry season than they do following the rains. Speaking generally, in the southern part of the Sonoran Desert, the heaviest rains fall in brief, hard-hitting storms during the hot months of July through September, sometimes extending into October (Hastings and Humphrey, 1969). With this moisture most plants come into full leaf, but with the end of the rains, many are soon leafless again. In ‘good’ years, gentle winter rains also come to the southern part of the Sonoran Desert and many species repeat the leaf cycle. Farther north, in south-central Arizona, where the total rainfall is about equally divided between summer and winter, Cercidium usually has two leafy seasons. All of the cercidiums flower at the height of the dry season, March through June, when for the most part, the trees are leafless or nearly so. Sometimes there is minor ‘off-season’ fall flowering. Keys to the taxa are included herein for both the vegetative and the flowering and fruiting stages.
Although no attempt is made here to resolve the long-standing controversy as to whether Cercidium should be regarded as distinct from Parkinsonia, some points brought out in this paper may add fuel to the fire. Watson (1876) argued for union of these two genera; Sargent (1889), principally on the basis of the legume, maintained them as distinct; Johnston (1924a), on other grounds and after some realignment of the taxa within the two genera, also argued for their maintenance as distinct genera; Britton and Rose in 1930 erected the genus Cercidiopsis for Cercidium microphyllum; Brenan (1963), on consideration of additional African taxa which he placed in Parkinsonia, felt that it is not possible to maintain Cercidium as distinct from Parkinsonia, but he refrained from making any nomenclatural changes in the American material. In accordance with Johnston’s delimitation, Cercidium in America is easily distinguished from Parkinsonia and is kept separate in recent floras. The question of generic relationships cannot be resolved without first making extensive comparative studies of these trees (their biology, morphology, cytology, genetics, etc.) from seedling stage to maturity. Lack of evidence for resolution of problems at the generic level, however, does not preclude our attaining a better understanding of relationships at the subgeneric level. For the purposes of this paper, I am following Johnston’s generic delimitation.
The following key summarizes the differences between the species of Cercidium and Parkinsonia aculeata Linnaeus.
Armature comprised of the first leaf developing at a node having a long-persistent, indurate petiole and rachis which terminates in a sharp, stout spine; leaves with pinnae’ 8 to 30 (-60) cm. long, the rachis 1 to 3.6 mm. wide, flattened, phyllodial, persistent; leaflets falling with drought, alternate and/or opposite, 10 to 40 on one
‘As used herein, the term ‘“‘pinnae’” refers to the secondary rachises plus leaflets of a bipinnate leaf. In many representatives of Caesalpinioideae the pinnae are reduced to a single pair.
CARTER: CERCIDIUM 19
Wot cL] =a \ ee eS cre a ae ASS SRE a is Sa ) | i EO ae Pes PaO [5 NE TR RCL Pe ee D SERET EE” REE ETE Cit ao ere ee Se ESS DE aes GEE California | eR ee TEE EE) FS ee ee, SNe ees bys Se Ye ee) 32°— 2a. Cercidium 24° —
E9 C.microphyllum C. praecox
--- C.X sonorae 114° Wo?
FicureE 1. Distribution of Cercidium microphyllum, C. praecox and C. X sonorae in the Sonoran Desert of northwestern Mexico and southwestern United States.
side of a given rachis; axillary leaf-bearing shoots usually 2 to 12 (-23) mm. long;
inflorescences developing with the leaves, the racemes much shorter than the Beeeay sew. ie Parkinsonia aculeata
pinnae; pedicels 10 to 21 mm. long (mean 14.4 mm.) Armature of one or two axillary thorns, or lacking; leaves with pinnae 0.3 to 6.5 cm.
20 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
\ :
Ly ul | nd Ye L = California bs, ey \ BB
Cercidium floridum Y
24°— 1 subsp. floridum (A subsp. peninsulare
Wie
Ficure 2. Distribution of Cercidium floridum subsp. floridum and C. floridum subsp. peninsulare in the Sonoran Desert of northwestern Mexico and southwestern United States.
long, the rachis less than 1 mm. wide, terete or sub-terete, not phyllodial, often falling with drought; leaflets opposite, 2 to 9 (-17) pairs; axillary leaf-bearing shoots so reduced that the leaves appear to arise in the primary leaf axil; inflorescences developing before the leaves or if with them, the racemes subequal to or exceeding the pinnae; pedicels 2.5 to 14 mm. long (mean 7.5 mm.) ......_. species of Cercidium
Vor. XL] CARTER: CERCIDIUM 21
Herbarium material of Cercidium belonging to the following institutions has been studied: British Museum (Natural History), London (BM); California Academy of Sciences, San Francisco, California (CAS); Dudley Herbarium, Stanford University, California (DS); Escuela Nacional de Ciencias Bioldgicas, Instituto Politécnico Nacional, México, D. F. (ENCB); Field Museum of Natural History, Chicago, Illinois (F); Gray Herbarium of Harvard University, Cambridge, Massachusetts (GH); Herbario Nacional del Instituto de Biologia, Universidad Nacional Autonoma de México, México, D. F. (MEXU); Instituto de Botanica, ‘‘Antonio José Cavanilles,’ Madrid (MA); Missouri Botanical Garden, St. Louis (MO); New York Botanical Garden (NY); Rancho Santa Ana Botanic Garden, Claremont, California (RSA); The Herbarium and Library, Royal Botanic Gardens, Kew (K); San Diego Museum of Natural History, California (SD); United States National Museum, Washington, D.C. (US); University of Arizona, Tucson (ARIZ) ; University of California, Berkeley (UC). Abbreviations for these institutions are those given by Lanjouw and Stafleu (1964). Collections made by the author will be widely distributed. Specimens cited include, in addition to those indicating limits of distribution, representative flowering and fruiting specimens as well as all specimens cited in illustrative material. I wish to express my appreciation to the directors and curators of the herbaria at these institutions for enabling me to study their collections.
In citing collections from the peninsula of Baja California, political sub- divisions are indicated as follows: Baja California Sur for the area from 28°N. southward to the tip of the peninsula, a territory of Mexico; Baja California [Norte] for the area from 28°N. northward to the Mexico-United States border, a state of Mexico.
Cercidium Tulasne Cercidium TutLasne, 1844, Arch. Mus. Paris, vol. 4, p. 133.
Trees or large shrubs with smooth greenish bark, the branches usually armed with axillary thorns, or having short, spine-tipped branches. Stipules slender, foliaceous, caducous. Clusters of dry axillary bud scales usually present. Leaves usually not long persistent, sessile or petiolate, bipinnate with one to three pairs of pinnae, the leaflets small, opposite; leaf rachis terete or sub-terete, 1 mm. or less in diameter and terminating in a slender foliaceous or semi-indurate bract. Flowers borne in condensed to open axillary racemes. Pedicels jointed, the joint being in the distal half. Calyx tube green, broadly campanulate, 1.5—2 mm. long, 3-4 mm. wide, the lobes 5, narrowly ovate, valvate or subvalvate, 4-7 mm. long, 2 mm. wide, yellow or greenish yellow. Petals 5, yellow or creamy white, pubescent at base, the upper (posterior) with a conspicuous claw which usually elevates the limb of this petal above the others, the base of the limb auriculate, the auricles often curved over the apex of the claw to form a channel; other four
22 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 3. Comparison of leaves, legumes, and seeds of Cercidium: A, C. microphyllum, the upper figure is a single pinna of a bipinnate, sessile leaf; B, C. X sonorae, the leaves show variation in petiole length from subsessile to short-petiolate (one pinna has fallen from a leaf in 4945); C, C. praecox; D, C. floridum subsp. peninsulare. Numbers refer to Carter, and/or Carter et al. collections; data for these are given under “Representative collections cited” for the respective taxa.
petals with shorter claws. Stamens 10, distinct; filaments pilose near the base; anthers versatile. Pods oblong to linear-oblong, or linear and torulose, coriaceous or papery. Seeds 1 to 4, flattened or subglobose.
Based on Cercidium spinosum Tulasne, 1844, p. 136: ‘““Regionem Amazonum (Bonplandi herb. propr. munc in herb. Mus. Paris), Colombiam prope Maracaibo (Plée, herb. No. 73), nec non prov. Oaxaca Novae Hispaniae propter Tehuacan (in Cordillera alt. 1700 metr. — Galeotti herb. No. 3212) habitat.”
Key TO CERCIDIUM IN THE SONORAN DESERT (Based on vegetative characters)
Branches spinescent at tip; bark yellow-green, horizontally striate on older branches; axillary thorns lacking; leaves sessile, the pinnae of the primary” leaves borne on a scale-like, semi-indurate rachis which is tardily deciduous, the leaflets minute, @'5=Syemmnl lon oo ae ee ce ee 1. C. microphyllum
2The term “primary”? leaf refers herein to the first leaf developing at a node in contrast to those
developing subsequently from a succession of buds produced on a reduced shoot in the axil of the primary leaf or primary leaf scar.
VoL. XL] CARTER: CERCIDIUM 23
Branches not spinescent at tip; axillary thorns usually present; leaves petiolate (sometimes subsessile in C. X sonorae), the leaflets larger, usually 3-12 mm. long. Pinnae usually bearing 4-12 (17) pairs of leaflets.
Nodes bearing one or two stout axillary thorns 2-18 (-—25) mm. long and having conspicuous clusters of dry, dark bud scales in the axils; bark waxy-coated on older branches and having a fine, quadrate-pustulate pattern; petioles Able TX Taal © Tn spe ee ere a ee Be Da Cra PiaeGox
Nodes bearing a single slender axillary thorn 2-11 (-20 mm.) long (or sometimes none); dry bud scales present at nodes but usually not conspicuous; bark on older branches smooth or inconspicuously horizontally-striate, the waxy coat inconspicuous or lacking; petioles O-3 (—12) mm. long __._ Sei Gu xe SOMOTCE
Pinnae usually bearing 2-4 pairs of leaflets; nodes without conspicuous clusters of dark axillary bud scales; bark of older branches horizontally striate _ 4. C. floridum
Branchlets of mature trees glabrous to glabrate; leaflets 4-8 mm. long, mostly 3 pairs pew pinna (Sonoran! Desert except Baja California), 2 Sil oy 1s ee eg Bee oe ee re i ee ee Ee 4a. C. floridum subsp. floridum
Branchlets villous or pilose, leaflets 6-15 mm. long, mostly 2 pairs per pinna; clusters of reddish brown glandular hairs in axil of primary leaves (confined to_southern Baya, California) 22 4b. C. floridum subsp. peninsulare
Key TO CERCIDIUM IN THE SONORAN DESERT (Based on flowering and fruiting characters)
Ovary glabrous at anthesis; legumes not strongly long-tapered at ends.
Inflorescences borne in clusters along older branches; petals deep yellow, the upper often orange-dotted near base of limb; mature legumes papery, flat, conspicuously net-veined; seeds flattened, gray-brown with brown mottling. 2. C. praecox
Inflorescences on terminal or subterminal branches; legumes when immature bearing conspicuous white dots (stomata) but these obscure in mature legumes, coriaceous, the veins inconspicuous; seeds oblong to ovate, flattened, brown with marginal area lighter and sometimes faintly mottled. 4. C. floridum
Inflorescences open, the racemes with rachis 1.0-4.5 (-7.0) cm. long, pedicels 6-12 (—20) mm. long; petals bright yellow, the upper sometimes orange- dotteda(Sonoranms Desert, except) Bajals California) y= ee ee eee 2 FeO A Se oe ee 4a. C. floridum subsp. floridum
Inflorescences somewhat congested, the racemes with rachis mostly 0.3-1.0 (—2.0) cm. long; pedicels 4-9 (-12) mm. long; petals deep or bright yellow, the upper not orange-dotted (confined to southern Baja California). eA 2 Awa Se ETA CO LOT UMUESUDS Dan DCNINSULANE
Ovary lightly pubescent to strigose at anthesis; legumes strongly tapered at both ends.
Legumes torulose, lightly striate-veined; petals light yellow® except for the upper one which is white or creamy-yellow but never orange-dotted; seeds sub-globose, DO We ee ee eee te ee ere ee ee ee 1. C. microphyllum
Legumes sinuate, conspicuously striate-veined; petals light yellow® except for the upper one which is variable (whitish or pale yellow and sometimes orange- dotted) ; seeds oblong to ovate, flattened, dark mottled brown. _... 3. C. X sonorae
3 Petal color is useful only with fresh material; use vegetative key to distinguish these two species if only dry material is available.
24 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
1. Cercidium microphyllum (Torrey) Rose and Johnston.
Cercidium microphyllum (Torrey) Roser AND JoHNsTOoN in I. M. JoHNston, 1924, Contr. Gray Herb., vol. 70, p. 66.
Parkinsonia microphylla Torrey, 1857, Pac. R. R. Rep., vol. 4, p. 82; 1859, Bot. Mex. Bound, p. 59.
Cercidiopsis microphylla BRITTON AND Rose, 1930, N. Amer. FI., vol. 23, p. 306.
Shrub or tree 3—4 m. tall and nearly as broad, occasionally up to 8 m., the branches usually ascending and often presenting a broom-like appearance; bark yellow-green with fine horizontal striations; branches pubescent (strongly strigose in Baja California), puberulent or pilose to glabrate, the branchlets rigidly divaricate, spinose-tipped; axillary thorns lacking, the nodes bearing few to many bud scales; leaves 1-2 per node, sessile [i.e. no petiole evident on the common rachis except in seedlings (fig. 7b) ], the single pair of pinnae (3-65 mm. long) of the primary leaves borne on a scale-like, indurate rachis which is some- times tardily deciduous after the pinnae fall; leaflets 1-7 (—13) pairs, broadly elliptical, obtuse to emarginate at apex, 0.5—5 mm. long, 0.75—1.5 (—2) mm. broad, puberulent when young; rachis of the raceme 0.2—4.5 cm. long, puberulent, bearing 1-6 (—10) flowers on pedicels 4-8 (—15) mm. long; calyx pubescent to glabrate; petals yellow except for the upper (posterior) whose limb is usually white but occasionally creamy or pale yellow; upper petal 7-10 mm. long, the limb 4—5 mm. long, 3.5—6.0 mm. wide, rhomboidal or broadly ovate, the claw 3—4 mm. long, subequal to the limb in length, the other four petals slightly shorter than the upper, the limb rhomboidal or lanceolate, the margins not overlapping one another at anthesis, the claw short, 1-2 mm. long; ovary at anthesis strigose to puberulent; legumes 3.5-11.0 cm. long, 7-9 mm. wide, strongly tapered basally and apically, torulose, circular in cross-section where the seeds are borne, longi- tudinally striate, not coriaceous, dehiscence irregular, the valves often breaking as well as the sutures opening; seeds 1-4, brown, sub-globose, 8 (—10) mm. long, 5—7 mm. wide.
Type. Mexican Boundary Survey: Diluvial banks of the Colorado, Ft. Yuma, 13 January 1854. A. F. Schott (NY, holotype; F!, date of 1855 is in error) [flowering specimen]; Williams’ River [Arizona], 12-22 February [1854], J. M. Bigelow (N.Y.) [in fruit].
REPRESENTATIVE SPECIMENS. MEXICO. GULF OF CALIFORNIA ISLANDS, BaJA CALIFORNIA Sur: S. end of Amortajada Bay, San José Island, 11 April 1952, R. Moran 3772 (DS, UC); Puerto Balandra, Isla Carmen, 23 March 1971, J. R. Hastings 71-131 (ARIZ, SD); Danzante Island, 7 April 1962, Moran 9228 (CAS, SD); Arroyo de los Chivos, NE. side of San Marcos Island, 29 March 1962, Moran 8981 (CAS, RSA, SD). Baja Carirornta [Norte]: main arroyo, San Esteban Island, 26 April 1966, Moran 13054 (DS); ca. % mi. S. of Refugio
VoL. XL] CARTER: CERCIDIUM 25
Bay, Isla Angel de la Guarda, 26 March 1963, Moran 10427 (DS, UC). Sonora: Tiburon Island, 5 May 1952, Moran 4064 (DS, SD, UC, US), 23 April 1966, Moran 12992 (ARIZ, CAS, RSA, SD).
PENINSULA, BAJA CALIFORNIA SuR: Rancho El Salto, Arroyo Coyote (ca. 24°47'N., 110°50'W.), altitude 525 m., 5 November 1971, Moran 19020 (14 mi. [22.4 km.] (by road) W. of San Luis Gonzaga, 21 October 1964, Turner & Hastings 64-380 (ARIZ); Agua Verde, 1 April 1911, J. N. Rose 16578 (US); Portus Escondido, 2 February 1842, Wosnessensky (GH); 55 km. E. of Villa Insurgentes on highway to Loreto, 3 May 1972, Carter 5670; 1 km. W. of Las Parras summit, road from Loreto to San Javier, 6 May 1972, Carter 5672; Rancho La Venta, ca. 16 km. westerly from Loreto on road to San Javier, 21 April 1962, Carter 4415; Rancho Aguajito, Arroyo Gua between Loreto and Rancho Sauce, 24 April 1955, Carter & Ferris 3446; Arroyo Gua, N. of Loreto, 7 November 1960, Carter 4110; Rancho Naucajoa (26°16’N., 111°36.5’W.), W. of Llanos de San Juan, Carter & Reese 4531; Coyote Bay [Cove], Concepcion Bay, 18 June 1921, Johnston 4172 (CAS, F, K, MO, UC, US); wash 25 mi. [40 km.] S. of San Ignacio, 19 April 1931, Wiggins 5434 (CAS, DS, GH, RSA); eastern bajada of Sierra Calvario, Systema de Sierra Viscaino, 10-15 March 1947, H.S. Gentry 7501 (DS). Baja CatirorniA [Norte]: ca. 5 mi. [8 km.] N. of Mision de San Borja along road to Bahia de Los Angeles, 17 May 1959, /. L. & D. B. Wiggins 14857 (CAS, DS); Cajén de Santa Maria, 12 May 1889, T. S. Brandegee s.n. (DS, UC); San Luis Gonzales [Gonzaga] Bay, 29 April 1921, Johnston 3348 (CAS, US); San Felipe Desert between El Cajon and Algodones, along eastern foot of Sierra San Pedro Martir, altitude 2500 ft., [800 m.], 10 May 1941, Wiggins 9845 (DS, UC, US). Sonora: island in harbor, Guaymas, 14 April 1921, Johnston 3084 (CAS, US); hills NW. of shrimp cannery, Guaymas, 6 April 1962, Carter 4363; 26 km. S. of Hermosillo on Guaymas road, 21 March 1934, Ferris 8759A (DS); between Hermosillo and Kino Bay, 21 December 1968, V. Rudd 3038, with P. Bauer & A. C. Fox (ARIZ, SD, US); Sonora Alta, 1830, Coulter 490 (K, 2 sheets, one lacking collection number) ; N. of bay, within 5 mi. [8 km.] of coast, vicinity of Libertad, 2 May 1928, E. H. Graham 3822 (DS); 7 mi. [11 km.] S. of Altar, 4 May 1928, Graham 3906 (DS); Pasa de San Luis [Poso de Luis on US sheet], 4 June 1894, E. A. Mearns 2697 |Int. Bound. Commission] (DS, US).
UNITED STATES. Arizona: Tumamoc Hill, Tucson Mts., Pima Co., 8 June 1938, H. S. Gentry 3777 (CAS); 5 mi. [8 km.] S. of Florence, Pinal Co., 26 July 1927, H. W. Graham s.n. (DS); Tempe Butte, altitude ca. 1300 ft. [415 m.], Maricopa Co., 4 May 1952, E. P. Killip 42150 (US); Government Springs, 3.5 mi. [5.6 km.] N. of Bumblebee, State Highway 69, Yavapai Co., 3 July 1940, Ferris 9902 (CAS, DS, GH, RSA, UC); 14 mi. [22 km.] NE. of Topock,
26 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Mohave Co., 10 June 1967, Carter 5243. CALIFORNIA: 1 mi. [1.6 km.] S. of Copper Basin Lake, Whipple Mts., San Bernardino Co., 22 April 1940, A. M. Alexander & L. Kellogg 1200 (DS, GH, RSA, US); base of Whipple Mts. adjacent to Colorado River 11 mi. |18 km.] above Earp on road to Parker Dam, San Bernardino Co., 15 March 1940, Wolf 9721 (RSA).
Coarse soil of plains and hillslopes, mostly below 600 m., but also between 700 and 800 m. in north central Sonora and southeastern Arizona. The trees leaf out following the rainy seasons, but the leaves are ephemeral. Flowering is mostly from March through May, the height of the Sonoran Desert dry season. Leaves, if present during flowering, are confined to non-floriferous branches. This is the most widespread of the taxa in Baja California where it is known as dipua and extends from a little south of Latitude 25°N. northward to the United States border. It is also abundant in west central Sonora and southwestern Arizona where it extends to Latitude 35°N.
Even when leafless, the branches are an important source of food for stock. Trees with their upper branches lopped off by machetes indicate that a traveller rested nearby and provided food for his mule, burro, or horse. In times of drought, the branches are also cut to feed cattle. The seeds are edible.
As an historical aside, it is interesting to note that Thomas Coulter collected both Cercidium microphyllum and C. floridum in “Sonora Alta” [vicinity of Hermosillo, Sonora] in 1830. According to the correspondence between Harvey and Bentham [Kew archives], some years after the packages of Coulter’s col- lections were received at Trinity College, Dublin, Harvey separated the legumes and sent them to Bentham at Kew for identification. Specimens of both taxa are annotated in Bentham’s hand as new species. At about that time, Asa Gray visited Kew, presumably saw all of the Coulter material, and subsequently published Cercidium floridum Benth. ex Gray. Five years later, John Torrey, in describing material collected on the Mexican Boundary Survey, named Parkinsonia microphylla, chose a Schott specimen as type, and made no mention of the much earlier Coulter collection to which Bentham had applied the same specific epithet.
The following collections are putative hybrids between Cercidium micro- phyllum and C. floridum subsp. floridum: P. Kamb 2014. Bottom of NE. caldera of Molina Crater, altitude ca. 950 ft. [310 m.], crater region NW. of Sierra Pinacate, Sonora, Mexico, 29 April 1951 (DS, UC). The leaves are petiolate. The leaflets are larger than typical for C. microphyllum and there are too many leaflets for it to be C. floridum. It has the stem pubescence and the strigose ovary of C. microphyllum and thorns similar to those of C. floridum. On both of the above cited sheets of this collection, the flowering branch is the putative hybrid and the separate fruiting branch—presumably from a different tree—is typical C. floridum subsp. floridum. Both of the putative parents are
Vor. XL] CARTER: CERCIDIUM 27
known to occur in this area (Hastings, Turner and Warren, 1972). A sheet at University of Arizona bears only a fruiting branch and this is typical C. floridum subsp. floridum.
C. B. Wolf 9722. Base of Whipple Mts., adjacent to Colorado River, 11 mi. [18 km.] above Earp on road to Parker Dam, altitude ca. 400 ft. [130 m.], San Bernardino Co., California, 15 July 1940 (DS, RSA). Collector’s note: “This lone tree looks like a typical C. microphyllum in olive-green color and shape, but the pods are rich brown, somewhat flattened, and not constricted between the seeds, which are larger and flattened. Leaflets somewhat larger than typical C. microphyllum and C. floridum. Both species grow here in abundance.” The valves of the pods are coriaceous, striate, and white-dotted. The stems are sparsely appressed-pubescent and bear short spines.
Papers treating these and other putative hybrids in Cercidium are published elsewhere (Carter, 1974; Carter and Rem, 1974).
2. Cercidium praecox (Ruiz and Pavon) Harms. (Figure 4.)
Cercidium praecox (Ruiz and Pavon) Harms, 1908, Bot. Jahrb., vol. 42, p. 91.
Caesalpinia [Sappania] praecox Ruiz anD PAvon, 1802, Fl. Peruv., vol. 4, pl. 376, plate only [Entire volume published by Consejo Superior de Investigaciones Cientificas, Madrid, 1957].
Caesalpinia praecox Ruiz AND PAvoNn, 1833, in HOOKER AND ARNOTT, Bot. Misc., vol. 3, p. 208. Cercidium spinosum TULASNE, 1844, Arch. Mus. Paris, vol. 4, p. 134.
Rhetinophloem viride KArSTEN, 1862, Fl. Columb., vol. 2, p. 25, pl. 113.
Cercidium viride KARSTEN, 1887, Bot. Jahrb., vol. 8, p. 346.
Cercidium plurifoliolatum M. MicHett, 1903, Mém. Soc. Phys. et Hist. Nat. Geneve, vol. 34, p. 269, pl. 18.
Cercidium unijuga Roser, 1905, Contrib. U. S. Nat. Herb., vol. 8, p. 301, 1905.
Cercidium Goldmanii Roser, 1905, Contrib. U.S. Nat. Herb., vol. 8, p. 301. [Goldman 735, type (US!): leaves glabrous; waxy coating of stems only slightly visible. |
Shrub or tree usually 2-4 m. tall, but up to 9 m. in forested areas of north- eastern Sonora, the crown usually rounded, or flat-topped and spreading in exposed habitats, erect and less branched in sheltered habitats; branches and trunk bright green to the base, the bark with a minute quadrate-pustulate pattern and bearing a heavy coating of wax, the branchlets glabrate to pubescent, the hairs usually appressed; axillary bud scales prominent; thorns axillary, stout, usually one but sometimes two per node, 2-18 (—25) mm. long (mean 8.6 mm.), often dark brown; leaves 1-3 (—6) per node, pubescent, the petiole (1—) 4-11 (—21) mm. long, bearing 1 (occasionally 2-3) pair of pinnae 0.44.5 cm. (—S.0, Wiggins 6473) cm. long; leaflets (3—) 5-9 (-17, Wiggins 6473) pairs, oblong, rounded at apex, 3-10 mm. long, 1.4-3.8 mm. wide; inflorescences borne in
28 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
clusters along mature branches, usually compact, 1-3 (4) per node, the rachis (1.5—) 2-11 (—15) mm. long, sparingly pilose, bearing 1-6 (—9) flowers on pedicels 7-10 mm. long; petals deep yellow, the upper often orange-dotted near base of limb; upper petal 9-11 mm. long, the limb 6-7 mm. long, 6-8 mm. wide, broadly ovate, the claw 3-5 mm. long, shorter than the limb, the other four petals slightly shorter than the upper, the limb broadly ovate and sometimes auriculate at base, the claw 1-2 mm. long; ovary glabrous at anthesis; legumes 3-6 (—8) cm. long, 0.6-1.0 cm. wide, flat and papery, not narrowed between the seeds, the veins conspicuous, forming an elongate-reticulate pattern; seeds 1—2 per pod, oblong, flattened, gray-brown with dark brown mottling, up to 1 cm. long, 3—4 mm. wide.
Type. Middle western Peru. Ruiz and Pavon plate. [cf. Johnston, 1924, p. 67.| (Ruiz and Pavon collections, MA!).
REPRESENTATIVE SPECIMENS. MEXICO. GULF OF CALIFORNIA ISLANDS, BAJA CALIFORNIA SuR: summit, Ildefonso Island (26°37'N., 111°27’W.), 17 May 1921, Johnston 3753 (CAS), 2 April 1962, Moran 9066 (RSA, SD, US); Tortuga Island (27°26N., 111°54’W.), 24 April 1952, Moran 4007 (DS), 30 March 1962, Moran 9016 (SD), 11 May 1921, Johnston 3592 (CAS, GH, K, UC, US). PENninsuta, Baja CALIFORNIA SuR: San José del Cabo, 26 March 1911, J. N. Rose 14466 (US); La Paz, 10 August 1944, Maximino Martinez s.n.
VoL. XL] CARTER: CERCIDIUM 29
(US); along dry washes between Médano and Venancio, 29 April 1931, Wiggins 5532 (DS, RSA, UC, US); between Rancho Segundo Paso and San Javier, altitude 300 m., 21 April 1962, Carter 4412; between Canon de Las Calaveras and La Tinaja, western side of Mesa de San Alejo, altitude 690 m. (ca. 25°51'N., 111°36’W.), 11 November 1961, Carter 4306; along old mission trail SE. of Comondu, altitude 420 m., 20 April 1955, Carter & Ferris 3424 (ARIZ, DS, GH, SD, UC, US); Cuesta de Los Encinos, SE. of Cerro Giganta, altitude 500 m., 29 March 1960, Carter & Ferris 4046 (UC), same tree, 9 November 1960, Carter 4146 (UC); 4 mi. [6 km.]| (by road) E. of San Lina (suburb of San Ignacio), 29 October 1963, Turner & Hastings 63-294 (ARIZ, DS, SD). Stnatoa: La Constancia, Munic. El Fuerte, December 1924 [sic! on original label at DS; US copied label is 1926], Jesus Gonzales Ortega 6200 (DS, GH, MEXU, US). Sonora: Alamos study area, 7.5 mi. [12 km.] W. of Alamos, 28 April 1967, R. D. Krizman 16 (ARIZ); Agua Caliente N. of Alamos, 2 November 1939, Gentry 4839 (ARIZ, MO, distributed as C. torreyanum and so cited by Gentry, 1942, p. 131); San Bernardo, Rio Mayo, 1 March 1935, Gentry 1377 (ARIZ, GH, MEXU, UC, distributed as C. torreyanum and so cited by Gentry, 1942, p. 131); Cerro de Bayajori, 12 mi. [19 km.] W. of Navajoa, 11 April 1948, Gentry 7947 (UC, US, distributed as C. floridum) ; island in Bay, Guaymas, 14 April 1921, Johnston 3078 (CAS, US); low hills and flats near tannery E. of Guaymas, 28 February 1933, Wiggins 6348 (DS, RSA, US); small valley 20 mi. [32 km.] N. of Guaymas, 8 March 1933, Wiggins 6473 [spines up to 30 mm. long, pinnae 45-50 mm. long, leaflets up to 15 pairs] (DS); 20 km. S. of Carbo junction, 3 May 1971, Carter, Hastings & Turner 5597; Horcasitas, 17 April 1932, Abrams 13360 (DS); 2 mi. [3 km.] S. of Los Hoyos, altitude 810 m., 23 April 1971, Carter, Hastings & Turner 5576; 4.2 km. S. of Los Hoyos, 4 July 1971, Hastings 71-199 (SD); Colonia Oaxaca, 24 July 1938, Stephen S. White 663 (ARIZ, GH).
Ranging from coastal plains up to bajadas, mesas, hills, and mountains at elevations up to 825 m. in Baja California and 1115 m. in the mountains of northeastern Sonora. Flowering is from March through May with the peak in April, before the leaves develop.
In Baja California, C. praecox extends from near the tip of the peninsula (23°N.) northward to the vicinity of San Ignacio (27° 25’ N.). On the main- land, it occurs in southern Mexico and is abundant in Sonora from a little south of Guaymas (27° 56’ N.) almost to the United States border (30° 54 N.). Occasional collections have been made as far south as 26°N. in Sinaloa. This is the only one of our Sonoran Desert species of Cercidium having a disjunct distribution. According to Johnston (1924a), in South America it occurs from extreme middle-western Peru to northern Venezuela.
30 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
During the months when the trees are leafless, C. praecox may be recognized readily by the bright green bark extending down to ground level, and the conspicuous accumulation of dark bud scales at the nodes. Development of new branch growth appears to be limited, and most seasonal growth (production of leaves and flowers) is axillary year after year on the old branches. In addition, the stout, often dark brown spines, which may reach a length of 2.6 cm., set it apart from the other taxa. It is the only one of our Cercidium species with bark having a quadrate-pustulate pattern. When in flower, the species may be recognized easily by the usually deep yellow flowers borne in close clusters along the length of the old branches.
The common name, palo brea, is derived from the fact that the waxy substance coating the bark, after being scraped from the branches and melted by heat, is used as a ‘gum’ for gluing together leather objects and furniture; thus it is used just like /a verdadera brea.
The following Mexican collections are putative hybrids between Cercidium praecox and Parkinsonia aculeata: edge of town, Coyuca, Mina, Guerrero, 11 May 1934, G. B. Hinton 6040 (BM, NY), 25 March 1937, Hinton 9968; (ARIZ, BM, K, MEXU, NY, RSA, TEX); near Los Hoyos, northeastern Sonora, 23 April 1971, Carter, Hastings & Turner 5575 (to be distributed). This material is discussed in a separate paper (Carter and Rem, 1974).
3. Cercidium X sonorae Rose and Johnston. (Figure 5.)
Cercidium X sonorae ROSE AND JOHNSTON, 1924 (pro sp.) stat. nov., zz I. M. Johnston, Contr. Gray Herb., vol. 70, p. 66. (April). [C. microphyllum (Torrey) Rose and Johnston x C. praecox (Ruiz and Pavon) Harms].
Cercidium molle I. M. JounsTon, 1924, Proc. Calif. Acad. Sci., IV, vol. 12, p. 1038. (May).
Spreading tree 4-8 m. tall, usually with lax branches (but sometimes the branches short and stiff), the bark smooth or sometimes faintly horizontally striate, green to yellow-green, sometimes with an inconspicuous, thin, waxy coating, branches short-villous, pilose, or sometimes glabrate; axillary thorns present or lacking, slender (2—) 5-11 (—20) mm. long (mean 6.5 mm.), variable as to presence on a given tree; bud scale clusters prominent to inconspicuous; foliage bright yellow-green, the leaves 1 to 3 per node, the petioles usually 1-3 mm. long, but varying from 0 to 12 mm.; pinnae 1 to 2 pairs, 0.7—5.0 cm. long, the leaflets (3—) 6-8 (-12) pairs elliptical-oblong, (1.5—) 2-3 (-—6.0) mm. long, (1.0—) 1.5 (—3.0) mm. wide; inflorescence usually open, the rachis of the racemes 0.5—4.0 cm. long, bearing 1 to 10 flowers on pedicels 6-10 (—14) mm. long; petals light yellow, except for the upper whose limb may be whitish, creamy, or creamy and yellow and often orange-dotted near the base of the limb; upper petal 10-12 mm. long, the limb 5—7 mm. long, 5-8 mm. wide, broadly ovate but with
VoL. XL] CARTER: CERCIDIUM 31
pointed apex, the claw 4-5 mm. long, shorter than the limb; the other four petals slightly shorter than the upper, the limb ovate, the claw short, 1.5-3 mm. long; ovary strigose at anthesis; legumes 3-8 cm. long, 7-10 mm. wide, slightly flattened and somewhat coriaceous, sinuate if more than one-seeded, long- tapering to each end, the surface with conspicuous longitudinal striations, dehiscing along the sutures or irregularly; seeds 1 to 2 (4), oblong to oblong- ovate or ovoid, dark mottled brown, flattened or thick, 9-10 (—12) mm. long, 4-5 mm. wide.
Type. Dry hills in the vicinity of Guaymas, Sonora, Mexico, 1910, Rose, Standley & Russell 12586 (US!).
REPRESENTATIVE SPECIMENS. MEXICO. BajA CALIFORNIA SuR: Arroyo San Ramon just W. of Rancho San Ramon (25°14.5’N., 111°17'W.), 21 October 1964, Carter 4818; Rancho Tasajera, ca. 3.5 km. NE. of San José de Agua Verde, 3 June 1965, Carter & Sharsmith 4936; Agua Verde Bay, 26 May 1921, Johnston 3877 (CAS!, type of C. molle, DS, GH, K, MO, UC, US); Puerto Agua Verde [Bahia Agua Verde], 5 June 1965, Carter & Sharsmith 4945, 23 August 1971, Carter 5610; Mision San Javier, 36 km. SW. of Loreto, 6 May 1972, Carter 5679; Arroyo Ranchito, ca. 9 km. SE. of Llanos de San Julio on road from San Javier to Comondti (26°02’N., 111°39'W.), 5 June 1963, Carter & Reese 4554; Cuesta de Los Encinos, SE. of Cerro Giganta, altitude ca. 500 m.,
32 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
29 March 1960, Carter & Ferris 4045, and from same tree 9 November 1960, Carter 4145; La Higuera, NE. base of Cerro Giganta, 18 October 1966, Carter & Sousa 5212;Arroyo Hondo, N. side of Cerro Giganta, altitude 540 m., 17 October 1966, Carter & Sousa 5199; Tortuga Island, 11 May 1921, Johnston 4409 (CAS, K, UC, US), 16 mi. [26 km.] from San Ignacio on road to Santa Rosalia, 10 March 1934, Ferris 8626 (US); 10.5 [17 km.] mi. ENE. of San Ignacio, 27.4°N., 112.8°W.., altitude 170 m., 17 October 1971, Turner & Hastings 71-134 (ARIZ). Sonora: N. of Guaymas (0.6 mi. [.96 km.] N. of junction with Bahia San Carlos road), 2 May 1971, Carter, Hastings & Turner 5595; 25 mi. [40 km.] N. of Guaymas, 2 April 1935, Shreve 7310 (MO; sheets at ARIZ and F bear both C. X sonorae and C. praecox); 31 mi. [50 km.] N. of Guaymas, 2 April 1935, Shreve 7313 (ARIZ, F, MO); El Pozo, 26 km. S. of Hermosillo on road to Guaymas, 21 March 1934, Ferris 8761 (DS, US); Torres, 10 February 1903, F. V. Coville 1664 (US); Sierra Lopez Rancho [37 mi., 59 km. NW. of Hermosillo], 13 April 1932, Abrams 13327 (DS,F).
Plains and hillslopes below 600 m. Flowering March to June with the peak in April; leaves usually present when trees in flower.
In Baja California, where it is known as palo estribo, Cercidium X sonorae occurs principally in the Sierra de la Giganta from about 25°15’N. (opposite the northern end of Isla San José) northward to the northern slopes of Cerro Giganta (26°10'N.). One outlying collection (Turner & Hastings 71-134) is from east of San Ignacio (27.4°N.). In Sonora, C. X sonorae is most abundant from near Guaymas northward to the vicinity of Carbé (i.e., from about 28° to 30° N.). On neither side of the Gulf of California is C. X sonorae as abundant as C. microphyllum and C. praecox, one or the other or both of which appear always to occur in the vicinity of trees of C. X sonorae. Throughout its range, but especially noticeable in Baja California, is the fact that C. X sonorae occurs only within the limits of the distributional overlap of C. microphyllum and C. praecox. In char- acters of bark surface, spininess, pubescence, flower color, and seed shape, C. X sonorae appears to be more variable than any of the other taxa in the group. In several morphological characters, C. X sonorae falls between C. praecox and C. microphyllum; furthermore, pollen of C. X sonorae stained with ‘cotton blue’ (aniline-blue-lactophenol) shows a much lower percentage of presumably viable grains than does the pollen of either of its putative parents. These facts lead to the hypothesis that C. X sonorae is of hybrid origin, the putative parents being C. microphyllum and C. praecox. Discussion of the hybrid origin of C. X sonorae and of other hybrids involving species of Cercidium is published else- where (Carter, 1974; Carter & Rem, 1974).
In 1965 while on a field trip to Agua Verde Bay in search of Johnston’s Cercidium molle, I was told that a large palo estribo tree comparing favorably to that described by Johnston (1924b) had been cut down some twenty years
Vor. XL] CARTER: CERCIDIUM 33
before. On a subsequent trip I explored the ‘huge amphitheater-like canyon” (Johnston, 1924b, p. 1056) south of the puerto of Agua Verde Bay and found only a few trees of palo estribo. Collections from one of these (Carter 5610) closely approach the type specimen of C. molle in its lack of spines, its pubescence, flower size, and leaflet size, but differ in the pinnae being shorter, and the leaflets more closely spaced than in the type. Considering the amount of variation occurring in C. X sonorae, however, C. molle falls well within the limits of that taxon. Had Johnston been able to explore further in the Sierra de la Giganta (instead of being limited to short sallies from aboard ship), he undoubtedly would have encountered trees of palo estribo similar to that which he described from across the Gulf near Guaymas as C. sonorae and would have realized the close affinity of the trees as they occur on both sides of the Gulf.
In his excellent discussion of Cercidium species in the “Vegetation of the Sonoran Desert” Shreve (1951, p. 145; 1964, p. 153) unfortunately confused C. sonore with the much more abundant C. praecox. The description and distribution given under the heading ‘“Cercidium sonorae” and the common name brea apply to C. praecox.
The strong, tough wood of Cercidium X sonorae is used to make stirrups, hence the name, palo estribo.
4. Cercidium floridum Bentham ex Gray. [Synonymy and references given under subspecies. |
Trees or shrubs 2.5—8.0 (—12) m. tall, bark of main branches above main trunk green, horizontally striate; leaves 1 or 2 per node, petiolate; pinnae 1 (occasionally 2) pair, bearing 2—4 pairs of leaflets; flowers yellow, the petals clawed, the claw of the upper petal longer than those of the other four petals; ovary glabrous at anthesis (occasionally a very few hairs present); legumes linear-oblong, flat, cuneate at base and apex, coriaceous, white-dotted (stomata) when immature, obscurely so when mature, inconspicuously reticulately-veined ; seeds oblong to ovate, somewhat flattened, brown, the lighter margin sometimes slightly mottled.
4a. Cercidium floridum Bentham ex Gray subsp. floridum.
Cercidium floridum BENTHAM ex GRAY, 1852, Plantae Wrightianae, vol. 1, p. 58, subsp. floridum.
Parkinsonia florida Watson, 1876, Proc. Amer. Acad., vol. 2, p. 135.
Parkinsonia torreyana Watson, 1876, Proc. Amer. Acad., vol. 2, p. 135.
Cercidium torreyanum SARGENT, 1889, Garden & Forest, vol. 2, p. 388.
Tree 4-8 (-12) m. tall, the crown spreading, branchlets of mature trees usually slender, flexuous or drooping, the trunk gray-green; bark with fine horizontal striations, often dark in the indentations; branches glabrous or
34 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
sparingly pubescent; thorns absent or 1.8—-5.4 mm. long (mean 3.6 mm.), slender, solitary; bud scales not accumulating at the nodes to form conspicuous dark clusters; foliage blue-green; leaves with petiole (1—) 2—5 (—11) mm. long bearing one pair of pinnae (occasionally more) 0.3-1.0 (—1.6) cm. long; leaflets mostly 3 pairs per pinnae, but often 2 and occasionally 4 pairs, oblong or obovate, sometimes emarginate, 4-8 mm. long, 2-5 mm. wide, slightly pubescent or glabrate; inflorescence open, glabrous or nearly so, borne on the younger branches; rachis of the racemes 1.0—4.5 (—7.0) cm. long, bearing 1-7 (-10) flowers on pedicels 6-12 (—20) mm. long; petals bright yellow, the upper sometimes orange-dotted near base of limb, the upper petal 9-15 mm. long, the limb 5—9 mm. long, 6-13 mm. wide, broadly ovate, cordate at base, the claw 3-5 mm. long, shorter than the limb, the other four petals slightly shorter than the upper, the limb broadly ovate, the claw short, 1.5—-2.0 mm. long; legumes linear to elliptic, not or only slightly narrowed between the seeds, 3-11 cm. long, 1.0—-1.5 cm. wide; seeds usually 3 per legume, 9-12 mm. long, 5—7 mm. wide.
Type. “Sonora Alta,” Mexico in 1830, Thomas Coulter (Trinity College, Dublin, Ireland; also 2 duplicate specimens bearing name in Bentham’s hand, K! )
The nomenclatural problem concerning Cercidium floridum and C. torrey- anum is ably discussed by Johnston (1924a) and Benson (1940).
REPRESENTATIVE SPECIMENS. MEXICO. Srnatoa: La Constancia, Munic. El Fuerte, December 1924 |sic! on original label at DS; US copied label is 1926], Jesus Gonzalez Ortega 6199 (DS, GH, MEXU, US); vicinity of San Blas, 22 March 1910, Rose, Standley & Russell 13203 (GH, US), 28 January 1927, M. E. Jones 23086 (RSA); SonorRA: vicinity of Guaymas, 23 April 1910, Rose, Standley & Russell 15038 (US); plain N. of Empalme, 6 April 1962, Carter 4359; 20 mi. [32 km.] from Guaymas on Hermosillo road, 20 March 1934, Ferris 8752 (DS, US); 3.5 mi. [5.6 km.] (by road) N. of Desemboque, 29 April 1964, Turner & Hastings 64-49 (ARIZ, DS, SD); El Alamo, near Magdalena, 25 May 1925, P. B. Kennedy 7010a (UC, US); W. side of Isla Tibur6én just N. of Punta Willard, 19 March 1962, Wiggins 17150 (DS); Isla Tiburon, 19 March 1962, Moran 8723 (RSA, SD), desértico espinosa, altitude 60 m., 3 May 1971, C. L. Diaz Luna 2266 (ENCB); Molina Crater, crater region NW. of Pinacate Crater, 29 April 1951, Kamb 2014 (ARIZ). |The sheets of this collection number at UC and DS are mixed collections, the branch bearing only flowers being a putative hybrid between Cercidium microphyllum and C. floridum.| Papago Tanks, Pinacate region, 15 March 1959, R. M. Turner 59-30 & C. H. Lowe 2979 (ARIZ, CAS); Pozo de Luis, 4 January 1894, Mearns 2696 [Int. Bound. Commission] (US). UNITED STATES. Arizona: Sabino Canyon, Santa Catalina Mountains, Pima Co., 25 May 1917, Shreve 5202 (ARIZ); vicinity of Coolidge Dam, 13 May 1935, Bassett Maguire 11321 (UC); Black Canon Road, 23 mi. |37 km.| N. of Phoenix, Maricopa Co., 21 October 1931,
VoL. XL] CARTER: CERCIDIUM 35
John W. Gillespie 8657 [2 sheets, one fl., one fr.] (DS, GH, UC, US); washes W. of Castle Dome Mts., Yuma Co., 22 April 1949, J. H. Thomas 396 (DS); Topock, Mohave Co., 24 May 1919, A. Eastwood 8886, 8887 (CAS, GH), 29 May 1950, J. T. Howell 26618 (ARIZ, CAS). CALIFORNIA: Whipple’s Expl. R. R. route, 35th parallel, 1853-54, Bigelow (K); 20 mi. [32 km.] NE. of Ogilby, Imperial Co., 6 April 1932, Munz & Hitchcock 12166 (F, GH, MO, RSA, UC); Box Canyon at Shafer’s Well, Chocolate Mts., Riverside Co., 14 June 1918, Ferris 977 (CAS, DS); 10 mi. [16 km.] W. of Coachella, Riverside Co., April 1905, H. M. Hall 5784 (ARIZ, UC, US); Palm Springs (Agua Caliente), desert base of San Jacinto Mt., Riverside Co., 4-13 April 1896, S. B. Parish 4115 (BM, GH, kK, MO, US); 4 mi. [6 km.] NW. of Desert Center on Aqueduct Road, Riverside Co., 10 April 1947, P. A. Munz 11722 (ARIZ, RSA, SD); ca. 5 mi. [8 km.] E. of Clark Lake in Borrego unit of Anza State Park, San Diego Co., 21 April 1955, Guy Fleming 45817 (DS, SD); near Needles, San Bernardino Co., 3 June 1929, H. L. Mason 5362 (DS, UC); base of Whipple Mts. adjacent to Colorado R., 2 mi. [3.2 km.] above Earp on road to Parker Dam, 15 July 1940, Wolf 9723 (RSA).
Flowering from March to June with the peak in April and with occasional off-season blooming from August to November.
The blue palo verde, as it is called in Arizona, occurs principally in fine soil along washes and on flood plains, for the most part at altitudes below 1100 m. Its water requirements appear to be greater than those of C. microphyllum. It occurs in the Colorado Desert area of southeastern California, in southern Arizona from 35°N. near the Colorado River, southeast to ca. 33°N. on the eastern border of Arizona and southward through the coastal and middle portion of Sonora to 27°30’N. Outlying collections have been made in coastal Sinaloa as far southward as 26°N. Cercidium floridum and C. microphyllum are the most widely distributed members of the genus within the Sonoran Desert, but C. floridum subsp. floridum is the only one restricted to the mainland. Although current floras indicate C. floridum subsp. floridum as occurring in Baja California, and Goldman (1916, p. 335) cites it as being there, no specimens have been seen which support such a distribution.
In most areas, the peak of the blooming season of Cercidium floridum is about two weeks in advance of that of C. microphyllum.
4b. Cercidium floridum Bentham ex Gray subsp. peninsulare (Rose) stat. nov. and comb. nov. Cercidium peninsulare Rose, 1905, Contrib. U.S. Nat. Herb., vol. 8, p. 301.
Shrub or tree 2.5—8 m. tall, openly branched, but with a dense, symmetrically- rounded crown; bark gray and scaly-fissured at base, bright green above, inconspicuously horizontally striate; branchlets not flexuous, having pilose or
36 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
sparse, appressed-villous hairs when young, glabrate to glabrous in age; a single axillary thorn usually present at a node, 2-3 (—7) mm. long (mean 3.87 mm.), slender to stout; axillary bud scales, when present, not dark and conspicuous; leaves with petiole 1-4 (—10) mm. long bearing 1 (rarely 2) pair of pinnae 0.3-1.7 (—2.1) cm. long, the leaflets mostly 2 pairs per pinna but many (25 percent) with 3 pairs, obovate and sometimes emarginate, 6-11 (—18) mm. long, 3—7 mm. broad, sparingly pilose to glabrate; inflorescences borne on old wood as well as on younger branchlets, more or less compact, the racemes with rachis sometimes almost absent, but usually 3-10 (—20) mm. long, pubescent, bearing 3—5 flowers on pubescent pedicels 4-9 (—12) mm. long; calyx sparingly pubescent to glabrous; petals deep or bright yellow, the upper 7.5—9.0 mm. long, the limb 5—7 mm. long, 5.0—-6.5 mm. wide, ovate or rhombic ovate, the claw 2 mm. long, the other four petals slightly shorter to subequal to the upper, limb ovate, the claw short, 1.5-2.0 mm. long; legumes 1.5—5.5 cm. long, 8-15 mm. wide, flat, the margins slightly constricted between the seeds, seeds 1-2 (3) per legume, 9-10 mm. long, 6-7 mm. wide.
Typr. La Paz, Lower California, Mexico, in 1890, Palmer 112 (US!).
REPRESENTATIVE SPECIMENS. MEXICO. IsLANps, BAJA CALIFORNIA SUR: arroyo leading eastward from old Ruffo Ranch, Isla Ceralvo, 16 April 1962, Wiggins 17758 (DS, ENCB); Arroyo de Agueda, NE. side of San José Is., 11 April 1962, Moran 9402 (CAS, RSA, SD); arroyo above Ensenada Ballena, Espiritu Santo Is., 21 April 1962, Moran 9634 (SD, US); S. end of Santa Cruz Is., 18 April 1962, Moran 9577 (SD, UC); arroyo above spring, Santa Catalina Is., 14 April 1952, Moran 3866 (DS), 10 April 1962, Moran 9369 (SD); Puerto Ballandra [ Balandra], Carmen Is., 21 May 1921, Johnston 3802 (CAS, DS, GH, K, UC, US); 4 June 1963, Carter & Reese 4546. PENINSULA, BAJA CALIFORNIA Sur: San José del Cabo, March—June 1897, A. W. Anthony 363 (F, GH, K, UC, US); Arroyo San Lazaro, ca. 10 mi. [16 km.] NW. of San José del Cabo, 2 May 1959, Thomas 7775 (ARIZ, CAS, DS, SD, UC, US); Arroyo del Salto, E. of La Paz, 30 March 1949, Carter 2595 (DS, UC); La Paz, 16 April 1899, E. A. Goldman 388 (GH, US! Type); Arroyo San Ramon W. of Rancho San Ramon (25°14.5’N., 111°17'W.), 25 October 1964, Carter +866; vicinity of Puerto Agua Verde, 4 June 1965, Carter & Sharsmith 4943, arroyo S. of Bahia Agua Verde, 5 June 1965, Carter & Sharsmith 4946: Arroyo Peloteado, W. of Rancho Peloteado (25°45’N., 111°30'W.), 21 April 1962, Carter 4410; San Javier, 21 April 1962, Carter 4414; Rancho Viejo, ca. 28 km. from Loreto on road to San Javier, 6 May 1972, Carter 5680; Rancho Quifi, on old mission trail SE. of Comondu, 20 April 1955, Carter & Ferris 3423 (DS, MEXU, SD, UC, US); alluvial arroyo margin, Las Cuevitas below Comondu, 12 April 1939, Gentry 4458 (ARIZ, GH, K, MO, UC, US); Valle de Los Encinos, S. side of Cerro Giganta, altitude 750 m., 8 November 1960, Carter 4134, 8 June 1963, Carter &
Vout. XL] CARTER: CERCIDIUM 37
Reese 4573; coastal strand, San Bruno (26°12.5’N., 111°23'W.), 1 June 1963, Carter & Reese 4537; Arroyo Mulegé, ca. 10 mi. [16 km.] W. of town on road to El Potrero, 11 April 1963, Wiggins 18231 (CAS, DS, K, MEXU, US); Picachos de Santa Clara, Desierto Viscaino, 5-10 November 1947, Gentry 7754 (DS); wash 16 mi. [26 km.| from San Ignacio on Calmalli road, 10 March 1934, erms.9037 (Ds, US).
Flowering is from March to June with the peak in April, the branches usually bearing leaves at the same time. The flowering period precedes that of its congeners in the same area.
It grows on coastal plains and along washes and canyons into mountain valleys from sea level up to 750 m. This tree, known locally as palo verde is the only taxon in Cercidium restricted to Baja California, where it is moderately common from near the tip of the peninsula northward to San Ignacio (27°25’N., 112°52'W.). As noted by Gentry on one of his collections (4458), this tree has a denser, leafier, more symmetrical crown than any other members of the genus. Also, it retains its leaves for a greater portion of the year than any of the others. The upper petal of C. floridum subsp. peninsulare is not as conspicuously elevated above the other four petals as it is in the other taxa.
DISCUSSION
In an attempt to understand the relationships between the Sonoran Desert taxa of Cercidium as set forth above, the following characters were of use and merit some discussion.
Bark. Inasmuch as most cercidiums retain only a few leaves during much of the year, or have minute leaves, photosynthesis also occurs in the green bark which is relatively smooth except for the lower trunks of large trees. In C.
38 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
praecox, the bark is bright green clear to the ground, whereas in the others the main trunk becomes grayish and often fissured. The bark of C. praecox differs also in having a minute quadrate-pustulate surface pattern (fig. 6a,b), easily visible with a hand lens or even to the naked eye, and in being conspicuously coated with a waxy substance. Shreve (1951, p. 145; 1964, p. 153) mentioned these characteristics, but he attributed them to C. X sonorae which he at that time mistook for C. praecox. The bark of C. micruphyllum and C. floridum, on the other hand, is finely horizontally striate (fig. 6c) and, especially in C. floridum subsp. floridum, often dark in the indentations, thus giving a ‘dirty-neck’ appear- ance. These striations are less conspicuous in C. floridum subsp. peninsulare. The bark of C. X sonorae is usually smooth, but sometimes horizontal striations are visible with a hand lens. On some specimens of C. X sonorae there is a thin, inconspicuous layer of waxy material which suggests C. praecox.
PUBESCENCE. Young branchlets of Cercidium show varying degrees of pubescence; older branchlets are glabrous to glabrate. In C. floridum subsp. floridum, branchlets are essentially glabrous, but sometimes with a few hairs, especially southward in its range. In C. floridum subsp. peninsulare, on the other hand, the young branchlets are conspicuously pubescent with pilose or sparsely appressed villous hairs. In C. microphyllum there is some correlation between geographic distribution and type of pubescence on the stems: all of the Baja California material is strongly strigose; the Sonoran collections studied are puberulent or pilose (especially in the interior); in Arizona and California some collections have glabrate branchlets. Branchlets of C. praecox vary from glabrate to pubescent. In C. X sonorae the branchlets may be strigose, short villous, or pilose, or occasionally the branchlets are glabrous.
The pubescence of the ovary at anthesis is of diagnostic value. In both subspecies of C. floridum and in C. praecox the ovary is glabrous; in C. micro- phyllum and C. X sonorae it is lightly pubescent to strigose at anthesis.
ARMATURE AND BRANCHING PATTERN. Most treatments of Cercidium refer to the thorns as “nodal” or “stipular,’’ but Kearney and Peebles (1960, p. 407) describe them as “rudimentary branches transformed into spines.” This concept (except for the use of the term “spine” instead of ‘“‘thorn”’) is in accordance with Blaser’s (1956) definition, which is followed herein, of a thorn being a reduced determinate shoot. Leaves of seedling plants and of young seasonal growth of Cercidium bear a pair of inconspicuous, slender, foliaceous stipules; these do not form spines, but are fugaceous. Thorns, when present, are the first shoots to develop in a primary (first to develop at a node) leaf axil. Their shoot nature is clearly shown in the early stages by the presence of a cluster of rudimentary leaves at the distal end of the developing thorn (fig. 7c). These soon disappear as the thorn increases in size and hardens (fig. 7d). Subsequent growth at a node results in either inflorescence, leaf, or shoot development from buds on an
Vor. XL] CARTER: CERCIDIUM 39
extremely shortened stem in the axil of the primary leaf. However, the positional relationship of the thorn primordia to the shoot, leaf, and flower primordia cannot be determined without more extensive morphological investigation.
In Cercidium microphyllum, none of the buds in a primary leaf axil develop
40 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
AN & ny : Ny ofc ry WAV C) WAS & © = \ tS o, ~ sS —\> S Xe. e \2. \y s NS Ne: S 3 ONC a\s e ~ AA ¢ 0" tA? C.microphyllum C. X sonorae C. praecox C. floridum subsp. floridum subsp. peninsulare
Ficure 8. Branching patterns in Cercidium, based on number and length of branchlets in the terminal 25 cm. of a stem.
into a short leafless thorn; rather, at the majority of nodes one bud develops into the stout, determinate, leafy branch, spinescent at the tip, which is so characteristic of this species. Cercidium praecox is consistently the thorniest of the Sonoran Desert members of the genus inasmuch as at each node one, and occasionally two, axillary buds develop into thorns. When two thorns occur at a node, one is always larger. In C. floridum subsp. floridum, comparatively few axillary buds develop,—the resulting mature tree having long, slender branchlets, a less divaricately branched appearance, and fewer thorns (especially in the northern part of its range) than do most other species. However, Shreve (1951, p. 142; 1964, p. 150) states that young trees of C. floridum less than 2 m. in height have very thorny twigs. Cercidium floridum subsp. peninsulare and C. X sonorae are variable in the number of axillary buds which develop into thorns. In many instances, C. X sonorae is thornless.
Cercidium microphyllum, which, as noted above, has no axillary thorns but bears only spine-tipped, determinate branchlets, has the greatest number of branchlets (mean, 8) in the terminal 25 cm. of a branch, whereas C. praecox, the thorniest of the species, has the fewest branchlets (fig. 8). In C. micro- phyllum, there is usually a gradual reduction in length of the branchlets toward the distal end of the branch in contrast to the other taxa where reduction in branchlet length often shows no correlation with position on a stem segment. Cercidium X sonorae has an intermediate number of branchlets in comparison with its putative parents and the branchlets are longer and more flexuous than those of either parent. The average number of branchlets developing in the terminal 25 cm. of the branches of the two subspecies of C. floridum is rather
Vor. XL] CARTER: CERCIDIUM 41
close: 5.4 branchlets in C. floridum subsp. floridum and 6.1 branchlets in C. floridum subsp. peninsulare, but the mean length of the branchlets diverges significantly. The flexuous branchlets of C. floridum subsp. floridum have a mean length of 13.3 cm. and the stouter and stiffer branches of C. floridum subsp. peninsulare have a mean length of 9.3 cm. (fig. 8). These means for number and length of branchlets help to indicate their relationship within the group, but in all cases, the range of these measurements within each species and the overlap with other species, is too great for use of the characters in keys.
LEAVES. The seedlings of all Sonoran Desert taxa of Cercidium have con- spicuous cotyledons, 23 to 25 mm. long in C. floridum and 6 to 18 mm. long in the others. In the seedlings, each of the first few nodes (usually up to five) bears a pinnate leaf with several pairs of large leaflets; following these, the bipinnate leaves more typical of each of the various taxa develop (figs. 3, 7a). The inconspicuous, foliaceous stipules are caducous. The pinnae (i.e., the secondary rachises with their leaflets) are short (0.3 to 6.5 cm. long) and the leaflets are opposite. A small, foliaceous or semi-indurate bract is borne at the apex of the leaf rachis and at the apex of each of the pinnae. Because of the need to discuss them separately, the first leaf developing at a node is herein designated as a ‘primary’ leaf and those developing subsequently in the axil as ‘axillary’ leaves. Buds produced in the axil of the primary leaf may develop into a thorn (a short determinate shoot), or into foliaceous or floral shoots. The foliaceous shoots are usually so reduced that the leaves borne on them appear to arise directly in the axil of the primary leaf or leaf scar; one to eight such leaves may be present at a time. Leaves are deciduous in most taxa, usually falling before the blooming period or with periods of drought.
Leaves of Cercidium microphyllum differ from those of the other taxa in being non-petiolate, except in the seedling stages wherein the primary leaf may have a slightly winged petiole from 1 to 4 mm. long which merges inconspicuously with the broadened rachis area where the pair of pinnae arises. In later stages, the primary leaf pinnae are borne on an indurate, scale-like structure, the basal portion of which usually is not distinguishable from the rachis as a petiole. The indurate terminal bract is also indistinguishable from the rachis. The two pinnae fall separately from this scale-like rachis and it remains on the branch for some time before falling and exposing the primary leaf scar. (Sometimes in C. microphyllum, the leaflets fall before the pinnae do.) In the other species of Cercidium the entire leaf abscisses at the base of the petiole. (Occasionally in C. floridum subsp. floridum the pinnae fall from the apex of the petiole, or from the rachis if the leaf bears two pairs of pinnae, and the petiole, or petiole and rachis remain on the stem for a short time.) In specimens of C. X sonorae with subsessile leaves, the shortened petiole and the rachis become somewhat indurate,
42 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
35 ‘ C. floridum \ ——-—subsp.floridum » 30 : subsp. peninsulare io: \ E 25 ; n e ol \ 20 ! D \ a \ E 15 \\ = \ \ fe) \ py \ a1) Aone ae eo—e. aN a Wa o. 9) fe) 15
Length of petiole (mm)
Number of samples
Ficure 9. Comparison of length of petioles and pinnae in Cercidium floridum subsp floridum and C. floridum subsp. peninsulare.
but they fall with the rest of the leaf instead of being tardily dehiscent as in C. microphyllum. In C. praecox and C. X sonorae, the scar of the primary leaf is conspicuous on younger branches; in C. floridum subsp. floridum it is
inconspicuous. In C. floridum subsp. peninsulare, the scar of the primary leaf is often obscured by a cluster of reddish-brown glandular hairs in the axil.
In Cercidium praecox and C. floridum the bract at the apex of the rachis tends to be linear and foliaceous rather than indurate. In specimens of C. X sonorae with petiolate leaves, the elongate apex of the rachis is usually slightly indurate, while in those having subsessile leaves the rachis is an indurate, long-tipped scale similar to that of C. microphyllum.
Cercidium floridum \eaves have pinnae usually less than 1 cm. long and
Vor. XL] CARTER: CERCIDIUM 43
mostly bearing 2 to 4 pairs of leaflets, whereas the other three taxa (C. micro- phyllum, C. praecox, and C. X sonorae) have pinnae up to 6.5 cm. long bearing 4 to 9 pairs of leaflets. At a significance level of 1 percent the means for both characters differ between members of the two groups.
As indicated by the graphs (figs. 9, 10), the two subspecies of C. floridum are similar in characters of petiole length, pinna length, and leaflet number. Each subspecies exhibits its greatest variability in length of pinnae. Mean length, however, is about the same for the two (7.8 mm. for C. floridum subsp. floridum, 7.9 mm. for C. floridum subsp. peninsulare) and the difference between the means is not statistically significant. At a 1 percent significance level the only leaf difference is the mean number of leaflet pairs. In the primary as well as axillary leaves, C. floridum subsp. floridum usually has 3 pairs (mode 3, mean 2.7) and C. floridum subsp. peninsulare, usually either 2 or 3 pairs (mode 2, mean 2.5). In all of these structures, the measurements show too much overlap to make them principal key characters. That these characters differ in degree rather than in kind supports the subspecific disposition of C. floridum peninsulare.
Among the other three taxa, petiole length is useful in recognizing entities: the leaves of C. microphyllum are non-petiolate, i.e., the pinnae are sessile (except in seedling stages); the leaves of C. praecox always have conspicuous petioles; those of C. X sonorae vary from no petioles up to petioles 12 mm. long (fig. 11). The mean petiole length for each taxon differs significantly (P = 0.99) from that of the other two. As with C. floridum the pinnae in each species exhibit great variability in length, but the means, although clustered, are not so close (table 1). At a 1 percent significance level the only difference occurs between those for C. X sonorae and C. praecox. As to the leaflet number, the members of this group overlap considerably and show no significant difference in mean number of pairs. Both in this character and in petiole length the mean for C. X sonorae is inter- mediate between that for C. microphyllum and C. praecox. In pinna length, however, the mean for C. X sonorae exceeds that of the other two taxa; but its value is close to the mean for C. microphyllum, and the two values do not differ significantly.
As suggested by the common name for Cercidium floridum, blue palo verde, leaf color has been utilized in identification, but because interpretations of color differ so markedly, no attempt has been made to include precise shades of green in the detailed descriptions. The following field observations, however, point up the foliage color differences in the taxa of Cercidium as they occur in Sonora (correspondence: Hastings to Carter, August 10, 1971).
“T made a trip to Sonora again week before last and the summer rains have
wrought their usual alchemy there. The parched country we saw in April is green and incredibly lush. Also incredibly muggy, buggy and mud‘ty.
[Proc. 4TH SER.
CALIFORNIA ACADEMY OF SCIENCES
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VoL. XL] CARTER: CERCIDIUM 45
C.floridum ——- | subsp. floridum —— [J subsp. peninsulare
Number of samples
! 25773; 495 2 pairs 3 pairs 4 pairs Pairs of teaflets
FicureE 10. Comparison of number of pairs of leaflets in Cercidium floridum subsp. floridum and C. floridum subsp. peninsulare.
“All four palo verdes were in full leaf and seeing them that way did a good bit to restore my ego. Not only is it possible easily to tell Cercidium floridum from C. microphyllum, [but also] C. praecox and C. X sonorae can be distinguished at a glance. Cercidium praecox shows up as blue—even bluer than C. floridum. C. X sonorae, on the other hand, is relatively yellow—not the nearly chlorotic yellow of
C. microphyllum, but about the color of the stems on both C. xX sonorae and C. praecox.”
INFLORESCENCE. In all of the Sonoran Desert taxa of Cercidium, except C. floridum subsp. peninsulare, when both flowers and leaves are present con- currently, the inflorescences exceed the leaves in length. In C. microphyllum, C. praecox, and C. X sonorae, the inflorescences normally develop before the leaves; if leaves are present, they are usually on separate branches. In C. floridum subsp. floridum, flowering precedes leafing on about 30 percent of the collections studied. In C. floridum subsp. peninsulare, where both flowers and leaves are usually present on a given branch, the inflorescences are equal or subequal to the leaves; occasionally, flowering precedes leafing.
The flowers are borne in compact or open racemes, which may be either single or fascicled at the nodes. For the most part, the inflorescences are borne in profusion on the terminal branchlets, but C. praecox departs strongly from this pattern in having its usually compact inflorescences strongly fascicled along the stems, none of which are as branched as in the other taxa. Shreve (1951, p. 145; 1964, p. 153) pointed out this striking characteristic of C. praecox, but here again, he mistakenly attributed it to C. X sonorae. In C. floridum subsp.
46 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
-—-C.microphyllum —: C.X sonorae — C.praecox
Number of samples
Number of samples
Ficure 11. Comparison of number of pairs of leaflets and length of petiole in Cercidium microphyllum, C. X sonorae and C. praecox.
peninsulare, some racemes are borne on older branches; therefore the trees do not have the strong terminal flowering aspect of C. floridum subsp. floridum.
Inflorescence rachis length varies both within the taxa and between them. In C. floridum subsp. floridum the rachises vary from 1.0 to 4.5 cm., while in C. floridum subsp. peninsulare they are usually 0.3 to 1.0 cm., but occasionally up to 2.cm. In C. microphyllum they are 0.2 to 4.5 cm. long, while in C. praecox they are 0.2 to 1.0 cm. long. Those of C. X sonorae (0.5—4.0 cm.) fall between those of C. microphyllum and C. praecox.
VoL. XL] CARTER: CERCIDIUM 47
The pedicel is jointed, with the joint in the distal half. The distance between joint and calyx varies less than that between joint and rachis. In total length of the pedicel, as in rachis length, C. floridum subsp. floridum and C. floridum subsp. peninsulare fall at opposite ends of the spectrum for the five taxa. Their respective means, 10.0 mm. and 5.9 mm., differ significantly at P = 0.99. So do
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Ficure 13. Comparison of petal shape and size in Cercidium: a, C. microphyllum (Carter 5683); b, C. X sonorae (Carter 5679); c, C. praecox (Carter 5678); d, C. floridum subsp. floridum (Turner, Arizona, pickled material); e, C. floridum subsp. peninsulare (Carter 5680). Numbers refer to Carter and/or Carter et al. collections; data for these are given under “Representative collections cited” for the respective taxa.
48 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
the means for C. microphyllum (6.0 mm.) and C. praecox (8.1 mm.). The average pedicel length for C. X sonorae (8.0 mm.) falls between those of its putative parents, but does not differ significantly from that of C. praecox (table 1).
As for the caesalpinoid flowers (figs. 12, 13), there is size variation within each taxon, but the flowers of C. floridum subsp. floridum are the largest in the group and those of C. microphyllum and C. floridum subsp. peninsulare are the smallest. Flowers of C. microphyllum differ also in having the limb of the long- clawed upper petal white, or occasionally cream or pale yellow and the other four petals rhomboidal or lanceolate rather than broadly ovate (figs. 12, 13). The upper, long-clawed petal of C. praecox nearly always bears a cluster of small orange dots near the base of the limb; these are also often present in C. floridum subsp. floridum, but they are lacking in C. floridum subsp. peninsulare. The upper petal of C. X sonorae is extremely variable. It may be white, creamy, or light yellow, and it may or may not bear orange dots.
The tricolpate pollen grains, which vary in diameter from 18 to 34 microns, are prolate spheroidal and supra-reticulate (Carter and Rem, 1974). The lumina are reduced in diameter and depth near the margin of the furrows.
LEGUMES AND SEEDS. The venation of the valves of Cercidium legumes is of diagnostic value: the valves of C. praecox are somewhat papery with conspicuous reticulate venation; those of C. microphyllum and C. X sonorae are striate— especially noticeable in the latter; and those of C. floridum are smooth with veins scarcely visible to the naked eye. The surface of the valves of immature legumes of C. floridum bear conspicuous white-dotted areas. These are stomatal pores; apparently the young fruits as well as the bark carry on photo- synthesis at a time when the trees are nearly leafless. Stomata are present also in the legumes of the other taxa, but they are somewhat obscured by the prominent venation.
Although Brenan (1963, p. 207, table) used the upper suture of the pod as a distinguishing character, this is more variable within our Sonoran Desert taxa than he indicated. The legumes are tardily dehiscent along the sutures or indehiscent; in Cercidium microphyllum and C. X sonorae the thin-walled valves also sometimes break irregularly. Differences in shape and size of legumes and seeds may be compared in figure 3. In characters of legume venation and shape, C. X sonorae approaches C. microphyllum more closely than it does C. praecox.
As to surface pattern, seeds of C. praecox and C. X sonorae are gray-brown with various degrees of brown mottling on the flat surfaces; seeds of C. micro- phyllum and C. floridum are brown with the marginal area somewhat lighter and sometimes faintly mottled. In all of the taxa the hilum and micropyle are subterminal; in some species they are in a slightly recessed or notched area
VoL. XL] CARTER: CERCIDIUM 49
Ficure 14. Comparison of seeds (cross section and position of hilum and micropyle) in Cercidium and Parkinsonia: CEFL, C. floridwm subsp. floridum, a, side view to indicate position of cross section (b) to entire seed (E. E. Schellenger 6, Chucawalla [Chuckawalla ] Bench, Colorado Desert, California, 25 July-14 August, 1903, UC); CESO, C. Xx sonorae, note variation in form of cotyledons (Carter & Sousa 5212); CEPR, C. praecox (Carter 4146) ; CEMI, C. microphyllum, note thick cotyledons (Ferris 9902); CEPE, C. floridum subsp. peninsulare (Carter 4821); PAAC, Parkinsonia aculeata (Harbison & Higgins, San Telmo, Baja California, 17 December 1953, UC); PAAF, Parkinsonia africana (Rodin 2156, Namib Desert near Swakop River, Southwest Africa, 31 October 1947, UC). Diagrams drawn after seeds had soaked in water overnight. Seeds, x 24, side view of apex, X 1%.
(fig. 14). As seen in cross section, the thick, oblong or subglobose seeds of C. microphyllum are distinct in having large cotyledons in comparison to the amount of endosperm (fig. 14). The other taxa, with more flattened and oblong or oblong-ovate seeds, follow much the same pattern in cross-section (fig. 14) as that illustrated by Boelcke (1946) for C. australe Johnston, the Argentinian species closely related to C. praecox.
CHROMOSOME NUMBERS. Chromosome counts of 2n = 28 have been reported for Cercidium floridum, C. microphyllum and C. X sonorae (Turner and Fearing, 1960). Although contraindicated in the publication, no voucher was preserved
50 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
for the C. X sonorae count (correspondence, B. L. Turner to Carter, 5 July 1972). It is probable, however, that the seeds provided for this count were those of C. praecox rather than of C. X sonorae (correspondence, R. M. Turner to Carter, 23 June and 18 August, 1972).
HYBRIDIZATION. Few putative hybrids between Cercidium microphyllum and C. floridum subsp. floridum have been noted, whereas C. X sonorae, which occurs in the area of overlap of C. microphyllum and C. praecox, is relatively abundant. A six-year record kept by W. G. McGinnies for trees in the foothills north of Tucson, Arizona, shows that C. floridum subsp. floridum blooms about two weeks earlier than C. microphyllum and C. praecox (the latter being a cultivated tree in that area). So, although there is some overlapping of the blooming period of C. floridum subsp. floridum and C. microphyllum in the Tucson area at least, the peak is reached at different times (correspondence, Hastings to Carter, 24 January 1972); no putative hybrids have been reported from this area. On the other hand, in the lower Colorado River basin between Arizona and Colorado, these two species do reach peak of bloom at approximately the same time and a small percentage of putative hybrids has been noted (pers. comm., Jones to Carter, September 1973). Ultraviolet light studies show that the flowers of these two species appear different to pollinating insects: C. microphyllum petals absorb UV light and all are “bee purple,” while in C. floridum subsp. floridum only the upper petal absorbs UV light and the other four reflect it. The pollinators appear to be highly selective in accordance with these ultraviolet patterns (pers. comm., C. E. Jones to Carter, September 1973). In the case of C. X sonorae, on the other hand, whose putative parents, C. microphyllum and C. praecox, bloom concurrently and have the same ultraviolet absorption patterns as the above mentioned taxa (Carter, 1974), it appears that their pollinators are not selective in their visits inasmuch as there is apparently ample cross-pollination. No putative hybrids have been noted between C. floridum subsp. peninsulare and the other three taxa occurring with it in southern Baja California. Normally this species blooms before its congeners; its ultraviolet absorption pattern is the same as that of C. floridum subsp. floridum and of C. praecox, i.e., the upper petal absorbs and the other four reflect ultraviolet light.
RELATIONSHIP BETWEEN CERCIDIUM AND PARKINSONIA. As indicated in the key at the beginning of this treatment, a number of characters serve to distinguish members of the New World genus Cercidium from Parkinsonia aculeata Linnaeus, the only member of the latter genus considered to be native to the western hemisphere. Treatments of these two genera fail to clarify the structure of the primary leaf and its relationship to thorns or spines; many floras merely state “stems armed” or ‘“‘stems unarmed.” As detailed above, armature in Cercidium, when present, consists of thorns which develop in the axil of a primary leaf.
VoL. XL] CARTER: CERCIDIUM 51
In Parkinsonia aculeata, on the other hand, there are no axillary thorns. The conspicuous armature consists of the short petiole and rachis of the primary leaf itself, which has become enlarged, thickened and indurate, plus the tip of the rachis which has developed into a long, stout, sharp spine. Hutchinson (1969, pp. 70-71, fig. 64E), misinterpreted this structure, calling it a stipule. Also, he failed to differentiate between primary and axillary leaves. The stipules are either early deciduous or develop into short, indurate lateral spines, which, however, may fall long before the indurate petiole and rachis. One to three pairs of long (10 to 60 cm.) pinnae are produced on the short leaf rachis. In both primary and axillary leaves the secondary rachises are conspicuously flattened (phyllodial). In her anatomical comparison of Parkinsonia aculeata and Cercidium torreyanum (C. floridum), Scott (1935) found that only in the rachises of the pinnae and in the pulvinae is there a marked difference between these two taxa. She did not include armature in her studies. The small, caducous leaflets borne along the margins of the pinna rachis may be either opposite or alternate on the same pinna. The long phyllodial pinnae usually remain until the onset of winter and lend the trees a graceful, drooping aspect. When pinnae of the primary leaves fall, the indurate, spinose-tipped petiole and rachis remain on the branch. On older branches, this structure (spine) also eventually falls, leaving a conspicuous scar at the node. Burkart (1952, fig. 36) illustrates the spinose nature of the leaf rachis. He does not point out, however, that these stout spines are restricted to the first leaf at a node of a main branch and are only slightly, if at all, developed on leaves of the usually short axillary branches which are from 2 to 12 (—23) mm. long in contrast to the completely reduced axillary shoots in Cercidium.
Until the recent work of J. P. M. Brenan (1963), only one other taxon has currently been considered as belonging to the genus Parkinsonia, the African species P. africana Sonder. The primary leaf of P. africana is similar in some respects to that of Cercidium microphyllum. In Parkinsonia africana a single pair (occasionally one of the pair aborts) of pinnae 4 to 17 cm. long arises from a small, bractlike, nearly sessile leaf rachis which has a short, slender, foliaceous tip; stipules are inconspicuous, caducous, bractlike structures; the pinnae are not flattened, but are elliptical in cross section and bear tiny opposite, caducous leaflets, as is the case in the axillary leaves. Troll (1939; p. 1611, fig. 1372; p. 1612, fig. 1373), considers these leaves to be a reduced form of the typical caesalpinoid leaf which is petiolate and bears several pairs of pinnae. Armature, when present, is a short, stout, determinate stem (thorn) in the axil of the primary leaf. Johnston (1924a, p. 63) erred by indicating in his key to Cercidium and Parkinsonia that both P. aculeata and P. africana have similar armature. Presence or absence of these axillary thorns is extremely variable in P. africana, as it is also in some taxa of Cercidium. In all of these characters except the
52 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
somewhat longer pinnae with leaflets caducous, Parkinsonia africana resembles Cercidium microphyllum more than it does Parkinsonia aculeata. Torrey (1859, p. 60) reached a similar conclusion.
The inconspicuous, bractlike rachis of the primary leaf in both Cercidium microphyllum and Parkinsonia africana might well be considered a rudimentary expression of the highly developed spine of the primary leaf of Parkinsonia aculeata. So, in this character, Cercidium microphyllum tends to resemble Parkinsonia more closely than it does its congeners with petiolate leaves (both primary and axillary) which fall entire. However, in the short pinnae which are not flattened and in the opposite leaflets which usually are not caducous, it falls within Cercidium.
It has not been possible to make careful examination of the other African trees placed in Parkinsonia by Brenan: P. anacantha Brenan and P. scioana (Chiovenda) Brenan |Peltophoropsis scioana Chiovenda]; so comparisons cannot be made regarding the structure of the primary leaves (matters which are not adequately treated in the descriptions of these two taxa).
There is a significant difference in pedicel length (P= .99) between Parkinsonia aculeata and P. africana, on the one hand, and the species of Cercidium studied, on the other. The mean pedicel length for the taxa of Cercidium ranges from 5.9 to 10.0 mm., with the ratio of total pedicel length to length from pedicel joint to calyx ranging from 3.5:1 to 4.3:1. In Parkinsonia aculeata the mean length is 14.5 mm.; the ratio, 6.4:1. In P. africana the mean pedicel length is 13.0 mm., not significantly different from that of P. aculeata. But the ratio is only 2.4:1, i.e., the joint occurs almost at the midpoint of the pedicel, whereas in P. aculeata it is relatively close to the calyx. In Cercidium, the joint is in the distal half of the pedicel, but never proportionately so close to the calyx as in Parkinsonia aculeata. Material was not available to carry the comparison to the other African taxa placed in Parkinsonia.
Pollen studies in relation to hybridization in Cercidium and Parkinsonia have shown that putative hybrids between taxa in Cercidium have a much higher percent of presumably viable pollen (as indicated by staining with aniline-blue lactophenol) than do the few known hybrids between Parkinsonia aculeata and species of Cercidium (Carter and Rem, 1974). In addition, pollen grains of putative hybrids between species of Cercidium are not malformed whereas those of putative hybrids between Parkinsonia aculeata and species of Cercidium are irregular in shape (Carter and Rem, 1974, fig. 1). No putative hybrids between C. microphyllum and Parkinsonia aculeata have been noted even though these two taxa sometimes occur in the same area.
As seen under the scanning electron microscope, pollen grains of the several species of Cercidium studied and those of Parkinsonia aculeata have similar
VoL. XL] CARTER: CERCIDIUM 53
supra-reticulate sculpturing (Martin and Drew, 1969, 1970; Carter and Rem, 1974). Pollen of Parkinsonia africana (fig. 15), on the other hand, although being reticulately sculptured, has many shallow lumina as well as the deep lumina characteristic of Cercidium and Parkinsonia aculeata. Furthermore, in P. africana the lumina are much smaller and the muri thicker than in the above- mentioned taxa. Pollen of Parkinsonia |Peltophoropsis| scioana (fig. 15) is triporate with three large pores; its sculpturing differs markedly in that the reticulate surface appears rugose and the lumina are small and the muri wide. Unfortunately, the material of P. anacantha available contained insufficient pollen for a scanning electron microscope preparation or for satisfactory light microscope examination.
The legumes of Cercidium microphyllum resemble those of Parkinsonia aculeata more than they do those of other species of Cercidium. They are thin- walled and strongly torulose, i.e., much constricted on either side of the 1 to 4 seeds; dehiscence is irregular, as well as occurring along the sutures. Legumes of C. X sonorae resemble those of C. microphylium in dehiscence. Those of other species of Cercidium appear to be indehiscent, or only slightly dehiscent. Legumes of Parkinsonia aculeata are sinuate or somewhat torulose and 1 to 5 seeded; otherwise they are similar to those of Cercidium microphyllum. Legumes in Parkinsonia africana material at hand are sinuate, flattened, and strongly dehiscent, while those of P. scioana and P. anacantha are flattened, but not constricted; those of P. scioana, at least, are indehiscent. Also, P. scioana is unique in the group in having long-funicled seeds. Although Cercidium micro- phyllum strongly resembles Parkinsonia aculeata in the character of its legumes, it stands apart from that species, and from other species of Cercidium, in a seed
54 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
character: the cotyledons are thick in relation to the endosperm (fig. 14). Also, in dehiscence of the pinnae from the scale-like leaf rachis, C. microphyllum ap- proaches Parkinsonia aculeata, but in the majority of characters it is more at home in Cercidium than in Parkinsonia.
As stated in the introduction, resolution of the generic relationship of Cercidium and Parkinsonia ,should be based upon extensive comparative biological, morphological, cytological, and genetic studies. On the basis of present information, however, one can say that Cercidium forms a discrete, easily recognizable unit confined to the Americas, with C. microphyllum most closely related to Parkinsonia. Study of Brenan’s comparative chart (1963, pp. 206, 207) shows that among the taxa included, Parkinsonia aculeata is the most discordant element. Furthermore, the American species of Cercidium included agree in more gross morphological characters than do the African taxa placed in Parkinsonia. Pollen of two African taxa, as seen under the scanning electron microscope is of two distinct types whereas in Cercidium it is all similar. Pollen studies (Carter & Rem, 1974) indicate that species of Cercidium are more closely related to each other than they are to Parkinsonia aculeata, a species thought to be native to America, but which has been introduced widely throughout the warmer parts of the world. I would like to suggest that Parkinsonia be considered a monotypic genus comprised of P. aculeata Linnaeus, and that the relationships of the three African species (P. africana, P. anacantha, and P. scioana) be considered further before accepting them as congeneric with Parkinsonia and Cercidium as proposed by Brenan.
ACKNOWLEDGMENTS
In conclusion, I wish to express my sincere appreciation to the various governmental departments in Mexico which have been instrumental in granting me permission to carry on field work in their country, especially those of the Director General de Aprovechamientos Forestales and the Consejo Nacional de Ciencias y Tecnologia as well as to Dr. Arturo Gdémez-Pompa, Jefe del Departamento de Botanica, Instituto de Biologia, Universidad Nacional Autonoma de México. In La Paz, Sr. Guillermo Arrambidez A., formerly Jefe de la Division Hidrométrica de Baja California, Secretaria de Recursos Hidraulicos, has through the years enabled me to time my field trips to take the best advantage of rains. Many residents of Loreto and the Sierra de la Giganta have facilitated my work by their friendship and cooperation.
Also, I am deeply indebted to the Belvedere Scientific Fund of the California Academy of Sciences and to the Director of the Academy, Dr. George Lindsay, for their generous support of my field work in Baja California during recent years. To Dr. Lincoln Constance and other members of the Staff of the
Vor. XL] CARTER: CERCIDIUM 55
Herbarium of the University of California, I owe special thanks for facilitating my work on Baja California materials. To Miss Nel Rem I am indebted for her assistance with studies on pollen and to Miss Charlotte Mentges for preparation of drawings and graphs. Drs. R. M. Turner and the late J. R. Hastings have shared with me the results of their extensive field work in the Sonoran Desert and generously aided me in many ways during the course of the study. And, last but not least, to the several fellow-botanists who have set forth with me from time to time on field trips in the Sierra de la Giganta, I give special thanks. They never knew just what lay ahead of them on these trips!
LITERATURE CITED
Benson, L. 1940. Taxonomic contributions. I. The native palo verdes of Arizona. American Journal of Botany, vol. 27, pp. 186-187. BrAser, H. W. 1956. Morphology of the determinate thorn-shoots of Gleditsia. American Journal of Botany, vol. 43, pp. 22-28. BoELckE, O. 1946. Estudio morfologico de las semillas de Leguminosas, Mimosoideas y Caesalpinioideas de interés agrondmico en la Argentina. Darwiniana, vol. 7, pp. 240-321 plus 11 plates. Brenan, J. P. M. 1963. Notes on African Caesalpinioideae: the genus Peltophoropsis Chiov. and _ its relationship. Kew Bulletin of Miscellaneous Information, vol. 17, pp. 203-209. Burkart, A. 1952. Las Leguminosas Argentinas, silvestres y cultivadas. Buenos Aires. xv + 568 pp. Carter, A. M. 1974. Evidence for the hybrid origin of Cercidium sonorae (Leguminosae: Caesalpin- ioideae) of northwestern Mexico. Madrofio, vol. 22, no. 5, pp. 266-272. Carter, A. M., and N. C. Rem 1974. Pollen studies in relation to hybridization in Cercidiwm (Leguminosae: Caesalpin- ioideae). Madrono, vol. 22, no. 6, pp. 303-311.
Gentry, H. S. 1942. Rio Mayo Plants. Carnegie Institution of Washington Publication 527, pp. vii + 328, 29 pls.
GoLpMAN, E. A. 1916. Plant records of an expedition to Lower California. Contributions from the United States National Herbarium, no. 16, pp. 1-371. Hastincs, J. R., and R. R. HumpuHrey (editors) 1969a. Climatological data and statistics for Baja California. Technical reports on the meteorology and climatology of arid regions, no. 18, iv + 96 pp. University of Arizona, Institute of Atmospheric Physics. ; 1969b. Climatological data and statistics for Sonora and northern Sinaloa. Technical reports on the meteorology and climatology of arid regions, no. 19, iv + 96 pp. University of Arizona, Institute of Atmospheric Physics. Hastincs, J. R., R. M. Turner, and D. K. WARREN 1972. An atlas of some plant distributions in the Sonoran Desert. Technical report
56 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
no. 21, xiii + 255 pp. University of Arizona, Institute of Atmospheric Physics. HutcHinson, J.
1969. Evolution and phylogeny of flowering plants. Academic Press, London, xxv ++
717 pp., 557 figs. Jounston, I. M.
1924a. Taxonomic records concerning American spermatophytes. 1. Parkinsonia and Cercidium. Contributions from the Gray Herbarium of Harvard University, n.s., no. 70, pp. 61-68. (April).
1924b. Expedition of the California Academy of Sciences to the Gulf of California in 1921. The botany (the vascular plants). Proceedings of the California Academy of Sciences, ser. 4, vol. 12, pp. 951-1218. (May).
Kearney, T. H., and R. B. PEEBLES
1960. Arizona flora. University of California Press, Berkeley and Los Angeles. 2nd ed. 1085 pp.
Lanjouw, J., and F. A. STAFLEU
1964. Index Herbariorum, Part 1, The herbaria of the World. Fifth edition. 251 pp. International Association for Plant Taxonomy. Utrecht, Netherlands.
Martin, P. S., and C. M. Drew
1969. Scanning electron photomicrographs of southwestern pollen grains. Journal of the Arizona Academy of Science, vol. 5, pp. 147-176.
1970. Additional scanning electron photomicrographs of southwestern pollen grains. Journal of the Arizona Academy of Science, vol. 6, pp. 140-161.
Muwz, P. A.
1959. A California flora. University of California Press, Berkeley and Los Angeles.
1681 pp. SARGENT, C. S.
1889. Notes upon some North American trees. V. Parkinsonia. Garden and Forest,
vol. 2, p. 388. Scott, F. M.
1935. The anatomy of Cercidium Torreyanum and Parkinsonia microphylla. Madrono, vol. 3, pp. 33-41. [In a later paper (Madrono, vol. 3, p. 190, 1936) Scott stated that in the above title and throughout the paper Parkinsonia aculeata should be substituted for P. microphylla.]
SHREVE, F.
1951. Vegetation of the Sonoran Desert. Carnegie Institution of Washington Publication, no. 591, xii + 192 pp., 37 plates, 27 maps. [Republished in 1964 as part of vol. 1, Shreve and Wiggins, Vegetation and flora of the Sonoran Desert. Stanford University Press, Stanford, California. ]
SHREVE, F., and I. L. Wiccins
1964. Vegetation and flora of the Sonoran Desert. 2 vols. Stanford University Press,
Stanford, California. 1740 pp. Torrey, J.
1859. Botany of the Boundary [United States and Mexican Boundary Survey], vol. 2,
pp. 30-270, 61 pls. TROLL, W.
1939. Vergleichende Morphologie die Héheren Pflanzen. Erster Band: Vegetationsorgane.
Zweiter Teil. Gebriider Borntraeger, Berlin.
VoL. XL] CARTER: CERCIDIUM 57
Turner, B. L., and O. S. FEARING 1960. Chromosome numbers in the Leguminosae. III. Species of the southwestern United States and Mexico. American Journal of Botany, vol. 47, pp. 603-608. Watson, S. 1876. Descriptions of new species of plants, chiefly Californian, with revisions of certain genera. Proceedings of the American Academy of Arts and Sciences, Boston vol. 11, pp. 121-148.
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 3, pp. 59-86; 4 figs.; 2 tables. October 30, 1974
THE COMPARATIVE MORPHOLOGY OF EXTRINSIC GASBLADDER MUSCULATURE IN THE SCORPIONFISH GENUS SEBASTES
(PISCES: SCORPAENIDAE)
By Leon E. Hallacher
San Francisco State University San Francisco, California 94132!
Aspstract: Phylogenetic relationships within the rockfish genus Sebastes (Scorpae- nidae) are not well understood. Study of variation in the structure of extrinsic gasbladder musculature within this group may help clarify understanding of evolutionary relationships. Eighty-two species of rockfishes were dissected for examination of gasbladder muscles. Possible evolutionary implications are discussed. The function of the gasbladder muscles is sound production, but they may also be used for sound reception.
INTRODUCTION
The rockfish genus Sebastes, of the family Scorpaenidae, contains approxi- mately 100 species, ranging in size from about 200 mm. to 1000 mm. standard length as adults (Phillips, 1957; Eschmeyer and Hureau, 1971). Unlike most species of scorpionfishes which are tropical in distribution, rockfishes inhabit cold temperate seas. They are found principally in the North Pacific from Japan to Mexico, although a few species can be found in the North Atlantic and in the temperate Southern Hemisphere (Eschmeyer and MHureau, 1971). Fifty species of rockfishes may occur at a single latitude with up to 10 species occurring at a single collecting station. This high degree of congeneric sympatry
1Present address: Department of Zoology, University of California, Berkeley, CA 94720. This study formed the basis of a Master’s thesis at California State University, San Francisco.
[59]
60 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
is of interest to community ecologists studying division of resources and isolating mechanisms.
Although rockfishes are important to commercial and sports fisheries, little is known about their biology. All rockfishes are believed to be generalized large- mouthed carnivores. They vary from benthic hole-dwelling solitary species to schooling species that live off the bottom in the kelp canopy or in open water. They are ovoviparous, giving birth to large numbers of yolk-sac larvae (Moser, 1967).
Field observations made in 1970 by the author indicated that several species can be stimulated to produce sounds under stress. Recent work by McInerney and Yearsley (in press) shows that rockfishes produce sounds during agonistic encounters. The sound producing mechanism consists of a pair of muscles, one on each side of the midline, originating on the occipital portion of the cranium and inserting on the gasbladder. Gasbladder muscles of this sort were first called ‘“‘extrinsic” by Dufosse (1874) because they inserted on the gasbladder at one end and elsewhere at the other end. Muscles with both points of insertion on the gasbladder wall, such as those found in triglids (Tower, 1908), were labeled by Dufosse as “intrinsic.” In this paper, muscles originating on the cranium and inserting on or near the gasbladder will be called extrinsic gasbladder muscles or, simply, gasbladder muscles.
The first major work which described these gasbladder muscles for a variety of scorpaenid fishes was that of Matsubara (1943). All the scorpaenids dissected by Matsubara have some type of muscle associated with the gas- bladder, although a variety of morphological patterns exist. For Oriental rockfishes, he showed that intrageneric variations in the structure of these muscles do exist. Dissections made on 82 species of rockfishes in the present study also reveal great variations in the patterns of the gasbladder muscles within Sebastes.
The species in the genus Sebastes are well known from several regional works (Matsubara, 1943; Phillips, 1957; Barsukov, 1964), but a worldwide review has not yet been attempted. One character that may be useful in an attempt to clarify phylogenetic relationships within this genus is the variation in the structure of the extrinsic gasbladder musculature.
As a result of the attempt to clarify the possible origin and evolution of the various rockfish muscle patterns, dissections were made on representatives from 23 other genera in the family Scorpaenidae and on fishes from 9 other families in the order Scorpaeniformes. The classification of groups within this order is uncertain (Greenwood ef al., 1966), and it is possible that an eventual overall survey of gasbladder musculature will aid in the understanding of phylogenetic relationships in the so-called mail-cheeked fishes.
Another character not surveyed in this study that may also prove useful
VoL. XL] HALLACHER: GASBLADDER MUSCLES 61
in clarifying phylogenetic relationships within Sebastes is variability in gas- bladder structure itself. The morphology of rockfish gasbladders appears to be diverse. Variability in the thickness of gasbladder walls, in attachment of the bladder, and in division of the bladder into two chambers is apparent (see also Matsubara, 1943, p. 126-147).
ACKNOWLEDGMENTS
Specimens used in this study were obtained from a variety of sources. Eastern Pacific material came principally from the California Academy of Sciences fish collection, although some material was obtained from the collection at California State University at San Diego and the Scripps Institution of Oceanography. Western Pacific material came from the California Academy of Sciences fish collection and from the Museum of Zoology of the University of Michigan.
Field collections were made during the course of this study to provide fresh material. I wish to thank the following people for their aid: Dr. George W. Barlow of the University of California at Berkeley who first advised the author of sound production by rockfishes; Dr. John S. Stevens of Occidental College and the crew of the research vessel Van Tuna, who provided specimens from Catalina Island off southern California; Dr. Lo-chai Chen of California State University at San Diego and the Scripps Institution of Oceanography for supplying a boat for collections made off La Jolla, California; Dr. Tomio Iwamoto of the California Academy of Sciences, who collected specimens off British Columbia from the research vessel G. B. Reed; Dr. William N. Eschmeyer of the California Academy of Sciences, who collected specimens off the Oregon coast from the Oregon State University vessel Vaquina; Dr. Eschmeyer, Dr. Chen, Christopher Tarp, Frederick Jones, and Ernest W. Iverson, who assisted in making collections from sportfishing boats; David W. Behrens, Kenneth R. McKaye, and divers from the University of California at Berkeley Diving Program, who aided the author in netting and spearing shallow water species; Daniel J. Miller and Robert N. Lea of the California Department of Fish and Game, who donated specimens; Dr. Reeve M. Bailey, Dr. Robert R. Miller, and staff of the Museum of Zoology of the University of Michigan, who provided the assistance to Dr. William N. Eschmeyer which resulted in the loan of Oriental specimens from their University collection; Dr. John E. McInerney and Dr. John H. Yearsley for allowing me to read their manuscript on sound production by rockfishes.
Additional help came from many persons. Dr. Richard Winterbottom of the National Museum of Canada provided information regarding nomenclature on the subject of gasbladder musculature. Daniel J. Miller, Robert N. Lea, Dr. Chen, Dr. Eschmeyer, and W. I. Follett assisted in identification of specimens. Pearl M. Sonoda, James E. Gordon, Beverly J. Wesemann, and David W. Behrens
62 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
provided curatorial assistance. Cherryl P. Pape assisted in the typing of the manuscript. Katherine Keeney Smith did all drawings.
This study was supported by National Science Foundation (NSF GB 34213), Dr. Chen and Dr. Eschmeyer co-principal investigators. The California Academy of Sciences provided working space and access to collections. Dr. Eschmeyer provided background information on scorpionfishes, and he, Dr. Chen, and Dr. Margaret G. Bradbury gave advice during the course of the study. I wish to thank the members of my thesis committee, Drs. Margaret G. Bradbury, William N. Eschmeyer, and Jerry W. Gerald, for their suggestions and comments. I would further like to acknowledge the following persons for their assistance in revising the manuscript for publication: Dr. Eschmeyer, Dr. Walter R. Courtenay, Lillian J. Dempster, and W. I. Follett.
METHODS
In this paper the term gasbladder has been applied to the structure more commonly referred to as the airbladder or swimbladder. This follows the terminology suggested by Lagler, Bardach, and Miller (1962). Since the gasbladder in rockfishes is physoclistous (no duct connects the gasbladder to the alimentary canal) and inflated by means of a gas gland, the term gasbladder seems more appropriate than either alternative expression. In accordance with this terminology, any specialized muscles associated with the gasbladder have been referred to by this author as gasbladder muscles rather than the more commonly used ‘swimbladder muscles’.
Extrinsic gasbladder muscles were exposed for examination by dissection. These muscles appear to have arisen from the ventral fibers of the epaxialis (the dorsal half of the lateral body musclature). All species have a pair of muscles, one on each side of the midline. They extend posteriorly from their origin on the occipital region of the cranium to the gasbladder or vertebral parapophyses where they insert by means of tendens. Since their position relative to skeletal structures is constant throughout the genus, one standard dissection technique was sufficient for all rockfish species and could be done with a scalpal and forceps. In preliminary dissections, the muscles from both sides of specimens were examined with no structural differences being observed. All subsequent dissections were made on the right side of specimens, with only the muscle on the right side being examined. A large patch of skin was removed, its perimeter extending along the rear of the cranium and pectoral girdle, posteriorly along the base of the spinous dorsal fin to its end, downward to the ventral half of the fish’s side and forward to the pectoral fin axil.
All exposed muscle tissue lying dorsal to the right dorsal ribs was removed. The right gasbladder muscle was exposed to view by gently breaking away the dorsal ribs. Care was taken while removing the first two dorsal ribs since the
Vor. XL] HALLACHER: GASBLADDER MUSCLES 63
gasbladder muscle lies just ventral to them. In most species of rockfishes the muscle passes between the second and third ventral ribs so that it is found on the visceral cavity side of the third through eighth ventral ribs.
Once the gasbladder muscle was exposed, all skeletal muscle tissue surrounding it was plucked away cautiously to expose the muscle from its point of origin on the cranium to its points of insertion on the gasbladder or vertebral parapophyses. The insertion points were determined by clearing away all intercostal muscle between the ventral ribs. Once the first eight or nine ribs were exposed, the dense tendon tissue of the gasbladder muscle was seen extending laterally just under the ribs. These whitish tendons were followed to their points of insertion on the vertebral parapophyses. In species where these tendons inserted first upon the gasbladder, an incision into the body cavity was made to examine posterior insertion sites.
Another method was sometimes used in species where the insertion points on the vertebral parapophyses were not readily determined from a dissection of lateral body musculature. The body cavity was entered from the ventral side and the viscera and the gasbladder were removed. Once the gasbladder was removed, the whitish tendons of the gasbladder muscle could be seen inserting on the vertebral parapophyses.
Dissections of other members of the order Scorpaeniformes were conducted in much the same manner since their gasbladder muscles were located in the same general area of the lateral skeletal musculature. The notable exceptions were members of the suborder Cottoidei. In species of this group, muscles termed “cranioclavical muscles” by Barber and Mowbray (1956) extend from the posterior cranium to the pectoral girdle. To expose these cranioclavical muscles, only musculature situated directly behind the cranium was removed.
At the onset of the study, radiographs were taken of dissected specimens to clarify the topographic relationships of the origin and insertion points of the extrinsic gasbladder musculature. Once the anatomy of these fishes became familiar, radiography was no longer required for the majority of species.
Specimens used in the study are listed in the appendix.
RESULTS
Dissection of 82 species of rockfishes reveals considerable variation among species in gross morphology of the extrinsic gasbladder musculature. In species where both sexes were examined no sexual dimorphism was observed. Several insertion points may be listed for one muscle because of the fact that in most cases, the striated muscle tissue of the gasbladder muscle ends in one to several tendons after the muscle passes under the third rib. In several species, only one or two tendons develop from the muscle, but later branch to insert on several different vertebral parapophyses. Table 1 summarises origin and insertion points for the gasbladder muscles of all rockfishes examined,
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TasLe 1. Muscle type and points of insertion of the gasbladder muscle for all species of Sebastes examined. Where more than one specimen of a species was examined, the information on each specimen examined is listed. Abbreviations for the points of insertion are as follows: gb represents the gasbladder, r-1 thru r-4 represent the first through the fourth ventral ribs, and v-7 through v-11 represent the seventh through the eleventh vertebrae. Muscle category includes major type (either Type I or Type II) and the subdivision within a type: a-3 = aleutianus-zacentrus, d-v = atrovirens-vexillaris, i-v =ijimae-vulpes, p = paucispinis, s = serriceps, t = taczanowskii, m = marinus.
Points of Insertion
Muscle Species Category gb r-l r-2 r-3 r-4 v-7 v-8 v-9 v-10—v-11 aleutianus I a-z x x x alutus I a-z x Xx x x " I a-z x x Xx x x auriculatus I a-z x x >< " I a-z x x x " I a-z x x x aurora I a-z x x x atrovirens Il a-v xX x x x " Il a-v x x x babcocki I a-z x x< x baramenuke I a-z x x x Xx borealis I a-z x x x brevis pinis I a-z x x x capensis I a-z x x carnatus Jul aay S< x x x caurinus jul way SK x x " If a-v x x x x chlorostictus I a-z x s< x chrysomelas UL A? SK x x ciliatus leal=z, < x x x constellatus I a-z x x x crameri I a-z x x x dallii I a-z x x x " I a-z x Xx x diploproa I a-z x x x elongatus I a-z x x x x em phaeus I a-z x " T a-z x x x ensifer I a-z x x entomelas I a-z x x eos I a-z < ~< exsul I a-z x x x x flameus It a% x x Xx flavidus I a-z x x x gilli I a-z x < x glaucus * az, < < <x goodei T a-z x x x
* Not examined. Information from Matsubara (1943).
VoL. XL] HALLACHER: GASBLADDER MUSCLES 65 TABLE 1. Continued Points of Insertion Muscle = Species Category gb r-1 r-2 r-3 r-4 v-7 y-8 v-9 v-10—-v-1l helvomaculatus I a-z x x x '" I a-z x x <x hopkinsi I a-z x x hubbsi I a-z x x ijimae II i-v x x imermis II i-v x x x x " II i-v x x x x ivacundus * I a-z x x x itinus * I a-z x x x jordani iga=Z x x joynert I a-z x x x Rawaradae * I a-z x x x x lentiginosus I a-z x x x levis I a-z x x x x x longis pinis I a-z XK x x macdonaldi I a-z >< x< xX xX x x x maliger Il Bay SK Xx x " lh asy S< x x marinus Im x x x x " Im x x x x " Im x x x matsubarae I a-z x x x x x x melanops I a-z x x x melanostictus * I a-z x x x x melanostomus I a-z x x x x x " IL OY x x x miniatus I a-z x x x mystinus I a-z See Table 2 nebulosus lita=v, xX x x nigrocinctus I a-z x x x NLVOSUS II i-v x x oblongus II i-v x x ovalis ita=z x x x x owstonti I a-z x x x pachycephalus II i-v x x x paucis pinis II p x x " II p x x phillipsi iga=Z x x pinniger I a-z x x x x proriger I a-z x x x x rastrelliger I a-z x x reedi az x x xX x rosaceus I a-z x x x rosenblatti I a-z x x x
* Not examined. Information from Matsubara (1943).
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TABLE 1. Continued
Points of Insertion Muscle = — ew Ese -
Species Category gb r-1 r-2 r-3 r-4 v-7 v-8 v-9 v-10_—v-1l ruberrimus I a-z x x x x rubrivinctus I a-z x x x x rufus I a-z x x x x saxicola IWwaez x x x schlegelii II i-v x x x
" II i-v x x x scythro pus I a-z Points of insertion decayed serranoides a=z x x x service ps TS; x x simulator I a-z x x smmensis I a-z x x steindachneri az x x taczanowskii 100% x< x x x thompsoni II i-v x x x trivittatus Il i-v. x x x umbrosus I a-z x x x variegatus lia=Z, x x x x vexillaris lita x x vulpes II i-v x x wakiya * I a-z x x x wilsoni I a=z x x sacentrus I a=z x x x
x x
" liga=7 x
* Not examined. Information from Matsubara (1943).
For the purpose of determining the extent of variation within a species, dissections were done on 17 specimens of the blue rockfish, Sebastes mystinus. These dissections revealed only slight variations in the size and shape of the extrinsic gasbladder muscles, but they did indicate great intraspecific variation in posterior insertion points (table 2). No apparent sexual dimorphism was found. The size range of this series, consisting of eight males and nine females, was 70 mm. to 376 mm.
On the basis of morphological variations, two basic divisions in gasbladder muscle structure are recognized for the genus Sebastes. These divisions have arbitrarily been designated Type I and Type II. Type I consists of all muscle variations that attach firmly to the pectoral girdle as they pass posteriorly from their origin on the occipital cranium to their points of insertion on the vertebrae (fig. 1). Type II is comprised of muscle variations that bypass the pectoral girdle as they pass posteriorly from their origin on the occipital cranium to their points of insertion on the gasbladder or vertebrae (fig. 2). Each division is further subdivided into subtypes on the basis of additional structural variations.
Vor. XL] HALLACHER: GASBLADDER MUSCLES 67
TaBlE 2. Muscle type and points of insertion of the gasbladder muscle for 17 specimens of the blue rockfish, Sebastes mystinus. Sex and standard length (S.L.) are listed for each specimen. Abbreviations for the points of insertion are the same as in table 1.
Points of
BHA at Insertion Insertion Sex sess. Muscle v-7 v-8 v-9 v-10 Stee Selb, Muscle v7 v-8 v-9 v-10 e PIP simon, IL Bey x x OF i 2mm la=z < x Gucolemm..) lea=z x > x OF is mma li a=z < < & 25il warn, Il we 4 x x ® 191 mm. I a-z x< K Cue Samim Lea=z x< x © 264 mm: 1 a=z < x< K CmecoOhmm=. 1 a=z SK OF 265 mm. Wl a=z SK 6 26/7 mm. I a-z x >< x< OF 294 mmr i a=z < x CueaSemm, 1 aqz x x OD 347 amma a=z x x< Gzs9 mm. Il a=z x x © 376 mm. I a=z x fe) 70 mm. I a-z < <
Type I contains three subdivisions that are nominally denoted: aleutianus- zacentrus, serriceps, and marinus. Type II contains four subdivisions: paucispinis, ijimae-vulpes, atrovirens-vexillaris, and taczanowskii.
Subdivision nomenclature was determined by simply listing all species with a particular type of musculature in alphabetical order. The first and last species names on the list were taken to denote the subdivision name for that particular muscle type. Hence, the aleutianus-zacentrus subdivision is the pattern in all species in alphabetical order from Sebastes aleutianus through Sebastes zacentrus that possess dorsoventrally flattened muscles, originating on the cranium, attaching to the cleithrum, and inserting on the vertebral parapophyses. Four of the subdivisions are found in only one species each, so only one species name is listed for each of these four systems.
Type I
All species with musculature of this major category have extrinsic gasbladder muscles that attach firmly to the pectoral girdle as the muscles pass posteriorly from their origin on the occipital cranium to their insertion points on the vertebral parapophyses. Subdivisions of this major category have been formed on the basis of additional structural variations.
ALEUTIANUS-ZACENTRUS SUBDIVISION. This was the basic muscle structure found in the genus Sebastes with 62 of the 82 species dissected having this morphology. (See fig. 1A). Unlike some of the other muscle types which are found only in one geographic region, this type occurred in species taken throughout the geographic range of the genus.
The gasbladder muscles of this group are dorsoventrally flattened with the point of origin being the cranium. The specific site of attachment to the cranium is only slightly variable, and is usually around the exoccipital-opisthotic suture,
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‘Y{BUI] PAVpue]s “WUT SOZ ‘sNULIDU Sajspgag WOT] epeU BUIMBICT “pa}assIp Sajspqgag’ JO satoeds Tg 19Y}O [[B UL SqII psy pue puosas ay} usaeMjaq 0} pasoddo se sqii YANoy pue PAY} oY} WoaMyzoq Suissed ogposnur (3) sutmoys adAjqns snulieul JO MaIA [eso :Q ‘ySue_ plepueys “wu [¢7Z ‘sdaId4as Sajspqag Woy speu SuMBIqE “avIqa}1aA yUaAes ayy Jo sisXydodeied 9y} UO SuUTjIasUt UOpUdy gjzuis oy} (J) pue ‘sqLI oy} 0} aposnw pazelTys Jo JUoUTYDe}e (2) Surmoys adAyqns sdaotites JO MIA [eSIOd : “Y}suUs] Plvpuvjs “WU SOT ‘“asyjasqsvA Sajspgag WO] speW SUIMLIG: “JUSWIRST] sjofepneg (Pp) pure ‘gIp4IS [erojoed 0} JUaWYIeWY (9) ‘UoT10d Uopus} (q) ‘Uor}10d 9posnu payeiys (8) Surmoys sapsnw Jappryqses pue uwnyod [ev1qo}19A adAqqns snajuacez-snuennege JO MaIA [esIoq :W ‘sedAjqns asoy} Ul Japprlgses oy} 0} yor}e JOU Op suOpUa} IY, “9P1IG2}19A IO SqII 9Y} UO UONAssUT IToYy} 0} WinTURID [eqidio90 9y} UO UTZIIO Mey} Wor Ap1o119}sod ssed sojosnu ay} Sse a[pas [etojood ay} 0} JUSWIYDLIAB IT}sMojOeIBYO Surmoys (A[UO opis YS) saosnur sappeqses J addy, sayspqag “T AYIA
Vor. XL] HALLACHER: GASBLADDER MUSCLES 69
although in some species the attachment site seems wholly on one or the other of these two bones, or on the opisthotic. As the muscle passes posteriorly, some of its fibers attach to the supracleithral bone near the distal attachment site of Baudelot’s ligament. Insertion by tendons occurs on the ribs or vertebral parapophyses.
The points of insertion, which tend to vary interspecifically, are listed in table 1 for all 62 species included in this muscle category. Unless otherwise stated in table 1, only one specimen of each species was dissected. The species of Sebastes examined, listed alphabetically, are as follows: aleutianus, alutus, auriculatus, aurora, babcocki, baramenuke, borealis, brevispinis, capensis, chlorostictus, ciliatus, constellatus, crameri, dallii, diploproa, elongatus, emphaeus, enstfer, entomelas, eos, exsul, flameus, flavidus, gilli, goodei, helvomaculatus, hopkinsi, hubbsi, jordani, joyneri, lentiginosus, levis, longispinis, macdonaldi, matsubarae, melanops, melanostomus, miniatus, mystinus, nigrocinctus, ovalis, owstoni, phillipsi, pinniger, proriger, rastrelliger, reedi, rosaceus, rosenblatti, ruberrimus, rubrivinctus, rufus, saxicola, scythropus, serranoides, simulator, sinensis, steindachneri, umbrosus, variegatus, wilsoni, and zacentrus.
Species of Sebastes dissected by Matsubara (1943) possessing this type of musculature are as follows: baramenuke, flameus, hubbsi, longispinis, matsubarae, owstoni, scythropus, and steindachneri: plus those examined by Matsubara but not by the author (Matsubara’s results are included in table 1): glaucus, iracundus, itinis, kawaradae, melanostictus, and wakijai.
SERRICEPS SUBDIVISION. This muscle arrangement is found only in the treefish, Sebastes serriceps, an eastern Pacific species. The gasbladder muscles of this species are very similar to those of the aleutianus-zacentrus subdivision, being dorsoventrally flattened muscles with the point of origin on the cranium. As the muscles in this species pass posteriorly, a portion of the muscle fibers attach to the supracleithral bone. The mode of insertion of the gasbladder muscles in S. serriceps is however quite different from that found in the species listed in the aleutianus-zacentrus group. In the treefish, the posterior margin of the gasbladder muscle inserts primarily on the second rib, without first becoming a tendon. Careful examination revealed one small tendon coming off the muscle that passed between the second and third ventral ribs and inserted to the parapophysis of the seventh vertebra (see fig. 1B).
MARINUS SUBDIVISION.! The extrinsic gasbladder muscles of Sebastes marinus are similar in overall structure to those found in the aleutianus-zacentrus group (fig. 1C). Each muscle originates on the occipital region of the cranium and attaches to the pectoral girdle as it passes posteriorly to insert by means of tendons to the vertebral parapophyses. The major difference between this species and all other rockfishes is that the muscle crosses between the third and fourth
1 See addendum.
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‘YqSue] plvpueys “ww ZZ] ‘wysmounzID} sayspqag WOT} spell SuIMVIG *9v1Gd}10A YWUaAVTA pure ‘y}U94 ‘yqurU ay} Jo saskydodezed ay} 03 yAasut yeyy Suopue} JOYS 9214} 94} (P) pue “Iappelqses ay} 0} ATJaIIpP ansst} sPsNUr pozelyys 94} JO JUouUYyse}e 9Yy} (9) Surmoys adAqqGns IDsMOUezZIRY JO MOIA [eSIOd :( “YyIsuZ] plepuejs “wu O¢E ‘snuzNDI Sajspgag WO} ape Surmeiq, ‘sashydodesed [eaqaz19a 94} 0} suopus} 9Y} JO UOTZAaSUT (q) pue ‘1appe[qses ay} 0} uoNs0d uopua} 94} JO JUSWUYIEL 94} (B) Surmoys adAjqns sie[[IxaA-suattAosqe JO MOIIA [BSIOdG :D “Y}sue] paepurys “WU TOT ‘sagjna saqspqag wory spew Summeiq ‘sarads 994} Ul Jeppe[qses oy} 0} JUIWIYIEI}e ON “avAqazIVA 24} UO S}1osuT Jey} UOTI0d uopuey ay} (9) Surmoys adajqns sadjna-aeun{t yo Mara [esiod :q ‘“Yyisue] prepuerjs “wu 6¢7 ‘seuzdsiannd Saqspgagy WOIf apeul SUIMPIC: ‘sjird OM} OFUT oNss}} IPSNU Pozel4}S 94} SOplAIp jeYy} vlosey (p) pue uowesT] sjojapneg (2) “P1G2}19A Y}U9} IY} JO sisXydodesed ay} 0} uopusy sty} Jo UOT}JaSUT 94} (q) “Iappelqses sy} 0} JUIWYIeI}e S,UOpUs} I[SUIs ay} (v) SutMmoys aposnur Jappe[qses pue uuimjoo [eiqoy19a adxjqns stutdstoned JO MOA [esIOod :W ‘Joppr[qses 10 aviqaziaA ay} UO UOT}JaSUT Toy} 0} wWntueId [epdI990 oY} UO UISIIO May} WOIY ATIOII0}s0d ssed saposnu 94} SB a[pils y[e10j29d oy} Jo ssedAq OYSHO}IVILYI IY} SuIMoys (ATUO apis }YSII) sapsnur Jopperqses J] eA], sayspqag °z aunorg
VoL. XL] HALLACHER: GASBLADDER MUSCLES 71
ventral ribs as it passes posteriorly towards the vertebrae, instead of passing between the second and third ventral ribs as in all other rockfish species examined. See table 1 for points of insertion in all Sebastes marinus dissected.
Type II
All species having muscular of this major category have extrinsic gasbladder muscles that bypass the pectoral girdle as the muscles pass posteriorly from their origin on the occipital cranium to their insertion points on the gasbladder or vertebral parapophyses. The muscles of this category tend to be more massive and cylindrical than those of the Type I category. Subdivisions of this Type II category are delineated by the author on the basis of additional structural variations.
PAUCISPINIS SUBDIVISION. This muscle arrangement is found only in the bocaccio, Sebastes paucispinis, an eastern Pacific species. In this species the gasbladder muscles have the point of origin on the cranium. As the muscles pass posteriorly, they do not make a connection with the pectoral girdle. This species, unlike any others in the genus, has the striated muscle tissue of the gasbladder muscles divided into two parts by a thin fascia (fig. 2A). The gasbladder muscles insert, each by means of a single tendon, on the posterior portion of the gasbladder. This tendon, while firmly fastened to the gasbladder wall, curves dorsally to anchor the gasbladder to the parapophysis of the tenth vertebra.
IJIMAE-VULPES SUBDIVISION. The gasbladder muscle pattern included in this group was found in nine species, all from the Orient. In these fishes, as in the rest of the genus, the extrinsic gasbladder muscles have the area of origin on the cranium, near the suture of the opisthotic and exoccipital. Like the muscles found in Sebastes paucispinis, these do not make a connection to the pectoral girdle as they pass posteriorly. The muscles have a striated portion that is slightly more massive than those described in previous groups. They are not quite as broad as the dorsoventrally flattened muscles of the aleutianus-zacentrus or serriceps subdivisions, but are of greater girth, being more cylindrical in shape. They appear to insert by tendons to the vertebral parapophyses without first attaching to the gasbladder (fig. 2B). Points of insertion for all species in this group are listed in table 1. The species of Sebastes falling into this group are as follows: ijimac, inermis, nivosus, oblongus, pachycephalus, schlegeli, thompsoni, trivittatus, and vulpes.
ATROVIRENS-VEXILLARIS SUBDIVISION. The muscles of this grouping and the next (taczanowskii subdivision) are the largest in the genus Sebastes. These gasbladder muscles originate on the cranium. As they pass posteriorly they do not make a connection to the pectoral girdle. They insert as tendons that attach firmly to the gasbladder before swinging up and attaching to the vertebral parapophyses (figs. 2C and 3). In several of the species, one of the two tendons
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poyenys [BoLIpurpAd 9
Q Oo
Vot. XL] HALLACHER: GASBLADDER MUSCLES 73
initially attaching to the gasbladder diverges so that three tendons are firmly attached to the gasbladder wall and anchor the gasbladder to three vertebral parapophyses. Points of insertion for all species in this group are listed in table 1. The species of Sebastes characterized by this muscle pattern are as follows: atrovirens, carnatus, caurinus, chrysomelas, maliger, nebulosus, and vexillaris.
TACZANOWSKII SUBDIVISION. This type is represented by one species, Sebastes taczanowskiu, an Oriental form. It is much like that of the previous group (atrovirens-vexillaris) except that the muscle attaches to the gasbladder wall as muscle rather than as tendon. Originating on the cranium, the muscle passes posteriorly to attach to the outer wall of the gasbladder. No connection to the pectoral girdle occurs. While connected to the gasbladder, the muscle passes posteriorly along the gasbladder to a point approximately under the sixth rib where it then diverges to form three tendons. These three tendons, while firmly attached to the gasbladder wall, curve dorsally to anchor the gasbladder to the parapophyses of the ninth, tenth, and eleventh vertebrae (fig. 2D).
OTHER SCORPAENIFORM DISSECTIONS
Dissection of specimens from 23 other genera within the family Scorpaenidae and nine other families within the order shows many to have muscles similar to the extrinsic gasbladder muscles found in the genus Sebastes. As might be expected, the highest degree of similarity in structure of these muscles occurs within families, but general similarity in structure does occur even between some families in different suborders, as is the case of the first group listed below. Dissections of scorpaeniform representatives were made primarily for general comparison and were not as precise as those done on rockfish specimens.
MUSCLE TYPES SIMILAR TO SEBASTES TypE I. All fishes placed in this group have two lateral muscles similar in structure to those found in Type I muscles as defined for the genus Sebastes. This structural form is the most widespread and common in the order, being predominant in four of the five suborders examined. The notable exceptions were members of the suborder Cottoidei.
The species listed below possess muscles that are dorsoventrally flattened like those in the Sebastes Type I category. These muscles, like those in the rockfish group have the point of origin on the occipital cranium. As _ the muscles pass posteriorly, they make a connection with the pectoral girdle. The points of insertion in all cases appeared to be the vertebral parapophyses. Unless the precise points of insertion were determined, however, species in the group are simply listed. The species examined are as follows:
Scorpaenidae: Ectreposebastes imus; Gymnapistes marmoratus; Helicolenus dact yl-
opterus; Inimicus cuvieri; Iracundus signifer; Minous monodactylus; Minous pictus; Neomerinthe beanorum; Parascorpaena species; Plectrogenitum nanum;
74 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 4, Other types of gasbladder musculature found in the order Scorpaeniformes. A: Dorsal view of the right extrinsic gasbladder muscle of Sebastiscus marmoratus showing (a) the broadened striated muscle tissue that inserts directly onto the gasbladder, (b) the pectoral girdle, and (c) Baudelot’s ligament. This muscle system is similar in morphology to the Sebastes Type II muscles. Drawing from specimen 131 mm. standard length. B: Dorsal view of gasbladder from A pistes species showing (d) the fully intrinsic muscle tissue on the anterior lobe of the gasbladder. Drawing made from specimen 109 mm. standard length. C: Dorsal view of right cranioclavical muscle in Cottus asper showing (e) the short striated muscle band. Drawing from specimen 110 mm. standard length.
Pontinus longispinis; Scorpaena agassizi, Scorpaena albobrunnea, Scorpaena brasiliensis, Scorpaena elongata, Scorpaena guttata, Scorpaena mystes, Scorpaena porcus, and Scorpaena russula, all with four tendons, one attaching to each of the parapophyses of the sixth, seventh, eighth, and ninth vertebrae; Scorpaenopsis species; Sebastosemus species; Setarches longimanus; Synanceia verrucosus.
Hexagrammidae: Hexagrammos decagrammus
Platycephalidae: Platycephalus species
Hoplichthyidae: Hoplichthys langsdorfi
MUSCLE TYPES SIMILAR TO SEBASTES TYPE II (TACZANOWSKII SUBDIVISION).
The species listed in this group have two large gasbladder muscles similar in over- all structure to the muscle system listed for Sebastes taczanowsku. These muscles are in most cases rather large and cylindrical, originating on the occipital portion of the cranium and inserting usually on the bladder (fig. 44). No connection to the pectoral girdle occurs. Species listed below are from the family Scorpaenidae. Once again, dissections were not as exact as for the genus Sebastes, and precise posterior attachment sites were not exactly determined. The species listed in this group are as follows: Scorpaenidae: Brachypterois serrulifer, Dendrochirus zebra, Macroscorpius pallidus,
Scorpaenodes parvipinnis, Pterois radiata, Sebastiscus marmoratus, Setarches guentheri.
VoL. XL] HALLACHER: GASBLADDER MUSCLES 75
INTRINSIC GASBLADDER MUSCLES
Dissections revealing gasbladders containing fully intrinsic gasbladder muscles were made on two species, Leptotrigla alata (Triglidae) and A pistes species (Scorpaenidae). Intrinsic gasbladder muscles have been widely examined in triglids (Tower, 1908; Fish, 1954; Tavolga, 1964; Evans, 1973), and Matsubara (1943) in his review of Japanese scorpaenids reported intrinsic musculature for A pistes carinatus. Figure 4B is a diagram of an A pistes gas- bladder.
CRANIOCLAVICAL MUSCLE SYSTEMS
The muscles of this group, found in members of the suborder Cottoidei, are stout cylindrical muscles connecting the pectoral girdle to the occipital region of the skull (fig. 4C). The point of attachment to the skull seems to correspond to the point of attachment of the extrinsic gasbladder muscles described for scorpaenids. Likewise, the point of attachment to the pectoral girdle seems to correspond to that in the scorpaenids. Species dissected are as follows:
Zaniolepididae: Zanzole pis latipinnis
Cottidae: Clinocottus analis, Cottus asper, Hemilepidotus jordani, Leptocottus
armatus, Myoxcephalus quadricornis, Scorpaenichthys marmoratus
Cottuniculidae: Cottunculus thompsoni
Agonidae: Occella verrucosa Cyclopteridae: Liparis florae.
DISCUSSION
TAXONOMIC VALUE OF GASBLADDER MUSCLE SYSTEMS. In all rockfish species examined, only one mode of origin for the gasbladder muscles was observed, that being on the occipital region of the cranium. In some species fibers of the striated muscle tissue attached to the supracleithral bone as the gasbladder muscles passed posteriorly, while in others no such attachment occurred. Two major muscle categories were recognized on the basis of the presence (Type I) or absence (Type II) of a pectoral girdle attachment. A total of seven subdivisions were recognized, three for Type I and four for Type II, on the basis of additional structural variations.
Because of the high degree of intraspecific variability (see table 2) and interspecific overlap in insertion sites (see table 1), these muscles were in most cases not useful in species delineations. However, of the 82 species examined, four had unique muscle types. These species are Sebastes serriceps, (Type 1), S. marinus (Type I), S. paucipinis (Type IL), and S. taczanowsku (Type II), and could easily be separated from other species on the basis of the morphology of their gasbladder muscles.
The use of these various muscle types in conjunction with other characters as a means of delineating subgenera seems in some instances useful. I’or example,
76 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
in the eastern Pacific subgenus Pteropodus, all species with the exception of one, had gasbladder muscles that were found only in this small group (atrovirens- vexillaris subdivision of Type II). Muscle morphology seems to support the subgeneric classification for the Pteropodus group. The one species in the Pteropodus complex that did not possess an atrovirens-vexillaris muscle system was S. rastrelliger. It had instead the more common aleutianus-zacentrus bladder muscles (Type I). Workers dealing with phylogenetic relationships in Sebastes should perhaps consider separating S. rastrelliger from the other species (S. atrovirens, S. carnatus, S. caurinus, S. chrysomelas, S. maliger, S. nebulosus, and S. vexillaris) in this group.
Two species of the subgenus Sebastodes, S. brevispinis and S. paucispinis, which are usually considered to be closely related, have major differences in the structure of the gasbladder musculature. Sebastodes brevispinis has the aleutianus-zacentrus muscles (Type I), and S. paucispinis (Type IL) possesses a unique form. These species seem to occur in similar habitats, so their general similarity may indicate ecological convergence from different ancestral lines.
The considerations given to the taxonomy of eastern Pacific subgenera in this section are in fact only brief speculations. Rearrangement of eastern Pacific subgenera lies far beyond the scope of this paper. Speculations about Oriental subgenera were completely omitted due to insufficient numbers of dissections on Japanese forms. It is probable that other workers, correlating the structure of gasbladder muscles with a variety of other taxonomic characters, will find the morphology of these gasbladder muscles useful in phylogenetic studies on the rockfish genus Sebastes.
ORIGIN AND EVOLUTION OF GASBLADDER MUSCULATURE WITHIN THE GENUS SEBASTES. In the course of a study in which one particular character is reviewed throughout a group, it is worthwhile to give consideration to the possible evolu- tionary origin and diversification of the structure concerned. The first worker to extensively examine the extrinsic gasbladder musculature of scorpaenids was Matsubara (1943). His hypothesis concerning the successive changes of the muscle bands was that ‘‘. . . the existence of well developed muscle is a primitive feature in the scorpaenoid fish.” Matsubara speculated that species such as Sebastes taczanowskii possessing well developed extrinsic gasbladder muscles represented the ancestral muscle condition. Conversely, species having muscles small in size and connected posteriorly to the axial skeleton by tendons as in the aleutianus-zacentrus subdivision were thought by him to possess the more recent evolutionary structure.
It is possible that these structures may have followed a mode of evolutionary differentiation opposite to that shown by Matsubara. This speculation is based upon the preponderance of small gasbladder muscles throughout the order.
If one assumes that the structural condition of the gasbladder muscle occurring
Vout. XL] HALLACHER: GASBLADDER MUSCLES
~r ~r
most frequently in these rockfishes is the type most likely to be the primitive state for the genus, then the Type I division, found in 62 of the 82 species examined, is representative of the ancestral condition for the genus. The gasbladder muscles in these species are dorsoventrally flattened, originating on the cranium, passing posteriorly to make a strong attachment to the pectoral girdle, and finally inserting by tendons to ribs or vertebrae. The other structural variants (Type II) which occurred much less frequently are probably derived states.
However, another interpretation of the widespread occurrence of the Type I muscle pattern is that this pattern arose originally from one of the less frequently occurring patterns. This seems unlikely, though, in view of the widespread occurrence of this Type I pattern throughout the rest of the family. A total of 24 species belonging to 15 genera in the family Scorpaenidae also display a similar structural condition. This suggests that the pattern did not arise in the genus Sebastes and that it is the primitive condition for the family.
The selective pressures that have caused this divergence of gasbladder muscle structure within the rockfishes are unknown. The work of McInerney and Yearsley (in press) indicates that rockfishes possessing the smaller gasbladder muscles that attach only to the ribs or vertebrae produce sounds that are very similar to those produced by rockfishes possessing the larger gasbladder muscles that attach directly to the gasbladder. In light of this information it does not seem probable that selection for increased sound producing ability was an important evolutionary force involved in the divergence of gas- bladder muscles within this lineage, although it may have been an important factor in the general elongation of gasbladder musculature in this group. What factors were involved in their subsequent divergence is a question open to speculation.
SPECULATIONS ON THE ORIGIN AND EVOLUTION OF GASBLADDER MUSCULATURE WITHIN THE ORDER SCORPAENIFORMES. It is of interest to look beyond the generic level to speculate on the possible stages of evolution of gasbladder musculature within the order Scorpaeniformes. The muscle pattern of most widespread occurrence in the genus Sebastes and the family Scorpaenidae is that in which the muscles are dorsoventrally flattened, inserting on the vertebrae by tendons. This muscle pattern (similar to the Sebastes Type I) is found in four of the six suborders recognized by Greenwood et al. (1966) for the order Scorpaeniformes. Again if one assumes that the structural condition of the gasbladder muscles that occurs most frequently in this group is the type most likely to be the primitive state for the order, then the dorsoventrally flattened muscles that insert to ribs of vertebrae by tendons are representative of the ancestral condition for the order.
A muscle condition exists within the order, however, that complicates the
78 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
picture somewhat. All species examined from the suborder Cottoidei were found to possess lateral musculature different in structure from the long gasbladder muscles found in other scorpaeniform fishes. They possess, instead, two stout cylindrical muscles that originate on the occipital portion of the cranium and insert on the pectoral girdle. These structures were referred to by Barber and Mowbray (1956) as cranioclavical muscles (fig. 4c), and these workers demonstrated that sounds are produced when these structures are vibrated. The points of attachment to the cranium and pectoral girdle are similar to those points of attachment of gasbladder muscles in scorpaenids. Both systems seem to have evolved from modifications of the ventral fibers of the epaxialis, the dorsal half of the lateral body musculature. Homologies between the cranioclavical muscles of cottoids and the elongate muscles of other scorpaeniform fishes are not clear.
A variety of possible evolutionary schemes can be envisioned to explain possible differences between cottoid musculature and that found in other scorpaeniform groups. For example, the members of the suborder Cottoidei may have retained muscles similar to original ancestoral structures from which other scorpaeniform groups eventually developed elongate gasbladder muscles. What selection pressures might be involved in this process of muscle elongation are not clear. Perhaps the lengthening of lateral musculature increased mobility of the pectoral girdle in scorpaenidlike fishes, aiding them in movements associated with swimming or moving around on the bottom. After this primary elongation, the gasbladder muscles in some species increased in size possibly to become efficient sound-producing structures.
It is possible, however, that the situation may have been the opposite. The loss of the gasbladder in members of the suborder Cottoidei suggests that they may be derived forms. Perhaps cottoid ancestors once possessed elongate gasbladder muscles similar to those in scorpaenids. With the loss of the gasbladder, this group of benthic fishes may have evolved a modified pair of muscles that are in some way adaptive to a benthic existence. This picture, in which cottids evolved from a scorpaenoid ancestor, coincides with that proposed by Quast (1965) on the basis of osteological evidence.
Another possibility is that the common ancestor of cottoids and scorpaenids had some sort of gasbladder musculature different from that found in either of the present-day groups. It is also possible that this major difference in gasbladder musculature between the cottoids and the scorpaenids reflects different phylogenetic origins for these two groups. In any event, the picture is at present hazy, and none of the previous hypotheses can be firmly defended. For the present this author must agree with the views expressed by Greenwood et al. (1966) and Gosline (1971). The former stated, ‘‘The scorpaeniform fishes represent a more or less typical example of the work that needs still to be done.”’;
Vor. XL] HALLACHER: GASBLADDER MUSCLES 79
the latter, “The classification of the group appears to be in an advanced state of confusion.”
FUNCTION OF THE EXTRINSIC GASBLADDER MUSCLE IN SEBASTES. The function of extrinsic and intrinsic gasbladder musculature in a wide variety of fish groups is that of sound production. Two review papers covering sound- producing mechanisms in fishes are those by Tavolga (1971) and Courtenay GLO 7Ly:
The morphology of extrinsic gasbladder musculature in rockfishes suggests that most species are probably capable of producing sounds. Sound production has been reported for the western Pacific species, Sebastes schlegeli (Protasov et al., 1965). Sounds have recently been recorded and analyzed for nine eastern Pacific forms, Sebastes caurinus, S. flavidus, S. maliger, S. melanops, S. mystinus, S. nebulosus, S. nigrocinctus, and S. paucispinis (McInerney and Yearsley, in press). These workers found that sounds were commonly produced during agonistic encounters between conspecifics, both in the laboratory and under natural conditions in the field. This author has heard sounds produced in the field by Sebastes atrovirens, S. carnatus, S. caurinus, S. chrysomelas, and S. nebulosus, but only under unnatural conditions of stress.
The morphology of gasbladder musculature in several other scorpaenid species suggests that these fishes may also produce sounds. Sound production has been reported in Sebastiscus marmoratus (Dotu, 1951), a species which possesses extrinsic gasbladder muscles that originate on the occipital region of the cranium and insert upon the swimbladder wall (fig. 4A).
The gasbladder muscles of rockfishes and other scorpaenids may also aid in sound reception. This seems possible in view of the large otoliths possessed by rockfishes, as well as the origin of the gasbladder muscles on the cranium in the general proximity of the otoliths. Whether rockfishes are receptive to sounds is not known at this time, and work remains to be done in this regard.
POSSIBLE ECOLOGICAL SIGNIFICANCES OF SOUND PRODUCTION. Field observa- tions made by the author indicate that some species of the subgenus Pteropodus are territorial at least part of the year, unlike other rockfishes occurring sympatrically. These species are usually solitary although field observations by the author suggests that they aggregate in pairs or small intraspecific groups during certain times of the year. The work of McInerney and Yearsley (in press) indicates that these territorial, hole-dwelling fishes, which possess the large Type II atrovirens-vexillaris gasbladder muscles (see figs. 2C and 3), produce sound during territorial defense, and as a fright response. Such uses of sound have been demonstrated for squirrelfishes (Winn and Marshall, 1963; Bright, 1972), and toadfishes (Gray and Winn, 1961; Winn, 1964, 1967).
Sounds produced by rockfishes might be species-specific and thus serve as isolating mechanisms between sympatric species. A high degree of congeneric
80 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
sympatry exists among rockfishes, with as many as 10 species being captured at one collecting station. One method used in species isolation might be species-specific sound emissions. The work of McInerney and Yearsley indicates that this is possible, but interestingly, they found that structurally similar species produce sounds that appear to be indistinguishable from each other. Additional audio-isolation studies by species of Sebastes would be valuable.
SUMMARY
Rockfishes from the large scorpionfish genus Sebastes have been studied only regionally. One morphological feature that may prove useful for clarification of phylogenetic relationships within Sebastes is variation in the structure of extrinsic gasbladder musculature. Eighty-two species of rockfishes were dissected for examination of their gasbladder muscles. For the purpose of clarifying gasbladder muscle relationships within the genus, dissections were made on representatives from 23 other genera in the family Scorpaenidae, and fishes from 9 other families in the order Scorpaeniformes.
Dissections revealed considerable intrageneric variation in gross morphology of the extrinsic gasbladder musculature. On the basis of these structural variations, two major structural divisions in gasbladder musculature have been recognized for the rockfish genus Sebastes, and have been designated Type I and Type II. Type I have been divided into 3 subdivisions (aleutianus- zacentrus, serriceps, and marinus) and Type II has been divided into 4 subdivisions, (paucispinis, ijimae-vulpes, atrovirens-vexillaris, and taczanowskii).
In most cases these muscles were not especially useful for characterizing species because of a high degree of intraspecific variability and interspecific overlap in insertion sites. However, four species (Sebastes marinus, S. paucispinis, S. serriceps, and S. taczanowskii) had species-specific gasbladder musculature, and therefore could be separated from other rockfish species on the basis of morphology of their gasbladder muscles.
Delineations of subgenera through the use of the various muscle types is in most cases not practical since the muscle patterns exhibit considerable overlap between proposed subgenera. However, in the case of the subgenus Pteropodus, gasbladder muscles morphology supports the subgeneric classification of this group.
Within the genus Sebastes the presence of large cylindrical gasbladder muscles may represent the more recent evolutionary condition. Conversely, the small dorsoventrally flattened gasbladder muscles found most commonly are probably more representative of the ancestral form for the genus. Possible explanations for the occurrence of cranioclavical muscles in fishes of the suborder Cottoidei are discussed.
The function of the large cylindrical gasbladder muscles found in several
VoL. XL] HALLACHER: GASBLADDER MUSCLES 81
rockfish species seems to be primarily for the production of sound (McInerney and Yearsley, in press). Gasbladder muscles may also be used for sound reception.
LITERATURE CITED
BarBER, S. B., and Mowsray, W. H. 1956. Mechanism of sound production in the sculpin. Science, vol. 124, pp. 219-220. BarsuKov, V. V.
1964. Key to the fishes of the family Scorpaenidae. Vsesoyuznyi Nauchno- Issledovatel’skii Institut Morskogo Rybnogo Khozyaistva i Okeanografii (VNIRO), Trudy, vol. 53, pp. 226-266. [All-Union Scientific Research Institute of Marine Fisheries and Oceanography (VNIRO), Proceedings. ] (English translation by N. Kaner and A. Mercado, Israel Program for Scientific Translations, Jerusalem, 1968.)
Bricut, I. J.
1972. Bio-acoustic studies on reef organisms. Pp. 45-69 im B. B. Collette and S. A. Earle (eds.), Results of the tektite program: ecology of coral reef fishes. Natural History Museum, Los Angeles County, Science Bulletin 14, 180 pp.
Courtenay, W. R., JR.
1971. Sexual dimorphism of the sound producing mechanism of the striped cusk-eel,
Rissola marginata (Pisces: Ophidiidae). Copeia, vol. 1971, no. 2, pp. 259-267. Dortu, Y.
1951. On the sound producing mechanisms of a scorpaenoid fish, Sebastiscus marmoratus. Science Bulletin of the Faculty of Agriculture of Kyushu University, vol. 13, pp. 286-288.
Durossg, A.
1874. Recherches sur les bruits et les sons expressifs que font entendre les poissons d’Europe, et sur les organes producteurs de ces phénoménes acoustiques, ainsi que sur les appareils de l’audition de plusieurs de ces animaux. Annales des Sciences Naturelles (Zoologie), Paris, Ser. 5, vol. 19, no. 5, pp. 1-53; vol. 20, no. 3, pp. 1-134.
EscHMEYER, W.N., and J. C. HurEAU
1971. Sebastes mouchezi, a senior synonym of Helicolenus tristanensis, with comments on Sebastes capensis and zoogeographical considerations. Copeia, vol. 1971, no. 3, pp. 576-579.
Evans, R. R.
1973. The swimbladder and associated structures in western Atlantic sea robins
(Triglidae). Copeia, vol. 1973, no. 2, pp. 315-321. FisH, M. P.
1954. The character and significance of sound production among fishes of the western North Atlantic. Bulletin of the Bingham Oceanographic Collection, art. 14, no. 3, pp. 1-109.
GosLine, W. A.
1971. Functional morphology and classification of teleostean fishes. University of
Hawaii Press, Honolulu, 208 pp. Gray, G. A., and H. E. Winn
1961. Reproductive ecology and sound production of the toadfish, Opsanus tau. Ecology,
vol. 42, pp. 274-282. GREENWOOD, P. H., D. E. Rosen, S. H. WeitzMaAn, and G. S. Myers 1966. Phyletic studies of teleostean fishes, with a provisional classification of living
82 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.
forms. Bulletin of the American Museum of Natural Hstory, vol. 131, art. 4, pp. 339-456. Lacter, K. F., J. E. Barpacu, and R. R. MILrer 1962. Ichthyology. John Wiley and Sons, Inc., New York. 545 pp. Matsupara, K. 1943. Studies on the scorpaenoid fishes of Japan. The Transactions of the Sigenkagaku Kenyusho, nos. 1 and 2, 486 pp. McInerney, J. E., and J. H. YEArsLey In press. Sound production in nine species of rockfish (genus Sebastes) . Moser, H. G. 1967. Reproduction and development of Sebastodes paucispinis and comparison with other rockfishes off southern California. Copeia, vol. 1967, no. 4, pp. 773-797. Puitiies, J. B. 1957. A review of the rockfishes of California (Family Scorpaenidae). California Department of Fish and Game, Fish Bulletin 104, 158 pp. Prorasoy, V. R., E. V. RoMANENKO, and Yu. D. PopiipaLin 1965. The biological significance of sounds produced by some fishes. Voprosy Ikhtiologii, vol. 5, no. 3, pp. 532-539. OuAsT EC: 1965. Osteological characteristics and affinities of the hexagrammid fishes, with a synopsis. Proceedings of the California Academy of Sciences, vol. 31, no. 21, pp. 563-600. Tavorca, W. N. 1964. Sonic characteristics and mechanisms in marine fishes. Pp. 195-211 in W. N. Tavolga (ed.), Marine bio-acoustics. Pergamon Press, New York, 413 pp. 1971. Sound production and detection. Pp. 135-205 im W. S. Hoar and D. J. Randall (eds.), Physiology of Fishes. Vol. 5. Academic Press, Inc., New York and London, XVI + 600 pp. Tower, R. W. 1908. The production of sound in the drumfishes, the sea robin and the toadfish. Annals of the New York Academy of Sciences, vol. 18, part 2, pp. 149-180. Winn, H. E. 1964. The biological significance of fish sounds. Pp. 213-231 in W. N. Tavolga (ed.), Marine bio-acoustics. Pergamon Press, New York, 413 pp. 1967. Vocal facilitation and the biological significance of toadfish sounds. Pp. 283-304 in W. N. Tavolga (ed.), Marine bio-acoustics. Vol. 2. Pergamon Press, New York, 353 pp. Winn, H. E., and J. A. MarsHALL 1963. Sound producing organ of the squirrel fish, Holocentrus rufus. Physiological Zoology, vol. 36, no. 1, pp. 34-44.
APPENDIX
Specimens examined in the course of this study are listed below by family. Abbreviations of depositories of specimens are as follows: CAS—California Academy of Sciences, San Francisco, California. BC—University of British Columbia, Vancouver, B.C. UMMZ— University of Michigan Museum of Zoology, Ann Arbor, Michigan. SIO—Scripps Institution of Oceanography, La Jolla, California. SU—Stanford University, collection now incorporated in the CAS collection. ZMK—Zoologisk Museum, Copenhagen, Denmark. The number of specimens examined from a particular lot and standard length in mm. are given in parentheses.
VoL. XL] HALLACHER: GASBLADDER MUSCLES 83
ORDER SCORPAENIFORMES SCORPAENIDAE Sebastes
aleutianus: CAS 15271 (1 specimen, 410 mm.), British Columbia, Vancouver Island, 10 September 1972.
alutus: CAS 15314 (1, 227), British Columbia, Vancouver Island, 11 September 1972; CAS 14888 (1, 341), southern Oregon, 26-28 April 1972.
atrovirens: CAS 14861 (1, 280), California, 3 April 1972; CAS 15061 (1, 232), California, 21 August 1972.
auriculatus: CAS 14472 (2, 229-357), California, 20 May 1972; CAS 14870 (1, 284), California, 15 April 1972.
aurora: CAS 15301 (1, 261), British Columbia, 10 September 1972.
babcocki: CAS 15286 (1, 362), British Columbia, 10 September 1972.
baramenuke: CAS 30088 (1, 300), Japan, 12-13 August 1929.
borealis: CAS 15272 (1, 406), British Columbia, 10 September 1972.
brevis pinis: CAS 14431 (1, 230), Alaska, Summer 197i.
capensis: CAS 17644 (1, 198), Chile, 21 September 1966.
carnatus: CAS 14732 (1, 249), California, 23 April 1972.
caurinus: CAS 14869 (1, 330), California, 15 April 1972.
chlorostictus: CAS 14705 (1, 269), California, 26 May 1972.
chrysomelas: CAS 14896 (1, 271), California, 30 July 1972.
ciliatus: CAS 30089 (1).
constellatus: CAS 14724 (1, 266), California, 27 May 1972.
crameri: CAS 15292 (1, 258), British Columbia, 11 September 1972.
dalla: CAS 14708 (1, 139), California, 27 May 1972; SU 4377 (1, 159), California, no date.
diploproa: CAS 14889 (1, 284), Oregon, 26-28 April 1972.
elongatus: CAS 14713 (1, 221), California, 26 May 1972.
emphaeus: CAS 15153 (1, 110), Oregon, 21 August 1972; BC66-135 (1, 122), British Columbia, 7 September 1966.
ensifer: CAS 17609 (1, 141, paratype), California, 26 May 1965.
entomelas: CAS 14865 (1, 207), California, 5 March 1972.
eos: CAS 17611 (1, 378), California, 3 August 1968.
exsul: CAS 17605 (1, 191), Gulf of California, 29 April 1969.
flameus: CAS 30090 (1, 290), Japan, Spring 1929.
flavidus: CAS 14880 (1, 343), California, 8 April 1972.
gilli: SIO 65-1-53B (1, 488), California, 9 January 1965.
goodei: SU 87 (1, 161), California, Santa Cruz Island, 7 February 1888.
helvomaculatus: CAS 15302 (1, 256), British Columbia, off Vancouver Island, 10 September 1972; CAS 17606 (1, 180), British Columbia, off Vancouver Island, 9-10 February 1967.
hopkinsi: CAS 15300 (1, 221), California, Santa Monica Bay, 16 September 1972.
hubbsi: CAS 30091 (1, 151), Japan, station H29-240, 9 August 1929.
ijimae: CAS 30080 (1, 192), Japan, no date.
inermis. CAS 30081 (1, 159), Korea, Seiskin, 15 September 1924; SU 7413 (1, 140), Japan, Misaki, no date.
jordani: CAS 26448 (1, 201), California, off Gaviota, 7 June 1953.
84
CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
joyneri: SU 7604 (1, 133), Japan, Tokyo, no date.
lentiginosus: CAS 17614 (1, 175, paratype), California, Cortes Bank, 16 May 1963.
levis: CAS 26594 (1, 123), California, Santa Monica, 21 June 1953.
longispinis: CAS 14281 (1, 136), Korea, 23 July 1959.
macdonaldi: CAS 17610 (1, 163), Baja California, off west coast, 16 January 1970.
maliger: CAS 14890 (1, 312), California, Farallon Islands, 15 July 1972; CAS 15410
(1, 275), British Columbia, spring 1972.
marinus: SU 34329 (3, 205-232), Nova Scotia, Sable Island Gulley, 18-20 July 1939; SU 1770 (1, 126), no data; SU 9870 (1, 203), Norway, no date.
matsubarae: SU 7393 (1, 231), Japan, Misaki, no date.
melanops: CAS 14726 (1, 332), California, off Point Reyes, 7 June 1972.
melanostomus: SIO 67-79-53 (1,194), Mexico, Baja California, 3 May 1967; SU 2459 (1, 95), southern California, no date.
miniatus: CAS 14878 (1, 396), California, off Farallon Islands, 8 April 1972.
mystinus: CAS 14712 (1, 342), California, off Santa Catalina Island, 26 May 1972; CAS 14474 (1, 222), California, Halfmoon Bay, 20 May 1972; CAS 14712 (1, 258), California, Santa Catalina Island, 26 May 1972; CAS 14717 (3, 231-294), California, Fort Ross Cove, 11 June 1972; CAS 14727 (1, 265), California, Point Reyes, 7 June 1972; CAS 14863 (1, 376), California, Halfmoon Bay, 5 March 1972; CAS 14871 (2, 260-289), California, Halfmoon Bay, 15 April 1972; CAS 14882 (2, 267-275), California, Farallon Islands, 8 April 1972; CAS 14894 (1, 251), California, Farallon Islands, 15 July 1972; CAS 15440 (2, 157-191), California, Santa Rosa Island, 21 September 1972; CAS 25900 (1, 112), California, Monterey Bay, 7 October 1953; SU 15097 (1, 70), California, Pacific Grove, 31 July 1948.
nebulosus: CAS 14720 (2, 277-278), California, Fort Ross Cove, 11 June 1972; CAS 14731 (1, 288), California, Farallon Islands, 8 April 1972.
nigrocinctus: CAS 28877 (1, 224), Canada, British Columbia, 10 June 1963.
nivosus: CAS 30082 (1, 175), Japan, station H29-299, Spring 1929.
oblongus: SU 7421 (1, 202), Japan, Matsushima, no date.
ovalis: CAS 14710 (1, 278), California, off Santa Catalina Island, 26 May 1972.
owstoni: CAS 30083 (1, 145), Japan, Toyama Bay, station H29-249, 12-13 August 1929.
pachycephalus: CAS 30241 (1, 157), Japan, Sea of Japan, station H29-224, 4 August 1929.
paucispinis: CAS 15299 (1, 259), California, Santa Monica Bay, 16 September 1972; CAS 14864 (1, 453), California, Halfmoon Bay, 5 March 1972.
phillipsi: SIO 65-153-53A (1, 320), California, off Newport Beach, 17 June 1965.
pinniger: CAS 15057 (1, 315), Oregon, Stonewall Bank, 21 August 1972.
proriger: CAS 15289 (1, 329), British Columbia, off Vancouver, 11 September 1972.
rastrelliger: CAS 14895 (1, 198), California, Halfmoon Bay, 28 July 1972.
reedi: CAS 15290 (1, 332), British Columbia, off Vancouver, 11 September 1972.
rosaceus: CAS 14875 (1, 227), California, Farallon Islands, 8 April 1972.
rosenblatti: CAS 14703 (1, 347), California, La Jolla, 25 May 1972.
ruberrimus: CAS 14560 (1, 288), California, off Point Reyes, 28 May 1972.
rubrivinctus: CAS 14704 (1, 287), California, La Jolla, 25 May 1972.
rufus: CAS 17612 (1, 339), California, Santa Catalina Island, 3 May 1968.
saxicola: CAS 14885 (1, 269), southern Oregon, 26-28 April 1972.
schlegelii: SU 7428 (1, 223), Japan, Hakodate, no date; SU 6274 (1, 88), Japan, Hakodate, no date.
scythropus: SU 7169 (1, 147), Japan, Misaki, no date.
serranoides: CAS 15806 (1, 285), California, Farallon Islands, 15 July 1972.
serriceps: CAS 17613 (1, 251), California, San Diego, 13 December 1972.
VoL. XL] HALLACHER: GASBLADDER MUSCLES 85
simulator: CAS 17607 (1, 235, paratype), Mexico, Guadalupe Island, 29 August 1956. sinensis: CAS 17608 (1, 115), Gulf of California, Ballenas Channel, 18 January 1968. steindachneri: SU 7422 (1, 166), Japan, Hakodate, no date.
taczanowskii: CAS 30084 (1, 122), Japan, Mutsu Bay, summer 1929.
thompsoni: CAS 30085 (1, 188), station H29-185-63.
trivittatus: CAS 30086 (1, 160), Japan, station H29-292C, 1 September 1929.
umbrosis: CAS 14707 (1, 194), California, La Jolla, 25 May 1972.
variegatus: SIO 63-946-53 (1, 220), Gulf of Alaska, 14 August 1960.
vexillaris: CAS 18461 (1, 245), California, La Jolla, 5 April 1945.
vulpes: CAS 30087 (1, 161), Japan, station H29-292C, 1 September 1929.
wilsoni: SU 2443 (1, 116), Oregon, no date.
zacentrus: CAS 15309 (1, 210), British Columbia, off Vancouver Island, 11 September 1972.
OTHER GENERA AND SPECIES
Apistes species: CAS 15975 (1, 109), Hong Kong, off Lema Island, 25 July 1958.
Brachy pterois serrulifer: CAS 15973 (1, 79), Hong Kong, off Lema Island, 25 July 1958.
Dendrochirus zebra: CAS 15971 (1, 100), Palau Islands, 23 September 1957.
Dendroscorpaena species: CAS 15965 (1, 107), Gulf of Thailand, 18 June 1961.
Ectreposebastes imus: CAS 30092 (1, 154), 300 miles west of Sumatra, 26 May 1966.
Gymnapistes marmoratus: CAS 30093 (1, 71), Western Australia, 8 February 1970.
Helicolenus dactylopterus: CAS 13941 (1, 137), Morocco, 15 July 1969.
Inimicus cuvieri: CAS 13553 (1, 131), Gulf of Thailand, 17-21 December 1960.
Tracundus signifer: CAS 24990 (1, 93), Hawaiian Islands, Oahu, 20 July 1969.
Macroscorpius pallidus: Fisheries Research Station Hong Kong, uncat. (1, 93), Indonesia, off Borneo, 6 June 1964.
Minous monodactylus: CAS 13907 (1, 79), Gulf of Thailand, 14 December 1960. M. pictus: CAS 13894 (1, 89), Hong Kong, 24 July 1958.
Neomerinthe beanorum: CAS 24386 (1, 105), Gulf of Mexico, 4 February 1967.
Parascorpaena armatus: CAS 15967 (1, 100), Gulf of Thailand, 27 May 1960.
Plectrogenium nanum: CAS 15704 (1, 51), Hawaiian Islands, Lanai, November 1967.
Pontinus longispinis: CAS 15968 (1, 157), Gulf of Mexico, off Yucatan Peninsula, 22 January 1969.
Pterois radiata: CAS 15970 (1, 115), Society Islands, Moorea, 15 May 1957.
Scorpaena agassizii: CAS 24380 (1, 127), South Atlantic, 27 April 1966. S. albobrunnea: CAS 15969 (1, 46), Palau Islands, 29 September 1958. S. brasiliensis: CAS 15964 (1, 119), Colombia, Gulf of Morrosquillo, 28 November 1968. S. elongata: CAS 24399 (1, 192), Angola, 18 March 1968. S. guttata: CAS 2463 (1, 193), Mexico, Guadalupe, 16 March 1932. S. mystes: SU 50171 (1, 106), Mexico, Sonora, 30 December 1955. S. porcus: CAS 13924 (1, 153), Yugoslavia, Piran Bay, 5 October to 1 December 1968. S. russula: CAS 13955 (1, 116), Peru, 7 September 1966.
Scorpaenodes parvipinnis: CAS 15972 (1, 71), Mariana Islands, Guam, 4 April 1959.
Scorpaenopsis species: CAS 15966 (1, 108), Mariana Islands, Guam, 25 January 1959.
Sebastiscus species: CAS 17643 (1, 186), Taiwan Strait, Formosa Bank, February 1972.
Sebastosemus species: ZMK uncat. (1, 100), Kermadec Island, Galathea-Ekspeditionen 1950-52 Station Number 675, 3 March 1952.
Setarches guentheri: CAS 15974 (1, 101), Caribbean, off San Andrés Islands, 20 November 1968. S. longimanus: Fisheries Research Station Hong Kong, uncat. (1, 118), Hong Kong, 23 August 1964.
Synanceia verrucosa: CAS 14966 (1, 134), Mariana Islands, Guam, 26 April 1959.
86 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
OTHER SCORPAENIFORM FISHES TRIGLIDAE
Leptotrigla alata: SU 21270 (1, 142), Japan, Nagasaki, no date.
HEXAGRAMMIDAE
Hexagrammos decagrammus: SU 55245 (1, 194), California, Moss Beach, 3 July 1949. Ophiodon elongatus: CAS 21750 (2, 187-202), California, San Pablo Bay, 28 July 1953.
ZANIOLEPIDIDAE Zaniole pis latipinnis: CAS 15979 (1, 160), California, off Avila, 19-20 October 1970.
PLATYCEPHALIDAE Platycephalus species: SU 60950 (1, 166), Hong Kong, Plover Cove, 6 January 1958.
HopLicHTHYIDAE Hoplichthys langsdorfi: SU 49455 (1, 145), Formosa, no date.
COTTIDAE Clinocottus analis: CAS 15980 (1, 124), California, off Santa Rosa Island, 25 January 1949. Cottus asper: CAS 20857 (1, 110), California, Waddell Creek, 24 February to 2 March 1935.
Hemilepidotus jordani: CAS 15568 (1, 149), Alaska, Shumagin Island, 10 August 1962.
Leptocottus armatus: CAS 18110 (1, 157), California, Marin County, 27 May 1945.
Mvyoxcephalus quadricornis: SU 49209 (1, 134), Alaska, near Barrow, 19 July 1951.
Scorpaenichthys marmoratus: SU 19391 (2, 125-198), California, Moss Beach, 27, November 1951.
COTTUNCULIDAE Cottunculus thompsoni: SU 9458 (1, 131), Off Delaware, no date.
AGONIDAE Occella verrucosa: CAS 15149 (1, 142), Oregon, 19 August 1972.
CYCLOPTERIDAE Liparis florae: SU 63602 (1, 153), California, Monterey County, 14 October 1955.
ADDENDUM
Recently William Eschmeyer was able to examine additional specimens from the North Atlantic. He informs me that two structural conditions exist for gasbladder musculature in North Atlantic species of Sebastes. The gasbladder muscles pass between ventral ribs 2-3 in S. marinus and S. mentella and between ventral ribs 3-4 in S. fasciatus and S. viviparus. Therefore the specimens reported here as S. marinus are probably S. fasciatus, and the muscle type here referred to as the marinus subdivision should be termed the fasciatus-viviparus subdivision.
PROCEEDINGS
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CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 4, pp. 87-92, 2 figs. October 30, 1974
A NEW GENUS AND SPECIES OF EUBLEPHARINE GECKO (SAURIA: GEKKONIDAE)
FROM BAJA CALIFORNIA, MEXICO
By Robert W. Murphy
California Academy of Sciences, San Francisco, California 94118
Apstract: <A new genus and species of eublepharine gecko, Anarbylus switaki, from central Baja California, Mexico is described and figured. Karyotype data is given.
During a photographic expedition through Mexico, Mr. Karl H. Switak of the California Academy of Sciences’ Steinhart Aquarium, collected an unusual ground gecko in central Baja California, Mexico. Cytological and morphological characters of this specimen are unique among eublepharine geckos. Thus the description of a new genus and species, on the basis of a single specimen, seems justified. All descriptive color references refer to those taken from the living animal and are compared with the standardizations of Ridgeway (1912).
Anarbylus Murphy, new genus
TyPeE-sPECIES. Anarbylus switaki Murphy.
DeEFINITION. A monotypic genus of the subfamily Eublepharinae; moderate in size; body largely covered with granular scales interspaced with enlarged keeled tubercles except on head, limbs, and venter. Skin soft, unattached to bones of skull. Eyes large, pupils vertically elliptical; eyelids well developed and functional, inner surface pigmented with black. All gulars equal in size. Ventral surface of digits sheathed with strongly peaked granular scales, some- times forming longitudinally parallel rows; claws partially hidden by a pair of lateral scales and a single elongate dorsal terminal scute. Ventral scales flat, imbricate. Preanal pores in angular series. Parietal single; retroarticular process
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88 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.
Ficure 1. CAS 139472, the holotype of Anarbylus switaki. Copyright photograph re- printed by permission of Karl H. Switak.
recurved. Karyotype composed primarily of metacentric chromosomes; diploid number 24.
Etymotocy. Anarbylus (treated as a masculine noun) from the Greek “with- out shoes” refers to the lack of transverse lamellae on ventral surface of digits.
Anarbylus switaki Murphy, new species. Switak’s Barefoot Gecko (Figures 1-2.)
Hototyre. CAS 139472, an adult male, from 5.5 miles west of San Ignacio (27° 27’ N., 112° 51’ W.) along Mexican Highway 1, Baja California Sur, Mexico, 500 feet elevation, collected by Karl H. Switak on 20 June 1974.
Diacnosis. Characters of the genus but also having nine crossbands of yellow spots between limb insertions and a single enlarged postnasal scale.
Etymo.tocy. This species is named for Karl H. Switak, Supervising Her- petologist of the Steinhart Aquarium, who discovered and collected the sole specimen.
Description. (All bilateral counts are given as left-right.) Head and snout uniformly covered above and below with juxtaposed circular granules which are larger on snout. Rostral pentagonal, two times wider than deep, two short
Vot. XL] MURPHY: NEW GECKO 89
sides contacting first supralabials; longer concave edges contact prenasals and are rounded at apex, in contact with 1 of 2 internasals. One prenasal on each side, convex distally and twice concave proximally to supranasals and nasals; supranasals number 1-1, as long as broad, lateral side concave, medial side convex; postnasals slightly smaller than supranasals, triangular, and number- ing 1-1; single nasal scale on either side twice as tall as wide; nostril round; subnasals absent; granules in contact with nasal series from rostral to supra- labials number 11-12. Imbricate scales with serrate edges border well developed functional eyelids and number 18-18 above, 19-19 below; inner surface of eye- lids black except outer border which is brownish drab with warm blackish brown serrate edges above and pallid neutral gray bordered with brownish drab below; pupils vertically elliptical; in life eye grayish black with dense light speckling on outer and inner area of iris, otherwise speckles few. Ear vertically oval, two times longer than wide, ventral edge anterior to dorsal. Enlarged supralabials number 9-8, decrease in size posteriorly, terminate beneath center of eye, the first two times longer than following scales. Infralabials number 12-12. Mental largest head scale, slightly narrower than rostral, quadrilateral, as wide as deep, posterior side narrower than anterior and comprising a short circular arc con- tacted by 9 gulars; 14 gulars contact mental plus first infralabials on both sides. First infralabial quadrilateral, longest side along mental, wider and much deeper than succeeding infralabials; remaining infralabials decreasing in size posteriorly.
Dorsum covered by granules of similar size with interspaced enlarged tubercles in irregular rows posterior to head; about 16 tubercles transverse on midbody; about 41 tubercles between limb insertions adjacent to middorsal line; tubercles keeled, increasing in size from neck to tail, some peaked posteriorly.
Dorsal and lateral granules and tubercles replaced ventrally with flat tri- angular scales, larger posteriorly, slightly larger than dorsal granules; short umbilical line interrupts regularity of adjacent scales; approximately 41 imbri- cate scales across venter, about 100 midventral scales from center of arm insertion to enlarged preanal scales; scales in preanal region further enlarged with 6 con- spicuous preanal pores; pore-containing scales obtusely angular in arrangement, pointing forward, two median scales in contact.
Arm scales juxtaposed, strongly peaked, base size equal to dorsal granules; tubercles absent; dorsum of hand covered with imbricate scales; palmar surface granular; third and fourth fingers longest, equal in length; ventral surface of digits without series of transverse lamellae; slightly enlarged peaked granules in longitudinal parallel rows, except on fifth finger, blend to palmar granules, numbering approximately 14 on fourth finger; fingers terminating in a pair of large shell-like lateral scales, capped by a long wedge-shaped terminal scute; claws clearly evident. Legs covered in granular and imbricate scales; tubercles absent; toes sheathed as fingers; fourth toe longest; 18 scale rows on fourth toe; claws clearly evident.
90 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Bifurcated cloacal spurs placed on each side of and slightly posterior to vent, 1.4 mm. long, 1.4 mm. wide at base; 2 postanal sacs present.
Tail base sheathed dorsally and laterally in granular scales with interspaced tubercles and ventrally in wide imbricate scales; regenerated tail begins 6.5 mm. posterior to vent, round in section, covered both dorsally and ventrally with granular scales of equal size to midbody granules, tubercles lacking.
PATTERN AND cCoLor. (In life.) Anterior portion of head with pallid neutral gray spots on mottled base of warm blackish brown and brownish drab; spots blend to sulphur yellow on head above ears. Lightly pigmented canthal ridges pallid neutral gray; ridges extend from eyes to rostral where they converge; second set of slightly darker snout lines lying adjacent to supralabials and extend- ing from postnasals to below eyes. Supralabials pigmented with alternating bands of warm blackish brown and brownish drab; infralabials identical to supralabials. Two crossbands of sulphur yellow spots transverse on parietal region. Neck lightly mottled with warm blackish brown on brownish drab with 3 distinct crossbands of 8 to 10 sulphur yellow spots. Pallid neutral gray gular and throat scales lack dark pigmentation.
On dorsum between limb insertions nine crossbands of sulphur yellow spots blend to pallid neutral gray laterally with warm blackish brown spots concen- trated near crossbands; first 2 crossbands form an ‘X’ pattern behind arm insertion; fifth crossband located at midbody, composed of fourteen spots. Light middorsal stripe extends from base of parietal to tail. Lateral pallid neutral gray spots form two longitudinal rows; each spot encircled, at least partially, by warm blackish brown pigmentation; some spots blend to venter and are not distinct from it; nine spots in both lateral rows offset anteriorly to nine dorsal crossbands. Ventral scales pallid neutral gray, immaculate.
Tail base with 2 crossbands of spots, one lateral and anterior to cloacal spurs, one adjacent to regenerated portion of tail, coloration identical to body cross- bands. Regenerated tail pale purplish gray with randomly dispersed large blackish brown spots.
Hands brownish drab, slightly lighter on tips of fingers; forearms lightly mottled; upper arm identical to lower but with heavier mottling and pallid neutral gray spots at insertion. Feet mottled at region of metacarpals; lower leg heavily mottled, few sulphur yellow spots occur on posterior surfaces; upper leg heavily mottled, containing 3 crossbands of sulphur yellow spots which fade to pallid neutral gray laterally, first crossband at leg insertion, second and third almost equidistant between insertion and knee.
Karyotype. Chromosome pattern (fig. 2) is a diploid complement of 24 chromosomes consisting of a graded series of 22 metacentric and 2 acrocentric chromosomes. When ranked in size sequence each 2 pairs of chromosomes are conspicuously larger than the following pair.
These data represent chromosome complements analyzed from 39 intestinal
VoL. XL] MURPHY: NEW GECKO 91
DAK RK x8
aA CR RY ORS
5 6 7 8
x % AK x x x x x
9 10 11 12
Ficure 2. The karyotype of Anarbylus switaki. Note the single pair of acrocentric chromosomes (no. 5).
cells taken from the holotype: cells were prepared by colchicine/hypotonic citrate technique. Centromeric position classification follows that of Levan, et al. (1964).
MEASUREMENTS. (In mm.) Snout-vent length, 87; tail length, 48 (regen- erated) ; head width, 16.4; head length, 26.7; width between eyes, 2.8; distance snout to eye, 10.5; distance snout to ear, 20.0; eye width, 5.0; internarial width, 2.9: fourth finger length, 4.3; fourth toe length, 5.3; arm length, 24; leg length, 31D
RANGE. Known only from type locality.
Remarks. Anarbylus switaki is readily distinguished from all other euble- pharine geckos in having a single enlarged postnasal scale. It is further dis- tinguished from Aeluroscalabotes, Coleonyx, and Eublepharis in not having enlarged transverse lamellae; from Aeluroscalabotes, Eublepharis, and Hemi- theconyx in having numerous small gulars in contact with the mental and in- fralabials; and from Holodactylus and Coleonyx (brevis-variegatus complex) in having tubercles.
92 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Though the karyotypes of only a few species of gekkonids are known, Gor- man (1973) speculated that the “typical karyotype” is sufficient to serve as a cytological definition. Such a definition consists of three elements: 1) a range in diploid number of 32 to 46; 2) a graded series of acrocentric chromosomes with no distinct break between macrochromosomes and microchromosomes; 3) large metacentric elements infrequent with the majority of two-armed chromo- somes having subterminal centromeres. The karyotype of Anarbylus switaki (fig. 2) does not agree with any criterion of Gorman’s definition. Such cyto- taxonomic data strongly support the view that this species is neither an aberrant Coleonyx nor can it be referred to any currently recognized genus within the subfamily. The preliminary results of a cytotaxonomic study of the eublepharine geckos, in which the karyotypes of Anarbylus switaki, Coleonyx brevis, C. variegatus, C. elegans, C. reticulatus, Eublepharis macularius, and Hemitheconyx caudicinctus have been examined, support the above conclusions.
ACKNOWLEDGMENTS
My appreciation is due Karl H. Switak for making available this unique gecko for study and for permission to reproduce the copyright photograph of the holotype. Alan E. Leviton and Robert C. Drewes kindly read, criticized, and improved the manuscript and assisted with the karyotyping. James E. Gordon provided radiographs of the specimen and Robert Dempster of the Steinhart Aquarium assisted with the photography of the karyotype.
LITERATURE CITED
GoRMAN, GEORGE C. 1973. The chromosomes of the Reptilia, a cytotaxonomic interpretation. pp. 349-424. In Chirelli, A. B., and E. Capanna, eds. Cytotaxonomy and Vertebrate Evolution. Academic Press, New York, 778 pp. Levan, A., K. Freoca, AnD A. A. SANBERG 1964. Nomenclature for centromeric positions on chromosomes. Hereditas, vol. 52, pp. 201-220. RIDGEWAY, ROBERT 1912. Color Standards and Color Nomenclature. Washington, D.C., 43 pp., 53 pls.
PROCEEDINGS
OF THE , Wi / CALIFORNIA ACADEMY OF SCIENCES,
FOURTH SERIES
Vol. XL, No. 5, pp. 93-107; 4 figs. February 18, 1975 TWO NEW BLIND SNAKES (SERPENTES: LEPTOTYPHLOPIDAE) FROM BAJA CALIFORNIA, MEXICO WITH A CONTRIBUTION TO THE BIOGEOGRAPHY OF PENINSULAR AND INSULAR HERPETOFAUNA
By Robert W. Murphy
California Academy of Sciences, San Francisco, California 94118
ApsTRACT: Two new subspecies of Leptotyphlops humilis trom the Gulf of California, Mexico are described. Biogeographical theories explaining the distribution of the herpetofauna of southern insular and peninsular Baja California, Mexico are re- viewed. Recent developments in continental drift form the basis of a new interpre- tation of the distribution of the isolated forms.
Recently two specimens of Leptotyphlops which appear to belong to L. humilis were received by the California Academy of Sciences. They were col- lected on Isla Santa Catalina, Gulf of California, Mexico by Bruce Feldhammer in March, 1972. These specimens are of considerable interest because they are the first records for the genus from Isla Santa Catalina which has a highly endemic herpetofauna. Of ten reptiles known on the island, all are endemic, so it is not surprising to find that this is indeed the case for these blind snakes. Another insular representative of Leptotyphlops was reported by Soulé and Sloan in 1966. They assigned a single specimen of L. Aumilis from Isla Carmen with seven pigmented rows of median dorsal scales to L. h. slevini, a subspecies described by Klauber (1931) as having only five lightly or moderately colored
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94 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
dorsal scale rows. Re-examination of the Soulé-Sloan specimen showed this blind snake to be so unique that reference to a distinct taxon seems justified. I take pleasure in naming the Santa Catalina population for Dr. Alan E. Leviton who has provided me the opportunity to study at the California Academy of Sciences. The Isla Carmen population is named in honor of Dr. George E. Lindsay who, as former Director of the San Diego Society of Natural History and current Director of the California Academy of Sciences, has played a most important role in biological research in Baja California.
The following abbreviations are used in this paper: CAS—California Acad- emy of Sciences; CAS-SU—California Academy of Sciences—Stanford Univer- sity Collection; LACM—Los Angeles County Museum of Natural History; MVZ—Museum of Vertebrate Zoology, University of California, Berkeley; SDSNH—San Diego Society of Natural History.
Leptotyphlops humilis levitoni Murphy, new subspecies.
Santa Catalina Island Blind Snake. (Figure 1.)
Hototype. CAS 135146, adult, from Isla Santa Catalina, Gulf of California, Mexico [26° 40’ N., 110° 47’ W.], collected by Bruce Feldhammer on 24 March 1972.
PaRATYPE. CAS 135147, adult, same locality data as the holotype.
Dracnosis. A subspecies of Leptotyphlops humilis (Baird & Girard) having a low dorsal scale count, seven pigmented scale rows, 12 scale rows around the tail, a low number of subcaudal scales, and no pigmentation around the mouth.
Description. Snout bluntly rounded; rostral elongate and wedge-shaped with widest point at level of nostrils, in contact with nasals and prefrontal; nasals completely divided; lower nasal elongate, wider dorsally, completely separating rostral and anterior supralabial; upper nasal elongate, wider ventrally, in contact with ocular, rostral, lower nasal, and prefrontal; single anterior supralabial with dorsal edge acutely terminating at lower level of eye; large ocular extending from central dorsal row to the mouth and completely separating supralabials; eye anterior in upper half of ocular; occipital and parietal elongate with the latter in contact with posterior supralabial; temporal smaller than other postoccipitals. First four dorsal median scales are hexagonal, almost as long as broad with approximate order of increasing size being 2nd-1st-4th-3rd; fifth dorsal median scale slightly broader than all others. Head widest at occipitals. Four infra- labials, anterior being minute (easily confused with mental) and posterior being largest. Mental very small, approaching triangular shape. Chin shields ir- regular, blending with ventrals at level of parietals.
Body almost cylindrical; head slightly distinct, narrower than mid-body diameter; tail slightly but distinctly diminished in diameter and terminating in
Vol. XL] MURPHY: NEW BLIND SNAKES 95
Ficure 1. Leptotyphlops humilis levitoni Murphy, new subspecies. A. Dorsal view of head. B. Lateral view of head.
a laterally compressed sharp spine. Body scales equal in size without con- spicuously enlarged dorsal or ventral scales.
Ground color (in 75% ethanol) of cinnamon or medium dark brown present on seven median dorsal scale rows; pigmentation sometimes involves as few as five rows anteriorly and as many as nine posteriorly. Pigmentation is applied evenly as a multiplicity of dots with a greater density on the terminal edge of each pigmented scale. An area surrounding the mouth lacks pigmentation, except for occasional spots, and includes: anterior two-thirds of rostral; lower nasals; lower one-fourth of the upper nasals; anterior supralabials; lower portion of the ocular and posterior supralabials from a line beginning at the top of the anterior supralabial to the angle of the jaw. Infralabials, mental, and chin shields lack pigmentation. Ventral color varies from light brown in the anterior half to cream in the posterior half.
MEASUREMENTS. Total length 212 mm.; tail length 9.6 mm.; body diameter 5.1 mm.; tail length into total length 23.1; average body diameter into total length 42.8. Fourteen scale rows around the body; 12 scale rows around the tail: 249 dorsal scales from rostral to spine; 14 subcaudal scales; anal plate not divided.
RANGE. Known only from type locality.
PARATYPE. The single paratype, an adult, adheres closely to the description of the holotype. Total length 214 mm.; tail length 8.5 mm. (tail somewhat constricted); body diameter 4.8 mm.; tail length into total length 25.2; average
96 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
body diameter into total length 43.2. Fourteen scale rows around the body; 12 scale rows around the tail; 250 dorsal scales from rostral to spine; 14 sub- caudal scales. The first four dorsal scales are hexagonal, almost as long as broad with approximate order of increasing size being Ist-2nd-3rd and 4th. Coloration is identical to the holotype.
Remarks. A blind snake of the species Leptotyphlops humilis (Baird & Girard) which differs from L. h. humilis, L. h. cahuilae, L. h. utahensis, and L. h. segregus in having fewer dorsal scale rows; from L. h. slevini and L. h. cahuilae in having 7 pigmented median dorsal scale rows; from L. h. segregus and L. h. tenuiculus in having 12 scale rows around the tail; and from L. h. dugesi in having fewer subcaudals and no pigmentation around the mouth.
Leptotyphlops h. levitoni appears to be most closely allied to L. h. dugesi, and on the basis of the number of dorsal scales, to L. h. slevini. However, the character of the number of dorsal scales may be misleading. Fox (1948) showed that temperature may be a major factor in determining the number of scales of Thamnophis couchi (= elegans) atratus. One may observe a similar situation in Leptotyphlops humilis, for the number of dorsal scales appears to increase in specimens from south to north (or warm to cold). This trend was noted first by Klauber (1940) and again by Hardy and McDiarmid (1969).
Leptotyphlops humilis lindsayi Murphy, new subspecies.
Lindsay’s Blind Snake. (Figure 2.)
Leptotyphlops humilis slevint KLAUBER (part), 1966, Trans. San Diego Soc. Nat. Hist., vol. 14, no. 11, pp. 137-156.
HototypeE. SDSNH 44386, adult female from Isla Carmen, Gulf of Cali- fornia, Mexico [25° 57’ N., 111° 12’ W.], collected by Charles E. Shaw and George E. Lindsay on 4 April 1962.
Diacnosis. A subspecies of Leptotyphlops humilis (Baird & Girard) having a low dorsal scale count, seven pigmented median dorsal scale rows, 12 scale rows around the tail, low number of subcaudal scales, and pigmented supra- labials.
Description. Head widest behind occipital; snout bluntly rounded; rostral elongate, widest point at level of nostril, in contact with nasals and prefrontal and terminating at anterior level of eye; lower nasal elongate, wider dorsally, completely separating rostral and anterior supralabial; upper nasal longer than wide with greatest width at level of eye, in contact with ocular, rostral, lower nasal, and prefrontal; single anterior supralabial with dorsal edge acutely termi- nating; large ocular extending from central dorsal scale row to the mouth, completely separating anterior and posterior supralabials; eye anterior in center of ocular; occipital and parietal elongate with latter in contact with posterior
Vol. XL] MURPHY: NEW BLIND SNAKES 97
Ficure 2. Leptotyphlops humilis lindsayi Murphy, new subspecies. A. Dorsal view of head. B. Lateral view of head.
supralabial; first three postoculars as elongate scales on right side, first two elon- gate on left side; temporals smaller than body scales. First four dorsal median scales hexagonal, almost as long as broad and nearly equal in size; fifth dorsal median scale not greatly enlarged. Four infralabials, anterior being greatly re- duced, posterior being largest; mental very small, almost triangular in shape; chin shields irregular anteriorly, blending with ventrals behind head.
Body nearly cylindrical, head only slightly distinct; tail slightly, but dis- tinctly, diminished in diameter and terminating in a laterally compressed sharp spine. Body scales equal in size without conspicuously enlarged dorsal or ventral scales.
The ground color (in 75% ethanol) of sayal or medium brown is usually present on seven dorsal median scale rows and occasionally involves as few as five anteriorly and as many as nine posteriorly. Body pigmentation is evenly dispersed as a multiplicity of dots except at the terminal edge of the scales where pigmentation appears denser; lateral rows may appear lighter than dorsal rows. Other body scales are varying shades of cream. Head pigmentation is as follows: posterior one-half of rostral; right lower nasal slightly pigmented; left lower nasal; upper nasals; dorsal one-half of anterior supralabials; oculars; dorsal and anterior portion of posterior supralabials; all dorsal scales. The lower jaw is void of pigmentation.
MEASUREMENTS. Total length 202 mm.; tail length 8 mm.; body diameter 4.9 mm. (may be small since the body cavity is completely opened); tail length into total length 25.3; body diameter into total length 41.2. Fourteen scale rows around the body; 12 scale rows around the tail; 243 dorsal scales from rostral to spine; 14 subcaudal scales; anal plate not divided.
98 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
RANGE. Known only from type locality.
Remarks. A blind snake of the Leptotyphlops humilis (Baird & Girard) complex differing from L. h. humilis, L. h. cahuilae, L. h. utahensis, and L. h. segregus in having fewer dorsal scales; from L. h. slevini and L. h. cahuilae in having seven pigmented median dorsal scale rows; from L. h. segregus and L. h. teniculus in having 12 scale rows around the tail; from L. A. levitoni in having pigmented upper lips; and from L. /. dugesi in having fewer subcaudals and no pigmentation in lower labials of adults.
One characteristic of L. h. lindsayi appears unique. The enlarged right third postocular (or postoccipital) has not been reported for any other sub- species of L. humilis. The occurrence of this phenotypic expression within a genetically isolated population seems significant.
In other characteristics, Leptotyphlops h. lindsayi is allied to L. h. levitoni and L. h. dugesi. Similarity between these three forms is seen in low dorsal scale counts, coloration, and similarity of scale patterns.
DISCUSSION
Savage (1960) presented a detailed hypothesis of southern peninsular Baja California invasion by northern herpetofaunal forms. According to Savage, immigration of northern forms, which presumably began in the Pliocene and occurred in four successive waves, replaced the early Tertiary fauna of the area. Savage contended that the first Pliocene wave ultimately formed the basic modern fauna of the Californian and San Lucan areas. A second wave of desert- adapted groups invaded the northern areas as aridity increased during the end of the Pliocene, while at the beginning of the Pleistocene a third wave invaded the northern portion of the peninsula as general temperature drops occurred; this third cluster produced the desert forms of the east and south peninsula. Finally, in the Pleistocene, the fourth or possibly several waves of desert forms invaded from the northeast at times of interglacial conditions. Overlap of San
>
Ficure 3. Early and late stages in the history of the San Andreas fault. A. twenty-five million years ago Baja California presumably nestled against mainland Mexico. The first section of oceanic rise between the Murray fracture zone and the Pioneer fracture zone has just collided with the continent. Trench deposits are uplifted and become part of the Coastal Ranges of California. The block containing the present San Francisco area (stippled) is about to start its long northward journey. A block immediately to the east (cross-hatched) becomes attached to the Pacific plate and eventually is jammed against the San Bernardino Mountains. B. Three million years ago the Gulf of California has started to open. As the peninsula moves away from mainland Mexico a series of rifts appear, fill with magma, and are offset by numerous fractures. Baja California may have been torn off in one piece or in slivers. (Reprinted from Anderson, The San Andreas Fault. Copyright © 1971 by Scientific American, Inc. All rights reserved.)
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Lucan forms moving north and the northern forms moving south then occurred in the central peninsula.
The plate tectonic conception of continental drift was founded in the late 1950’s. Many American scientists were reluctant to accept the concept until the late 1960’s (Wilson, 1972). This may account for the fact that Savage failed to include the distribution and origin of the insular faunas, which could not be readily explained by his northern origins hypothesis. Thus, Savage overlooked one important possibility, that much of the San Lucan and insular herpetofauna did not arrive by Pliocene and Pleistocene invasion from the north but rather by traversing the Gulf on the Peninsula as the latter broke away from mainland Mexico. In fact, Streets, in 1877 (pp. 41-42), suggested ‘“. . geological changes . . . since the post-Tertiary period,’ to explain the insular occurrence of reptiles on Pacific and Gulf islands. This concept is best exemplified by the presence of Bipes biporus, Natrix valida celaeno, Ctenosaura hemilopha, Pseudemys scripta nebulosa, and Eridiphas slevini, the latter possibly representing a link between Leptodeira and Hypsiglena (Leviton & Tanner, 1960).
Since the discussion by Savage (1960), Anderson (1971) evaluated current information on the San Andreas Fault and presented his ‘“Two-fault hypothesis” (fig. 3). According to Anderson, before the Pacific plate began moving north- west the “San Francisco” area was located near the present location of Ensenada in northern Baja California and present day “Baja California” presumably nestled against mainland Mexico. Twenty-five million years ago the northern Pacific plate separated from Bahia Sebastian Vizcaino (near Scammon’s Lagoon) and began its northwest journey of up to 700 miles. Twenty million years later (four to six million years ago) the southern portion of the Pacific plate was torn from mainland Mexico forming the Gulf of California. Eventually, after 300 miles of northwest migration, the southern Pacific plate rammed into the northern Pacific plate and formed a single unit. It should be noted that the peninsula may have split from mainland Mexico in several fragments. Anderson (1971, p. 60), in illustrating the early and late stages of the history of the San Andreas Fault, pictures the southern tip of Baja California as being a separate land mass as late as three million years ago (fig. 3).
Auffenberg and Milstead (1965) noted that very little taxonomic differentia- tion occurred between the Pliocene and Recent, particularly at the species level. They further state that the greatest effects of the Pleistocene on Baja California were those of changes in sea-level and not climatic change. The sea-level changes (such as a drop of approximately 110 meters and estimated rise of 30 meters | Flint, 1971]) had little effect on Baja California topography except for creating and drowning many islands (Durham & Allison, 1960). On this basis Soulé and Sloan (1966) placed many islands in a category of probable recent or young shallow-water islands, including therein Tiburon, San Marcos,
Vol. XL] MURPHY: NEW BLIND SNAKES 101
Coronados, San Jose, San Francisco, and Espiritu Santo as well as many small coastal and satellite islands and possibly Carmen, Monserrate, and Danzante (fig. 4).
In view of recent information and concepts, it is possible to suggest the fol- lowing interpretation of the distribution and interrelationships of the forms of Leptotyphlops humilis. Prior to the separation of Baja California from mainland Mexico L. humilis, as well as many other forms of reptiles and am- phibians, occurred in both areas. The southern Pacific plate began moving northwest and carried the blind snakes with it. The Cape San Lucan region parted from mainland Mexico, possibly from a location further south, as a separate unit, free of the peninisula, and remained isolated until mid-Pleistocene. The population of L. Aumilis isolated on this island evolved into a group having only five pigmented median dorsal scale rows. This group is presently known as L. h. slevini. As the peninsula was torn away from mainland Mexico two southern groups of islands were formed and have remained without further contact: Isla Santa Catalina and islas San Diego and Santa Cruz. Evidence for this is the high percentage of endemic reptiles present on these islands. If these islands had had recent contact with the mainland, then mainland forms would be expected to occur. In addition, the existence of Crotalus catalinensis, allied to C. atrox (Klauber, 1956, 1972), on Isla Santa Catalina and C. atrox on Isla Santa Cruz is better explained by continental drift than by the swimming or rafting suggested by Klauber (1956, 1972). Sator angustus, which occurs on San Diego and Santa Cruz presents the same problem [see below]. Lepto- typhlops humilis is presently known from one of these “old” islands, Isla Santa Catalina.
Another group of islands was formed later. During the Pleistocene, tectonic shifts and sea-level changes formed the young, shallow-water islands (as noted earlier) including Carmen, Monserrate, and Danzante. These islands share a large percentage of their herpetofauna with the Baja California mainland. During Pleistocene glaciation and lower water-levels, reptiles could easily pass from the peninsula to these islands. There is evidence that some of these islands have undergone recent uplifting (Anderson, 1950; Wilson & Rocha, 1955; Soulé & Sloan, 1966) indicating recent faulting and instability. If this dual theory of island formation is true then L. #. levitoni has remained isolated from the parental stock since mid-Pliocene, L. 4. lindsayi being isolated sometime during the Pleistocene.
The presence of Sator grandaevus on Isla Cerralvo and its absence from the San Lucan region of mainland Baja California has been a subject for interesting speculation for a long time. On those islands on which Sator occurs, no other sceloporine lizards are known. The occurrence of the old southern forms of rep- tiles, as Eridiphas slevini, on both Isla Cerralvo (Soulé, 1961) and the southern tip of mainland Baja California, would seem to indicate that these two regions had
BAJA CALIFORNIA
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“GULF OF CALIFORNIA : (MAR DE CORTEZ)
ANGEL de la GUARDA PARTIDA(N) RAZA
SALS|PUEDES TIBURON
SAN ESTEBAN SAN LORENZO (N-S)
S.P NOLASCO
SAN MARCOS
MEXICO \LDEFONSO CORONADOS CARMEN FARALLON DANZANTE MONSERRATE inevageies SANTA CATALINA a SANTA CRUZ SAN Se SAN DIEGO SAN FRANCISCO
CAYO and COYOTE PARTIDA (S) and ESPIRITU SANTOS CERRALVO
Ficure 4. A map of the Gulf of California indicating the islands discussed in this paper.
Vol. XL] MURPHY: NEW BLIND SNAKES 103
a similar origin. Seemingly, Isla Cerralvo was separated from the San Lucan region before the San Lucan plate joined with the southern Pacific plate in the mid-Pleistocene. When these two plates joined, the sceloporine lizards of the north invaded the San Lucan region and pressured the San Lucan form of Sator to extinction. After the San Lucan plate joined the Pacific plate, the sea-level dropped briefly allowing invasion of Isla Cerralvo by northern iguanids. How- ever, the length of time for invasion was not sufficient to allow invading scelop- orines to overrun the Cerralvo populations of Sator. Islas San Diego and Santa Cruz may have had a similar southern origin and have been moving northwest since the San Lucan plate rammed the southern Pacific plate.
The relationships between Isla Cerralvo and islas San Diego and Santa Cruz have never been understood. If these islands had a similar southern origin, and if they were only recently separated, it seems reasonable to expect that any Leptotyphlops discovered would be of similar origin and therefore would have only five pigmented median dorsal scale rows. Unfortunately, we have no evi- dence at this time to test this speculation.
Leptotyphlops humilis slevini, as mentioned earlier, was long separated from other populations of L. humilis. By the time the San Lucan plate joined the rest of the peninsula, the local population of L. Aumilis had undergone a substantial change, that of reduction in the number of pigmented median dorsal scale rows. The occurrence of this subspecies on Isla Cerralvo (Banks & Farmer, 1963) provides further evidence that this island is part of the San Lucan plate.
Only one other subspecies, L. h. cahuilae, is known to have five pigmented median dorsal scale rows. Klauber (1940) postulated that the Sonoran L. x. cahuilae became isolated for a considerable length of time, possibly by Lake Cahuilla, and then remet and interbred with L. h. humilis. He also noted that different desert-dwelling populations of L. h. cahuilae, as that of the Vizcaino Desert in central Baja California, either evolved parallelly with Sonoran L. /. cahuilae or represent a residue of a former intrusion from Sonora. Continental drift adds two additional possibilities; either the Vizcaino Desert population traversed the gulf or immigrated from the south. Two specimens of L. h. cahuilae (CAS 103465 and LACM 2167) from Bahia de Los Angeles, Baja California Norte, [28° 55’ N., 113° 32’ W.], indicate that the ranges of Sonoran and Vizcaino Desert L. hk. cahuilae are continuous. Seemingly this would eliminate Klauber’s anticipation of the isolated Vizcaino Desert population. However, current data are not adequate to determine the “‘slevini-cahuilae” subspecies relationships.
Of the nine subspecies of L. humilis currently recognized two are definitely allied; L. A. utahensis is certainly derived as a northern extension of L. 4. humilis, L. h. utahensis being distinct in having a divided fourth and an enlarged fifth anterior median dorsal scale. The occurrence of five L. h. humilis xX utahensis intergrades (CAS 89570, 89577, 89591-89593) from Inyo County,
104 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
California extends the expected range of intergradation, as predicted by Klauber (1940), by approximately 100 miles.
As noted earlier, L. h. dugesi, L. h. levitoni, and L. h. lindsayi also appear to be quite closely related for all three share the characteristics of a low number of dorsal scales, 12 scale rows around the tail, a low ratio of body length to body diameter, 7 pigmented median dorsal scale rows, and a medium to dark brown coloration. L. 4. slevini shares some of these characteristics but differs in having 5 pigmented median dorsal scale rows and a light brown coloration. Based on the transgulfian migration theory, L. /. slevini forms a unique taxon within the L. humilis complex and, though closely related, is not a member of the “dugesi-levitoni-lindsayi’ combination.
In their discussion of the biogeography and distribution of the herpetofauna on the islands in the Gulf of California Soulé and Sloan (1966) commented on two trends of insular reptiles; gigantism in lizards and dwarfism in snakes (with noted exceptions). The ratio of body length to body diameter for L. h. levitoni and L. h. lindsayi is less than those of any other subspecies of L. humilis. Such a low ratio of body length to body diameter indicates that the two insular worm snakes have heavier bodies than their mainland relatives. Once the population structure is better known these statistics can be further interpreted as being attributable to gigantism, an effect of dwarfism, or the result of insufficient data.
A Revisep Key To THE SuBSPECIES OF THE WESTERN BLIND SNAKE, LEPTOTYPHLOPS HUMILIS
Lape hive pismentededorsall smecamie Salle 0 vv Siem eee eee ee 2 bs seven! to) nine) pigmented dorsal median scale: ols) | eee eee een 3
2a. Dorsal scale rows less than 270; Cape region of Baja California Sur; Isla Cerralvo,
(Gwili oi Calhbiorma, Iie) ~~ L. h. slevini Klauber b. Dorsal scale rows more than 280; central Baja California north to southern California and Arizona; northern Sonora, Mexico _..... L. h. cahuilae Klauber Sa, ISN CeAlle Tos eieoiael tell ols ek eee 4 bs 4Ewelve. scale. tows: around ‘tail 22 =. 3 we. eee 5 4a. Dorsal scale rows less than 250; San Luis Potosi —— L. h. tenuiculus (Garman) b. Dorsal scale rows more than 250; central Coahuila, Mexico north to southeastern Arizonasandethioughy trans=ecosmel exc sy seen ean L. h. segregus Klauber
5a. Dorsal scale rows more than 280; fourth mid-dorsal scale often divided; fifth dorsal
much wider than sixth; southwestern Utah _- L. h. utahensis Tanner b. Dorsal scale rows less than 280; fifth mid-dorsal scale not much wider, if any, Gham iSixth, "ye eh el ee 6
6a. Dorsal scale rows greater than 257; central Baja California, Mexico north to southern Nevada southeast to south-central Arizona; Cedros Island — eel ok SE io ors als) ett i te Be Be Ae L. h. humilis (Baird and Girard)
Vol. XL] MURPHY: NEW BLIND SNAKES 105
AD ILOW CHAM ASAlS Sip Ie TCT te Clie | ees ee Se eee ek 8 b. Lower nasals not pigmented; Isla Santa Catalina, Gulf of California, Mexico —_ Re ES Oe ko L. h. levitoni Murphy
8a. Subcaudals 15 or less; infralabials not pigmented in adults; Isla Carmen, Gulf of (Callitormiayes Mie xd C0 ig eee ee eee, ee ee ee ee L. h. lindsayi Murphy
b. Subcaudals usually more than 15; infralabials of adults often pigmented; southern Sonora southeast to Colima; Guanajuato, Mexico — L. h. dugesi (Bocourt)
MATERIAL EXAMINED
L. h. cahuilae (13). CattrorniA. Imperial Co.: 5 mi. N. Winterhaven (MVZ 63562) ; Riverside Co.: Palm Springs, China Canyon (MVZ 71090). ARIZONA: Yuma Co., (CAS-SU 5697). MEXICO. Baja CaLirorniA DEL Norte: 5 mi. N. San Felipe, Playa del Sol Camp (CAS 136368-136369); 6 mi. N. San Felipe (LACM 36504); Puncta San Felipe, +50 ft. (MVZ 50168); Bahia de Los Angeles (CAS 103465, LACM 2167); Sierra Juarez (CAS 85069); Base of grade below Alaska (CAS-SU 11567); San Jose, 2300 ft. (MVZ 9637). Sonora: 56 mi. E. San Luis (LACM 9033).
L. h. dugesi (12). MEXICO. Srnatoa: Mazatlan (CAS-SU 1776); 11 mi. N. Culiacan (LACM 6773); 20 mi. S. Culiacan, Mexico Highway 15 (LACM 51564); 0.5 mi. N. Terroros (LACM 6773). Nayarit: Tepic, LaLoma Motel (CAS-SU 19243); 5 mi. E. Santa Cruz (CAS 134054); between San Blas and Mexico Highway 15 (LACM 8730-8731); 18.5 mi. E. (by road) San Blas (MVZ 71333). Jatisco: Jocotepec (CAS 85470); 14.5 mi. E. Tapalpa (LACM 37327-37328).
L. h. humilis (28). Cattrornta. San Diego Co.: (CAS 58160, 62992, 64442); Palomar Mountain (CAS 58132); Ladrillo Station (Rose Canyon, CAS 53933); Scissors Crossing, Highway 78 (MVZ 64478); 1.2 mi. E. Scissors Crossing (CAS-SU 19763); 2 mi. E. Scissors Crossing (CAS-SU 19766); 0.5 mi. E. Scissors Crossing (CAS-SU 19764-19765); 1 mi. ENE. Scissors Crossing, 2200 ft., San Felipe Valley (MVZ 79215); Lemon Grove (MVZ 10189). San Bernardino Co.: 9 mi. W. Earp (MVZ 51268); Twenty-nine Palms (MVZ 56491). Inyo Co.: Inyo Mountains, N. side of Daisy Canyon, 3900 ft. (CAS 89594) ; Nelson Mountains, Grapevine Canyon, 4500 ft. (CAS 89568). Los Angeles Co.: near mouth of Big Santa Anita Canyon (MVZ 74653). Riverside Co.: mouth of Whitewater Canyon, E. canyon wall (MVZ 80920). Arizona. Pima Co.: Tucson (CAS 33525, 33835-33836, 91533); Santa Catalina Mountains (CAS 35849). Pinal Co.: S. Florence Junction (CAS 84135). MEXICO. Baya CarirorntA DEL Norte: San Ignacio, 500 it. (MVZ 10667); 15 mi. E. San Telmo de Arriba (CAS 123717); 7 mi. S. Tecate (CAS 134800). CuimuaHuA: 3 mi. NW. Chilmahma, 0.5 mi. W. main highway (MVZ 57331).
L. h. levitoni (2). MEXICO. Gulf of California, Isla Santa Catalina (CAS 135146-135147). L.h. lindsayi (1). MEXICO. Gulf of California, Isla Carmen (SDSNH 44386).
L. h. slevini (16). MEXICO. Baja CacirorniA Sur: Gulf of California, SW. side of Isla Cerralvo (CAS 93009); La Paz (CAS 129644-129645, 134772-134223; MVZ 45386; LACM 25061); 7 km. S. La Paz (CAS 134801-134802); Eureka (MVZ 11850-11851); San Jose del Cabo (CAS-SU 4118, paratype); vicinity of Cabo San Lucas (CAS-SU 14032-14033) ; Santa Anita (CAS-SU 6025); vicinity of Cape San Lucas (CAS-SU 16061).
L. h. humilis * cahuilae intergrades (191). CAttrornta. Imperial Co.: Colorado River, Laguna Dam (MVZ 63543-63548, 63550-63556, 63560-63561, 70457); Laguna Dam, Laguna
106 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Island (MVZ 105162-105172, 85237-85238, 93113, 93115); Laguna Dam, Potholes (MVZ 63549, 50228-50233; CAS 80392-80459, 80461-80468, 80470-80532); 3 mi. S. Laguna Dam (MVZ 93114); about 15 mi. E. Glamis on Highway 78-S (MVZ 84701). Riverside Co.: 1 mi. SE. Cabazon, base of San Jacinto Mountains (MVZ 74652); 1.5 mi. ENE. Cabazon, base of San Jacinto Mountains (MVZ 74647-74651); Palm Springs, China Canyon (MVZ 70458). Inyo Co.: Death Valley, Coweruls, 4 mi. N. Furnace Creek Dam, 150 ft. (MVZ 19284); Death Valley, Cow Creek (MVZ 35360). San Bernardino Co.: Merango Valley, 4 mi. N. Riverside Co. line (MVZ 63571). MEXICO. Sonora: 136 mi. E. Mexicali, Mexico Highway 2 (LACM 20309); 23 mi. SSE. Sonoyta (LACM 25168-25169).
L. h. humilis < wutahensis intergrades (5). CAtrrorntA. Inyo Co.: Nelson Mountains, Grapevine Canyon, 4630 ft. (CAS 89592-89593); Nelson Mountains, Grapevine Canyon, 4480 ft. (CAS 89570); Saline Valley, S. end, Racetrack Valley Road, 2100 ft. (CAS 89591) ; Saline Valley, S. end, Lower Grapevine Canyon, 4000 ft. (CAS 89577).
ACKNOWLEDGMENTS
I thank the following, and their institutions, for the use of specimens: Alan E. Leviton (California Academy of Sciences); David B. Wake and Steven B. Ruth (Museum of Vertebrate Zoology, University of California, Berkeley) ; Robert L. Bezy and John W. Wright (Natural History Museum of Los Angeles County); and Thomas H. Fritts (San Diego Natural History Museum). For editorial and typing assistance I thank my wife Wrennie Murphy and Christine M. Brennan. A special thanks to Jean Durham for her assistance in locating geologic and paleontologic references and to Ute L. von Doyen for her assistance in preparation of the maps. To both Alan E. Leviton and Lawrence W. Swan I am indebted for critical evaluation and suggestions. Credit for figure 3 should go to W. H. Freeman and Company who kindly granted permission to reproduce the figure from Scientific American, vol. 225, no. 5, Dp. 60:
LITERATURE CITED
ANDERSON, C. A. 1950. 1940 E. W. Scripps cruise of the Gulf of California. Part I. Geology of islands and neighboring land areas. Geological Society of America Monograph 43, 53 pp. ANDERSON, D. L. 1971. The San Andreas Fault. Scientific American, vol. 225, no. 5, pp. 52-68. AUFFENBERG, W., AND W. W. MILsTEAD 1965. Reptiles in the Quaternary of North America, pp. 557-568. Im: H. E. Wright, Jr. and D. G. Frey [ed.]. The Quaternary of the United States. Princeton Univer- sity Press, Princeton. Banks, R. C., anp W. M. FARMER 1963. Observations on reptiles of Cerralvo Island, Baja California, Mexico. Herpeto- logica, vol. 18, no. 4, pp. 246-250. DurHaM, J. W., AND E. C. ALLISON 1960. The geologic history of Baja California and its marine faunas. Systematic Zoology, vol. 9, pp. 47-91.
Vol. XL] MURPHY: NEW BLIND SNAKES 107
Frint, R. F. 1971. Glacial and Quaternary Geology. John Wiley and Sons, Inc., New York, xiv + 892. Fox, W.
1948. Effect of temperature on development of scutellation in the garter snake, Thamnophis elegans atratus. Copeia, 1948, no. 4, pp. 252-262. Harpy, L. M., anp R. W. McD1Armip 1969. The Amphibians and Reptiles of Sinaloa, Mexico. University of Kansas Publi- cations of the Museum of Natural History, vol. 18, no. 3, pp. 39-252. Krauser, L. M. 1931. Notes on the worm snakes of the southwest, with descriptions of two new sub- species. Transactions of the San Diego Society of Natural History, vol. 6, no. 23, pp. 332-352. 1940. The worm snakes of the genus Leptotyphlops in the United States and northern Mexico. Transactions of the San Diego Society of Natural History, vol. 9, no. 18, pp. 87-162. 1956. Rattlesnakes: Their habits, life histories, and influence on mankind. University of California Press, Berkeley. 2 vol., xxix + 1476 pp. 1972. Rattlesnakes: Their habits, life histories, and influence on mankind. 2 ed. University of California Press, Berkeley. 2 vol., xxx + 1533 pp. Leviton, A. E., anp W. W. TANNER 1960. The generic allocation of Hypsiglena slevini Tanner (Serpentes: Colubridae). Occasional Papers of the California Academy of Sciences, no. 27, 7 pp. Savace, J. M. 1960. Evolution of a peninsular herpetofauna. Systematic Zoology, vol. 9, no. 3-4, pp. 184-211. STREETS, T. H. 1877. Contributions to the natural history of the Hawaiian and Fanning Islands and Lower California, made in connection with the United States North Pacific surveying expedition, 1873-1875. Bulletin of the U.S. National Museum, no. 7, 172 pp. Sour, M. E. 1961. Eridiphas slevini (Tanner) on Cerralvo Island, Gulf of California, Mexico. Her- petologica, vol. 17, p. 61. SouLkt, M. E., anp A. J. SLOAN 1966. Biogeography and distribution of reptiles and amphibians on the islands in the Gulf of California, Mexico. Transactions of the San Diego Society of Natural History, vol. 14, no. 11, pp. 137-156. Witson, I. F., anp V. S. RocHa 1955. Geology and mineral deposits of the Boleo copper district, Baja California, Mexico. U.S. Geological Survey Professional Paper 273, 134 pp. Witson, J. T. 1972. Preface and introductions. Jn: Continents Adrift: Readings from Scientific American. W. H. Freeman and Co., San Francisco. iv + 172 pp.
4
regina, tote BE
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- i
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PROCEEDINGS
OF THE i Wee CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 6, pp. 109-141; 4 figs.; 9 tables. February 18, 1975
THE ROCKFISHES, GENUS SEBASTES (SCORPAENIDAE), OF THE GULF OF CALIFORNIA, INCLUDING THREE NEW SPECIES, WITH A DISCUSSION OF THEIR ORIGIN
ay Lo-chai Chen Department of Zoology, San Diego State University, San Diego, California 92182
Apstract: Seven species of Sebastes, herein described and compared, occur in the Gulf of California at the near-tropical fringe of the range of the genus in the northeastern Pacific. Six of the species are endemic to the Gulf. Three of these, (S. spinorbis, S. varispinis, and S. peduncularis) are described as new. Only S. macdonaldi occurs also on the outer coast of Baja California.
Sebastes sinensis and S. cortezi are mesopelagic as well as benthic. Sebastes cortezi, like S. diploproa of the outer coast, inhabits floating vegetation as prejuve- nile; it undergoes extreme morphological change on leaving its surface habitat.
The distribution pattern of these seven species seems to be related to water tem- perature and dissolved-oxygen level.
Multiple invasion from the outer coast and subsequent radiation within the Gulf may both have contributed to the species diversity within the Gulf. Sebastes diplo- proa of the outer coast probably came from reciprocal invasion by S. cortezi from the Gulf.
INTRODUCTION
The rockfish genus Sebastes is represented by many species in the North Pacific. Only a few others are known from the North Atlantic and from the southern hemisphere. Chen (1971) listed 65 species of the genus for the Ameri- can coast of the North Pacific. Since then there have been three additions: S. borealis Barsukov (1970), S. variegatus Quast (1971), and S. rufinanus Lea and Fitch (1972).
[109]
110 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
The first species of Sebastes recorded from the Gulf of California were described as Sebastichthys sinensis in 1890 and Scorpaenodes cortezi in 1938. Lavenberg and Fitch (1966) reported specimens from the Gulf but did not assign species names. The next record was that of Chen (1971), who reported the occurrence of S. macdonaldi and described S. exsul from the Gulf. I here report seven species from the Gulf, of which three are described as new (two of these three were included, but not specifically named, in my 1971 list of 65 American North Pacific species).
With these additions, with another new species to be described by Lea and Fitch (personal communication), and with the placement of S. vexillaris in the synonymy of S. caurinus (Chen, in preparation), the number of known species of Sebastes in the eastern North Pacific is now 69.
ACKNOWLEDGMENTS
I thank Drs. Carl L. Hubbs and Richard H. Rosenblatt for reading the manuscript and for advice in naming the species. I thank those persons who supplied materials or loaned specimens in their care. In alphabetical order of the names of their institutions, they are: W. E. Eschmeyer, California Academy of Sciences; R. J. Lavenberg, Los Angeles County Museum; H. G. Moser, National Marine Fisheries Service; R. H. Rosenblatt, Scripps Institution of Oceanography; B. W. Walker and J. E. Bleck, University of California, Los Angeles; and B. G. Nafpaktitis, University of Southern California. This study was supported by NSF grant GB 34213.
METHODS
The terminology and methods in this report follow those of Chen (1971).
Abbreviations for the listed collections are: CAS, California Academy of Sciences; LACM, Los Angeles County Museum; SDSC, San Diego State University; SIO, Scripps Institution of Oceanography; UCLA, University of California, Los Angeles.
Key To THE ROCKFISHES, GENUS SEBASTES, OF THE GULF OF CALIFORNIA
la Lower edge of lachrymal with three or four small, spinelike projections; lateral-line pores more than 50; second anal spine shorter than third; lateral surface of lachry- mal with a spine in specimens longer than 10 cm. (fig. la) 9 S. macdonaldi
1b Lower edge of lachrymal with only two projections which may be spinelike and some- times split; lateral-line pores fewer than 50; second anal spine longer than third;
lateral surface of lachrymal without a spine (except in S. spinorbis) 2 2a. _Supraocular ;spines; présent. —_-.- e ee 3 2b- Supraocular’ spines absent) 2. eee 4
3a Lateral surface of lachrymal and/or orbital edge of first and/or second suborbital with spines: Pa les takers onminst yelleanclie29—330 (hice el|[)) eee S. spinorbis
3b Lateral surface of lachrymal and orbital edge of first and second suborbitals without spines) Ex) dl nakersionmninstaeeill=-anchyrs2—3i/a0 (il caeel|C) eee S. exsul
VoL. XL] CHEN: SEBASTES 111
4a Scales mostly cycloid, occasionally with weakly developed ctenii; dorsal soft-rays more often 11 than 12 (mouth and gill-cavity linings dusky in specimens 4 or 5 cm. long, jet-black in larger ones; ventral lachrymal projections pointing backward; rakers
Om ise eileen 2OL8s)) Gines, Gl eunGl Blo) ee S. sinensis 4b Scales mostly ctenoid, with ctenii evident in specimens 4 or 5 cm. long; dorsal soft- has NOS Orso WA-qdaein, il see Se eee zee 5
5a Mouth and gill-cavity linings black; lower jaw slightly projecting; dorsal spines some-
times 12 (20%); (rakers on first gill-arch 30 or more) (fig. 2a) S. varis pinis’ 5b Mouth and gill-cavity linings may be dusky, never black; jaws subequal; dorsal IS PDTTN ES marc TN OS Come cal OieLy Swe eie wm tapers eee a s ©
6a Head width in S.L. 6.5 or less; (rakers on first gill-arch 30 or fewer; anterior lachrymal projection directed forward in specimens longer than 6 cm.) (figs. Tes kee eenaG PAG) = a I A Rea Se Dre ee NE 8 ee S. cortezz
6b Head width in S.L. more than 6.5 (fig. 2e) ___ Se ee et S. peduncularis*
Sebastes exsul Chen, 1971. (Figure 1c.) Sebastes exsul CHEN, 1971, p. 27 (type locality: Gulf of California, at 28° 59’ N., 113°
25254 W)
Diacnosis. Sebastes exsul can be differentiated from all other species of Sebastes by the following combination of characters: (1) the color pattern of the subgenus Sebastomus (body pink or red, with white blotches at tip of opercle, bases of fourth, eighth, and last dorsal spines and last dorsal ray, and also one below ninth dorsal spine just above lateral line); (2) the cranial spine pattern (with nasal, preocular, supraocular, postocular, tympanic, and parietal spines) ; (3) the meristics (D. XIII, 12-13; A. III, 6; P; 17; rakers on first gill-arch 32-37; lateral-line pores 35-43); (4) a nearly vertical anal profile; (5) a narrow interorbital divided by frontal ridges into three grooves; (6) the oral and the gill-cavity linings white; (7) subequal jaws; (8) scaled mandibles; (9) short dorsal spines (orbit length in fourth dorsal spine less than 1.2); and (10) absence of spines on the lateral surface of lachrymal and the orbital edge of the first and second suborbitals.
Description. A detailed description was given by Chen (1971) and is supplemented here.
Principal caudal rays 14; vertebrae (in 8 specimens) 26 (11 + 15). Each one of the two extrinsic gas-bladder muscles (2 specimens examined) extends backward from an origin on or near the opisthotic, passes medial to the cleithrum to which it has a membranous connection, then splits into three tendons which pass between the second and third ribs, running straight along the inner side of succeeding ribs and inserting respectively on the parapophyses of the 7th, 8th, and 9th centra. The last tendon is split in one specimen, with the additional branch attached to the parapophysis of the 10th centrum. There is no direct connection between the tendons and the gas-bladder wall.
13-5 cm. juveniles only.
112 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Vor. XL] CHEN: SEBASTES 113
For morphometrics and meristics, see tables 1, 7, 8, and 9.
Remarks. This species is most likely to be confused with S. umbrosus, S. rosenblatti, and S. spinorbis. It can be distinguished from S. umbrosus by its lack of the conspicuous honeycomblike color pattern, from S. rosenblatti by its higher average number of gill rakers, and from both by its short dorsal spines. From S. spinorbis, S. exsul can be distinguished by the lack of spines on the lateral surface of the lachrymal and the orbital edge of the first and second sub- orbitals, by having only 17 pectoral rays, by having more rakers on the first gill-arch (32-37 instead of 29-33), by lacking spines on the lower edge of the gill-cover, and by having clear fin membranes and white oral and gill-cavity linings.
Since describing the species in 1971, I have acquired 21 additional specimens, all from the type locality. Of these specimens, one (SIO 69-437, 180 mm.) resembles S. spinorbis in having a spine on the lateral surface of each lachrymal, but it is regarded as S. exsul because the spine is weak and because it agrees with S. exsul in having 16-17 pectoral rays, 33-34 rakers on the first gill-arch, no spines on the lower edge of gill-cover, the fin membranes clear, and the oral and the gill-cavity linings white.
RANGE. Known only from the vicinity of the type locality, 28° 59’ N., 1S 92'5.5 Wi
MATERIAL EXAMINED. A total of 36 specimens, 150-212 mm.
Holotype. SIO 68-1, 171 mm., adult female, from 28° 59’ N., 113° 25.5’ W., 2.5 miles N. of Punta Roja, Bahia de los Angeles.
Paratypes. All from vicinity of the type locality. SIO 62-241, 7 (158-212); SI@ 168-3, 7 (150-202).
Other specimens. All from vicinity of the type locality. SIO 69-318, WNGITO=188 SIO 69-437) 14 (156=197)— CAS 17605, 1 (184).
Sebastes spinorbis Chen, new species. (Figure 1b.)
Dracnosis. Sebastes spinorbis can be differentiated from all other species of Sebastes by the following combination of characters: (1) the color pattern and the cranial spine pattern of the subgenus Sebastomus (see diagnosis of S. exsul); (2) presence of a spine on the lateral surface of the lachrymal and/or the orbital edge of the first and second suborbitals; (3) 18 pectoral rays; (4) 29-33 gill-rakers; and (5) dusky to black pectoral membrane.
Ficure 1. a). Sebastes macdonaldi, adult male, 412 mm., SDSC 74-12. b). S. spinorbis, holotype, adult male, 225 mm., SIO 69-318. c). S. exsul, holotype, adult female, 171 mm., SIO 68-1. d). S. sinensis, adult female, 149 mm., LACM 30065-1. e). S. cortezi, adult female, 207 mm., LACM 33939-1.
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{[Proc. 4TH SER.
Vot. XL] CHEN: SEBASTES 115
TaBLE 1. Morphometrics (standard length/body part measurement) of Sebastes exsul and Sebastes varispinis. Characters are in sequence as given for the subgenus Sebastomus by Chen (1971), with the most frequently negatively allometric one placed above and the most frequently positively allometric one placed below. “-+-” means significant positive allometry and “— means significant negative allometry.
S. exsul S. varis pinis size range, mm. 151-212 33-53 Character no. 36 20 2nd anal spine 5.1 — 6.3 4.3 -— 5.5 Gill-raker at angle 18 — 31 159 — 2 3rd anal spine 6.1 — 7.6 5.6 — 7.5 2nd dorsal soft-ray 5.7 — 6.9 5.1 - 7.4 1st anal spine 10.0 -12.8 8.4 -11.3 Orbit length 7.1 — 8.4 8.4 -10.0 Total length 0.81— 0.84 0.80— 0.83 4th dorsal spine 6.6 — 8.1 4.6 — 6.2 1st anal soft-ray 4.7 — 5.4 5.0 — 6.2 Anal-fin base 6.6 — 8.1 5.9 — 7.5 Pelvic-fin length 4.1 -— 4.9 4.2 — 4.9 Predorsal length 2.55 — 2.8 2.8 — 3.2 Pectoral-fin length Sel S37 3.3 -— 3.8 Lower peduncle length 4.8 — 5.6 4.8 — 5.9 Soft-dorsal base 4.4 — 5.0 4.1 - 5.1 Dorsal-fin incision 12 — 16 7.6 -13.8 Caudal-peduncle depth 9.5 -11.2 10.2 —11.6 Upper peduncle length 6.6 — 8.4 6.2 — 7.6 Spinous-dorsal base 2.5 — 2.9 2.8 — 3.3 Snout length 8.7 -11.4 (+) 11.0 -13.2 Prepelvic length 2.1 — 2.5 (+) 2.3 — 2.7 Head length 2.3 — 2.4 (+) 2.6 — 3.0 Upper-jaw length 4.6 — 5.1 (+) 5.6 — 6.8 Preanal length 14 - 1.5 1.4 - 1.5 Body depth lV MES (aE) Cech Head width 4.6 — 54 (+) 6.3 — 8.0 Lachrymal width 27 — 38 50 — 62
Interorbital width 13 —- 17 (+) 14 — 16
DESCRIPTION: Wy XGMIT13=14- A. II. 6; Py 18) (1l6—18). C. 14= rakers on first gill-arch 29-33 (9-10 + 20-23); lateral-line pores 33-41; vertebrae (in 4 specimens) 26 (11 + 15).
Base of skull straight. Interorbital concave, divided by frontal ridges into
<
Ficure 2. a). Sebastes varispinis, holotype, subsurface juvenile, 49 mm., LACM 8821-11. b). S. sinensis, juvenile, 46 mm., LACM 30064-3. c). S. cortezi, transformed juvenile, 51 mm., SIO 68-90. d). S. cortezi, surface prejuvenile, 38 mm., SIO 62-239. e). S. peduncu- laris, holotype, juvenile, 43 mm., LACM 8818-6.
116 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER
three grooves. Nasal, preocular, supraocular, postocular, tympanic, parietal, and, frequently, nuchal spines present, sharp, and strong. Parietal ridges well developed, elevated. Lower edge of gill-cover where interopercular and suboper- cular meet often with one or two spines. Preopercular spines equally spaced, the upper two directed backward, the lower two or three downcurved. Lachrymal projections two, spinelike, directed backward, sometimes bifurcate; lateral sur- face of lachrymal above and between lachrymal projections and orbital edge of first and second suborbitals each often bearing a sharp, backward-directed spine. Supracleithral, cleithral, and the two opercular spines well developed.
Symphyseal knob round, directed downward; jaws subequal. Maxillary reaching to vertical from posterior edge of orbit.
Scales ctenoid. Upper and lower lips and snout between and before nasal spines scaleless; branchiostegals with or without scales, head scaled elsewhere. Fin rays scaled; fin membranes scaled basally, most extensively on anal and caudal.
Dorsal spines increasing in length to 4th, then decreasing gradually to 12th, 13th spine longer than 12th; soft dorsal high, with the short posterior rays when depressed reaching to about midpoint of caudal peduncle. Second anal spine stronger and longer than third, its tip extending to about two-thirds the length of first anal soft-ray when depressed; tips of posterior anal soft-rays, when depressed, exceeded by tips of anterior anal soft-rays. Profile of anal nearly vertical. Caudal truncate. Origin of pectoral below third or fourth dorsal spines; pectoral pointed, with longest ray (usually 11th) notably longer than adjacent rays; pectoral extending beyond tip of pelvic to above anus. Origin of pelvic below 4th dorsal spine; tip of pelvic reaching to anus or falling short by one-fifth of pelvic-to-anal distance. Origin of anal below 3rd or 4th dorsal soft-ray, insertion of anal below 9th to 11th dorsal soft-ray.
Extrinsic gas-bladder muscles (one paratype examined) as in S. exsul.
Color in life pink to red. Body with six white blotches as in S. exsul. Dark green dusky patches on top and sides of head, above lateral line, and along basal part of dorsal fin, those on the back sometimes disrupted into vermiculations. Fins pink, conspicuously dusky on membranes of pectoral, soft-dorsal, and caudal. Scales on side of body with dusky margin. Gill cavity and mouth lining dusky. Peritoneum blackish. In preserved condition dusky marks become black and pink or red fade.
For morphometrics and meristics, see tables 2, 7, 8, and 9.
Erymotocy. The name spinorbis, from the Latin, refers to the spines along the lower edge of the orbit.
Discussion. Because only four specimens are available and because the several distinguishing characteristics are subject in the genus to some variation, it is difficult on the basis of single characters to differentiate S. spinorbis from S. exsul. The spine on the lateral surface of lachrymal, although not found
VoL. XL] CHEN: SEBASTES 117
TasLeE 2. Morphometrics (standard length/body part measurement), meristics (‘*” signifies left and right counts), and head spines of Sebastes spinorbis.
Holotype Paratypes
Character SIO 69-318 SIO 69-437 CAS 30689 Standard length (mm.) 225 259 241 21325 Morphometrics:
2nd anal spine 5.9 5.9 5.9 5.6 Gill-raker at angle Dil 24 25 25 3rd anal spine 6.4 6.9 6.4 Holl 2nd dorsal soft-ray 5.9 6.5 6.1 6.0 1st anal spine Teal WE 11.0 11.4 Orbit length 8.7 8.2 8.7 8.1 Total length 0.83 0.83 0.83 0.81 4th dorsal spine 8.0 7.9 (oll 7.4 1st anal soft-ray 4.6 5.0 4.7 4.7 Anal-fin base eS 7.6 1S) 7.4 Pelvic-fin length 4.2 4.5 4.2 4.2 Predorsal length Zi 2.6 2.6 2.6 Pectoral-fin length S6 3.4 Sil 33 Lower peduncle length Sy) 5.8 Sail 5.3 Soft-dorsal base 4.8 4.8 4.8 4.5 Dorsal-fin incision 15.3 15.0 15.8 14.6 Caudal-peduncle depth 9.9 10.4 9.9 9.7 Upper peduncle length 7.6 8.0 Coll eS Spinous-dorsal base DH 2.8 Dell Bell Snout length 10.1 10.0 9.2 9.7 Prepelvic length 2.6 2.4 DS 2.4 Head length 7 gl OBS, De) 2.3 Upper-jaw length 4.8 4.4 4.7 4.6 Preanal length 1.40 1.34 1.41 1.39 Body depth 2.6 2.4 2.6 25 Head width 4.6 41 4.5 4.3 Lachrymal width 30 30 28 30 Interorbital width 15 14 13 16 Meristics: Lateral-line pores * 38-38 37-36 33-35 41-40 Dorsal soft-rays 13 14 13 13 Pectoral rays * 18-18 18-18 18-18 16-18 Anal soft-rays 6 6 6 6 Gill-rakers * 30-31 31-32 33-32 29-29 Head spines *:
Nuchal 1-1 0-0 0-1 0-0 Edge of lower gill cover 2-2 1-0 2-2 2-2 Lateral surface of lachrymal 1-1 1-1 1-1 0-1 Orbital edge of 1st suborbital 0-1 0-0 1-0 0-0 Orbital edge of 2nd suborbital 1-1 0-0 0-0 0-0
118 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
in any other species of subgenus Sebastomus, is variably present in Sebastiscus marmoratus (personal data), and does not develop in Sebastes macdonaldi until the fish reaches 10 cm. The spines along the orbital edge of the first and second suborbitals develop only in adults in Sebastes aleutianus. The higher pectoral-ray count (significantly different from that of S. exsul at 0.01 level, rank-sum test), the lower gill-raker number (significantly different from that of S. exsul at 0.002 level, rank-sum test), the presence of spine(s) on lower edge of gill cover, and the dusky fin membranes, all are variable and overlap with S. exsul. The separation of the two species is based on: (1) the non- obligatory correlation of the above characters (higher pectoral-ray count and wider pectoral base, or larger eye and narrower lachrymal would be obligatorily correlated); (2) the larger size of the specimens of S. spimorbis; and (3) the co-occurrence of the two forms (taken at the same stations from the same depths. )
MATERIAL EXAMINED. Four specimens, 214-259 mm., two males and two fe- males, all mature, but with gonads in resting stage.
Holotype. SIO 69-318, 225 mm., male, from 28° 58.5’ N., 113° 26.5’ W., 2.5 miles NE. of Punta Rojas, Bahia de los Angeles, collected by hook and line from bottom at 130-160 m., by H. G. Moser on 29 April 1969.
Paratypes. All from the type locality at depths between 130-160 m. SIO 69-437, 1 (259); CAS 30689, 2 (214-241).
Sebastes sinensis (Gilbert, 1890).
(Figures 1d and 2b.)
Sebastichthys sinensis GILBERT, 1890, p. 81 (type locality: Albatross Station 3015, at 29° 19? 00” Ne 11122"'50") 00” Wo).
Pteropodus sinensis: EIGENMANN & BEESON, 1893, p. 670 (relationships); 1894, p. 397 (species analysis).
Sebastodes sinensis: CRAMER, 1895, p. 600 (species analysis). JoRDAN & EVERMANN, 1896, p. 431 (placed in subgenus Hispaniscus) ; 1898, pp. 1776 & 1813 (key and description). Husss, 1951, p. 129 (designated as type species of subgenus Allosebastes).
Hispaniscus sinensis: JORDAN, EVERMANN, & Clark, 1930, p. 368 (name only).
Sebastes sinensis: CHEN, 1971, pp. 63 & 77 (meristics and distribution).
Remarks. M’Clelland (1843) described Sebastes sinensis from China. The ‘sinensis’ of M’Clelland is now considered to be a junior synonym of Sebastiscus marmoratus and is not to be confused with the present species. Since the ‘sinensis’ of Gilbert has been placed into the genus Sebastes previously (Chen, 1971), according to the subsection (ii) added in 1972 to the Article 59 (b) of the International Code of Zoological Nomenclature, the ‘sinensis’ of Gilbert is not to be rejected for its secondarily homonymous condition.
Dracnosis. Sebastes sinensis can be differentiated from all other species of Sebastes by the following combination of characters: (1) the cranial spine pattern (with sharp nasal, preocular, postocular, tympanic, parietal, and occasionally nuchal spines); (2) the meristics (D. XIII, 11 (11-13); A. III,
Vout. XL] CHEN: SEBASTES 119
5—6: P, 18 (17-19); rakers on first gill-arch 29-33); (3) division of the inter- orbital by frontal ridges into three shallow grooves; (4) sharp, spinelike, back- ward-directed lachrymal projections; (5) the oral and the gill-cavity linings jet-black; and (6) predominantly cycloid rather than ctenoid squamation.
DESCRIPTION: (Da ub Tie Gui 13) eA SI 6" (5-6): Px 18) (L7=19)); C. 14; rakers on first gill-arch 29-33 (8-10 + 20-23); lateral-line pores 38-43 (36-46); pyloric caeca (in 6 specimens) 9-12; vertebrae (in 25 specimens) 26 Guibseaey.
Base of skull straight. Interorbital concave, divided by frontal ridges into three shallow grooves. Nasal, preocular, postocular, tympanic, parietal, and occasionally nuchal spines present, sharp, and moderately strong. Parietal ridges well developed and elevated. Lower edge of gill-cover spineless. Upper three or four preopercular spines parallel, pointing straight backward or obliquely upward, lower one or two preopercular spines tend to point downward and be more widely spaced. Lachrymal projections two, spinelike, directed backward, occasionally bifurcate. Supracleithral, cleithral, and the two opercular spines well developed.
Symphyseal knob inconspicuous in small specimens, directed downward in large ones; jaws subequal. Maxillary ending behind vertical from posterior edge of pupil but before margin of orbit.
Most scales cycloid, occasionally with weakly developed ctenii. Lips and snout between and before nasal spines scaleless; mandibles and maxillaries scaleless or rarely with patches of fine scales, skin smooth to touch; branchios- tegals scaleless; head scaled elsewhere. Fin rays scaled; fin membranes scaled basally, most extensively on anal and caudal.
Dorsal spines increasing in length to 4th, then decreasing gradually to 12th, 13th spine longer than 12th; soft dorsal high, with the short posterior rays when depressed reaching to near midpoint of caudal peduncle. Second anal spine stronger and longer than third, its tip often reaching to or beyond tip of first anal soft-ray when depressed; tips of posterior anal soft-rays, when de- pressed, exceeded by tips of anterior anal soft-rays Profile of anal nearly vertical. Caudal truncate. Origin of pectoral below 3rd or 4th dorsal spines; margin of pectoral fin rounded, 9th to 11th rays longest; pectoral extending beyond tip of pelvic to above anus. Origin of pelvic below 4th or 5th dorsal spine; tip of pelvic reaching to anus (small specimens) or falling short by one-fourth of pelvic-to-anal distance (large specimens). Origin of anal below 4th or 5th dorsal soft-ray, insertion of anal below 9th to 10th dorsal soft-ray.
Extrinsic gas-bladder muscles (7 specimens examined) as in S. exsul, except that each muscle sometimes splits into only two instead of three tendons, inserting respectively on parapophyses 8 and 9, 9 and 10, or 8, 9, and 10.
Color in life pink or red, with light to heavy dusky patches on back and sides. Membranes of the edge of spinous dorsal and distal part of pectoral
120 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
TasLe 3. Morphometrics of Sebastes sinensis. Explanations as table 1.
size range mm. 44-49 51-99 102-152
Character no. 6 25 27 2nd anal spine — 3.8 — 4.4 3.6 — 4.5 4.1 — 5.2 Gill-raker at angle — 18 — 23 17 — 23 18 — 32 3rd anal spine — 48-57 4.7 — 5.6 4.8 — 5.8 2nd dorsal soft-ray — 5.0 — 5.8 Sell = (0-7) 5.5 — 6.5 1st anal spine 8.8 —10.8 8.2 -10.9 8.4 -11.5 Orbit length ++ 6.9 — 7.3 6.8 — 8.1 6.3 -— 8.1 Total length — 0.78- 0.81 0.79— 0.83 0.80— 0.84 4th dorsal spine — 4.4 — 5.2 4.1 — 5.3 4.6 — 5.6 Ist anal soft-ray 4.7 — 5.9 4.7 — 5.7 4.6 — 5.5 Anal-fin base — 510) = 9723 6.3 — 7.6 6.7 — 8.8 Pelvic-fin length — 41-45 4.0 — 4.5 4.1 — 4.7 Predorsal length BA) = Dall 2.6 — 2.8 2.55 — 2.9 Pectoral-fin length — 3.4 — 3.6 3.2 — 3.7 3.2 — 3.8 Lower peduncle length — 4.9 — 5.5 48-58 Holl = 6.2 Soft-dorsal base 4.7 — 5.4 44 -— 5.8 44 — 5.6 Dorsal-fin incision — 7.0 — 8.0 6.4 — 9.9 7.7 -10.1 Caudal-peduncle depth 11.4 -12.5 10.8 -13.1 10.2 —13.1 Upper peduncle length — 6.7 — 7.5 6.2 — 7.5 6.6 — 8.0 Spinous-dorsal base 2.7 — 3.0 2.6 — 3.0 2.6 — 3.2 Snout length + 11.6 —13.0 10.5 —12.4 9.8 -11.8 Prepelvic length + 2.4 — 2.6 Be D7 2.3 — 2.6 Head length + 2.4 — 2.5 24 — 2.5 2.3 — 2.5 Upper-jaw length + 5.3) — 5:8 5.0 — 5.7 4.8 — 5.6 Preanal length + 5 1.4 — 1.5 13 — 1.4 Body depth + 3.0 — 3.2 2.7 — 3.3 2.7 = 13 Head width + 5.6 — 6. 5.4 — 6.6 5.0 — 6.5 Lachrymal width + 46 -— 53 41 — 52 39 — 52
Interorbital width + 16 - 17 15 — 19 14 = 17
relatively clear in juveniles but becoming conspicuously black in adults; other fin membranes dusky over pink. Specimens in alcohol pale, with dusky to black patches on the back and on various fin membranes and on the edge of the dorsal. Peritoneum and oral and gill-cavity linings jet-black.
For morphometrics and meristics, see tables 3, 7, 8, and 9.
RANGE. The range of this species is rather limited. In spite of the numerous collections, S. sinensis is definitely known only within a range of about one degree square (28° 35’ to 29° 49’ N., 112° 50’ to 113° 59’ W., see Material examined). The only exception is from a questionable record (CAS SU 123, 31° 227Ni, 114° 07" 45” W., 34 m:, 18:4°%C,, 25 March 1880).
S1zE. S. sinensis is a small fish. Females as small as 99 mm., collected 18 January 1968, carry eyed embryos. The largest female examined measures 152 mm. and the largest male 147 mm.
VoL. XL] CHEN: SEBASTES 1
bo —
DeptH. Most specimens examined were collected with otter trawls at depths ranging from 290 m. to 654-670 m. Some of the specimens, however, were collected with midwater trawls far above the bottom (in one case at a depth of less than 500 m. where the bottom is at 1400 m. The circumstance that these specimens, both juveniles and adults, possess characteristics not normal to epipelagic prejuveniles of Sebastes, indicates that S. sinensis occurs meso- pelagically.
MATERIAL EXAMINED. A total of 102 specimens from 46 to 150 mm.
Holotype. USNM 43085, 150 mm., adult male, from Albatross Station 3015, ZOE NIAOOM IN. 112.507 00" W.,. 290) m.
Other material. SDSC 72-28, 10 (59-133), 29° 48.7’ N., 113° 57.2’ W. SIO 63-805 158 (104=13)5)), 29> 43-07 IN. 113° 58:0" Wisitor 29° 40:22 Ni, 113°°55.47 Wi SI@iG3=905 26 (94-1149), 290-93 059N. 1S > 55.59 We to 29243250 Ne 113 59.07 Nye SI@ 2638-10215) ((55=62))29> 00 50N. Ise 7.54 We to 287 S6slo N= 113” 12.1’ W., to 28° 47.4’ N., 113° 08.0? W. LACM 8821-12, 5 (58-117), 28° 58’ Netlose des We LACM 8818-8..12. (106-156), 28° 55° Ne I22 505° W. AGIs 0064-3514 (46-128). 28° 46045" Ni, 113~ 06%007 Ws to28e 35° 367 NGI 527 56% W. LACM 30065-1, 22 (47-148), 28° 35” Ni. 112° 52” Wi, to 29-7037 007 Nz 113> 21” 30% W. CAS SU 123,11 (119), Aldcizoss Station 3026 (Giltee22-oN,. 114° 077-45” W.)-
Sebastes cortezi (Beebe and Tee-Van). (Figures le, 2c, and 2d.) Scorpaenodes cortezi BEEBE & TEE VAN, 1938, p. 304 (type locality: 24° 55’ N., 110°
20’ W.).
Sebastes cortezi was first described on the basis of suface-living prejuveniles. In the original description the species was assigned to the genus Scorpaenodes because of the poorly developed palatine teeth in the type. The pointed posterior end of the suborbital stay, however, indicates clearly that the species is of the genus Sebastes, and it was so listed by Chen (1971).
Diacnosis. The combination of the cranial spine pattern (with sharp nasal, preocular, postocular, tympanic, parietal, and occasionally nuchal spines) and the forward direction of the spinous lachrymal projection in adults differentiates this species from all other species of the genus except S. diploproa. This species is most similar to S. diploproa and S. sinensis. It differs from the former in lacking dentigerous knobs and a conspicuous symphyseal knob and in having fewer rakers on the first gill-arch (31 or fewer). From S. sinensis it differs in having an anteriorly directed, spinelike lachrymal projection, a dusky rather than jet-black gill-cavity lining, ctenoid rather than cycloid scales, and more often 12 rather than 11 dorsal soft-rays.
Surface-living prejuveniles of S. cortezi can be distinguished from those of other species of Sebastes by the following combination of characters: (1)
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the cranial spine pattern (with skin-covered and therefore blunt-looking spines) ; (2) flat or slightly concave interorbital with inconspicuous frontal ridges; (3) the meristics (D. XIII, 12 (11-12); A. III, 6 (5-6); P; 18 (17-19); rakers on first gill-arch 27-31); (4) small eyes (8.4-10.0 in SL in 28-43 mm. speci- mens); and (5) a distinctive color pattern (fig. 2d).
DESCRIPTION. —D: XIII, 12 (11-12); A. IH, 6 (5=6)3 Py 189(17=E C. 14; rakers on first gill-arch 27-31 (8-10 + 18-22), with the first 3-8 rakers on each limb of first arch often rudimentary; lateral-line pores 33—45; pyloric caeca (in 3 specimens) 9-10; vertebrae (in 33 specimens) 26 (11 + 15).
Base of skull straight. Nasal, preocular, postocular, tympanic, parietal, and occasionally nuchal spines present. Parietal ridges well developed and elevated. Lower edge of gill-cover spineless. Supracleithral, cleithral, and both opercular spines well developed.
Symphyseal knob inconspicuous; jaws subequal. Maxillary ending behind vertical from posterior edge of pupil but before margin of orbit.
Scales ctenoid. Lips and snout between and before nasal spines scaleless; mandibles and maxillaries scaleless or with patches of fine scales, skin smooth to touch; branchiostegals scaleless; head scaled elsewhere. Fin rays scaled; membranes scaled basally, most extensively on anal and caudal.
Dorsal spines increasing in length to 4th or 5th, then decreasing gradually to 12th, 13th spine longer than 12th; soft dorsal high, with the short posterior rays when depressed reaching to only about midpoint of caudal peduncle. Second anal spine stronger and longer than third, its tip reaching to tip (transformed juveniles) or falling short by one-fourth of the length (surface- living prejuveniles and large adults) of first anal soft-ray when depressed. Caudal truncate. Origin of pectoral below 2nd or 3rd dorsal spine; margin of pectoral fin rounded, 9th to 11th rays longest; pectoral extending beyond tip of pelvic to near anus. Origin of pelvic below 3rd or 4th dorsal spine; tip of pelvic reaching to anus (transformed juveniles) or falling short by one-fourth of pelvic-to-anal distance (surface-living prejuveniles and large adults). Origin of anal below 2nd dorsal soft-ray, end of anal below 10th to 11th dorsal soft-ray.
Extrinsic gas-bladder muscles (4 specimens examined) as in S. exszl, except that each muscle splits variably into three to six tendons, with variable insertions. Of the four specimens examined, the tendons insert on parapophyses SOO 4 65 75, 83 4, 6, 7,8, 9: sands 5,00.) 5,slosn/ Oo ehespectiviely:
In other respects the surface-living prejuveniles differ from the transformed stages that live below the surface. The interorbital is flat rather than being concave and divided by frontal ridges into three grooves. Cranial and fin spines are so well covered with skin to appear blunt, rather than being superficially sharp. The preopercular spines are spaced equally and radiate, rather than having the two upper ones directed backward and the three lower ones down- curved. The two lachrymal projections are blunt rather than becoming spine-
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Taste 4. Morphometrics of Sebastes cortezi. “*” signifies significant difference (analysis of covariance, P < .005) between prejuveniles and transformed specimens. Other explanations
as table 1. Prejuveniles Transformed specimens size range mm. 28-40 51-91 104-144 160-198 208-255 Character no. 20 4 5 4 2 2nd anal spine* — 5.8 — 8.1 4.0 — 5.0 44 -— 5.9 5.1 — 6.4 5.9 — 6.9 Gill-raker at angle — 18 — 27 19 = 23 20 — 34 23 = 42 34 — 42 3rd anal spine * — 7.0 — 9.6 4.9 — 7.0 5.2 — 6.7 6.0 — 7.3 6.5 — 7.6 2nd dorsal soft-ray — 56) = Osfl 5.7 — 6.0 5.8 — 6.3 6.3 — 6.6 5.9 — 6.7 Ist anal spine * — 10.5 -19.3 8.6 —10.3 8.8 —11.2 10.0 —13.1 11.8 -17.5 Orbit length * — 8.4 -10.1 6.6 — 7.1 6.6 — 8.0 7.3 — 7.6 8.4 -— 8.5 Total length — 0.78- 0.81 0.79-— 0.81 0.80— 0.81 0.81— 0.82 0.82— 0.84 4th dorsal spine * — 68 = Tal 5.0 — 5.8 5.6 — 7.4 6.6 — 7.3 7.1 — 7.4 1st anal soft-ray * 5.2 — 6.7 4.7 — 6.0 49 — 5.5 4.9 — 5.7 53} Anal-fin base — 6.1 — 7.2 6.0 — 8.1 7.2 — 8.9 7.6 — 8.5 75 — 7.6 Pelvic-fin length — 4.5 -— 5.2 4.2 — 5.0 4.5 — 5.4 4.9 — 5.3 4.9 — 5.2 Predorsal length 2.7 — 3.0 2.7 — 2.8 2.5 — 2.8 2.6 2.6 — 2.7 Pectoral-fin length * 3.6 — 4.3 3.4 -— 3.7 3.3 — 3.6 3.4 -— 3.7 3.6 — 3.7 Lower peduncle length * Aer 5.1 -— 5.7 4.7 — 5.6 5.2 — 5.5 iP) Soft-dorsal base 4.1 — 5.0 4.7 — 4.9 4.4 -— 5.3 4.6 — 4.9 4.8 — 4.9 Dorsal-fin incision * — 1 > 22 8.9 -11 11 -— 17 Lit = US 12 — 14 Caudal-peduncle depth * 8.7-10.3 11.5 —12.6 11.3 -13.0 11.4 -12.2 10.9 -11.7 Upper peduncle length Gil = 75 6.8 — 7.8 7.0 — 8.0 7.1 — 8.0 Tal = 758 Spinous-dorsal base ++ Dh = BO 2.9 — 3.1 2.6 — 2.9 2.6 — 2.8 2.55 — 2.8 Snout length 9.3 -11.4 10.2 —12.4 9.5 -11.9 10.1 —11.4 9.9 -10.7 Prepelvic length 2.4 — 2.7 2.4 - 2.5 2.4 — 2.7 24 = 231 2:4 — 2.5 Head length * 2.6 — 2.8 2.4 DO = BS DL = AS DBO) r= CS) Upper-jaw length 55) = 08 Sal = 5 4.9 — 5.8 5.0 — 5.3 4.7 — 5.2 Preanal length 1.5 — 1.6 1.4 -— 1.5 (eS aleS 1.4 - 1.5 1.4 Body depth * + PE ee Sith enw eS ie BYR el ho a RS Head width + 5.1 — 6.5 5.6 — 5.8 5s) = Ss) 4.8 -— 5.2 4.2 — 4.7 Lachrymal width + 36 — 55 40 - 51 35 = uy 38 — 41 31 — 40 13 - 15
Interorbital width * 139— 16 15 — 19 15, —)20 Wh = iy
like, with the anterior one directed forward and both occasionally bifurcate. Tips of posterior anal soft-rays, when depressed, exceed rather than are ex- ceeded by tips of anterior anal soft-rays. Profile of anal slanting slightly posteri-
orly, rather than being vertical.
Life color of surface juveniles, as quoted from the original description: “Body dark reddish-brown with a yellowish tinge, paler below, the entire body covered with black punctulations and irregular greenish-yellow blotches, the latter especially marked posteriorly. Dorsal fin dark brown, the membranes of the spinous dorsal black, the soft dorsal irregularly blotched with lighter. Caudal fin yellowish-green, without pattern. Pectoral fin brownish, yellow
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40
20
BODY PART LENGTH BODY PART LENGTH fo)
a
20 50 100 200 STANDARD LENGTH
Ficure 3. A) Head width and caudal peduncle depth and B) second anal spine length and orbit length of Sebastes peduncularis (x), and prejuvenile (open circles) and transformed (solid dots) S. cortezz.
toward the tip and dusky at the base. Pelvic fins black at base, yellow at tips and with scarlet along the anterior edge. Anal fin black at base, yellow on outer half and with a small scarlet patch at base of the first spine.” Specimens in alcohol light brown, with dusky marks in pattern as illustrated (fig. 2d). Peri- toneum and oral and gill-cavity linings slightly dusky.
Life color of adults pink, with trace of dusky. Specimens in alcohol pale, with trace of dusky on back, on various fin membranes, and on edge of dorsal. Peritoneum dusky to jet-black. Oral and gill-cavity linings slightly dusky.
For morphometrics and meristics, see tables 4, 7, 8, and 9.
Discussion. In fishes, transition from epipelagic to benthic stages often is associated with a growth infection, resulting in altered body form, as demon- strated in Pseudupeneus maculatus by Caldwell (1962) and in Sebastolobus species by Moser (1974). The surface-living prejuveniles (28-43 mm.) and the transformed specimens (51-255 mm.) of Sebastes cortezi differ markedly in appearance, as is indicated in the species description. In addition, the surface- living prejuveniles have a number of significantly different morphometrics, par- ticularly the heavier caudal peduncle, the smaller eye, and the shorter dorsal and anal spines (table 4, fig. 3). Many of the same differences, however, also exist (personal observation and George Boehlert, personal communication) between epipelagic prejuveniles and benthic forms in S. diploproa, a species the adults
Vot. XL] CHEN: SEBASTES 125
of which are most similar to the 15 transformed specimens herein called S. cortezi. It is on the basis of the known ontogenetic changes in the closely related S. diploproa and the identical meristics (tables 7, 8, and 9) that I consider the 15 specimens to be the older stage of S. cortezi.
RANGE. Chen (1971) in giving the range of this species as 26° N., 111° 49’ W. to 29° N., 113° 30’ W., overlooked the type locality and listed the trans- formed specimens of the species as species “a.” The corrected range of S. cortezi should be 24° 55’ N., 110° 20’ W. to 29° 43.9’ N., 113° 58.0’ W.
DeptH. Ten of the 15 transformed specimens examined, including the several largest and the several smallest, were collected by otter or beam trawls at depths ranging from 200-270 to 800-1100 m. The other five specimens (91-160 mm. long), however, were collected with midwater trawls in areas with 1000-1060 and 1400-1700 m. of water. The exact depth of capture of these specimens cannot be determined but presumably they were mesopelagic.
Size. The largest male examined measures 255 mm.; the largest female examined, 207 mm. This 207 mm. female was collected on 2 December 1967; its ovary contains large ova.
MATERIAL EXAMINED. A total of 93 specimens.
Surface-living prejuveniles: 78 (15.5—43).
Holotype. CAS SU 46503 (formerly Dept. Trop. Res. N. Y. Zool. Soc. 24889-A), 43.4 mm., taken at surface in weed, at 24° 55’ N., 110° 20’ W., 8 April 1936.
Other specimens. SIO 62-239, 77 (15.5-39.8), 29° N., 11326307 Wee also taken at surface in weed.
Transformed specimens: 15 (51-255).
SHO68-89*2 (61),.29° 43.0’ N., 113° 58:0’ W. to 29° 40.2’ N., 113° 55.4 W. Si@ros-90- 5, (51-198), 29° 39.5’ N., 113° 55.5’ W. to 29° 43.5’ N., ise 597 W._ SIO 59-205. 42(91—139), 28° 45.0’ N., 113° 03.4’ W. LACM 33939-1, PatA=207)~ 28°r 38 Nes 113° 00’ ‘We to 28° 33’ N., 112° 50° W. LACM BGWO5-6 1 (160) 928° 35’ 26" Ni 1129 52756” W: to 29° 037 00” N., (i lpn 7 be BO We bACMi883 7-3. 14(255), 26° 59:17 N., 111° 48.9" W.
Sebastes varispinis Chen, new species. (Figure 2a.)
Diacnosis. Sebastes varispinis can be differentiated from all other eastern North Pacific species of Sebastes by the following combination of characters: (1) the cranial spine pattern (with sharp nasal, preocular, postocular, tym- panic, parietal, occasionally nuchal, and, rarely, coronal spines); (2) presence of frontal ridges; (3) the meristics (D. XIJ-XIII, 11-12 (11-13); A. III, 5-6 (5-7): P, 18 (17-19); rakers on first gill-arch 29-33); (4) slightly protruding lower jaw; (5) small eyes (orbit 8.4-10.0 in S. L. in 33-53 mm. specimens) ; (6) slender caudal peduncle (peduncle depth 10.3—11.6 in S. L. in 33-53 mm. speci-
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mens); (7) the oral and the gill-cavity linings black; and (8) a color pattern as illustrated (fig. 2a).
Although the present species description is based on juveniles and no adults are known, the black oral and gill-cavity linings and the protruding lower jaw (two characters that are believed to persist into the adult stage), together with the tendency for reduction of the number of dorsal spines from 13 to 12, and the ctenoid rather than cycloid scales will differentiate adults of this species from those of other species of Sebastes in the Gulf.
DescriPTIon. D. XIIJ-XIII, 11-12 (11-13); A. III, 5-6 (5-7); P, 18 (17-19); C. 14; rakers on first gill-arch 29-33 (8-10 + 20-23); lateral-line pores 37-44; pyloric caeca (in 2 specimens) 9; vertebrae (in 32 specimens) D6 (aie =e ey,
Base of skull straight. Interorbital divided by frontal ridges into three shallow grooves. Nasal, preocular, postocular, tympanic, parietal, occasionally nuchal, and rarely coronal spines present, sharp, and moderately strong; many of the specimens have pterotic spines although this is probably a juvenile characteristic (Moser, 1972). Parietal ridges well developed and elevated. Lower edge of gill-cover spineless. Upper four preopercular spines directed backward, lowermost preopercular spine directed downward and often more widely spaced. Lachrymal projections two, spinelike, directed downward and backward. Supracleithral, cleithral, and the two opercular spines well developed.
Symphyseal knob inconspicuous. Tip of lower jaw projecting beyond that of upper. Maxillary reaching to between verticals from mid-orbit and posterior margin of pupil.
Scales ctenoid, many of the specimens examined have scales with a single spine. Mandibles, maxillaries, branchiostegals, and the head before eyes scale- less; head scaled elsewhere.
Dorsal spines increasing in length to 4th or 5th, then decreasing gradually to 11th, 12th spine often slightly longer than 11th, 13th spine nearly twice as long as 11th; soft dorsal high, with the short posterior rays when depressed reaching to near midpoint of caudal peduncle. Second anal spine stronger and longer than third, its tip nearly reaching to tip of first anal soft-ray when depressed; tips of posterior anal soft-rays, when depressed, sometimes exceeded by tips of anterior anal soft-rays. Profile of anal nearly vertical. Caudal trun- cate. Origin of pectoral below 2nd to 4th dorsal spine; margin of pectoral fin rounded, 8th to 11th rays longest; tip of pectoral extending to anus. Origin of pelvic below 3rd and 4th dorsal spines; tip of pelvic reaching to three-fourths to four-fifths of the length of the pelvic-to-anal distance. Origin of anal below 3rd or 4th dorsal soft-ray, insertion of anal below 9th to 11th dorsal soft-ray.
Extrinsic gas-bladder muscles (2 specimens examined) as in S. exsul, except that each muscle splits into only two instead of three tendons and these insert respectively on the parapophyses of the 9th and 10th centra.
VoL. XL] CHEN: SEBASTES 27
Specimens in alcohol light brown, with dusky to black patches on body and various fins (fig. 2a). Pectoral and caudal membranes clear. Peritoneum jet-black. Oral and gill-cavity linings black.
For morphometrics and meristics, see tables 1, 7, 8, and 9.
Erymotocy. The name varispinis, from the Latin, refers to the variation in the number of dorsal spines.
MATERIAL EXAMINED. A total of 61 specimens, 33-56 mm., all juveniles col- lected with midwater trawls between surface and 500 m. (bottom at 1400 m.).
Holotype. LACM 8821-11, 49 mm., 28° 58’ N., 113° 11.4’ W.
Paratypes. LACM 8821-8, 60 (33-56) collected with the holotype.
Sebastes peduncularis Chen, new species.
(Figure 2e.)
D1acnosis. Juveniles of S. peduncularis can be differentiated from those of other eastern North Pacific species of Sebastes by the following combination of characters: (1) the cranial spine pattern (with sharp nasal, preocular, postocular, tympanic, and parietal spines); (2) the meristics (D. XIII, 12-13; A. III, 6; P, 18; rakers on first gill arch 29-30); (3) presence of frontal ridges; (4) subequal jaws; (5) small eyes (orbit length 8.0-10.0 in S. L. in 43-47 mm. specimens); (6) deep caudal peduncle (peduncle depth 9.0—-10.0 in S. L. in 43-47 mm. specimens); (7) narrow body (head width 7.1—7.7 in S. L. in 43-47 mm. specimens); and (8) a color pattern as illustrated (fig. 2e).
Drscriprion, DO. XI 12-13. A. DE 6; Pi 18 (7-18). 14; rakers on first gill arch 29-30 (8-9 + 21); lateral-line pores 39-43; vertebrae (in 2 specimens) 26 (11 + 15).
Base of skull straight. Interorbital slightly concave, divided by frontal ridges into three concave grooves. Nasal, preocular, postocular, tympanic, and parietal spines present, sharp, and moderately strong. Parietal ridges well developed and elevated. Lower edge of gill cover spineless. Upper four pre- opercular spines directed backward, lowermost preopercular spine directed downward and spaced wider. Lachrymal projections two, sharp but not spine- like. Supracleithral, cleithral, and the two opercular spines well developed.
Symphyseal knob inconspicuous; jaws subequal. Maxillary reaching to beyond vertical from posterior margin of pupil.
Scales ctenoid, with 2—3 ctenii on the scales of both specimens.
Dorsal spines increasing in length to 4th, then decreasing gradually to 11th, 12th spine nearly as long as 11th, 13th spine longer than 12th; soft dorsal high, with the short posterior rays when depressed reaching to near midpoint of caudal peduncle. Second anal spine stronger and longer than third, its tip reaching to about three-fourths the length of first anal soft-ray when depressed; tips of posterior anal soft-rays, when depressed, exceed tips of anterior soft-rays. Profile of anal slanting somewhat posteriorly. Origin of pectoral below 3rd dorsal spine;
128 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
TaBLe 5. Morphometrics (standard length/body part measurement) and meristics (“*” signifies left count and right count) of Sebastes peduncularis.
Paratype Holotype Character LACM 8818-11 LACM 8818-6 Standard length 48.0 44.0 Morphometrics: 2nd anal spine 6.1 6.3 Gill-raker at angle 25 24 3rd anal spine 74 7.1 2nd dorsal soft-ray 6.2 Byes) 1st anal spine 13.0 12.6 Orbit length 10.0 9.8 Total length 0.84 0.79 4th dorsal spine 6.9 6.5 1st anal soft-ray 5.8 5.9 Anal-fin base 6.7 6.3 Pelvic-fin length 4.8 4.6 Predorsal length Sail 2.9 Pectoral-fin length 4.2 3.9 Lower peduncle length 5.1 4.9 Soft-dorsal base 4.7 5.74 Dorsal-fin incision 16 13 Caudal-peduncle depth 10.0 9.0 Upper peduncle length 7.6 6.1 Spinous-dorsal base 3.0 2.9 Snout length 12.0 11.6 Prepelvic length 2.6 2.6 Head length 2.8 2.7 Upper-jaw length 6.9 6.1 Preanal length 1.48 1.49 Body depth 3.0 3.0 Head width Tol Yagil Lachrymal width 60 55 Interorbital 15.5 14.2 Meristics: Lateral-line pores * 43-40 39-— Dorsal soft-rays 13 12 Pectoral rays * 18-18 18-17 Anal soft-rays 6 6 Gill-rakers * 30-30 29-29
margin of pectoral fin rounded, 10th ray longest; tip of pectoral extending to near anus. Origin of pelvic below 4th dorsal spine; tip of pelvic reaching to two- thirds the length of the pelvic-to-anal distance. Origin of anal below 3rd or 4th dorsal soft-ray, end of anal below 10th or 11th dorsal soft-ray.
Vor. XL] CHEN: SEBASTES 129
Specimens in alcohol light brown with faint marks in pattern as illustrated (fig. 2e). Peritoneum dusky. Oral and gill-cavity linings slightly dusky.
For morphometrics and meristics, see tables 5, 7, 8, and 9.
Discussion. Considering the morphological changes involved in meta- morphosis from the surface-living prejuvenile stage to the juvenile stage in S. cortezi, it is very difficult to establish the validity of S. peduncularis without a series of specimens of different sizes. Meristics do not differentiate this form from S. cortezi, S. varispinis, or S. sinensis. The two specimens of S. peduncularis agree with prejuveniles of S. cortezi in having short dorsal and anal spines and pectoral fins, small eyes and head, deep caudal peduncle, and broad inter- orbital (fig. 3); and agree with transformed juveniles of S. cortezi in having frontal ridges and sharp cranial and fin spines. These two specimens are intermediate in size between specimens of S. cortezi of the the two contrasting stages. Since the two specimens were captured in a midwater trawl and as they show only a trace of the dark vertical bands characteristic of surface-living juveniles, they probably were in the process of transforming to the Juve- nile stage. However, although the two specimens have a mixed combination of the prejuvenile and juvenile characteristics of S. cortezi, none of the characteristics are intermediate between the two stages. That the two specimens of S. peduncularis are specifically distinct from S. cortezi is suggested by their characteristic color pattern (fig. 2e) not intermediate between the two stages of S. cortezi, by their narrow head (fig. 3), and by the dorsal soft-ray count of 13 and the gill-raker count of 30 in the paratype, two rather unusual counts for SE NCORLEZT.
The distinction between S. peduncularis and S. varispinis is much more apparent, as the specimens of the two species available are probably in a similar stage of development. Compared with juveniles of S. varispinis, the two specimens of S. peduncularis have shorter fin spines, gill-rakers, and pectoral fins; deeper caudal peduncle and body; and much lighter pigmentation in the gill cavity and oral linings. Furthermore, they lack the protruding lower jaw of S. varis pinis.
From S. sinensis, S. peduncularis can be distinguished by having ctenoid rather than cycloid scales.
Erymotocy. The name peduncularis, from the Latin, refers to the deep caudal peduncle of this species.
MATERIAL EXAMINED. Only two specimens are known.
Holotype. LACM 8818-6, 43 mm., juvenile, collected with midwater trawl from 28° 55’ N., 112° 50.5’ W., midway between southern tip of Tiburon Island and Angel de la Guarda Island, the water depths were 440-450 m. and the trawl was operated between bottom and surface (Lavenberg & Fitch, 1966).
Paratype. LACM 8818-11, 47 mm., collected together with the holotype.
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Sebastes macdonaldi (Eigenmann & Beeson). (Figure 1a.)
Sebastodes proriger (not of Jordan & Gilbert): EicenmMANN & EIGENMANN, 1890, p. 15 (description).
Acutomentum macdonaldi EIGENMANN & BEESON, 1893, p. 669 (type locality: San Diego). JorpDAN, EverMANN, & CLARK, 1930, p. 366 (name only).
Sebastodes macdonaldi: CRAMER, 1895, p. 594 (species analysis). JORDAN & EveRMANN, 1898, p. 1786 (species analysis, description). PuHitiips, 1957, p. 90 (key, description, range, depth, size).
Sebastes macdonaldi: CHEN, 1971, p. 75 (range). Moser, 1972, pp. 941-958 (development, distribution).
Dracnosis. The presence of a spine on the lateral surface of lachrymal (in specimens larger than 10 cm.) and three or four spines on the lower edge of lachrymal and the high lateral-line pore count (> 50) differentiate this species from all other species of Sebastes.
DeEscripTIon. D. XIII, 13 (12-14); A. III, 7; P; 19 (18-20); C. 14; rakers on first gill-arch 36-42 (10-13 + 25-29); lateral-line pores 52-58; pyloric caeca (in 3 specimens) 8-11; vertebrae (in 12 specimens) 26 (12 + 14).
Base of skull straight. Interorbital flat in juveniles, slightly convex in adults, without frontal ridges. Nasal, preocular, postocular, tympanic, parietal, and frequently supraocular (nearly 50%) and nuchal (nearly 30%) spines present, sharp, and moderately strong but recumbent. Parietal ridges well developed, low. Lower edge of gill-cover spineless. Preocular spines equally spaced, the two upper ones directed backward, the three lower ones downcurved. Lachrymal projections three or four, spinelike; lateral surface of lachrymal with a retrorse spine above the second lachrymal projection near edge of orbit in specimens longer than 10 cm. Upper edge of second suborbital with a small spine in specimens less than 5 cm. Supracleithral, cleithral, and the two opercular spines well developed.
Symphyseal knob conspicuous and directed forward in juveniles, incon- spicuous in adults; lower jaw definitely projecting, entering dorsal profile of head. Maxillary ending behind vertical from posterior edge of pupil but before margin of orbit.
Scales ctenoid, small. Head fully scaled, with the exception of the lips. Fin rays scaled; membranes scaled basally, most extensively on anal and caudal.
Dorsal spines increasing in length to 4th, then decreasing gradually to 12th, 13th spine longer than 12th; soft dorsal high, with the short posterior rays when depressed reaching to only about one-third the distance between dorsal insertion and caudal base. Second anal spine shorter than 3rd, tip of 3rd spine reaching to about one-fifth (large specimens) to one-half (small specimens) the length of first anal soft-ray when depressed; tips of posterior anal soft-rays reaching to about one-third the distance between anal insertion and caudal base, exceed-
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CHEN:
Vou. XL]
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132 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
ing tips of anterior anal soft-rays when fin depressed. Profile of anal slightly slanting posteriorly. Caudal slightly forked. Origin of pectoral below 3rd or 4th dorsal spine; pectoral pointed; 10th or 11th ray longest; pectoral extending to beyond tip of pelvic, with tip reaching to anus in small specimens but not in large specimens; origin of pelvic below 3rd to 4th dorsal spine; tip of pelvic extending to one-half (large specimens) to three-fourths (small specimens) of pelvic-to- anal distance; origin of anal below 2nd to 3rd dorsal soft-ray, insertion of anal below 10th to 11th dorsal soft-ray.
Extrinsic gas-bladder muscles (2 specimens examined) as in S. exsul, except that each muscle splits into four tendons which insert respectively on parapoph- yses 6, 7, 8, and 9.
Color in life chocolate brown on back, grading to dull red beginning from below the conspicuous light lateral line. Cheek with three oblique dark bands, one behind eye, one below eye, and one on posterior part of maxillary. Dorsal and caudal fin membranes wholly dark brown; pectoral membrane black, mixed with trace of red; pelvic and anal membranes red, with trace of black. Peritoneum black. Oral and gill-cavity linings slightly dusky. Juveniles smaller than 10 cm. have conspicuous dark vertical bands on body in pattern similar to that of the other species illustrated in figure 2.
For morphometrics and meristics, see tables 6, 7, 8, and 9.
RANGE. S. macdonaldi occurs from 36° 18’ N., 122° 04’ W. to 23° 24.0’ N., 111° 11.5’ W. along the outer Pacific coast (Chen, 1971) and from 26° 59.1’ N., 111° 48.9’ W. to 28° 58’ N., 113° 11.4’ W. in the Gulf of California (Moser, 1972). It is the only species of Sebastes known to occur both inside and out- side the Gulf of California. Chen (1971) suggested that the distribution of the species may be continuous around the tip of Baja California. A comparison made between materials from the two sides of Baja California (tables 6, 7, 8, and 9), however, discloses significant differences in a number of characters. The Gulf sample has longer lower jaw and anal spines (p << 0.005, analysis of co- variance) and a higher frequency of 20 pectoral rays and 12 dorsal soft-rays (p < 0.01, Chi-square test). These suggest isolation of the two populations.
DeEpTH AND size. Phillips (1957) gave the maximum depth for the species as 65 fms. (130 m.) and (1968) the maximum size for the species as 26 in. (~ 53 cm. S. L.). I herein report a new depth record of 350 m. (SIO 65-64).
MATERIAL EXAMINED. A total of 84 specimens, 33-532 mm.
Gulf of California. LACM 8821-9, 6 (33-46); LACM 8837-4, 1 (227); SIO 73-1, 3 (135-176); SIO 73-2, 3 (174-180).
Outer coast. SIO 54-153, 2 (310-318); SIO 65-126, 4 (478-532); SIO 65-194, 8 (272-322); SIO 65-217, 9 (258-316); SIO 65-227, 11 (60-72); SIO 65-230, 2 (385—405): SIO 68-5, 1 (307); SDSC 72-52, 1 (85); SDS@ 72-305 33 (145-238).
133
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136 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
RELATIONSHIPS, ZOOGEOGRAPHY, AND SPECIATION
Although at least seven species of Sebastes have been recognized there, with but a few exceptions their known occurrence in the Gulf of California is restricted to the area between 27° N. and 30° N., centering around the vicinity of Canal de las Ballenas west of Isla Angel de la Guarda and Isla Tiburén (fig. 4). Within this area, however, some of the species, such as S. macdonaldi, occur in great abundance, as is found in the catches of the 1972 FAO Hake Resource Survey of the Alejandre Humboldt (AH 72-03, David Kramer, personal communication). Walker (1960) listed a number of temperate species restricted to, or most common in, this area.
This pattern of distribution seems to be related to the distribution of water temperature and dissolved oxygen in the Gulf of California. At the tip of Baja California there is an oceanic front, at times with a sharp temperature gradient of from 20° C. in the northwest to 27° C. in the southeast across a distance of only one degree of latitude (Cromwell & Reid, 1956; Griffiths, 1965; and Reid e¢ al., 1958). As the warmest water in which pelagic Sebastes larvae have been found is 18.08° C. (CCOFT 5604, 127G. 40, Scripps Inst., Oceanogr., 1963; Oceanic observations of the Pacific, 1956), this oceanic front forms a surface barrier probably not transgressible by Sebastes. The surface temperature southeast of the front fluctuates seasonally, with a minimum consistently above 18° C. This condition usually extends northward into the Gulf, with the excep- tion of the vicinity of Canal de las Ballenas where the winter surface tempera- ture (14° C.) is significantly lower, probably caused by the strong local tidal mixing (Roden, 1964). This appears to be the only area in the Gulf where conditions favor the survival of Sebastes during their pelagic juvenile stage.
Although most parts of the Gulf have surface water too warm for Sebastes, adults of Sebastes capable of submerging should find suitable temperatures at depth from Ballenas Channel to the south. South of a submarine ridge south of the Ballenas Channel, however, an oxygen-minimum layer extends in from the North Pacific between 200 m. and 1000 m., with an oxygen concentration of from 0.5 ml./l. to as low as <0.1 ml./l. (Roden, 1964), lower than the lethal limit for most of the fishes as compiled by Doudoroff and Shumway (1970). This makes deep-water transgression from the outer coast into the upper Gulf also a difficult task for Sebastes. The only area in the Gulf with habitats favorable to Sebastes is thus the vicinity of Ballenas Channel where surface water is cold enough in the winter to allow a successful pelagic life for the larvae and where at depth (150-600 m.) the temperature is probably low enough (12-16° C.) throughout the year for the adults, and the oxygen level (> 1.0 ml./l.) is adequate.
The high surface temperature and the oxygen-minimum layer in deep water in the south not only limit the distribution of Sebastes in the Gulf but probably also create a dispersal barrier contributing to speciation in Sebastes. Recent
VoL. XL] CHEN: SEBASTES 137
Ficure 4. Distribution of Sebastes in the Gulf of California. Open circles: surface col- lections, included are a number of collections from CCOFI (courtesy of H. G. Moser) and UCLA not listed in this report. Solid dots: mesopelagic or benthic collections.
geological studies (Larson, 1972; and Elders ef al., 1972) suggest that the Gulf of California originated about four million years ago as a result of northwestward rifting of Baja California and southern and central California along the San Andreas Fault. The first successful invasion of the Gulf by Sebastes was apparently a much more recent event, after the development of an area in the Gulf such as the present Ballenas Channel, with conditions favorable to Sebastes and during a period of oceanic cooling allowing surface transgression (rather than bottom, assuming the oxygen-minimum layer has existed since the origin of the Gulf). Repeated Pleistocene glaciation is an apparent mechanism providing opportunities for such invasion from the outer coast into the Gulf and the reciprocal events, contributing to the diversity of Sebastes both inside
138 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
and outside the Gulf. An analysis of the relationships among the species will shed some light on the history of such events.
The seven species in the Gulf seem to represent three sets of species. S. mac- donaldi is an independent lineage and is represented also by population(s) in the outer Pacific waters. The low degree of differentiation of the population of this species in the Gulf from its parental population(s) on the outer coast sug- gests that the present disjunction in distribution originated rather recently. S. macdonaldi is the species of Sebastes with the southernmost known limit (23-2410! N-) (Chen, 1971):
The other six species are all Gulf of California endemics. Of the six, S. exsul and S. spinorbis form a pair, with vermiculations and/or dusky marks on the back and with dusky scale margins characteristic of the wmbrosus-lentiginosus and the chlorostictus-rosenblatti-eos complexes of the subgenus Sebastomus recently reviewed by Chen (1971). The close relationship between S. exsul and S. umbrosus is also supported by the identical hemoglobin electropherograms demonstrated by Sharp (1973). At the present, off southern California, Sebastomus is represented by four sets of species (constellatus, rosaceus-helvo- maculatus-simulator, ensifer-notius, and umbrosus-lentiginosus-chlorostictus- rosenblatti-eos, three of which contain shallow water species (constellatus, rosaceus, and umbrosus) which are more likely to accomplish surface transgres- sion. The present number of species in these species groups suggests that group divergence is not a very recent event. The exsul-spinorbis pair does not represent a separate species group but rather belongs to one of the four sets, suggesting that this lineage was distinct prior to the invasion of the Gulf. Sebastes exsul and S. spinorbis seem more similar to each other than to any other species outside the Gulf, suggesting that the two had a common immediate ancestor. This, if true, would suggest that this lineage represents a single invasion of the Gulf, and thus, the divergence of the two species took place in the Gulf after the invasion. It is interesting to note that the S. capensis complex in the southern hemisphere also shares with the wmbrosus-eos complex the characteristic vermicu- lations or dusky marks on the back and the dusky scale margins. Although it is possible that the exsul-spinorbis and the capensis complexes have the same origin, and that successful invasion of the Gulf and crossing of the tropics by Sebastomus were accomplished by the same ancestor species using the same climatic event, multiple crossings cannot be ruled out.
The four remaining species are related to the eleven species that I regard as comprising the subgenus Allosebastes Hubbs, 1951, namely S. diploproa, S. saxicola, S. semicinctus, S. dallii, S. zacentrus, S. wilsoni, S. emphaeus, S. vari- egatus, S. rufinanus, and S. proriger of the outer coast and S. scythropus of the western North Pacific. Shared characters are nasal, preocular, postocular, tympanic, parietal, and occasionally nuchal spines present; a characteristic banded color pattern (fig. 2), at least during the juvenile stage of their life
VoL. XL] CHEN: SEBASTES 139
(except perhaps S. scythropus), and a pair of extrinsic gas-bladder muscles which originates near the opisthotic, passes medial to the cleithrum to which it has a membranous connection, then each member of the pair splits typically into two or three tendons which pass between the second and third ribs, run straight along the inner side of succeeding ribs, and insert respectively on the parapophyses of the 8th to 10th centra, without any direct connection between the tendons and the gas-bladder wall.
Of these four species, S. cortezi and S. sinensis, which together with S. diploproa of the outer coast form a species group, all have rather similar body configuration, large eyes, uniformly red body with some dusky on back, and epipelagic juveniles charcterized by blunt cranial spines. In this group, S. cortezi and S. diploproa may be called a species pair; both have anteriorly-directed lachrymal projections (not found in any other species of Allosebastes), and both have extra tendons in the extrinsic gas-bladder muscles in addition to the typical Allosebastes pattern. S. sinensis probably is the species bridging the gap between the cortezi-diploproa pair and the closest species of the outer coast, probably S. saxicola.
The affinities of the two remaining species, S. varispinis and S. peduncularis, cannot be ascertained but they both are definitely close to the s¢mensis-diplo proa- cortezi complex. It is clear that the history of the four species of Adlosebastes in the Gulf is more complex than that of the two referred to Sebastomus, and suggests a possible multiple invasion of Adlosebastes species from the outer coast into the Gulf, either by the same ancestral stock at different periods, or by different parental species either simultaneously or not; after successful inva- sion(s), it appears that radiation took place in the Gulf and there probably was (were) successful reciprocal invasion(s) from the Gulf to the outer coast, as suggested by the cortezi-diploproa pair.
The relationships and zoogeographic history outlined above are very speculative. If further studies support the hypothesis that species differentiation took place within a limited area in the Gulf, it would suggest that speciation in Sebastes does not necessarily require the presence of a geographic barrier. This helps to explain the origin of the high diversity of Sebastes in the North Pacific, with approximately 100 species, most of which are extensively sympatric (Chen, 1971).
LITERATURE CITED
BarRSUKOV, V. V. 1970. Species composition of genus Sebastes in the North Pacific and description of a new species. Doklady Biological Sciences, Proceedings of the Academy of Sciences of the USSR, Biological Sciences Sections, vol. 195, pp. 760-763, 1 fig. BEEBE, W., AND J. TEE-VAN 1938. Eastern Pacific Expeditions of the New York Zoological Society, XV. Seven
140 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
new marine fishes from Lower California. Zoologica, vol. 23, part 3, no. 15, pp. 299-312, 5 figs., 3 pls. CaLpwe Ll, M. C.
1962. Development and distribution of larval and juvenile fishes of the family Mullidae of the western North Atlantic. Fishery Bulletin, United States Fish and Wildlife Service, vol. 62, no. 213, pp. 403-457, 43 figs.
CHEN, L.
1971. Systematics, variation, distribution, and biology of rockfishes of the subgenus Sebastomus (Pisces, Scorpaenidae, Sebastes). Bulletin of the Scripps Institu- tion of Oceanography, vol. 18, 107 pp., 16 figs., 6 pls.
CRAMER, F. 1895. On the cranial characters of the genus Sebastodes (rock-fish). Proceedings of the California Academy of Sciences, ser. 2, vol. 5, no. 1, pp. 573-610, pls. 57-70. CROMWELL, T., AND J. L. Rem, Jr. 1956. A study of oceanic fronts. Tellus, vol. 8, no. 1, pp. 94-101, 6 figs. Doupororr, P., AND D. L. SHumMWway
1970. Dissolved oxygen requirements of freshwater fishes. FAO Fisheries Technical
Paper, no. 86, 291 pp., 2 figs. EIGENMANN, C. H., ANd C. H. BEESON
1893. Preliminary note on the relationship of the species usually united under the generic name Sebastodes. American Naturalist, vol. 27, pp. 668-671.
1894. A revision of the fishes of the subfamily Sebastinae of the Pacific coast of America. Proceedings of the United States National Museum, vol. 17, no. 1009, pp. 375-407.
EIGENMANN, C. H., AnD R. S. EIGENMANN 1890. Additions to the fauna of San Diego. Proceedings of the California Academy of Sciences, ser. 2, vol. 3, pp. 1-24. Expers, W. A., R. W. Rex, T. Maipav, P. T. Ropinson, AND S. BIEHLER 1972. Crustal spreading in Southern California. Science, vol. 178, pp. 15-24, 7 figs. GILBerT, C. H.
1890. A preliminary report on the fishes collected by the steamer Albatross on the Pacific coast of North America during the year 1889, with descriptions of twelve new genera and ninety-two new species. Proceedings of the United States National Museum, vol. 13, no. 797, pp. 49-126.
GrifFituHs, R. C.
1965. A study of ocean fronts off Cape San Lucas, Lower California. United States Fish and Wildlife Service, Special Scientific Report—Fisheries, no. 499, 54 pp., 37 figs.
Hupss, C. L.
1951. Allosebastes, new subgenus for Sebastodes sinensis, scorpaenid fish of the Gulf of California. Proceedings of the Biological Society of Washington, vol. 64, pp. 129-130.
Jorpan, D. S., anp B. W. EveRMANN
1896. A check list of the fishes and fish-like vertebrates of North and Middle America. United States Commission of Fish and Fisheries, Report of the Commissioner, 1895, part 21, appendix 5, pp. 207-584.
1898. The Fishes of North and Middle America. Bulletin of the United States National Museum, vol. 47, part 2, pp. 1241-2183.
Jorpan, D. S., B. W. EvERMANN, AND H. W. CLARK 1930. Check list of the fishes and fish-like vertebrates of North and Middle America
VoL. XL] CHEN: SEBASTES 141
north of the northern boundary of Venezuela and Colombia. Report of the United States Commissioner of Fisheries, 1928, part 2, 670 pp. Larson, R. L.
1972. Bathymetry, magnetic anomalies, and plate tectonic history of the mouth of the Gulf of California. Geological Society of America Bulletin, vol. 83, no. 11, pp. 3345-3360, 15 figs.
LAVENBERG, R. J., AND J. E. FitcH
1966. Annotated list of fishes collected by midwater trawl in the Gulf of California, March-April 1964. California Fish and Game, vol. 52, no. 2, pp. 92-110, 6 figs.
Lea, R. N., ano J. E. Fitcu
1972. Sebastes rufinanus, a new scorpaenid fish from California waters. Copeia,
1972, no. 3, pp. 423-427, 2 figs. M’CLELLAND, J.
1843. Description of a collection of fishes made at Chusan and Ningpo in China, by Dr. G. R. Playfair, Surgeon of the Phlegethon, War Steamer, during the late military operations in that country. Calcutta Journal of Natural History, vol. 4, pp. 390-413.
Moser, H. G.
1972. Development and geographic distribution of the rockfish, Sebastes macdonaldi (Eigenmann and Beeson, 1893) family Scorpaenidae, off southern California and Baja California. Fishery Bulletin, United States Department of Commerce, vol. 70, no. 3, pp. 941-958, 5 figs.
1974. Development and distribution of larvae and juveniles of Sebastolobus (Pisces, family Scorpaenidae). Fishery Bulletin, United States Department of Com- merce, vol. 72, no. 4, pp. 865-884.
Puituirs, J. B.
1957. A review of the rockfishes of California (family Scorpaenidae). Fish Bulletin, California Department of Fish and Game, no. 104, 158 pp., 66 figs.
1968. Review of rockfish program. California Department of Fish and Game, MRO reference no. 68-1, 27 pp., 8 figs.
OwuASt eC:
1971. Sebastes variegatus sp. n. from the northeastern Pacific Ocean (Pisces, Scorpaenidae). Fishery Bulletin, United States Department of Commerce, vol. 69, no. 2, pp. 387-398, 5 figs.
Rew, J. L., Jr., G. E. Ropen, anp J. G. WYLLIE
1958. Studies of the California Current System. Progress Report, California Coopera- tive Oceanic Fisheries Investigations, 1 July 1956 to 1 January 1958, pp. 27- Si, 2s) Wes
Ropen, G. I.
1964. Oceanographic aspects of Gulf of California, pp. 30-58, figs. 1-14. Jn Marine Geology of the Gulf of California—A Symposium. The American Association of Petroleum Geologists, Memoir no. 3.
SHarp, G. D.
1973. An electrophoretic study of hemoglobins of some scombroid fishes and related forms. Comparative Biochemistry and Physiology, vol. 44, no. 2B, pp. 381- 388, 5 figs.
Wacker, B.
1960. The distribution and affiniities of the marine fish fauna of the Gulf of California.
Systematic Zoology, vol. 9, nos. 3 & 4, pp. 123-133, 1 fig.
PROCEEDINGS
OF THE CALIFORNIA ACADEMY OF-SCIENCES _ FOURTH SERIES
Vol. XL, No. 7, pp. 145-208; 176 figs. August 8, 1975
THE LARGER MOTHS OF THE GALAPAGOS ISLANDS (GEOMETROIDEA: SPHINGOIDEA & NOCTUOIDEA)
By
Alan H. Hayes
British Museum (Natural History)
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 7, pp. 145-208; 176 figs. August 8, 1975
THE LARGER MOTHS OF THE GALAPAGOS ISLANDS (GEOMETROIDEA: SPHINGOIDEA & NOCTUOIDEA)'
By Alan H. Hayes British Museum (Natural History)
ABSTRACT: Ejighty-eight of the 90 species of Epiplemidae, Sphingidae, Arcti- idae, and Noctuidae now known to occur on the Galapagos Islands are illustrated. Twelve of these are described as new. Four new subspecies are described and six species-group names are newly placed in synonymy. The world distribution of the species is summarized and new biological data is given for some of the species.
INTRODUCTION
The Galapagos were discovered by Fray Tomas de Berlanga, Bishop of Panama, on a voyage to Peru in 1535. Thereafter, for nearly three centuries, the islands received only occasional or temporary visitors, becoming suc- cessively the haunt of buccaneers, whalers, and sealers. In 1832, with the dis- solution of the Spanish-American Empire, the archipelago was annexed by Ecuador and the first settlement was established, on Floreana Island. Today, the southern slopes of the islands of San Cristobal, Santa Cruz, and Isabela are inhabited and a small colony continues on Floreana. Altogether over five thou- sand people live in the islands, mainly engaged in fishing or subsistence farming, or connected in one way or another with tourism.
The Galapagos Islands have a unique place in the history of science because of the visit of Charles Darwin in 1835 and the subsequent role his observa- tions there played in the formulation of his ideas on organic evolution, which culminated in 1859 with the publication of the Ovigin of Species. The extraor- dinary indigenous wildlife of the islands suffered a rapid decline in the years
1 Contribution No. 171 of the Charles Darwin Foundation.
[145]
146 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
following their settlement, as a result both of exploitation and of introduction of plants and animals brought by man. These threats continue, although they are being alleviated by protective legislation and programs.
In 1959, all uninhabited areas of the islands were declared territory of a National Park by the Government of Ecuador and the current protection laws were brought into force. At the same time, a newly created international organization, the Charles Darwin Foundation for the Galapagos Isles, was entrusted with the task of coordinating scientific research in the islands and advising the Government on conservation policy and the development of the National Park.
The Galapagos Islands are the tops of huge shield volcanos composed mainly of basalt. Dating studies have shown that the majority of lavas on the surface today came from eruptions during Pleistocene and Recent times. Vol- canic activity continues and there have been eruptions on five of the islands in the past hundred years.
Approximately two-thirds of the land area are lava-strewn wastelands with a sparse xerophytic vegetation dominated by cacti and thorn-scrub. Only near the summits and on humid southern slopes of the higher islands does dense vegetation occur, composed typically of Scalesia woodland, a lush al- though somewhat seasonal undergrowth, and a limited number of epiphytic species. Rainfall is irregular, but in most years there is the possibility of heavy showers during the hot season, January to April, when daytime temperatures reach 35° C. on the coast. The cool, or garuéa, season, lasting for the remaining months of the year, is characterized by generally more cloudy weather and steady southeasterly winds, when corresponding temperatures drop to 22° C. or even lower.
The names of the islands have been a source of confusion. Ecuadorian names (used in this paper) and the equivalent English names are shown on the map. In fact, some of the islands have several names (see Bowman, 1966, p. Xvii).
Recent collecting has yielded many additions to the species recorded in Insects of the Galdpagos Islands by Linsley and Usinger. Dr. F. H. Rindge of the American Museum of Natural History has recently published on the Geometridae and a check list is included here. The Pyralidae and Micro- lepidoptera of the islands await further study.
An attempt has been made to include all known museum material. Fore- wing measurement is taken from the center of the mesothorax to the apex of the forewing. Species identifications are based as far as possible on com- parisons between Galapagos specimens and the type material. Where the type of a particular species has not been traced the identifications have been made by comparing Galapagos examples with material from the type-locality as far as this is known. The heading ‘Distribution’ refers to distribution within the
VoL. XL] HAYES: GALAPAGOS MOTHS 147
MARCHENA (BINDLOE)
r) GENOVESA (TOWER)
SANTIAGO (JAMES) > e
@ RABIDA (JERVIS) FERNANDINA SAN CRISTOBAL
SANTA CRUZ en CHATHAM (NARBOROUGH) a (INDEFATIGABLE)
& SANTA FE (BARRINGTON)
2 SHB, ESPANOLA FLOREANA (HOOD)
(CHARLES)
ISABELA (ALBEMARLE)
9° 90°
Map 1. Main group of islands forming the Galapagos Archipelago. Darwin (Culpepper) and Wolf (Wenman) lie to the northwest.
islands based on material studied by the author. A proportion of the species under consideration are widespread in distribution and reference to other faunistic works yields useful data; as these works may not be available to readers I have abstracted much of this data. The bibliography lists all pub- lications referred to in the text and in addition cites other scientific and general works relevant to the study of the Galapagos fauna.
Of the 90 species dealt with in this paper 28 are endemic. However, Galapagos populations of the 62 remaining species often differ from those of the mainland and 16 are sufficiently distinct to warrant subspecific status.
The following abbreviations have been used for depositories: AMNH— American Museum of Natural History, New York; BMNH—British Museum (Natural History), London; CAS—California Academy of Sciences, San Fran- cisco; CMP—Carnegie Museum, Pittsburgh; CU-—Cornell University, Ithaca; IRSNB—Institut Royal des Sciences Naturelles de Belgique, Brussels; IZ—
148 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Institute of Zoology, University of Uppsala; LACM—Los Angeles County Museum; LS—Linnean Society, London; MCZ—Museum of Comparative Zoology, Boston; MNHN—Museum National d’Histoire Naturelle, Paris; NM—Naturhistorisches Museum, Vienna; NR—Naturhistoriska Riksmuseum, Stockholm; RSM—Royal Scottish Museum, Edinburgh; UM—wUniversity Museum, Oxford; USC—University of Southern California, Los Angeles; USNM—National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C.; ZSBS—Zoologische Sammlung des Bayerischen Staates, Munich. Unless otherwise stated all figured specimens are in the BMNH.
ACKNOWLEDGMENTS
Thanks to the tremendous efforts made in rearing and collecting specimens for the British Museum (Natural History) by Dr. R. Perry (formerly Director of the Charles Darwin Research Station) and Dr. Tj. de Vries (of the Zoologisch Museum, Amsterdam on grants from the Netherlands Foundation for the advancement of Tropical Research and the World Wildlife Fund) excellent series have been built up to complement the historic material studied by Walker, that collected by Rollo H. Beck for Lord Rothschild, and the St. George Expedi- tion material collected by C. L. Collenette and Miss C. E. Longfield. Through the kindness of L. A. Berger I have examined the Institut Royal des Sciences Naturelles de Belgique material collected by J. and N. Leleup. I am most grate- ful to Dr. P. H. Arnaud, Jr. who made available to me the 11,387 specimens in the collections of the California Academy of Sciences amassed by D. Q. Cavagnaro, F. (P) Leon, E. G. Linsley, R. O. Schuster, D. W. Snow, I. L. Wiggins, F. X. Williams, and M. Willows (Jr.). E. C. Pelham-Clinton of the Royal Scottish Museum has been of great assistance in the identification of the Edinburgh University Expedition (1968) material. The material deposited in the National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C.; the American Museum of Natural History, New York; the Museum of Comparative Zoology, Boston (collected by Dr. R. Silberglied) ; and the Carnegie Museum, Pittsburgh, has also been examined. J. P. Donahue, Los Angeles County Museum of Natural History, has mailed me material from that institution and has also been kind enough to examine and photograph relevant Allan Hancock Foundation material studied by Prof. A. G. Richards, Jr. at the University of Southern California, Los Angeles. Dr. P. I. Persson of the Naturhistoriska Riksmuseum, Stockholm, and Dr. W. Dierl of the Zoologische Sammlung des Bayerischen Staates, Munich, loaned me their Gala- pagos material. Special thanks are due to Dr. E. L. Todd, United States Depart- ment of Agriculture, and Prof. J. G. Franclemont, Cornell University, for their great help and advice concerning identification of Noctuidae. Dr. Perry and Dr. Tj. de Vries have placed their comprehensive notes concerning early stages and foodplants with me. I wish to acknowledge the extensive help given by my
VoL. XL] HAYES: GALAPAGOS MOTHS 149
colleagues at the British Museum (Natural History). I am indebted to P. V. York of this museum for taking the photographs.
CuHECK-LIsT OF SPECIES EPIPLEMIDAE
Epiplema becki Hayes, new species.
GEOMETRIDAE (recently covered by Dr. F. H. Rindge (1973) and not included in the present work)
Cyclophora impudens (Warren) Disclisioprocta stellata (Guenée) Hydria affirmata (Guenée) Eupithecia leleupi Herbulot
E. perryvriesi Herbulot Perizoma (?) perryi Rindge Semiothisa cruciata cruciata Herbulot S. cruciata isabelae Rindge
S. cerussata Herbulot Thyrinteina infans Herbulot
T. umbrosa Herbulot Sphacelodes vulneraria (Hiibner) Oxydia lignata (Warren).
SPHINGIDAE
Agrius cingulatus (Fabricius)
Manduca sexta leucoptera (Rothschild & Jordan), new combination M. rustica calapagensis (Holland), new combination Erinnyis alope dispersa Kernbach
E. ello encantada Kernbach
E. obscura conformis Rothschild & Jordan
Enyo lugubris delanoi (Kernbach), new combination Pachygonia drucei Rothschild & Jordan
Eumorpha fasciata tupaci (Kernbach), new combination E. labruscae yupanquii (Kernbach), new combination Xylophanes norfolki Kernbach
X. tersa (Linnaeus)
Hyles lineata florilega (Kernbach), new combination.
ARCTUODAE
Utetheisa ornatrix (Linnaeus) U. devriesi Hayes, new species U. galapagensis (Wallengren)
U. perryi Hayes, new species.
NOcTUIDAE
NOCTUINAE Agrotis consternans Hayes, new species A, ipsilon (Hufnagel)
150 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
A. subterranea williamsi (Schaus), new combination Peridroma saucia (Hubner)
Psaphara conwayi (Richards), new combination
P. interclusa Walker, revived combination
Anicla oceanica (Schaus), new combination.
HELIOTHINAE Heliothis cystiphora (Wallengren) H. virescens (Fabricius).
HADENINAE
Mythimna solita (Walker)
M. latiuscula (Herrich-Schaffer) Pseudaletia sequax Franclemont P. cooperi (Schaus).
ACRONICTINAE
Magusa erema Hayes, new species
Trachea cavagnaroi Hayes, new species Cropia infusa (Walker)
Callopistria floridensis (Guenée)
Catabena seorsa Todd
Neogalea esula longfieldae Hayes, new subspecies Spodoptera eridania (Stoll)
S. latifascia (Walker)
S. dolichos (Fabricius)
S. frugiperda (Smith)
S. roseae (Schaus)
Elaphria encantada Hayes, new species Platysenta mobilis (Walker), revived species P. sutor (Guenée)
P. ruthae (Schaus)
Agrotisia williamsi (Schaus).
ACONTIINAE
Ozarba consternans Hayes, new species Bagisara repanda (Fabricius), new combination Eublemma recta (Guenée), new combination Amyna insularum Schaus
Heliocontia margana (Fabricius)
Spragueia creton Schaus
Ponometia indubitans (Walker).
EUTELIINAE Paectes arcigera (Guenée).
SARROTHRIPINAE Characoma nilotica (Rogenhofer).
CATOCALINAE
Mocis incurvalis Schaus
M. latipes (Guenée)
Celiptera remigioides (Guenée)
Zale obsita (Guenée), revived species.
Vou. XL] HAYES: GALAPAGOS MOTHS 151
PLUSIINAE
Autoplusia egena galapagensis (Schaus) Argyrogramma verruca (Fabricius) Pseudoplusia includens (Walker).
OPHIDERINAE
Melipotis acontioides producta Hayes, new subspecies M. indomita (Walker)
M. harrisoni Schaus
Ascalapha odorata (Linnaeus)
Letis mycerina (Cramer)
Rivula asteria Druce, new combination
Glympis toddi Hayes, new species
Anomis editrix (Guenée)
A. illita Guenée
A. luridula professorum Schaus, new status
A. erosa Hiibner
Plusiodonta clavifera (Walker)
Gonodonta biarmata evadens Walker, new status G. fulvangula Geyer
Metallata absumens contiguata Hayes, new subspecies Bendis formularis Geyer
Anticarsia gemmatalis Hubner
A. prona (Moschler)
Psorya hadesia Schaus
Epidromia zetophora Guenée
E. zephyritis Schaus.
HYPENINAE
Sorygaza variata Hayes, new species
Hypena vetustalis (Guenée)
H. microfuliginea Hayes, new species
Peliala fuliginea Hayes, new species
Ophiuche lividalis (Hiibner)
O. minualis constans Hayes, new subspecies.
Family EPIPLEMIDAE
Epiplema becki Hayes, new species. (Figures 16, 17, 168-170.)
Description. Male 8.5 mm. Patagia brown. Ground color of forewing white; costal margin and spot on posterior margin brown; fringes and traces of reticulate pattern brown; undersurface brown. Outer margin of hindwing with two short pointed processes; fringes brown; postmedial line brown; under- surface white. Does not resemble any American species known. Superficially resembles the Samoan Epiplema amygdalipennis Warren. Genitalia as figured.
Female 9.5 mm. Similar to but larger than male. Reticulate brown pattern prominent. The star shaped signum on the bursa is a distinctive feature of the genitalia.
152 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
DISTRIBUTION. Endemic species.
HototypE. Male. Fernandina (Narborough), 2-5 April 1906, F. X. Williams. CAS.
PaRATYPES. ‘Isabela (Albemarle)’, Volcan Sierra Negra (= Santo Tomas), Corazon Verde, 360 m., January 1971. R. Perry & Tj. de Vries, BMNH; ‘(N. Albemarle) Isabela’, 11 April 1902, R. H. Beck ex. Rothschild Bequest, BMNH. Both specimens are females.
BroLocy. No data available.
Family SPHINGIDAE
Agrius cingulatus (Fabricius).
(Figure 1.)
Sphinx cingulata Fasrictus, 1775, Systema Entomologiae, p. 545. Type material: America (not found by Zimsen, 1964, p. 519).
Herse cingulata (Fabricius): KERNBACH, 1962, Opuscula Zoologica, Miinchen, vol. 63, p. 1. Kernbach did not separate Galapagos specimens as a subspecies. DISTRIBUTION. Widely distributed in the neotropics. Galdpagos Islands:
Baltra, April; Floreana, March, April; Isabela, March-May, August; San
Cristobal, February, March; Santa Cruz, January—June, October-December;
Wolf, February. AMNH, BMNH, CAS, CMP, IRSNB, MCZ, NR, ZSBS. BroLtocy. Found from sea-level to summits of principal volcanos; visits
flowers of Opuntia, Ipomoea, and Cacabus miersii during the day. Foodplants.
Ipomoea species occur in the archipelago, upon which the larvae feed in the
U.S.A. Larvae. Not reared by Perry and de Vries but a description appears
in Williams (1911).
Manduca sexta leucoptera (Rothschild & Jordan), new combination. (Figure 2.)
Protoparce leucoptera ROTHSCHILD & JORDAN, 1903, Novitates Zoologicae, vol. 9, Suppl., p. 79. Holotype, female (examined): Galapagos: Chatham [San Cristébal], BMNH. Protoparce sexta leucoptera Rothschild & Jordan: KERNBACH, 1962, Opuscula Zoologica,
Miinchen, vol. 63, p. 2.
The stronger yellow lateral areas on the abdomen and the less conspicuous
reniform spot separate this insect from the lighter forms of Manduca rustica calapagensis.
>
FicureE 1. Agrius cingulatus (Fabricius), female, Santa Cruz (xX 1). FicurE 2. Manduca sexta leucoptera (Rothschild & Jordan), female, Santa Cruz (x 34). Ficure 3. M. rustica calapagensis (Holland), male, Santa Cruz (x °4). Ficure 4. Eumorpha labruscae yupanquii (Kernbach), male, Santa Cruz (x-94). Ficure 5. Erinnyis alope dispersa Kernbach, female, paratype, Santa Cruz (x1). Ficure 6. Eumorpha fasciata tupaci (Kernbach), female, Santa Cruz (xX 1).
Vou. XL]
UL
154 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
DISTRIBUTION. Endemic subspecies of the widespread neotropical insect. Galdpagos Islands: Floreana, January, February; Isabela, March-May; San Cristobal, March; Santa Cruz, January-June. AMNH, BMNH, CAS, CMP, IRSNB, MCZ, ZSBS.
BroLocy. Common in years with prolonged rainy season; wings when at rest held higher above body than in M. rustica calapagensis. Foodplant. Physalis pubescens. Larva. Green, pale to white dorsally; seven oblique lateral stripes blackish followed by yellow; anal horn red.
Manduca rustica calapagensis (Holland), new combination. (Frontispiece and figure 3.) Protoparce calapagensis HOLLAND, 1889, Proc. U.S. Nat. Mus., vol. 12, p. 195. Holotype
(examined): Galapagos: Charles [Floreana] ; USNM.
Protoparce rustica calapagensis Holland: RoruHscuip & JorDAN, 1903, Novitates Zoologicae,
vol. 9, Suppl., p. 85.
Protoparce rustica calapagensis ab. nigrita ROTHSCHILD & JORDAN, 1903, Novitates Zoologicae,
vol. 9, Suppl., p. 86.
Protoparce postscripta CLARK, 1926, Proc. New Eng. Zool. Club, vol. 9, p. 70. Protoparce rustica calapagensis Holland: KrErNBACH, 1962, Opuscula Zoologica, Miinchen, vol.
635) pe 4:
This insect exhibits considerable variation ranging from dark brown through gold to white in basic coloration. Williams (1911) and Kernbach (1962) give excellent coverage concerning this and other Galapagos Sphingidae.
DISTRIBUTION. Endemic subspecies of the widespread neotropical insect. Galdpagos Islands: Baltra, April; Espanola, February; Floreana, January— April; Genovesa, February—April; Isabela, February—April, July; San Cristobal, February—April; Santa Cruz, January-July. AMNH, BMNH, CAS, CMP, IRSNB, IMeZ NR. USNM> ZSBS.
BioLocy. Widespread, usually the most abundant sphingid. Foodplants. Clerodendrum molle, Cordia lutea, Cordia leucophlyctis. Larva. Variable; green to purplish with yellow granules; oblique stripes purple edged with white; anal horn yellowish green. The darker forms appear to be prevalent at times of great abundance of these larvae, when foodplants are virtually stripped of foliage (see frontispiece).
Erinnyis alope dispersa Kernbach. (Figure 5.)
Erinnyis alope dispersa KERNBACH, 1962, Opuscula Zoologica, Miinchen, vol. 63, p. 9. Para- type female (examined): Galapagos: Santa Cruz; BMNH.
Although very close to the mainland species I have found that aedeagus dif- ferences mentioned by Kernbach separate it.
DISTRIBUTION. Endemic subspecies of the widespread neotropical insect. Galapagos Islands: Santa Cruz, March-May. BMNH, CAS, ZSBS.
VoL. XL] HAYES: GALAPAGOS MOTHS 155
Biotocy. A single adult was taken on Santa Cruz on 1 March 1967. Sub- sequently, a mature larva was found on 16 March, and the adult reared from this emerged on 5 April 1967. Larva. Buff, darker above with indistinct transverse markings; pink between Ist and 2nd segments; prominent dark spot with inner, lighter ring on 3rd; stigmata with yellowish discs; anal horn short, slightly curved, buff. Foodplants. Carica papaya, species of Jatropha and Allamanda are listed as foodplants by Hodges (1971) and Kimball (1965).
Erinnyis ello encantada Kernbach. (Figures 13 & 14.) Erinnyis ello encantada KrERNBACH, 1962, Opuscula Zoologica, Miinchen, vol. 63, p. 10.
Paratype female (examined): Galapagos: Santa Cruz; BMNH.
Sexually dimorphic. Kernbach states this subspecies is smaller and lighter in coloration than mainland representatives.
DISTRIBUTION. Endemic subspecies of the widespread neotropical insect. Galdépagos Islands: Floreana, March, May; Isabela, March—May; San Cristobal, February—March, July; Santa Cruz, January—June, August, October. AMNH, BMNH, CAS, CMP, IRSNB, MCZ, RSM, ZSBS.
BioLocy. Foodplant. Hippomane mancinella. Larva. From the Galapagos Islands Curio (1965) describes 3 types of larva of this subspecies which vary in color, pattern, and behavior.
Erinnyis obscura conformis Rothschild & Jordan. (Figure 12.)
Erinnyis obscura conformis RoTHscHiItp & JoRDAN, 1903, Novitates Zoologicae, vol. 9, Suppl., p. 369. Holotype, male (examined): Galapagos: Albemarle [Isabela]; BMNH.
The males lack the dark line on the forewing found in Erinnyis obscura obscura Fabricius.
DISTRIBUTION. Endemic subspecies of the widespread neotropical insect. Galdpagos Islands: Baltra, April; Floreana, March—May; Isabela, January— June, August; Pinzon, April; Santa Cruz, January—April, June, August—October. AMNH, BMNH, CAS, CMP, IRSNB, MCZ, NR, RSM, ZSBS.
BioLocy. Foodplant. Sarcostemma angustissima. Larva. Gray; anal horn short. Williams (1911) refers to a second type which is pale green.
Enyo lugubris delanoi (Kernbach), new combination. (Figures 7 & 8.)
Epistor lugubris delanoi KERNBACH, 1962, Opuscula Zoologica, Miinchen, vol. 63, p. 11. Paratype male (examined): Galapagos: Santa Cruz; BMNH.
A smaller insect than the nominate subspecies. DIsTRIBUTION. Endemic subspecies of the widespread neotropical insect.
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Vor. XL] HAYES: GALAPAGOS MOTHS 157
Galdépagos Islands: Isabela, no month; Santa Cruz, January-June, December. AMNH, BMNH, CAS, MCZ, ZSBS.
Brotocy. A species more of the damper inland regions of the islands. Food- plant. Cissus sicyoides from Williams (1911).
Pachygonia drucei Rothschild & Jordan. (Not figured.) Pachygonia drucei RoTHSCHILD & JORDAN, 1903, Novitates Zoologicae, vol. 9, Suppl., p. 411.
Holotype, male (examined): [Panama]: Chiriqui; BMNH.
Doubtfully included, based on the specimen mentioned below which may well have been taken on the Cocos Islands.
DISTRIBUTION. Ecuador, Panama, and Honduras. Included here on the basis of one male specimen labeled Galapagos & Cocos Islands. A. J. Drexel. BMNH.
Brotocy. No data available.
Eumorpha fasciata tupaci (Kernbach), new combination. (Figure 6.) Pholus fasciatus tupaci KERNBACH, 1962, Opuscula Zoologica, Miinchen, vol. 63, p. 12. Holotype (photograph examined): Galapagos: Santa Cruz; ZSBS. Kernbach states that the forewing stripes show a more conspicuous pink tinge in this subspecies. DISTRIBUTION. Endemic subspecies of the widespread neotropical insect. Galapagos Islands: Santa Cruz, February, April-May. BMNH, CAS, ZSBS. BrioLocy. Two fresh adults were taken on Santa Cruz in February 1967;
there were no subsequent records for this species. Onograceae are listed as foodplants in the U.S.A.
Eumorpha labruscae yupanquii (Kernbach), new combination. (Figure 4.)
Pholus labruscae yupanquii KERNBACH, 1962, Opuscula Zoologica, Miinchen, vol. 63, p. 13. Holotype (photograph examined): Galapagos: Santa Cruz; ZSBS.
As stated by Kernbach the male genitalia are more heavily chitinized than those of the nominate subspecies.
<
Ficure 7. Enyo lugubris delanoi (Kernbach), male, Santa Cruz (x 1). Ficure 8. E. lugubris delanoi (Kernbach), female, Santa Cruz (x1). Ficure 9. Xylophanes tersa (Linnaeus), male, Santiago (<1). Ficure 10. X. tersa (Linnaeus), melanic male, Santiago (<1). Ficure 11. X. norfolki Kernbach, female, Santa Cruz (x 1). Ficure 12. Erinnyis obscura conformis Rothschild & Jordan, male, Santa Cruz (x1). Ficure 13. E. ello encantada Kernbach, male, Santa Cruz (xX 1; CAS). Ficure 14. E. ello encantada Kernbach, female, Santa Cruz (xX 1).
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DisTRIBUTION. Endemic subspecies of the widespread neotropical insect. Galépagos Islands: Floreana, February, November; Santa Cruz, January—June, August. AMNH, BMNH, CAS, IRSNB, MCZ, RSM, ZSBS.
Brotocy. Adults not uncommon; inland and coastal regions of main islands. Larvae were not found by Perry and de Vries.
Xylophanes norfolki Kernbach. (Figure 11.)
Xylophanes norfolki KERNBACH, 1962, Opuscula Zoologica, Miinchen, vol. 63, p. 14. Holo-
type, male (photograph examined): Galapagos: Santa Cruz; ZSBS.
The characteristic forewing pattern of this endemic species readily separates it from Xylophanes tersa.
DisTRIBUTION. Endemic species. Galdpagos Islands: Santa Cruz, January— June, July. AMNH, BMNH, CAS, IRSNB, ZSBS.
BroLocy. Adults have so far only been taken on Santa Cruz where the species is generally scarce and mainly restricted to inland regions. Larvae were not found by Perry and de Vries.
Xylophanes tersa (Linnaeus). (Figures 9 & 10.)
Sphinx tersa LINNAEUS, 1771, Mantissa Plantarum, vol. 2, p. 538. Type material: Maryland,
Jamaica, Antigua; not traced.
One female reared by F. X. Williams and located in the California Academy of Sciences was the only known specimen. Dr. Tj. de Vries has recently taken specimens on Santiago including melanic examples (figure 10).
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: San Cristébal, February; Santiago, November. BMNH, CAS.
Brotocy. Kimball lists Spermacoce (Rubiaceae) as foodplant in Florida.
Hyles lineata florilega (Kernbach), new combination. (Figure 15.)
Celerio lineata florilega KERNBACH, 1962, Opuscula Zoologica, Miinchen, vol. 63, p. 16.
Holotype (photograph examined): Galapagos: Santa Cruz; ZSBS.
A small but strikingly marked subspecies.
DISTRIBUTION. Endemic subspecies of the almost cosmopolitan insect. Galdépagos Islands: Baltra, April; Espanola, April; Floreana, January, March— May; Isabela, April; San Cristobal, February; Santa Cruz, January—June, October; Santiago, February, March, AMNH, BMNH, CAS, CMP, IRSNB, MCZ, RSM, USNM, ZSBS.
Brotocy. Adults. Widespread, diurnal, seasonally common. Foodplants.
Vor. XL] HAYES: GALAPAGOS MOTHS 159
Portulaca oleracea, Commicarpus tuberosus. Larva. Green with variable black and yellow markings; some purple near spiracles; anal horn long, curved, yellowish to red.
Family ARCTIIDAE
Utetheisa ornatrix (Linnaeus). (Figures 18 & 19.)
Phalaena (Noctua) ornatrix LINNAEUS, 1758, Systema Naturae (10th Ed.), vol. 1, p. 511. Type material: America; IZ.
This day-flying species is, I believe, the insect mentioned by Eibl-Eibesfeldt (1960, p. 99).
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Gardner near Espanola, April; Isabela, April, June, August; San Cristobal, January, February, April-June; Santa Cruz, January—April, June, September, October, December; Santiago, March, April AMNH, BMNH, CAS, IRSNB, MCZ, RSM, USNM, ZSBS.
Brotocy. More restricted than U. galapagensis yet conspicuous and abundant at times in open, moister areas. Larvae were not collected by Perry and de Vries, but species of Crotalaria, the host-plants elsewhere, are widespread in the archipelago.
Utetheisa devriesi Hayes, new species. (Figures 24-26, 175, & 176.)
DEscriIPTION. Male 20.5 mm. Antenna bipectinate. Palpus dark brown with some gray scaling. Forewing gray with dark brown scaling at margin and on median line. Hindwing gray with brown scaling at margin. Genitalia: the uncus structure distinguishes this species.
Female 22 mm. Antenna simple. Similar to male in maculation but with broader forewing. Genitalia: the considerable spicular ornamentation at the base of the ductus bursae distinguishes the genitalia.
Larger than, but closely related to U. galapagensis. The prominent darker shade on the median line also separates this species. Lacks the yellowish buff coloration of U. perryt.
DISTRIBUTION. Endemic species.
Hototype. Male. Pinta, Highlands, southern slope at approximately 500 m.; Zanthoxylum forest with undergrowth of Tournefortia shrub; 13-15 October 1973. BMNH.
PARATYPES. Same data, BMNH (27 specimens).
OTHER MATERIAL. A melanic male with the same data as the types (BMNH) although excluded from the type-series is tentatively placed here (fig. 26).
Brotocy. No data available.
160 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Vor. XL] HAYES: GALAPAGOS MOTHS 161
Utetheisa galapagensis (Wallengren). (Figures 22 & 23.)
Euchelia galapagensis WALLENGREN, 1860, Wiener entomologische Monatschrift, vol. 4, p. 161. Holotype, female (examined): Galapagos; NR.
The more grayish coloration and genitalic differences separate this species from U. perryi. A smaller moth than U. devriesi.
DISTRIBUTION. Endemic species. Galdpagos Islands: Baltra, April; Fer- nandina, February; Floreana, January, February, July; Isabela, January, March, April, August; Pinta, September, October; San Cristébal, February, September; Santa Cruz, January—December; Santiago, January, July, November, December. AMNH, BMNH, CAS, CU, IRSNB, MCZ, RSM, USNM.
Biotocy. Adults. Abundant generally in coastal and upland regions of main islands. On Santa Cruz often flying before dusk around plants of Scalesia affinis. Foodplants. Tournefortia pubescens, T. psilostachya. Larva. Grayish and brownish black with buff markings shading to pale buff below.
Utetheisa perryi Hayes, new species. (Figures 20, 21, 171, & 172.)
DeEscripTion. Male 16.5 mm. Antenna bipectinate. Palpus covered with dark brown and yellowish buff scales. Head, thorax, and forewing yellowish buff. Forewing irroration dark brown and black. Two such areas forming reniform spot. Hindwing yellowish cream, margin with dark brown markings and strongly marked discal spot. Genitalia: the uncus formation is diagnostic.
Female 16 mm. Antenna simple. Similar to male in basic coloration. Scaling on forewing forming longitudinal streak in some specimens.
The yellowish buff coloration separates this species from U. devriesi and U. galapagensis.
DISTRIBUTION. Endemic species.
HototyPe. Male. Santiago, 580 m., November/December 1970. J. Villa & J. Black. BMNH.
PaRATYPES. Isabela, January, February, October; Santiago, November, December; Santa Cruz, February, March, June. AMNH (2 specimens), BMNH (43 specimens), CAS (5 specimens).
Brotocy. Collected in transition and humid zones.
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Ficure 15. Hyles lineata florilega (Kernbach), male, Santa Cruz (x 2). Ficure 16. Epiplema becki Hayes, new species, holotype, male, Fernandina (x 2; CAS). Ficure 17. E. becki Hayes, new species, paratype, female, Isabela (xX 2). FicuRE 18. Utetheisa ornatrix (Linnaeus), male, Santa Cruz (x 2). Ficure 19. U. ornatrix (Linnaeus), female, Santa Cruz (xX 2).
162 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 20. Utetheisa perryi Hayes, new species, holotype, male, Santiago (xX 2). FIGURE 21. U. perryi Hayes, new species, paratype, female, Santiago (X 2). Ficure 22. U. gala- pagensis (Wallengren), male, Santiago (X 2). Ficurr 23. U. galapagensis (Wallengren), female, Santiago (xX 2). Ficure 24. U. devriesi Hayes, new species, holotype, male, Pinta (<2). Ficure 25. U. devriesi Hayes, new species, female, paratype, Pinta (x 2). FIGURE 26. U. devriesi Hayes, new species, melanic male, Pinta (X 2).
VoL. XL] HAYES: GALAPAGOS MOTHS 163
Family NocTuIDAE NOcTUINAE
Agrotis consternans Hayes, new species. (Figures 27, 28, 173, & 174.)
DescripTION. Male 15 mm. Antenna strongly bipectinate. Palpus dark brown. Thorax dark brown. Forewing midbrown with basal, antemedial, and postmedial lines buff edged with black. A broad black streak runs from the reniform through the orbicular spot. Hindwing cream with gray postmedian line and marginal shade. Genitalia as figured.
Female 16.5 mm. Similar to male in basic coloration. Costal margin yellowish buff, prominent.
Allied to A. bosgi Kohler, A. fasicola Dyar, and A. lutescens Blanchard, the orbicular and reniform spot and the strongly bipectinate male antenna separate this species.
DISTRIBUTION. Endemic species.
Hototypr. Male. Pinta, 630 m., November 1970. BMNH.
ParatypPEs. Isabela, August; Pinta, October, November; San Cristobal, April; Santa Cruz, March—June, November; Santiago, November. BMNH (92 specimens), CAS (29 specimens), MCZ (2 specimens), ZSBS (1 specimen).
Biotocy. No data available.
Agrotis ipsilon (Hufnagel). (Figures 31 & 32.) Phalaena ipsilon HUFNAGEL, 1766, Berlinisches Magazin, vol. 3, p. 416. Type material: Germany: Berlin; not traced. A well known species. DIsTRIBUTION. Almost cosmopolitan. Galdpagos Islands: Isabela, March; San Cristobal, April; Santa Cruz, January—July. AMNH, BMNH, CAS, MCZ. Brotocy. Not reared on the Galapagos Islands. Another general feeder. Kimball and Zimmerman give good coverage of crop damage, etc. Known as the Greasy Cutworm in the U.S.A.
Agrotis subterranea williamsi (Schaus), new combination.
(Figures 29 & 30.)
Euxoa williamsi Scuaus, 1923, Zoologica, vol. 5, p. 32. Lectotype female (examined): Galapagos: Indefatigable [Santa Cruz]; USNM.
Feltia annexa (Treitschke): RicHarps, 1941, Allan Hancock Pacific Expedition, vol. 5, py Zoe.
Scotia galapagosensis KOHLER, 1961, Anales de la Sociedad Cientifica Argentina, vol. 172, pp. 71-72. New synonym. The strongly pectinate male antenna and very dark forewing of the female
separate this subspecies. Hindwing of both sexes with some brown scales. DisTRIBUTION. Endemic subspecies of the widespread new world species.
164 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 27. Agrotis consternans Hayes, new species, holtype, male, Pinta ( X 2). Ficure 28. A. consternans Hayes, new species, paratype, female, Santa Cruz (Xx 2). FIGURE 29. A. subterranea williamsi (Schaus), male, Santiago (X 2). Ficure 30. A. subterranea williamsi (Schaus), female, Santa Cruz (x 2). Ficure 31. A. ipsilon (Hufnagel), male, Santa Cruz (x 2). Ficure 32. A. ipsilon (Hufnagel), female, Santa Cruz (X 2).
VoL. XL] HAYES: GALAPAGOS MOTHS 165
Galdpagos Islands: Baltra, April; Fernandina, February; Isabela, March, April, August; Pinta, May, October; Santa Cruz, January—July, October; Santiago, March, April, July, November. AMNH, BMNH, CAS, IRSNB, MCZ, USNM.
Biotocy. Larvae not found on the islands. Tietz (1972, p. 622) records the larvae of the nominate subspecies, the Granulate Cutworm, on a wide variety of plants.
Peridroma saucia (Hibner). (Figure 40, Mexican specimen.) Noctua saucia Htsner, [1808], Sammlung europdischer Schmetterlinge, vol. 4, fig. 378.
Type material: Europe; not traced.
Peridroma margaritosa (Haworth) sensu RicHarps, 1941, Allan Hancock Pacific Expedition, vol. 5, p. 235. Misidentification.
I have only examined 3 very worn specimens from the Galapagos Islands. These specimens have been identified on genitalic characters.
DIsTRIBUTION. Europe, North Africa, Asia, North America, South America, Hawaii. Galdpagos Islands: Floreana (Charles), 1300 ft., 1939, USC (2 ¢, ee).
BroLocy. Larvae not reared on the Galapagos Islands. Another general feeder, known as the Variegated Cutworm; Tietz (1972) lists a wide variety of foodplants including many crop species.
Psaphara conwayi (Richards), new combination. (Figures 37 & 38.) Peridroma conwayi RicHArps, 1941, Allan Hancock Pacific Expedition, vol. 5, p. 235.
Holotype, male (photograph examined): Galapagos: Charles [Floreana]; USC.
This species is closely allied to Psaphara interclusa Walker. I am taking the genus out of synonymy to contain these two species which on genitalic evidence merit separation from Peridroma. Genitalia figured by Richards (1941).
DisTRIBUTION. Endemic species. Galdpagos Islands: Floreana, no month; Isabela, January; Pinta, October; Santa Cruz, April. BMNH, CAS, USC.
Biotocy. No data available.
Psaphara interclusa Walker, revived combination.
(Figures 35, 36, & 163.)
Psaphara interclusa WALKER, 1857, List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, vol. 11, p. 607. Holotype, male (examined): West Coast of America [Galapagos]; BMNH.
The type is labeled ““W. Coast of America, Kellett and Wood, 1850-12.” Naval records (Seemann, 1853) disclose that H.M.S. Herald, captained by Sir Henry Kellett C. B., and H.M.S. Pandora, captained by Lt. Wood, visited the Galapagos Islands 6-16 January 1846, anchoring at Floreana, San Cristobal, and Santiago islands. The only additional material is a pair collected by Silberglied near ‘El Junco’ crater lake on San Cristébal, the female of which
[Proc. 4TH SER.
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VoL. XL] HAYES: GALAPAGOS MOTHS 167
is deposited in the BMNH. Both specimens match the type and this strongly suggests that the latter originated from the archipelago. Much historic material in the BMNH is labeled ‘“W. Coast of America” and a proportion appears to have been taken on the Galapagos Islands.
DISTRIBUTION. Endemic species. Galdpagos Islands: San Cristobal, April. BMNH, MCZ.
BioLtocy. No data available.
Anicla oceanica (Schaus), new combination. (Figures 33 & 34.) Lycophotia oceanica ScHaus, 1923, Zoologica, vol. 5, p. 32. Holotype, female (examined) :
Galapagos: South Seymour [Baltra]; USNM.
Lycophotia oceanica Schaus: Topp, 1973, Proc. Ent. Soc. Washington, vol. 75, p. 35 (Type specimen discussed).
A common insect on the Galapagos Islands. Sexes much alike but specimens show some variation in basic coloration from pinkish to grayish brown. Allied to Anicla infecta Ochsenheimer.
DISTRIBUTION. Endemic species. Galdpagos Islands: Baltra, April; Isabela, March, April, August; Pinta, October; Pinzon, April; Santa Cruz, January— April, June, August-October, December; Santiago, March, July. AMNH, BMNH, CAS, IRSNB, MCZ, RSM, USNM.
Brotocy. No data available.
HELIOTHINAE
Heliothis cystiphora (Wallengren)
(Figures 41 & 42.)
Anthoecia cystiphora \WALLENGREN, 1860, Wiener Entomologische Monatschrift, vol. 4, p. 172. Holotype, female: ‘Panama’; NR.
Anthoecia inflata WWALLENGREN, 1860, Wiener Entomologische Monatschrift, vol. 4, p. 172.
Anthoecia onca WALLENGREN, 1860, Wiener Entomologische Monatschrift, vol. 4, p. 172. Sexually dimorphic, the male possessing two prominent sensory patches on
the forewing. A common species, well represented in collections. DisTRIBUTION. Neotropical species. Galdpagos Islands: Baltra, April;
Espafola, May; Fernandina, February, April; Floreana, February—May;
Gardner near Espanola, February; Genovesa, April; Isabela, February—April;
e
Ficure 33. Anicla oceanica (Schaus), male, Santa Cruz (X 2). Ficure 34. A. oceanica (Schaus), female, Santa Cruz (x2). Ficure 35. Psaphara interclusa Walker, holotype, male, ‘W. Coast of America’ (Galapagos) (X 2 reversed). Ficure 36. P. interclusa Walker, female, San Cristobal (x 2). Ficure 37. P. conwayi (Richards), male, Isabela (X 2). Ficure 38. P. conwayi (Richards), female, Pinta (X 2). Ficure 39. Heliothis virescens (Fabricius), female, Santa Cruz (xX 2). Ficure 40. Peridroma saucia (Hiibner), male, Mexico (X 2).
168 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
San Cristobal, February, March; Santa Cruz, January—June; Santiago, March, April. AMNH, BMNH, CAS, IRSNB, LACM, MCZ, USNM.
BioLocy. A fast-flying species, visiting flowers during day; common in March 1969 at plants of Encelia hispida in the inland parts of Santa Fé. Food- plant. Sporobolus virginicus. Larva. Head yellow with black spots. Body with central gray stripe bordered by yellow, reddish brown, and white; over- lain with black spots; undersurface yellowish green.
Heliothis virescens (Fabricius). (Figure 39.) Noctua virescens FaBrictus, 1781, Species Insectorum, vol. 2, p. 216. Type material: [Virgin
Islands]: St. Crux [St. Croix]; not traced.
Some variation in the hindwing coloration is common in series of this moth.
DIsTRIBUTION. Widely distributed in the neotropics. Galdpagos Islands: Floreana, July; Genovesa, April; Isabela, January—April, August; Pinta, May, October; San Cristobal, February, April; Santa Cruz, January—April, June, July, October, December; Santiago, July. AMNH, BMNH, CAS, IRSNB, MCZ.
Briotocy. Foodplants. Passiflora foetida, Scalesia affinis. Larva. Head yellow. Body yellowish to brownish green merging into emerald below; some orange dorsally; black median and lateral lines; prominent black spots at bases of setae. Known as the Tobacco Budworm in the U.S.A. where it has been recorded on an extensive variety of plants.
HADENINAE Mythimna solita (Walker). (Figure 43.) Leucania solita WALKER, 1856, List of the Specimens of Lepidopterous Insects in the Collec- tion of the British Museum, vol. 9, p. 99. Holotype, male (examined): BMNH.
The longitudinal streak on the forewing and the white hindwing are features of this species.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: San Cristébal, April; Santa Cruz, February, March, May, June, October-December. AMNH, BMNH, CAS, IRSNB, MCZ.
Biotocy. Foodplant. Sporobolus virginicus. Larva. Head gray with brown reticulation. Body reddish brown with darker markings and diffuse white lines.
Mythimna latiuscula (Herrich-Schaffer). (Not figured.)
Leucania latiuscula HERRICH-SCHAFFER, 1868, Korrespondez-Blatt des Zoologisch-Minera- logischen Vereines in Regensburg, vol. 22, p. 148. Type material: Cuba. A specimen of this species was mailed to me after I had prepared the plates. It is figured by Draudt in Seitz, Macrolepidoptera of the World, vol. 7, pl. 24.
VoL. XL] HAYES: GALAPAGOS MOTHS
Ficure 41. Heliothis cystiphora (Wallengren), male, Floreana (X 2). cystiphora (Wallengren), female, Isabela (xX 2). FicurE 43. Mythimna solita (Walker), male, Santa Cruz (X 2). Ficure 44. Pseudaletia sequax Franclemont, male, Santa Cruz
(X 2). Ficure 45. P. cooperi (Schaus), male, Santa Cruz (xX 2). Ficure 46. P. cooperi (Schaus), female, Santa Cruz (xX 2).
FicuRE 42. dH.
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It is more drab than M. solita and lacks the longitudinal white streak on the forewing.
DisTRIBUTION. Widely distributed in the neotropics. Galdpagos Islands: San Cristobal, ‘El Junco’ crater lake, 700 m., April. MCZ.
Brotocy. Not reared on the Galapagos Islands. Tietz (1972) lists grami- naceous foodplants.
Pseudaletia sequax Franclemont. (Figure 44.)
Pseudaletia sequax FRANCLEMONT, 1951, Proc. Ent. Soc. Washington, vol. 53, p. 70. Holotype, male (examined): Jalapa, Mexico; USNM.
Genitalia compared with Franclemont’s figure and paratypes in the BMNH.
DISTRIBUTION. Widespread neotropical species. Franclemont gives full coverage. Galdpagos Islands: Isabela, February; San Cristdbal, April; Santa Cruz, January—April, June, August, October, December. AMNH, BMNH, CAS, IRSNB, LACM, MCZ, RSM.
Brotocy. No data available. Related species are general feeders.
Pseudaletia cooperi (Schaus).
(Figures 45 & 46.)
Cirphis cooperi ScHAUS, 1923, Zoologica, vol. 5, p. 33. Holotype, female (examined): Galapagos: Indefatigable [Santa Cruz]; USNM.
Pseudaletia cooperi Schaus: FRANCLEMONT, 1951, Proc. Ent. Soc. Washington, vol. 53, p. 64.
Both sexes figured.
DISTRIBUTION. Endemic species. Galdpagos Islands: Isabela, March, August; Pinta, October; San Cristdbal, April; Santa Cruz, January—June, September— November; Santiago, November. BMNH, CAS, IRSNB, MCZ, USNM.
BroLtocy. No data known.
ACRONICTINAE
Magusa erema Hayes, new species. (Figures 53, 54, & 155.)
Magusa orbifera Walker sensu ScHAus, 1923, Zoologica, vol. 5, p. 24 [Misidentification].
DeEscripTION. Male 21.5 mm. Antenna simple. Palpus, head, thorax, and forewing dark brown. Prominent white apical mark on forewing. Abdomen and hindwing grayish brown. Genitalia as figured. Valve process short.
Female 20 mm. Similar to male but with broader forewing.
Although reddish brown coloration occurs on the forewing of some specimens, a series of this insect does not exhibit the extreme variation shown by its nearest
VoL. XL] HAYES: GALAPAGOS MOTHS 171
relative Magusa orbifera Walker. The short valve process of the male genitalia also separates M. erema.
DISTRIBUTION. Endemic species.
HototyrPe. Male. Santa Cruz, December 1968, R. Perry and Tj. de Vries. BMNH.
ParATYyPES. Isabela, April; San Cristobal, April; Santa Cruz, January— May, July, August, October, December; Santa Fé, April. AMNH (1 specimen), BMNH (16 specimens), CAS (1,058 specimens), IRSNB (6 specimens), MCZ (143 specimens), RSM (3 specimens).
Biotocy. Foodplant. Scutia pauciflora. Larva. Green with paired, in- terrupted lines of gray-black and yellowish and creamy white stripes; some purple around legs and prolegs.
Trachea cavagnaroi Hayes, new species. (Figures 49, 50, & 156.)
DESCRIPTION. Male 17 mm. Antenna strongly bipectinate. Palpus dark brown and buff. Forewing with dark and mid brown patterning. Burnished orbicular and reniform spots. Marginal band and discal spot of hindwing gray. Remainder of hindwing yellowish gold. Genitalia as figured.
Female 19-22 mm. Antenna simple. Broader winged but similar to male in maculation. Conspicuous buff pattern in apical area and bordering terminal line of forewing.
Only one male and two females of this species have been taken. Provisionally placed in the genus Trachea. Does not resemble any known species.
DISTRIBUTION. Endemic species.
Hototyre. Male. Santa Cruz, Horneman Farm, 220 m., 3 May 1964. D. Q. Cavagnaro. CAS.
PaRATYPES. Santa Cruz, Grasslands, 750 m., 10 April 1964. D. Q. Cavagnaro. CAS. (2 females.)
BroLocy. No data known.
Cropia infusa (Walker).
(Figures 47 & 48.)
Decelea infusa WALKER, [1858] 1857, List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, vol. 13, p. 1116. Holotype, female (examined): America but no data; BMNH.
Few specimens of this species have been collected on the Galapagos Islands. Although somewhat smaller than mainland examples, no striking genitalic dif- ferences separate Galapagos Islands specimens.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Espanola, May; Fernandina, February; Gardner near Espafiola, April; Genovesa, Feb-
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Vor. XL] HAYES: GALAPAGOS MOTHS 173
ruary, March; Isabela, March; Santa Cruz, January, June; Santiago, July. AMNH, BMNH, CAS, MCZ. Brotocy. No data known.
Callopistria floridensis (Guenée).
(Figure 51.)
Eriopus floridensis GuENEE, 1852, in Boisduval & Guenée, Histoire Naturelle des Insectes. Lépidoptéres, vol. 6, p. 292. Holotype, male (examined): [U.S.A.]: Florida; BMNH. Identical with mainland specimens.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Isabela, January, August; Pinta, October; San Cristobal, April; Santa Cruz, January— July. AMNH, BMNH, CAS, IRSNB, MCZ.
Brotocy. Not reared on the Galapagos Islands. Known as the Florida Fern caterpillar in the U.S.A.
Catabena seorsa Todd.
(Figure 52.)
Catabena seorsa Topp, 1972, Jour. Washington Acad. Sci., vol. 62, no. 1, p. 38. Holotype, male (examined): Galapagos: Santa Cruz; CAS. Todd (1972) gives full coverage. Schaus (1923, p. 25) referred to a specimen now located in AMNH as a Catabena species? too poor to identify.
DIsTRIBUTION. Endemic species. Galdpagos Islands: Marchena, September, November; Espanola, September; Floreana, April, June, July; Genovesa, Feb- ruary—April; Isabela, January, March, August, October, November; Pinzon, April; San Cristébal, February; Santa Cruz, January—May, October, December. AMNH, BMNH, CAS, IRSNB, MCZ, USNM, ZSBS.
BroLtocy. Arid zones, generally. Foodplant. Lantana peduncularis. Larva. Head gray with black and orange markings. Body gray and black with fine lines of orange, yellow, and white. Pupation is within a parchmentlike cocoon.
Neogalea esula longfieldae Hayes, new subspecies. (Figures 55 & 56.)
The extensive fuscous margin of the hindwing separates Galapagos speci- mens from the nominate subspecies.
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Ficure 47. Cropia infusa (Walker), male, Espafiola (x 2). Ficure 48. C. infusa (Walker), female, Santiago (x2). Ficure 49. Trachea cavagnaroi Hayes, new species, holotype, male, Santa Cruz (x 2; CAS). Ficure 50. 7. cavagnaroi Hayes, new species, paratype, fe- male, Santa Cruz (x 2; CAS). Ficure 51. Callopistria floridensis (Guenée), male, Isabela (<2). Ficure 52. Catabena seorsa Todd, female, Santa Cruz (x 3). Ficure 53. Magusa erema Hayes, new species, holotype, male, Santa Cruz (x 2). Ficure 54. M. erema Hayes, new species, paratype, female, Santa Cruz (x 2).
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DISTRIBUTION. Endemic subspecies. The widespread neotropical species has been introduced to Australia, Hawaii, and Norfolk Island in an endeavour to control Lantana.
Hototype. Male. 17 mm. Isabela (Albemarle), Tagus Cove, 150 ft., 3 August 1924, St. George Expedition, C. L. Collenette, BM. 1925-488, BMNH.
PARATYPES. Same data as holotype, 4 August 1924, 1 female, BMNH; same data as holotype, 7 August 1924, 1 female, BMNH: Isabela, Punta Albemarle, March 1970, R. Silberglied, BM. 1970—567, 1 female, BMNH; Isabela, Punta Albemarle, March 1970, R. Silberglied, 4 males, 1 female, MCZ.
BioLocy. Not reared on the Galapagos Islands. Tietz lists Lantana and Verbena as foodplants for the nominate subspecies.
Spodoptera eridania (Stoll). (Figures 57 & 58.)
Phalaena (Noctua) eridania Stott, 1781, in Cramer, Uitlandsche Kapellen, vol. 4, p. 133, pl. 358, figs. E & F. Type material: Surinam; not traced.
A variable insect that is a common pest on the mainland.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Isabela, March; San Cristobal, April; Santa Cruz, February—July. AMNH, BMNH, CAS, MEZ;
Biotocy. Foodplants. Amaranthus viridis, Portulaca oleracea, Cryptocar pus pyriformis, Cissampelos pareira, Ipomoea pes-caprae. Larva. Head reddish brown. Body gray, streaked green and reddish brown with black markings; lateral line black. Tietz lists a very wide range of foodplants. This species is known as the Southern Armyworm in the U.S.A.
Spodoptera latifascia (Walker). (Figures 59 & 60.)
Prodenia latifascia WALKER, 1856, List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, vol. 9, p. 195. Holotype, male (examined): Jamaica; UM.
Sexually dimorphic. DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Flore-
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Ficure 55. Neogalea esula longfieldae Hayes, new subspecies, holotype, male, Isabela (x 2 reversed). Ficure 56. N. esula longfieldae Hayes, new subspecies, paratype, female, Isabela (X 2). Ficure 57. Spodoptera eridania (Stoll), male, Santa Cruz (xX 2). FIGURE 58. S. eridania (Stoll), female, Santa Cruz (Xx 2). Ficure 59. Spodoptera latifascia (Walker), male, Santa Cruz (xX 2). Ficure 60. S. latifascia (Walker), female, Santa Cruz (xX 2). Ficure 61. Spodoptera frugiperda (Smith), male, Santa Cruz (X 2; CAS). Ficure 62. S. frugiperda (Smith), female, Santa Cruz (x 2; AMNH).
VoL. XL]
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176 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.
ana, May; Isabela, April; San Cristobal, February; Santa Cruz, March, April, October, December. BMNH, CAS, IRSNB.
BioLtocy. Not reared on the Galapagos Islands. A pest of citrus on the mainland. Kimball and Tietz give foodplants.
Spodoptera dolichos (Fabricius). (Figures 65 & 66.) Noctua dolichos Fasrictus, 1794, Entomologia Systematica, vol. 3, no. 2, p. 95. Type ma-
terial: Americae meridionalis; not found by Zimsen, 1964, p. 570.
In common with some other members of the genus, this species is of economic importance on the mainland.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Floreana, May; Santa Cruz, January—June, October, December. AMNH, BMNH, CAS, IRSNB.
BroLocy. Larvae reared on Cryptocarpus pyriformis have been referred to this species. Polyphagous.
Spodoptera frugiperda (Smith).
(Figures 61 & 62.)
Phalaena frugiperda J. E. SmitH, 1797, in Abbot & Smith, The Natural History of the Rarer Lepidopterous Insects of Georgia, vol. 2, p. 191, pl. 96. Type material: U.S.A., Georgia; not traced.
Another sexually dimorphic species of Spodoptera.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Floreana, April; San Cristobal, April; Santa Cruz, April, June; Santiago, March. AMNH, CASS MEz:
BioLocy. Not reared on the Galapagos Islands. A very general feeder.
Spodoptera roseae (Schaus).
(Figures 63 & 64.)
Trachea roseae ScHAUS, 1923, Zoologica, vol. 5, p. 33, pl. 1, fig. 4. (but proposed as ‘Trachaew an incorrect spelling). Holotype, male (examined): Galapagos: Indefatigable [Santa Cruz]; USNM.
Laphygma roseae (Schaus): RicHArps, 1941, Allan Hancock Pacific Expedition, vol. 5, p. 239.
Spodoptera roseae (Schaus): Linstry & Usincrr, 1966, Proc. Calif. Acad. Sci., vol. 33, no. 7, p. 160.
Richards (1941) dealt with this species in detail.
DISTRIBUTION. Endemic species. Galdpagos Islands: Floreana, April; Isabela, January, March; Pinta, October; San Cristébal, April; Santa Cruz, January— July, October, December; Santiago, March, AMNH, BMNH, CAS, IRSNB, MCZ, USNM, ZSBS.
BioLocy. No data available.
VoL. XL]
Ficure 63. Spodoptera roseae (Schaus), male, Santa Cruz (x 2). Ficure 64. S. roseae (Schaus), female, Santa Cruz (xX 2). Ficure 65. Spodoptera dolichos (Fabricius), male, Santa Cruz (x 2). Ficure 66. S. dolichos (Fabricius), female, Santa Cruz (x 2). FIGURE 67. Agrotisia williamsi (Schaus), male, Santa Cruz (X 2). FicurE 68. Platysenta ruthae
(Schaus), male, Santa Cruz (X 2).
178 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Elaphria encantada Hayes, new species. (Figures 72, 73, 161, & 162.) Elaphria dubiosa (Schaus) sensu HAyeEs, 1972, Pan-Pacific Entomologist, vol. 48, p. 104.
Misidentification.
Description. Male 11 mm. Antenna simple. Palpus clothed with a mix- ture of dark brown and buff scales. Forewing with prominent orbicular and reniform spots. Claviform spot elongate. Variable in forewing color and pat- terning. Hindwing brown and buff. Genitalia as figured. Larger than average Pinta specimens cannot be separated structurally.
Female 10.5 mm. Similar to male. Variable.
Nearest relative is Elaphria chalcedonia Hibner. Elaphria encantada lacks the prominent apical spot of related species and can also be separated on genitalic structure.
DISTRIBUTION. Endemic species.
HototypPe. Male. Santa Cruz, December, 1968. R. Perry and Tj. de Vries. BMNH.
PaRATYPES. Espanola, February; Fernandina, February; Floreana, May; Isabela, January, March, August, September; Pinta, April, May, September, October; Pinzon, February, April, December; Santa Cruz, January—August, October-December; Santiago, March, April, July, August, November. AMNH (13 specimens), BMNH (331 specimens), CAS (853 specimens), IRSNB (46 specimens), MCZ (87 specimens), RSM (5 specimens), USC (1 specimen), ZSBS (1 specimen).
BrioLtocy. No data available.
Platysenta mobilis (Walker), revived species.
(Figure 71.)
Perigea mobilis WALKER, [1857] 1856, List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, vol. 10, p. 277. Holotype, male (examined):
St. Domingo; BMNH.
Confusion has arisen in the literature and collections over this insect which has in the past been incorrectly determined as Perigea apameoides Guenée. Some specimens lack the white reniform spot.
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Isabela, April; Santa Cruz, April-June; Santiago, April. AMNH, BMNH, CAS:
Brotocy. No data available.
Platysenta sutor (Guenée). (Figures 69 & 70.)
Perigea sutor GUENEE, 1852, in Boisduval & Guenée, Histoire naturelle des Insectes. Lépidop- téres, vol. 5, p. 231. Holotype, male (examined): Brazil; BMNH.
Vor. XL] HAYES: GALAPAGOS MOTHS 179
Perigea apameoides GUENEE, 1852, in Boisduval & Guenée, Historie naturelle des Insectes. Lépidoptéres, vol. 5, p. 229. (Lectotype designated by Viette, 1951, Bulletin Mensuel de la Société Linnéenne de Lyon, vol. 20, p. 160.) MNHN. New synonym. Lectotype figured here—figure 70.
Perigea ebba ScHAUsS, 1923, Zoologica, vol. 5, p. 36. New synonym.
Size and coloration of this species varies. The Guenée name P. apameoides has been incorrectly used in the literature and collections for P. mobilis. To avoid confusion, and acting as first reviser under the International Code of Zoological Nomenclature, Article 24, I am regarding ‘apameoides’ as a synonym of P. sutor although P. apameoides has page priority. Subsequent to Viette’s lectotype designation of P. apameoides, a paralectotype female from Coll. Guérin has been traced in the BMNH. It is conspecific with the lectotype and also bears the data “I. St. Thomas.”
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Baltra, April; Isabela, January, March; Pinzon, April; San Cristobal, April; Santa Cruz, January—June, October, December; Santiago, March, April, No- vember. AMNH, BMNH, CAS, IRSNB, MCZ, USNM.
BioLtocy. No data available. Kimball lists Wedelia and Tagetes (Com- positae) as foodplants in Florida.
Platysenta ruthae (Schaus).
(Figure 68.)
Perigea ruthae Scuaus, 1923, Zoologica, vol. 5, p. 35. Lectotype, male (examined): Galapagos: Albemarle [Isabela]; USNM.
Platysenta ruthae (Schaus): RicHArps, 1941, Allan Hancock Pacific Expedition, vol. 5, p. 237. Richards (1941) dealt with this species in detail. Widespread in the Islands. DISTRIBUTION. Endemic species. Galdpagos Islands: Baltra, April; Floreana,
April, July; Isabela, March, April, August; Pinta, October; Pinzon, April;
Santa Cruz, January—June, August, October-December; Santiago, March, April,
July. AMNH, BMNH, CAS, IRSNB, MCZ, USNM, ZSBS.
Biotocy. No data available.
Agrotisia williamsi (Schaus), new combination. (Figure 67.) Harrisonia williamsi ScHaus, 1923, Zoologica, vol. 5. p. 36. Lectotype, male (examined): Galapagos: South Seymour [Baltra]; USNM. Richards (1941) figured the genitalia of this species. I am indebted to Prof. J. G. Franclemont for pointing out that this species is congeneric with A. sub- hyalina Hampson, the type species of Agrotisia Hampson 1908. Widespread, seasonally abundant species. DISTRIBUTION. Endemic species. Galdpagos Islands: Baltra, April; Es-
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Ficure 69. Platysenta sutor (Guenée), male, Santa Cruz (X 2). Ficure 70. Perigea apameoides (Guenée), lectotype, male, I. St. Thomas, Virgin Islands, synonym of P. sutor Guenée (X 2; MNHN). Ficure 71. Platysenta mobilis Walker, male, Santa Cruz (X 2). Ficure 72. Elaphria encantada Hayes, new species, holotype, male, Santa Cruze Ficure 73. E. encantada Hayes, new species, paratype, female, Santa Cruzer
Vout. XL] HAYES: GALAPAGOS MOTHS 181
panola, February; Floreana, April; Gardner near Espanola, April; Isabela, March; Pinta, May; Santa Cruz, January—June, August-December; Santiago, July. AMNH, BMNH, CAS, IRSNB, MCZ, RSM, USNM, ZSBS.
Brotocy. No data available.
ACONTIINAE
Ozarba consternans Hayes, new species. (Figures 75-78, & 158.)
DeEscripTION. Male 8.5 mm. Antenna simple. Forewing blackish brown. Hindwing lighter shade of brown. Genitalia as figured. Unlike any known species.
Female 10 mm. Broader winged but similar to male.
Great variation in forewing maculation exists within this species (see figures). Provisionally placed in the genus Ozarba.
DISTRIBUTION. Endemic species.
Ho.totyvpe. Male. Santa Cruz, May 1970, R. Perry and Tj. de Vries.
PaRATYPES. Espanola, April; Floreana, April; Pinta, October; Santa Cruz, February, March, May, June; Santiago, March. BMNH (5 specimens), CAS (17 specimens), MCZ (17 specimens), ZSBS (5 specimens).
BroLtocy. No data available.
Bagisara repanda (Fabricius), new combination. (Figure 81.) Bombyx repanda Fasricius, 1793, Entomologia Systematica, vol. 3, no. 1, p. 462. Lecto- type (photograph examined): Americae meridionalis Insulis; UZM. McDunnough (1938) and Kimball (1965) listed this species as Atethmia subusta Hubner. Draudt in Seitz used the combination Bagisara subusta. DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Isabela (humid zone) Volcan Chico, June 1970, J. Gordillo; BMNH. BioLtocy. No data available. A specimen from Barbados in the BMNH was reared on Sida glomerata.
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Ficure 74. Eublemma recta (Guenée), male, Santa Cruz (Xx 2). Ficure 75. Ozarba con- sternans Hayes, new species, holotype, male, Santa Cruz (x 2). FIGURE 76. O. consternans Hayes, new species, paratype, female, Santa Cruz (XX 2). FicuRE 77. O. consternans Hayes, new species, paratype, male, Santa Cruz (x 2; CAS). Ficure 78. O. consternans Hayes, new species, paratype, female, Santa Cruz (x 2; CAS). Ficure 79. Amyna insularwm Schaus, male, Santa Cruz (x 2). Ficure 80. A. insularum Schaus, male. Santa Cruz (X 2). FIGURE 81. Bagisara repanda (Fabricius), female, Isabela (x 2).
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Eublemma recta (Guenée), new combination. (Figure 74.)
Micra recta GUENEE, 1852, in Boisduval & Guenée, Histoire naturelle des Insectes. Lépidop- teres, vol. 6, p. 245. Holotype, male (examined): neotropical but stated to be “Sierra Leone?” by Guenée; USNM.
Somewhat variable in basic coloration.
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Genovesa, April; Santa Cruz, September. AMNH, BMNH.
BroLtocy. Foodplant. Ipomoea triloba. Larva. Reddish brown. Pupation within silken cocoon. Forbes (1954) states the larvae feed on the buds and seeds of Convolvulus and sweet potato in the U.S.A. (Species listed as Eublemma obliqualis Fabricius).
Amyna insularum Schaus. (Figures 79 & 80.)
Amyna insularum ScHAUS, 1923, Zoologica, vol. 5, p. 37. Lectotype, male (examined): Galapagos: Indefatigable [Santa Cruz]; USNM.
Two forms of this insect are recognized. One possesses a prominent white reniform spot. Both forms are figured. Well represented in collections.
DISTRIBUTION. Endemic species. Galapagos Islands: Baltra, April; Espanola, April; Fernandina, February; Floreana, January, April, July; Gardner near Espanola, April; Genovesa, no month; Isabela, January, March—June, August; Pinzon, April, June; Rabida, June; San Cristobal, February—April; Santa Cruz, January—December; Santa Fé, April; Santiago, January, February—April. AMNH, BMNH, CAS, LACM, IRSNB, MCZ, RSM, ZSBS, USC.
BrioLtocy. Widespread, coastal regions; abundant; individuals at beginning of season may be smaller. At rest the wings are held flat and extended back- wards, so that the insect assumes a characteristically triangular outline. Food- plants. Alternanthera echinocephala, A. filifolia. Larva. Pale green with darker lines.
Heliocontia margana (Fabricius). (Figures 82 & 83.)
Pyralis margana Fapricius, 1794, Entomologia Systematica, vol. 3, no. 2, p. 257. Type ma- terial: Americae Insulis; UZM.
Sexually dimorphic.
DIsTRIBUTION. Widely distributed in the neotropical region. Galdpagos Islands: Baltra, April; Espanola, April; Floreana, April; Gardner near Espanola, April; Genovesa, February; Isabela, March; San Cristobal, February, April; Santa Cruz, February, April, May, August; Santiago, February, March. AMNH, BMNH, CAS, MCZ, ZSBS, USC.
Vor. XL] HAYES: GALAPAGOS MOTHS 183
BroLocy. Foodplants. Sida species, Abutilon depauperatum. Larva. Variable, green with lighter and darker lines.
Spragueia creton Schaus. (Figures 84-86.) Spragueia creton SCHAUS, 1923, Zoologica, vol. 5, p. 38. Lectotype, male (examined):
Galapagos: Genovesa (Tower); USNM.
Spragueia plumbeata ScHAUusS, 1923, Zoologica, vol. 5, p. 38. Spragueia creton Schaus: Topp, 1972, Jour. Washington Acad. Sci., vol. 62, no. 1, p. 36.
Todd has given full coverage of this sexually dimorphic species, placing S. plumbeata in synonymy.
DISTRIBUTION. Endemic species. Galdpagos Islands: Baltra, April; Espanola, February, April; Floreana, April; Gardner near Espanola, April; Genovesa, February, April; Isabela, March; San Cristobal, April; Santa Cruz, March, May; Santiago, February, Marchh AMNH, BMNH, CAS, MCZ, USNM, ZS BS.
Brotocy. No data available.
Ponometia indubitans (Walker).
(Figures 92-94.)
Nonagria indubitans WALKER, 1857, List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, vol. 11, p. 712. Holotype, male (examined): [Brazil]: Para; BMNH.
Another sexually dimorphic, variable species. Further material of this species is needed.
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Gardner near Hood, April; Genovesa, February; Isabela, March; San Cristobal, February; Santa Cruz, May. BMNH, CAS, MCZ, USNM.
Biotocy. Foodplant. Waltheria ovata. Larva. Head greenish brown with black etchings. Body grayish green with black lines and markings. Pupa. Thin- walled, yellowish brown.
EUTELIINAE
Paectes arcigera (Guenée).
(Figures 89-91.)
Ingura arcigera GUENEE, 1852, in Boisduval & Guenée, Histoire Naturelle des Insectes. Lépidoptéres, vol. 6, p. 312. Holotype, female: [Virgin Islands]: Ile Saint-Thomas; not traced.
Paectes indefatigabilis ScHAUsS, 1923, Zoologica, vol. 5, p. 38. New synonym.
Paectes isabel ScHaus, 1923, Zoologica, vol. 5, p. 39. New synonym.
Sexually dimorphic, specimens exhibit considerable variation. The new synonymy has been established by genitalia preparations. Well represented in collections.
184 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
FIGURE 82.
FIGURE 83.
Vor. XL] HAYES: GALAPAGOS MOTHS 185
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Baltra, April; Floreana, April, October; Gardner near Espanola, April; Genovesa, no month; Isabela, March—May, August; Pinta, May; Pinzon, April; San Cristobal, April; Santa Fé, April; Santa Cruz, January—August, October, December; Santiago, March. AMNH, BMNH, CAS, IRSNB, MCZ, USNM.
BroLtocy. On wing throughout the period December 1968 to May 1969, with a peak of abundance from mid March to early April; coastal areas, late afternoons, to be seen among foliage of Cryptocarpus pyriformis. Foodplant. Bursera graveolens. Larva. Green with pale lines; pale yellow on Ist segment and with 4 pigment spots at bases of setae.
SARROTHRIPINAE
Characoma nilotica (Rogenhofer). (Figures 87 & 88.)
Sarrothripa nilotica ROGENHOFER, 1882, Verhandlungen der Zoologisch-botanischen Gesell- schaft in Wien, vol. 31, p. 26. Holotype (examined): Egypt; NM.
Variable species.
DISTRIBUTION. Pantropical species. Galdpagos Islands: Fernandina, Jan- uary; San Cristébal, April; Santa Cruz, January, February, June, September— November; Santiago, July. AMNH, BMNH, CAS, IRSNB.
Biotocy. Larval foodplant Laguncularia racemosa.
CATOCALINAE Mocis incurvalis Schaus. (Figures 95-97.)
Mocis incurvalis ScHaus, 1923, Zoologica, vol. 5, p. 41. Lectotype, male (examined):
Galapagos: Indefatigable [Santa Cruz]; USNM.
Sexually dimorphic. Some variation in female specimens.
DISTRIBUTION. Endemic species. Galdpagos Islands: Baltra, April; Fer- nandina, April; Isabela, March, April, August; Pinta, May, October; San Cristébal, April; Santa Cruz, March—July, September, October. AMNH, BMNH, CAS, IRSNB, MCZ, RSM, USNM, ZSBS.
Brotocy. No data available. Gramineae are listed as important among foodplants of related species.
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(<2; IRSNB). Ficure 89. Paectes arcigera (Guenée), male, Santa Cruz (X 2). FIGURE 90. P. arcigera (Guenée), male, Santa Cruz (X 2). Ficure 91. P. arcigera (Guenée), female, Santa Cruz (x2). Ficure 92. Ponometia indubitans (Walker), male, Isabela (<2; CAS). Ficure 93. P. indubitans (Walker), male, Floreana (xX 2). Ficure 94. P. indubitans (Walker), female, Santiago (xX 2).
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Ficure 95. Mocis ncurvalis Schaus, male, Santa Cruz (xX 1). FicurE 96. M. incurvalis Schaus, female, Santa Cruz (x 1). Ficure 97. M. incurvalis Schaus, female, Santa Cruz (X 1). Ficure 98. Celiptera remigioides (Guenée), female, Santa Cruz (x1). FIGURE 99. Mocis latipes (Guenée), female, Santa Cruz (X11). Ficure 100. M. latipes (Guenée), female, Santa Cruz (1). Ficure 101. Melipotis acontioides producta Hayes, new subspecies, holotype, male, Santa Cruz (x 1). Ficure 102. WM. acontioides producta Hayes, new subspecies, paratype, female, Santa Cruz (x1). Ficure 103. M. indomita (Walker), male, Santa Cruz (x 1). Ficure 104. M. indomita (Walker), female, Santa Cruz (<1). Ficure 105. M. harrisoni Schaus, male, Santa Cruz (<1). FIGURE 106. M. harrisoni Schaus, female, Santa Cruz (x 1). FicurEe 107. Zale obsita (Guenée), male, Santa Cruz (x 1). FicurE 108. Z. obsita (Guenée), male, Santa Cruz (x 1; CAS). Ficure 109. Z. obsita (Guenée), female, Santa Cruz (Xx 1). FicurE 110. Z. obsita (Guenée), female, Santiago (x 1).
Vou. XL] HAYES: GALAPAGOS MOTHS 187
Mocis latipes (Guenée).
(Figures 99 & 100.)
Remigia latipes GuENEE, in Boisduval & Guenée, 1852, Histoire Naturelle des Insectes. Lépidoptéres, vol. 7, p. 314. Lectotype, male (examined): Guadeloupe; BMNH. Variable species.
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Baltra, April; Floreana, May; San Cristébal, April; Santa Cruz, April-June. AMNH, BMNH, CAS, MCZ, USC.
Brotocy. Larvae not yet found on the Galapagos Islands. In the U.S.A. this species feeds on grasses and other crops.
Celiptera remigioides (Guenée).
(Figure 98.)
Ophiodes remigioides GUENEE, 1852, in Boisduval & Guenée, Histoire Naturelle des Insectes. Lépidoptéres, vol. 7, p. 230, pl. 21, fig. 5. Syntype male (examined): neotropical but stated to be Central India; BMNH.
The hindwing of this species varies from yellow to fuscous.
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Genovesa, March; Santa Cruz, February—April, July, October, December. AMNH, BMNH, CAS.
BroLocy. No data available.
Zale obsita (Guenée), revived species. (Figures 107-110.) Homoptera obsita GUENEE, 1852, in Boisduval & Guenée, Histoire Naturelle des Insectes.
Lépidoptéres, vol. 7, p. 12. Holotype, female (examined): Brazil; BMNH.
Zale species, viridans group RicHarps, 1941, Allan Hancock Pacific Expedition, vol. 5, p. 243.
Richards figured the male genitalia of this variable species which was pre- viously synonymized with Z. viridans. I am indebted to Dr. E. L. Todd for help with this identification.
DisTRIBUTION. A neotropical species. Galdpagos Islands: Santa Cruz, February—April, June, October, November; Santiago, November. AMNH, BMNH, CAS, IRSNB, MCZ.
Brotocy. No data available.
PLUSIINAE
Autoplusia egena galapagensis (Schaus). (Figure 111.) Syngrapha egena galapagensis ScHAuS, 1923, Zoologica, vol. 5, p. 41. Holotype, female (examined): Galapagos: James [Santiago]; USNM. This is the dullest colored of the three species of this subfamily known from the Galapagos Islands.
188 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
DISTRIBUTION. Endemic subspecies of the neotropical species. Galdpagos Islands: Baltra, April; Isabela, May, September; Pinzén, June; San Cristobal, April, June; Santa Cruz, February, May, June; Santiago, April. AMNH, BMNH, CAS, USNM.
Brotocy. No data available. In the U.S.A. the nominate subspecies feeds on Phaseolus (Leguminosae).
Argyrogramma verruca (Fabricius). (Figure 112.)
Noctua verruca FaAsrictus, 1794, Entomologia Systematica, vol. 3, pt 2, p. 81. Type ma- terial: Americae meridionalis Insulis; UZM.
This is the smallest species of this subfamily found on the Galapagos Islands.
DISTRIBUTION. Widespread in the neotropics, extending to Canada. Gald- pagos Islands: Floreana, January; Isabela, January; Santa Cruz, February, April. AMNH, BMNH, CAS.
BroLtocy. No data available. Foodplants in the U.S.A. include Sagittaria and Calendula.
Pseudoplusia includens (Walker). (Figure 113.)
Plusia includens WALKER, [1858] 1857, List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, vol. 12, p. 914. Type female (examined): St. Domingo; BMNH.
Previously treated as Phalaena oo (Stoll) by some workers, but this is a junior primary homonym. DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands:
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Ficure 111. Autoplusia egena galapagensis (Schaus), male, Santa Cruz (xX 1). FIGURE 112. Argyrogramma verruca (Fabricius), male, Isabela (X 1). Ficure 113. Pseudoplusia includens (Walker), female, Santa Cruz (x 1). FicurE 114. Ascalapha odorata (Linnaeus), male, Santa Cruz (x %; CAS). Ficurr 115. A. odorata (Linnaeus), female, Santa Cruz (X %). Ficure 116. Letis mycerina (Cramer), male, Panama (xX %4). Ficure 117. L. mycerina (Cramer), female, Santa Cruz (X %4). Ficure 118. Epidromia zetophora Guenée, male, Isabela (X 1). FicurE 119. Psorya hadesia Schaus, male, Santa Cruz (Xx 1). FIcURE 120. P. hadesia Schaus, female, Santa Cruz (x 1). Ficure 121. P. hadesia Schaus, female, Punta Suarez, Espafola (1). Ficurre 122. Epidromia zephyritis Schaus, male, Santa Cruz (<1). Ficure 123. E. zephyritis Schaus, female, Santa Cruz (x1). Ficure 124. Anticarsia prona (Moschler), male, Isabela ( 1). FicurE 125. A. prona (Moschler), female, Isabela (X 1). FiGuRE 126. A. gemmatalis Hiibner, male, Santa Cruz (1). FicureE 127. A. gemmatalis Hiibner, male, Santa Cruz (X 1). Ficure 128.4. gemmatalis Hiibner, female, Santa Cruz (xX 1).
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Baltra, April, June; Isabela, March, May; Floreana, no month; San Cristobal, February, April; Santa Cruz, February—April; Santiago, April. AMNH, BMNH, CAS, MCZ, USNM, USC. |
Biotocy. Abundant some years, particularly from February to May. Di- urnal, coming especially to flowers of Clerodendrum molle and Cordia species. Foodplants. Cordia leucophlyctis, Tournefortia psilostachya, Heliotropium angiospermum, Lantana peduncularis, Mentzelia aspera. Larva. Smooth; pale green with thin white lines. Pupa. Greenish, thin-walled in flimsy cocoon in coiled underside of leaf of host plant.
OPHIDERINAE
Melipotis acontioides producta Hayes, new subspecies. (Figures 101 & 102.)
Galapagos specimens can be separated by the extended marginal band on the posterior margin of the hindwing.
DISTRIBUTION. Endemic subspecies of the widespread neotropical species.
Hototypr. Male. 26 mm., Santa Cruz, February 1970, R. Perry and Tj. de Vries, BM 1970-170.
PaRATYPES. Espafola, April; Floreana, April; Isabela, March; Santa Cruz, January-March, August-November. AMNH (31 specimens), BMNH (27 specimens), CAS (63 specimens), IRSNB (7 specimens), MCZ (25 specimens), RSM (2 specimens).
BioLocy. Foodplant. Parkinsonia aculeata. Larva. A pattern of gray- black markings outlined with white; some reddish brown on side. The larvae have the habit of lying pressed to the central part of the leaf.
Melipotis indomita (Walker).
(Figures 103 & 104.)
Bolina indomita WALKER, [1858] 1857, List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, vol. 13, p. 1161. Holotype, female (examined): Brazil; BMNH.
Sexually dimorphic. This insect has recently reached Hawaii.
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Baltra, April; Espanola, April; Floreana, January, February—July, October; Genovesa, April; Isabela, March, April, August; Pinta, October; Pinzon, April, June; San Cristobal, February, April, July; Santa Cruz, January—June, August, October, December; Santa Fé, April, July; Santiago, March, July. AMNH, BMNH, CAS, IRSNB, MCZ, USNM, ZSBS, USC.
BroLtocy. A common species, found in all months except at prolonged dry periods; adults come to flowers at dusk in the rainy season. Foodplant. Prosopis jubiflora. Larva. Head shiny brown. Body greenish white with red and gray- brown markings; more reddish laterally; underside pale, unmarked. Larvae are
VoL. XL] HAYES: GALAPAGOS MOTHS 191
found during the day in litter or under rocks at the base of the plants. Pupa. Pupation takes place in a loose cocoon of particles of soil and litter.
Melipotis harrisoni Schaus. (Figures 105 & 106.) Melipotis harrisoni Scuaus, 1923, Zoologica, vol. 5, p. 42. Lectotype, male (examined):
Galapagos: South Seymour [Baltra]; USNM.
Sexually dimorphic. The prominent white spots on the hindwing separate this species from M. indomita.
DIsTRIBUTION. Endemic species. Galdpagos Islands: Baltra, January, Feb- ruary, April, November; Floreana, April; Isabela, January—April, August; Pinzon, April, December; Rabida, June; San Cristobal, February; Santa Cruz, January—October, December; Santiago, January, July. AMNH, BMNH, CAS, IRSNB, LACM, MCZ, RSM, USNM, ZSBS, USC.
BroLtocy. Foodplants. Acacia macracantha, Acacia rorudiana. Larva. Head gray bordered with black. Body pale greenish gray with darker markings; some purplish suffusion.
Ascalapha odorata (Linnaeus). (Figures 114 & 115.) Phalaena (Bombyx) odorata LinNAEUS, 1758, Systema Naturae (10th Ed.), vol. 1, p. 505.
Type material (examined): America; LS.
Sexually dimorphic. Has been seen on some of the smaller and drier islands such as Genovesa and Pinzon. Generally seen at dusk. I believe this to be the species mentioned by Nelson (1968) as a saturniid. Galapagos specimens are usually smaller than those from the mainland.
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Floreana, April, May; San Cristébal, April; Santa Cruz, March—June, October. AMNH, BMNH, CAS, MCZ, USC.
BroLtocy. No data available. Tietz lists Acacia and Cassia among its food- plants in the U.S.A.
Letis mycerina (Cramer).
(Figures 116 & 117.)
Phalaena (Attacus) mycerina CRAMER, 1777, Uitlandsche Kapellen, vol. 2, p. 115, pl. 172, fig. B. Holotype, female: Surinam; not traced.
A fairly large, sexually dimorphic noctuid which may have been dismissed as A. odorata previously. I have illustrated a male from Panama since this species is only represented from the Islands in the BMNH by a female.
DisTRIBUTION. Neotropical species. Galdpagos Islands: Santa Cruz, April, June. AMNH, BMNH.
Biotocy. No data available.
192 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Rivula asteria (Druce), new combination. (Figure 144.)
Thalpochares asteria DrucE, 1898, Biologia Centrali-Americana, Heterocera, vol. 2, p. 497, pl. 95, fig. 25. Holotype, female (examined): Teapa, Mexico; BMNH.
Rivula ? dubiosa ScHAus, 1923, Zoologica, vol. 5, p. 44. New synonym. Lectotype, female (examined): Galapagos, Indefatigable [Santa Cruz]; USNM.
I have illustrated a male of this species which is rare in collections. Galapagos material J have examined has been in poor condition. The genus Rivula is used provisionally until revisionary work is undertaken.
DISTRIBUTION. Mexico, Bolivia, Galapagos archipelago. Galdpagos Islands: Santa Cruz, March, April, June. AMNH, BMNH, CAS.
BioLocy. No data available.
Glympis toddi Hayes, new species. (Figures 140 & 141.)
DESCRIPTION. MALE 12 mm. Antenna simple. Palpus gray brown. Slight concavity in costal margin of forewing. Forewing narrow, fuscous, and darker brown. Antemedial band angled. Postmedial band well-defined. Undersurface orange, yellow, and fuscous. Postmedial line of hindwing prominent orange. Abdominal tufts reduced.
Female 14 mm. Broader winged but similar to male.
Both sexes highly variable in forewing maculation. Differs from the nearest species Glympis incusalis Grote by the differing forewing maculation and more fuscous hindwing.
DISTRIBUTION. Endemic species.
Hototyre. Male. Floreana, Asilo de la Paz, 360 m., January 1971.
PaRATYPES. Floreana, April; Isabela, March, September; Pinta, May, Sep- tember; Santa Cruz, January-March, May—June, August-September. AMNH (4 specimens), BMNH (133 specimens), CAS (86 specimens), IRSNB (11 specimens), MCZ (7 specimens), RSM (5 specimens), ZSBS (8 specimens).
BioLocy. No data available.
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FicurE 129. Anomis editrix (Guenée), male, Floreana (XX 2). Ficure 130. A. illita (Guenée), male, Santa Cruz (x2). Ficure 131. A. luridula professorum Schaus, male, Isabela (X 2). FicuRE 132. A. erosa Hiibner, male, Floreana (xX 2). FicuRE 133. Bendis formularis Geyer, male, Mexico (xX 2). Ficure 134. B. formularis Geyer, female, Santa Cruz (X 2). Ficure 135. Gonodonta fulvangula Geyer, female, Santa Cruz (x 2; AMNH). FIcuRE 136. G. biarmata evadens Walker, male, Santa Cruz (X 2). FicuRE 137. G. biarmata evadens Walker, female, Santa Cruz (xX 2).
Vor. XL] HAYES: GALAPAGOS MOTHS 193
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Anomis editrix (Guenée). (Figure 129.) Gonitis editrix GUENEE, 1852, in Boisduval & Guenée, Histoire Naturelle des Insectes.
Lépidoptéres, vol. 6, p. 404. Type material (examined): Haiti; BMNH.
One of four species in this genus found on the Galapagos Islands. Lacks yellow coloration but possesses an angled margin to the forewing. Variable species.
DISTRIBUTION. Widespread in the neotropical region. Galdpagos Islands: Floreana, January; Santa Cruz, June. AMNH, BMNH.
Biotocy. No data available.
Anomis illita Guenée. (Figure 130.)
Anomis illita GUENEE, 1852, in Boisduval & Guenée, Histoire Naturelle des Insectes. Lépidop-
teres, vol. 6, p. 400. Lectotype, female: Brazil; MNHN.
Possesses the straightest forewing margin of the four species of Anomis found on the Galapagos Islands.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Santa Cruz, March-July. AMNH, BMNH, CAS.
Brotocy. Foodplant. Hibiscus tiliaceus. Larva. Head green, spotted black. Body pale, purplish brown; anal plate green. The single larva found was feed- ing on the flower of the above hostplant.
Anomis luridula professorum Schaus, new status. (Figure 131.)
Anomis professorum ScHAUS, 1923, Zoologica, vol. 5, p. 42. Lectotype, male (examined):
Galapagos: Chatham [San Cristébal]; USNM.
Variable species. I am reducing ‘professorum’ to subspecific status. Genitalia match those of mainland specimens but Galapagos specimens are smaller and less strongly marked.
DISTRIBUTION. Endemic subspecies of the widespread neotropical species. Galdpagos Islands: Baltra, April; Floreana, March, August; Genovesa, April; Isabela, February—April, August; Rabida, June; San Cristobal, February, April; Santa Cruz, February, April—July; Santiago, April. AMNH, BMNH, CAS, LACM, USNM, ZSBS.
BioLtocy. No data available.
Anomis erosa Hibner. (Figure 132.)
Anomis erosa HUBNER, 1823, Zutrage zur Sammlung Exotischer Schmetterlinge, vol. 2, p. 19, figs. 287-288. Type female: [U.S.A.] Savannah; not traced.
VoL. XL] HAYES: GALAPAGOS MOTHS 195
Forewing yellow and gray with angled margin.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Floreana, January; Santa Cruz, March, June. AMNH, BMNH, CAS, MCZ, USC.
BioLtocy. No data available. Foodplants in the U.S.A. are members of Malvaceae.
Plusiodonta clavifera (Walker). (Figures 138 & 139.) Tafalla clavifera WALKER, 1869, Characters of undescribed Lepidoptera Heterocera, p. 43. Type material (examined): [Honduras] Limas; BMNH. Sexually dimorphic. DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Santa Cruz, February—-May, September-November. BMNH, CAS, ZSBS. BroLocy. No data available.
Gonodonta biarmata evadens Walker, new status.
(Figures 136 & 137.)
Gonodonta evadens WALKER, [1858] 1857, List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, vol. 12, p. 955. Lectotype, female (examined): [Galapagos] ‘W. Coast of America’; BMNH.
Gonodonta biarmata galapagensis Topp, 1959, Tech. Bull. Agric. Res. Serv. U.S. Dept. Agric., no. 1201, p. 20, new synonym. Holotype, male (examined): Galapagos, Inde- fatigable [Santa Cruz]; AMNH.
Facts relating to the origin of the two female specimens studied by Walker, labelled ‘““W. Coast of America,” (already mentioned under Psaphara interclusa,) strongly suggested they were taken on the Galapagos Islands. This is confirmed as they match Galapagos specimens.
DISTRIBUTION. Endemic subspecies of the widespread neotropical species. Galdpagos Islands: Fernandina, February; Floreana, no month; Isabela, March; Pinta, October; San Cristobal, January, April, June; Santa Cruz, January— July, November, December. AMNH, BMNH, CAS, MCZ, MRAC, USNM, USC.
BioLocy. The adults of some species of Gonodonta have caused injury to citrus crops by piercing the fruit. Todd (1959, p. 19) gives foodplants of nominate subspecies.
Gonodonta fulvangula Geyer. (Figure 135.) Gonodonta fulvangula GrveEr, 1832, in Hiibner, Zutrage zur Sammlung Exotischer Schmetter- linge, vol. 4, p. 32, figs. 737, 738. Type material: Monte Video; not traced. Only one specimen has so far been taken. DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Santa Cruz, Horneman Farm, 200 m., 27 June 1965, Mrs. J. DeRoy, 1 female, AMNH.
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BroLocy. No data available. Todd’s revision of this genus quotes an 1882 reference to Araticu as the foodplant. This name refers to species of Annona and possibly to A. montana.
Metallata absumens contiguata Hayes, new subspecies. (Figures 145 & 146.)
I am separating the Galapagos population of this extremely variable insect as a distinct subspecies. In each specimen examined the reniform spot on the forewing is fused or in very close proximity to the postmedial fascia. This is in direct contrast to mainland specimens where the two markings are always sepa- rated by at least 1 mm.
DISTRIBUTION. Endemic subspecies of the species widespread in the neo- tropical region.
HototyPe. Male. 14.5 mm. Floreana, Asilo de la Paz, 360 m., January 1971. BMNH.
ParaTyPEs. Floreana, April; Isabela, August; Marchena, November; San Cristébal, April; Santa Cruz, February—July, September, October. AMNH (16 specimens), BMNH (15 specimens), CAS (31 specimens), IRSNB (1 speci- men), MCZ (1 specimen), ZSBS (14 specimens).
Brotocy. No data available.
Bendis formularis Geyer. (Figures 133 & 134.)
Bendis formularis GEYER, 1837, in Hubner, Zutrage zur Sammlung Exotischer Schmetter- linge, vol. 5, p. 26, figs. 903-904. Type female: Brazil; not traced.
I have seen only three specimens from the Galapagos Islands.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Floreana, no month; Santa Cruz, June, September. AMNH, BMNH, USC.
Biotocy. No data available.
Anticarsia gemmatalis Hiibner. (Figures 126-128.)
Anticarsia gemmatalis Htpner, 1818, Zutrage zur Sammlung Exotischer Schmetterlinge, vol. 1, p. 26, figs. 153-154. Type material: Surinam; not traced.
An extremely variable, common species.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Baltra, March, April; Fernandina, April; Floreana, February; Genovesa, March, April; Isabela, March—May; Pinta, October; San Cristobal, February, April; Santa Cruz, January—July, October. AMNH, BMNH, CAS, IRSNB, MCZ, USNM, ZSBS.
BroLocy. Found in the arid and humid zones of the main islands; abundant
Vor. XL] HAYES: GALAPAGOS MOTHS 197
at times; diurnal to an extent. Foodplants. Cryptocarpus pyriformis, Piscidia carthagenensis, Rhynchosia minima. Larva. Green, paler lines edged with black. In the U.S.A. this species is known as the Velvetbean Caterpillar. It feeds on a wide range of plants.
Anticarsia prona (Moéschler). (Figures 124 & 125.)
Thermesia prona MOscu er, 1880, Verhandlungen der Zoologisch-botanischen Gesellschaft in Wien, vol. 30 (Abh.), p. 443. Type material: [Panama] Chiriqui; [Venezuela] Puerto Cabello; Surinam, Paramaribo; not traced.
Sexually dimorphic.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Isabela, January, August; Santa Cruz, February—July. AMNH, BMNH, CAS, ZSBS.
Brotocy. No data available.
Psorya hadesia Schaus. (Figures 119-121.)
Pa
Psorya hadesia Scuaus, 1923, Zoologica, vol. 5, p. 44. Lectotype, female (examined): Galapagos: South Seymour [Baltra]; USNM.
A single female in the BMNH from Punta Suarez, Espanola (fig. 121) differs in its lighter brown and white coloration. Introduced goats are destroying the recorded foodplant on Espanola and this interesting form may well be en- dangered.
DISTRIBUTION. Endemic species. Galdpagos Islands: Baltra, April, Septem- ber; Espafiola, May; Floreana, April, May, July; Isabela, March, April, August; Pinzén, December; Santa Cruz, January—December; Santiago, July. AMNH, BMNH, CAS, IRSNB, MCZ, RSM, USNM, ZSBS.
BroLocy. Foodplant. Maytenus octogona. Larva. Head and body uniform green.
Epidromia zetophora Guenée. (Figure 118.)
Epidromia zetophora GUENEE, 1852, in Boisduval & Guenée, Histoire Naturelle des Insectes.
Lépidoptéres, vol. 7, p. 326, pl. 23, fig. 5. Holotype, male: Brazil; not traced.
Variable species. Only one male of this species has been taken on the Islands. It was collected in 1932 by M. Willows, Jr. of the Templeton-Crocker Expedition.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Isabela, May. CAS.
BroLtocy. No data available.
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Epidromia zephyritis Schaus. (Figures 122 & 123.)
Epidromia zephyritis ScHAuS, 1923, Zoologica, vol. 5, p. 43. Holotype, female (examined) : Galapagos: Indefatigable [Santa Cruz]; USNM.
Another variable species.
DISTRIBUTION. Endemic species. Galdpagos Islands: Baltra, no month; Floreana, April, July; Isabela, January, March, July-September; Pinzon, April; San Cristébal, April, October; Santa Cruz, January—December; Santiago, March, July. AMNH, BMNH, CAS, IRSNB, MCZ, RSM, USNM, ZSBS.
Brotocy. Arid and coastal areas where its foodplant occurs. Foodplant. Scutia pauciflora. Larvae collected on the mangrove Laguncularia racemosa, at Punta Espinosa, Fernandina in September 1970, although not reared, have been referred to this species. Larva. Whitish with extensive gray and brown markings.
HYPENINAE
Sorygaza variata Hayes, new species. (Figures 142, 143, 164, & 165.)
DescripTIoN. Male 10 mm. Antenna simple. Palpus prominent, clothed with buff and dark brown scales. Forewing cream, orange-brown, and darker brown. Postmedial band and antemedial band crenulate. Darker scaling of subterminal line forming cresent at apex. Prominent dark spotting forming adterminal line. Genitalia figured.
Female 9.5 mm. Broader winged but similar to male.
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Ficure 138. Plusiodonta clavifera (Walker), male, Santa Cruz (x 2; CAS). Ficure 139. P. clavifera (Walker), female, Santa Cruz (x 2). FicuRE 140. Glympis toddi Hayes, new species, holotype, male, Floreana (XX 2). FicurRE 141. G. toddi Hayes, new species, paratype, female, Floreana (x 2). FicurE 142. Sorygaza variata Hayes, new species, holo- type, male, Santa Cruz (xX 2; CAS). Ficure 143. S. variata Hayes, new species, paratype, female, Isabela (<2). Ficure 144. Rivula asteria (Druce), male, Santa Cruz (X 2; CAS). Ficure 145. Metallata absumens contiguata Hayes, new subspecies, holotype, male, Floreana (x 2). FicurE 146. M. absumens contiguata Hayes, new subspecies, paratype, male (x 2). Ficure 147. Hypena microfuliginea Hayes, new species, holotype, male, Isabela (X 2). Ficure 148. H. microfuliginea Hayes, new species, paratype, female, Floreana (X 2). Ficure 149. Peliala fuliginea Hayes, new species, holotype, male, Pinta (X 2). FicuRE 150. P. fuliginea Hayes, new species, paratype, female, Pinta (x2). Ficure 151. Hypena vetustalis (Guenée), female, Floreana (> 2). Figure 152. Ophiuche lividalis (Hiibner), male, Floreana ( 2; CAS). Ficure 153. O. minualis constans Hayes, new subspecies, holo- type, male, Santa Cruz (Xx 2; CAS). Ficure 154. O. minualis constans Hayes, new sub- species, paratype, female, Pinta (x 2).
VoL. XL] HAYES: GALAPAGOS MOTHS 199
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[55 156
157 wes lige
Ficure 155. Magusa erema Hayes, new species, paratype, male, genitalia. FIGURE 156. Trachea cavagnaroi Hayes, new species, holotype, male, genitalia (CAS). Ficure 157. Hypena microfuliginea Hayes, new species, paratype, male, genitalia. Ficure 158. Ozarba consternans Hayes, new species, paratype, male, genitalia (CAS).
Vor. XL] HAYES: GALAPAGOS MOTHS 201
Variable species.
Nearest species is Sorygaza didymata Walker. The forewing of S. variata has a more evenly rounded postmedial band and is a more strongly colored species. The diagnostic dorsal process of the male genitalia (see figure) also separates S. variata.
DISTRIBUTION. Endemic species.
Hototyre. Male. Santa Cruz, Horneman Farm, 220 m., 5 April 1964, D. Q. Cavagnaro. CAS.
ParaAtypPEs. Isabela, January, August; Santa Cruz, April, May. BMNH (7 specimens), CAS (15 specimens).
BroLocy. No data available.
Hypena vetustalis (Guenée). (Figure 151.)
Bomolocha vetustalis GUENEE, 1854, in Boisduval & Guenée, Histoire Naturelle des Insectes. Lépidoptéres, vol. 8, p. 35. Holotype, female (examined): Haiti; BMNH.
Known only from female specimens. Further material may prove a species- complex is involved.
DISTRIBUTION. Widespread neotropical species. Galdpagos Islands: Floreana, January; Isabela, August; Santa Cruz, May, June, July. AMNH, BMNH.
Biotocy. No data available.
Hypena microfuliginea Hayes, new species. (Figures 147, 148, & 157.)
Description. Male 13.5 mm. Antenna simple. Palpus prominent, heavily scaled. Midbrown forewing with prominent orbicular spot. Postmedial fascia with two rounded extensions terminad. Hindwing drab brown.
Female 14 mm. Broader winged but similar to male.
Nearest species is Hypena vetustalis Guenée but H. microfuliginea can also be confused with the larger Peliala fuliginea. However, H. vetustalis has a straight median line. Palpus structure and male genitalia also separate H. microfuliginea from P. fuliginea.
>
Ficures 159 and 160. Rivula asteria (Druce) male genitalia (CAS). Ficurres 161 and 162. Elaphria encantada Hayes, new species, paratype, male, genitalia. FicuRE 163. Psaphara interclusa Walker, holotype, male, genitalia.
Ficures 164 and 165. Sorygaza variata Hayes, new species, holotype, male, genitalia (CAS). Ficures 166 and 167. Peliala fuliginea Hayes, new species, paratype, male, genitalia. Ficures 168 and 169. Epiplema becki Hayes, new species, holotype, male, genitalia (CAS). FicurEe 170. E. becki Hayes, new species, paratype, female, genitalia.
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Vor. XL]
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DISTRIBUTION. Endemic species.
HototyPe. Male. Isabela (Albemarle) 200 ft., 7 August 1924, St. George Expedition, C. L. Collenette. BMNH.
PaRATYPES. Floreana, January, April; Isabela, August; Santiago, July. BMNH (3 specimens), ZSBS (1 specimen).
Brotocy. No data available.
Peliala fuliginea Hayes, new species. (Figures 149, 150, 166, & 167.)
DescripTION. Male 16 mm. Antenna simple. Palpus prominent, thinly scaled. Tornus of forewing less strongly angulate, rounded. Postmedial fascia with two rounded extensions terminad. Dark brown coloration throughout with prominent orbicular spot. Hindwing midbrown. Genitalia figured.
Female 15 mm. Broader winged but similar to male. Somewhat variable in maculation.
Peliala fuliginea has rounded extensions on the postmedial fascia of the forewing which separates it from Hypena vetustalis. Palpus structure and genitalia also separate P. fuliginea from H. microfuliginea.
DISTRIBUTION. Endemic species.
HototypPe. Male. Pinta, 630 m., November 1970, R. Perry & Tj. de Vries. BMNH.
ParaATYPES. Floreana, January; Isabela, April, August, September; James, July; Pinta, October, November; Santa Cruz, February—August, December. AMNH (49 specimens), BMNH (28 specimens), CAS (73 specimens), ZSBS (4 specimens).
Brotocy. No data available.
Ophiuche lividalis (Hiibner). (Figure 152.) Pyralis lividalis HUBNeER, 1790, Beitrage zur Geschichte der Schmetterlinge, vol. 2, no. 4, pp. 86-87, pl. (4) 1, fig. E. Holotype: Italy, Florence; not traced. Based on a specimen in the CAS and a worn example in the ZSBS. DISTRIBUTION. Almost cosmopolitan. Galdpagos Islands: Floreana, April. CAS, ZSBS. BioLocy. No data available.
>
Ficures 171 and 172. Utetheisa perryi Hayes, new species, paratype, male, genitalia. Ficures 173 and 174. Agrotis consternans Hayes, new species, holotype, male, genitalia. Ficures 175 and 176. Utetheisa devriesi Hayes, new species, paratype, male, genitalia.
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Ophiuche minualis constans Hayes, new subspecies. (Figures 153 & 154.)
The median line on the forewing of Galapagos specimens is always straighter than that of mainland specimens.
DISTRIBUTION. Endemic subspecies of the neotropical species.
Hototyre. Male. 8 mm. Santa Cruz, Horneman Farm, 220 m., 5 March 1964, D. Q. Cavagnaro. CAS.
ParatyPes. Floreana, January, April; Isabela, September; Pinta, October, November; Santa Cruz, February—June, October. AMNH (6 specimens), BMNH (49 specimens), CAS (32 specimens), MCZ (1 specimen), MRAC (1 specimen), ZSBS (1 specimen).
Briotocy. No data available.
BIBLIOGRAPHY
ARNAUD, P. H. 1970. List of the scientific publications and insect taxa described by Francis Xavier Williams (1882-1967). Occasional Papers of the California Academy of Sciences, no. 80, pp. 1-33. BEEBE, W. 1923a. Williams Galapagos expedition. Zoologica, N.Y., vol. 5, pp. 1-20, 10 figs. 1 map. 1923b. Notes on Galapagos Lepidoptera. Zoologica, N.Y., vol. 5, pp. 51-59. Bowman, R. I. 1966. The Galapagos. Proceedings of the Galapagos International Scientific Project of 1964. University of California Press, 318 pp. Brower, D. 1968. Galapagos. The Flow of Wildness. 2 volumes. Sierra Club. Pp. 158 & 160. Butter, A. G. 1877. Account of the zoological collection made during the visit of H.M.S. “Peterel” to to the Galapagos Islands. X. Lepidoptera, Orthoptera, Hemiptera. Proceedings of the Zoological Society of London, pp. 86-91. CHAMPION, G. C. 1924. The insects of the Galapagos Islands. The Entomologist’s Monthly Magazine, vol. 60, pp. 259-260. Crark, B. P. 1926. A revision of the Protoparce of the Galapagos Islands. Proceedings of the New England Zoological Club, vol. 9, pp. 67-71. Conway, A, AND F. Conway 1948. The Enchanted Islands. Geoffrey Bles, London, 254 pp. Curio, E. 1965. Die Schutzanpassungen dreier Raupen eines Schwarmers (Lepidopt., Sphingidae) auf Galapagos. Zoologische Jahrbiicher. Abteilung fiir Systematik, Bd. 92, pp. 487-522, pls. Darwin, C. 1859. Origin of Species. John Murray, London, ix + 502 pp., 1 pl. Ecxuarnt, R. C. 1972. Introduced plants and animals in the Galapagos Islands. BioScience, vol. 22, pp. 585-590, figs.
VoL. XL] HAYES: GALAPAGOS MOTHS 207
ErsL-EIBeEsFELptT, I. 1960. The Noah’s Ark of the Pacific. Macgibbon & Kee, London, 192 pp. Forzes, W. T. M. 1941. The position of Utetheisa galapagensis (Lepidoptera, Arctiidae). Journal of the New York Entomological Society, vol. 49, pp. 101-110. Hopces, R. W. 1971. In Dominick, R. B., et al., The moths of America north of Mexico, fasc. 21, Sphingoidea. E. W. Classey Limited & R.B.D. Publications Inc., 14 pls., 158 + xii pp. Howarp, L. O. 1890. Scientific results of explorations by Steamer Albatross, no. 5. Annotated catalogue of the insects collected in 1877—’88. Proceedings of the United States Na- tional Museum, vol. 12, pp. 185-216 (1889). JorpDAN, K. 1939. On the constancy and variability of the differences between the old world species of Utetheisa (Lepid.; Arctiidae) Novitates Zoologicae, vol. xli, pp. 251-291, figs. KERNBACH, K. 1962. Die Schwarmer einiger Galapagos-Inseln (Lep. Sphingidae). Opuscula Zoologica, Minchen, vol. 63, pp. 1-19, abb. 1-12. 1964. Die Schwarmer einiger Galapagos-Inseln (II) (Lep. Sphingidae). Mitteilungen der Deutschen Entomologischen Gesellschaft, vol. 23, p. 88. KIMBALL, C. P. 1965. The Lepidoptera of Florida. Arthropods of Florida and neighboring land areas, vole eps 363: LinsteEy, E. G., anp R. L. USINGER 1966. Insects of the Galapagos Islands. Proceedings of the California Academy of Sciences, vol. 33, no. 7, pp. 113-196. McDunnoucu, J. 1938. Check list of the Lepidoptera of Canada and the United States of America, pt. 1, Macrolepidoptera. Memoirs of the Southern California Academy of Sciences, vol. 1, pp. 1-272. Mutter, R. C. 1965. Islands of the Tortoise. Pacific Discovery, vol. 18, no. 5, pp. 1-2. (this issue of Pacific Discovery is devoted entirely to the Galapagos Islands.) NELSON, B. 1968. Galapagos: islands of birds. Longmans, Green, London, 338 pp. Nve, I. W. B. 1975. Generic names of moths of the world, vol. 1. Noctuoidea (part). British Mu- seum (Natural History), 568 pp., 1 pl. Parkin, P., D. T. Parkin, A. W. Ewinc, ann H. A. Forp 1972. A Report on the Arthropods collected by the Edinburgh University Galapagos Islands Expedition, 1968. Pan-Pacific Entomologist, vol. 48, no. 2, pp. 100-107. Perry, R. 1969. Conservation problems in the Galapagos Islands. Micronesica, vol. 5, no. 2, pp. 275-281. 1972. The Galapagos Islands. Dodd, Mead & Co. New York, 92 pp. Ricuwarps, A. G. 1941. The Noctuid moths of the Galapagos from the collections of the Allan Hancock Foundation. Report of the Allan Hancock Pacific Expedition, vol. 5, pp. 233- 251, 4 pls.
208 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.
Rinpce, F. H.
1973. The Geometridae (Lepidoptera) of the Galapagos Islands. American Museum
Novitates, no. 2510, pp. 1-31. ROTHSCHILD, W., AND K. JORDAN
1903. A revision of the lepidopterous family Sphingidae. Novitates Zoologicae, vol. IX,
supplement, cxxxv + 972 pp., 67 pls. ScHaus, W.
1923. Galapagos Heterocera with descriptions of new species. Zoologica, vol. 5, pp.
21-48. SEEMANN, B.
1853. Narrative of the voyage of H.M.S. Herald during the years 1845-1851 under command of Captain Henry Kellett, R.N., being a circumnavigation of the globe and three cruises to the arctic in search of Sir John Franklin. Reeve & Co. London, 2 vols., 322 and 302 pp.
SLEVIN, J.R.
1959. The Galapagos Islands. A history of their exploration. Occasional Papers of
the California Academy of Sciences, vol. 25, pp. 1-150. Tre1z, H. M.
1972. An Index to the described life histories, early stages and hosts of the Macro- lepidoptera of the Continental United States and Canada. 2 vols. The Allyn Museum of Entomology, Sarasota, Florida, 1041 pp.
Topp, E. L.
1959. The fruit piercing moths of the genus Gonodonta Hiibner (Lepidoptera, Noctuidae). Technical Bulletin of the United States Department of Agriculture, no. 1201, pp. 1-52, 12 pls.
1972a. Descriptive and synonymical notes for some species of Noctuidae from the Galapagos Islands (Lepidoptera). Journal of the Washington Academy of Sciences, vol. 62, no. 1, pp. 36-40, figs.
1972b. A note on the generic transfer of “Catabena” esula (Druce) and new synonymy (Lepidoptera : Noctuidae). Proceedings of the Entomological Society of Wash- ington, vol. 74, no. 2, p. 260.
1973. The types of some noctuid moths from the Galapagos Islands described by William Schaus in 1923. Proceedings of the Entomological Society of Wash- ington, vol. 75, no. 1, pp. 33-39.
Wiccins, I. L., anp D. M. Porter. 1971. Flora of the Galapagos Islands. Stanford University Press, 998 + xx pp., figs. WitiaMs, F. X.
1911. Expedition of the California Academy of Sciences to the Galapagos Islands 1905-1906. III. The butterflies and hawk-moths of the Galapagos Islands. Pro- ceedings of the California Academy of Sciences, vol. 1, pp. 289-322, 2 pls., 1 in color.
ZIMMERMAN, E. C.
1958. Macrolepidoptera. Insects of Hawaii, Honolulu, vol. 7, 542 pp.
1969. Francis Xavier Williams. Pan-Pacific Entomologist, vol. 45, pp. 135-146. ZIMSEN, E.
1964. The Type Material of I. C. Fabricius. Munksgaard. Copenhagen, 656 pp.
PROCEEDINGS |
OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES so aes
Vol. XL, No. 8, pp. 209-230; 7 figs. August 8, 1975
NEW SPECIES AND NEW COMBINATIONS OF FERNS FROM CHIAPAS, MEXICO
By Alan Reid Smith
Herbarium, Department of Botany, University of California, Berkeley, California 94720
In preparation for a pteridophyte flora of the state of Chiapas, Mexico, I find it necessary to describe the following new species and make several new combinations. Most of the new species were discovered after intensive field investigations by Dr. Dennis Breedlove and myself; these explorations have resulted in approximately 4000 new collections of pteridophytes, many from previously uncollected or poorly collected areas of Chiapas. In circumscribing the new species I have relied most heavily on herbarium holdings of Mexican and Central American ferns in Dudley Herbarium, Stanford University (DS), University of California Herbarium (UC), United States National Museum (US), New York Botanical Garden (NY), Philadelphia Academy of Science (PH), Field Museum of Natural History (F), and Universidad Nacional de México (MEXU). I thank curators of these herbaria for making their collec- tions available for study. Isotypes and duplicates of Breedlove collections will be distributed to MEXU, NY, and US. I also thank Charlotte Mentges for preparing the illustrations.
Asplenium breedlovei A. Reid Smith, sp. nov. (Figure 1, A-B.)
Plantae terrestres; rhizomata breve repentia, ca. 4 mm. diametro, frondibus paucis (1-3) praedita; stipites atropurpurei vel atrobrunnei, impoliti, glabri, ca. 2 mm. diametro, laminas fere aequantes; rhachides leviter flexuosae, basi fuscatae, apicem versus viridescentes; frondes basi fere quadripinnatae, usque 35 cm. longae, 15 cm. latae, deltoideae, apicem versus gradatim reductae, non
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proliferae; pinnae usque ca. 10-jugae, atrovirides, non articulatae, alternae, ascendentes, usque 10 cm. longae, 3 cm. latae, axibus virellis; segmenta penultima cuneiformia, maximam partem 2—5-loba; segmenta ultima usque 1.5 mm. lata, apice acuta vel subacuta; sori secus venas segmentorum ultimorum usque ca. 3 mm. longi; indusia integra vel leviter erosa, ca. 0.6 mm. lata, fulva.
Ho.iotype. Mexico. Chiapas: municipio Villa Corzo, east base of Cerro Tres Picos near Cerro Bola, SW. of Colonia Agronimos Mexicanos, 1500-1800 m., Breedlove 30041 (DS).
REMARKS. Superficially, this species resembles more dissected members of the A. radicans complex, but the stipe and rachis are dull, the lamina is deltoid, and the pinnae ascend at an angle of about 60° from a slightly flexuose, non- proliferous, non-flagelliform rachis. The relationship to A. cristatum Lamarck is perhaps closer, but the general architecture of the fronds and the creeping rhizomes of A. breedlovei adequately distinguish it from that species.
Asplenium munchii A. Reid Smith, sp. nov. (Figure 7, A-B.)
Plantae terrestres; rhizomata suberecta, basibus stipitum abscondita, cau- dices usque 1.3 cm. diametro; frondes numerosae, caespitosae, stipites 4-9 cm. longi, ca. 1 mm. diametro, brunnei vel plumbei, non lustrati, glabri; laminae tripinnatifidae, (17)25-30 cm. longae, (3.5)5-7 cm. latae, apicem versus attenuatae et proliferae; pinnae usque ca. 25-jugae, plerumque alternae, sessiles, maximae in medio, 4 cm. longae, 1.5 cm. latae, gradatim reductae sursum et deorsum (infimae deflexae), inaequilaterae, pinnulis grandioribus acroscopicis; pinnulae ad angulum 45-60° costis, maximae profunde 5-lobae; segmenta ultima ca. 1 mm. lata, apice rotundata vel acutiuscula; laminae membranaceae, glabrae; sori secus venas segmentorum ultimorum positi, usque 3 mm. longi, indusiis 0.6 mm. latis, integris, fulvis.
Ho.otyPe. Mexico. Chiapas: San Pablo, Miinch 114 (DS).
ParatyPes. Mexico. Chiapas: municipio Tenejapa, sumidero of Yochib, paraje Koltol Te’, Breedlove 6222 (DS); 4 mi. N. of Jitotol, Thorne & Lathrop 41783 (DS); municipio La Independencia, Las Margaritas to Campo Alegré, Breedlove 33648 (DS).
Remarks. Allied to A. sessilifolium Desvaux, but much more dissected than that species and occurring at lower elevations (1300-2300 m.). The
Ficure 1. A-B. Asplenium breedlovei, Breedlove 30041, DS: A, ultimate segment; B, plant. C-E. Asplenium soleirolioides, Breedlove & Smith 31820, DS: C, pinnae; D, frond apex; E, plant. F—G. Asplenium olivaceum, Hatch & Wilson 149, US: F, plant; G, pinnae.
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blades of A. munchii are similar in dissection to large fronds of A. myriophyllum (Swartz) Presl, which always lacks buds and has less oblique pinnules.
Asplenium olivaceum A. Reid Smith, sp. nov. (Figure 1, F-G.)
Rhizomata compacta, frondibus numerosis erectis vel arcuatis praedita; stipites nigrescentes, glabri, nitidi, ca. 0.6 mm. diametro, laminis 0.2—0.6 plo breviores; frondes usque 15 cm. longae, 1.5 cm. latae, non proliferae, segmento ovali terminali; pinnae usque 15-jugae, olivaceae, articulatae, oppositae, ob- longae, usque 8 mm. longae, 5 mm. latae, secus marginem superiorem integrae vel leviter undulatae, apice rotundatae; sori 1-2 per pinnam, 2—3 mm. longi, soro primario parallelo margini inferiori, interdum soro secundario secus venam acropetam; indusia plus minusve integra, usque 0.9 mm. lata, albida vel cinerascentia.
Hototype. Guatemala. Alta Verapaz: Senahu, summit of Cerro Sillab, limestone cliffs, 6000 ft. [1800 m.], Hatch & Wilson 149 (US).
Paratype. Mexico. Chiapas: municipio La Trinitaria, Lagos de Monte Bello, along Comitan River at its sumidero, limestone rocks, 1300 m., Breed- love & Smith 22379 in part (DS).
Remarks. Perhaps most closely related to 4. heterochroum Kunze, which was collected with the paratype cited. It can be distinguished easily from that species by the peculiar olivaceous color of the fronds, number and placement of the sori, and the oblong, entire (or nearly so) pinnae.
Asplenium soleirolioides A. Reid Smith, sp. nov. (Figure 1, C-E.)
Rhizomata minuta, radicibus paleisque abscondita, frondibus numerosis arcuatis vel decumbentibus praedita; stipites rhachidesque castaneae, glabrae, nitidae, ca. 0.3 mm. diametro; frondes usque 9 cm. longae, 5 mm. latae, apicem versus proliferae et radicantes; pinnae infra gemmam usque 15-jugae, sub- virides, articulatae, plerumque alternae, rotundatae vel irregulariter ovales, usque 2.5 mm. longae, 2 mm. latae, venis 2—3-jugis; sori 1-2 per pinnam, grandis- simi secus venam inferioram in quoquo segmenta; indusia integra, usque 0.6 mm. lata, fulva.
HototyPe. Mexico. Chiapas: municipio La Grandeza, 10 km. E. of El Porvenir along road from Huixtla to Siltepec, 2800 m., Breedlove & Smith 31820 (DS).
Remarks. Named after the monotypic genus Soleirolia (Urticaceae), which it resembles in habit and general shape of the segments.
I am unable to suggest close relatives for this peculiar species, but it is certainly a member of A. trichomanes group. The pinnae of A. soleirolioides are much thinner in texture than in most other species of this alliance.
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Ctenitis bullata A. Reid Smith, sp. nov. (Figure 2, E-H.)
Rhizomata suberecta, basibus stipitum et paleis abscondita, caudices ca. 2.5 cm. diametro; stipites usque ca. 40 cm. longi, 4 mm. diametro, brunneoli, basin versus paleis usque 1 cm. longis patentibus vel reflexis subulatis atrobrun- neis nitentibus; rhachides paleis similaribus sed brevioribus; laminae atrovirides, usque ca. 45 cm. longae, 25 cm. latae, basi bipinnatae vel tripinnatifidae, sursum profunde bipinnatifidae; pinnae usque ca. 20-jugae, alternae vel sub- oppositae, basi latissimae, aequilaterae, usque ca. 13 cm. longae, 3.3 cm. latae, usque 6 mm. petiolulatae, pinnae distales sessiles; pinnulae usque 1 mm. petiolulatae, serratae, pinnulae pinnarum distaliorum sessiles serratae vel in- tegrae; venae usque 10-jugae, simplices vel furcatae, margine terminatae; costae subtus paleis numerosis bullatis castaneis, pilis paucis longis (usque 0.7 mm.) articulatis, et pilis numerosis brevibus glandiferis (ca. 0.1 mm. longis) ; pagina laminarum super glandiferae, pilis articulatis costis venisque: pagina laminarum infra glabra; sori inframediales, ca. 1 mm. diametro; indusia persistentia, rotundato-reniformia, ca. 0.6—-0.8 mm. diametro, glandifera, fulva, margine erosa vel glanduloso-ciliata.
HototypPe. Mexico. Chiapas: municipio La Trinitaria, Lagos de Monte Bello, 1400 m., Breedlove 25339 (DS).
PaRATYPES. Same locality, 1600 m., Breedlove 14968 (DS); same locality, 1300 m., Breedlove & Smith 22331 (DS); same locality, Breedlove 38909 D'S):
Remarks. Most closely related to C. strigilosa (Davenport) Copeland, known from Veracruz (type) and Guatemala, but differing in its much larger fronds, eglandular tissue below, presence of long, articulate hairs on costae and costules below, larger indusia, and costal scales decidedly more bullate (scarcely bullate in C. strigilosa).
Ctenitis baulensis A. Reid Smith, sp. nov. (Figure 2, I-K.)
Rhizomata suberecta, basibus numerosis stipitum veterum abscondita, caudices 2—3 cm. diametro; frondes caespitosae, stipites usque 20 cm. longi, ca. 2 mm. diametro, basi fuscati, sursum straminei vel viridi-straminei, dense squamosi, paleis usque 3 mm. longis, 1.5 mm. latis, complanatis usque sub- bullatis, fulvis (apicem versus fuscatis); laminae pinnato-pinnatifidae, usque 20 cm. longae, 13 cm. latae; pinnae usque ca. 10-jugae, sessiles, maximae 7 cm. longae, 2.5 cm. latae, fere ad costas profunde incisae; segmenta obliqua, maxima 12 mm. longa, 4.5 mm. lata, apicem obtusum vel acutiusculum versus integra usque denticulata, approximata vel etiam imbricata; paria basalia segmentorum leviter reducta pinnis infimis; venae usque 9-jugae, marginem supra sinum attingentes; costae infra parce glandiferae, dense squamosae, paleis basi fulvis
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Ficure 2. A-D. Ctenitis thelypteroides: A, segment, Breedlove 33280, DS: B, lower pinna, Lundell 18115, US; C, rhizome and stipe bases, Breedlove 33280, DS; D, frond apex, Lundell 18115, US. E-H. Ctenitis bullata: E, rhizome, Breedlove & Smith 22331, DS; F,
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et manifeste bullatis, apice fuscatis et attenuatis, costae supra pilis densis Ctenitidis ca. 0.5 mm. longis; paginae laminarum utrinque glabrae vel infra pilis paucis appressis glandulosis; sori inframediales, exindusiati, usque 1 mm. diametro.
Hototyree. Mexico. Chiapas: municipio Cintalapa, 16 km. NW. of Rizo de Oro, SE. of Cerro Baul, 1600 m., Breedlove & Smith 21812 (DS).
PARATYPE. Same locality, Breedlove & Smith 31328 (DS).
ReMARKS. Most closely related to Dryopteris tonduzi (Christ) C. Chris- tensen, from Guatemala and Cost Rica, differing from that in the exindusiate, inframedial sori, approximate segments, and smaller, more decidedly bullate- scaly fronds. Also related to C. nigrovenia (Christ) Copeland, but differing in the more or less glabrous blade surfaces, inframedial sori, and approximate segments.
Ctenitis thelypteroides A. Reid Smith, sp. nov. (Figure 2, A-D.)
Rhizomata breve repentia vel oblique erecta, basibus stipitum maximam partem abscondita, usque 4 mm. diametro; stipites usque 28 cm. longi, 2 mm. diametro, basi paleis patentibus lanceolatis castaneis usque 5 mm. longis; rhachides paleis paucis similaribus; laminae pinnato-pinnatifidae, usque ca. 30 cm. longae, 16 cm. latae; pinnae usque 15-jugae, petiolulatae usque 2 mm., usque 9 cm. longae, 2 cm. latae, profunde incisae fere costis; segmenta obliqua ca. 3 mm. lata, apice acuta vel obtusa, serrulata, segmentis basilaribus quam segmentis distalibus leviter reductis et congestioribus; venae usque 8-jugae, infimae marginem 1—2.5 mm. supra sinum attingentes; costae infra glanduliferae, paleis dispersis lineari-lanceolatis castaneis non bullatis integris usque 1.5 mm. longis, supra pilis densis Ctenitidis usque 0.5 mm. longis; pagina laminarum infra glabra vel glandulis inconspicuis dispersis appressis tubularibus luteolis; sori usque 6-jugi, plerumque ad apices segmentorum limitati, 0.5-1.0 mm. diametro; indusia ca. 0.3 mm. diametro, fulva, persistentia.
Hototypre. Mexico. Chiapas: municipio Las Margaritas, eastern side of Laguna Miramar E. of San Quintin, 366 m., Breedlove 33280 (DS).
PaRATYPES. Mexico. Chiapas: jct. of Rio Perlas and Rio Jataté at San Quintin, 200 m., Sohns 1696 (US). Guatemala. Peten: ca. 4.5 mi. NEE. of Pucté on La Libertad trail, Lundell 18115 (US).
REMARKS. Closely related to C. nigrovenia (Christ) Copeland, but distin-
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segment, Breedlove 14968, DS; G, upper pinna, Breedlove 14968, DS; H, lowermost pinna, Breedlove 14968, DS. I-K. Ctenitis baulensis: I, lamina, Breedlove & Smith 21812, DS; J, rhizome and stipe bases, Breedlove & Smith 31328, DS; K, segment, Breedlove & Smith 21812, DS.
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guished by the smaller sori localized at the tips of the segments, the reduced basal pinna segments that are more crowded, and the habit of the rhizome.
Ctenitis ursina A. Reid Smith, sp. nov. (Figure 3, C—E.)
Rhizomata ignota (probabiliter suberecta, ampla); stipites 50 cm. vel plures longi, usque ca. 1 cm. diametro, paleis patentibus lanceolatis integris vel parce denticulatis lustratis brunneis usque ca. 1 cm. longis, 1 mm. latis perdense vestiti; rhachides, costae costulaeque similes stipitibus, sed paleis parvioribus et minor densis et coloratis pallidioribus, denique paleis bullatis costis costulisque; laminae quadripinnatifidae, usque 75 cm. longae, 70 cm. latae, deltoideae; pinnae ca. 10—12-jugae, usque ca. 37 cm. longae, 13 cm. latae, aequilaterae; pinnulae usque ca. 20-jugae per pinnam, usque 7 cm. longae, 1.5 cm. latae, apicem versus pinnarum adnatae decurrentes, basin versus leviter reductae; segmenta profunde serrata vel integra; venae prae margine terminatae; axes supra pilis densis Ctenitidis ca. 0.5 mm. longis; paginae utrinque laminarum glabrae; sori ca. 1 mm. diametro; indusia_ persistentia rotundato-reniformia fulva, in centro leviter fuscata, ad marginem erosa vel elanduloso-ciliata.
Horotype. Mexico. Chiapas: municipio La Trinitaria, E. of Laguna Tzikaw, Monte Bello National Park, 1300 m., Breedlove & Smith 32274 (DS —2 sheets).
PARATYPE. Same locality, Breedlove 35261 (DS).
Remarks. Most closely related to C. melanosticta (Kunze) Copeland, differing by the densely scaly, even shaggy, stipes and by the indusia not so distinctly bicolorous.
Dryopteris futura A. Reid Smith, sp. nov. (Figure 3, F-I.)
Rhizomata non visa (probabiliter erecta, crassa); stipites usque 45 cm. longi, 5 mm. lati, basin versus dense paleacei, paleis ovato-lanceolatis, usque 2.5 cm. longis, 5 mm. latis, integris, brunneis (sursum fulvis), concoloribus sublustratis; rhachides stramineae, glabrae vel parce stipitato-glandulosae; laminae infra tripinnato-pinnatifidae usque 4-pinnatae, usque ca. 40 cm. longae, 30 cm. latae, deltoideae; pinnae oppositae usque suboppositae, ca. 12-jugae, ad
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Ficure 3. A-B. Dryopteris munchii, Miinch 116, DS: A, lowermost pinna; B, ultimate segment. C—E. Ctenitis ursina, Breedlove ¢& Smith 33274, DS: C, ultimate segment; D. pinnule; E, stipe base. F-I. Dryopteris futura, Breedlove & Smith 32058, DS: F, lower- most pinna; G, ultimate segment; H, fourth pinna from base; I, stipe base.
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angulum 45—60° ascendentes, infimae deltoideae, usque 25 cm. longae, 15 cm. latae, petiolulatae usque 1.5 cm., basiscopicae, anadromae, ceterae catadromae sublanceolatae; pinnae secundariae suboppositae usque alternae, lanceolatae, leviter basiscopicae usque fere aequilaterae, infimae petiolulatae usque 5 mm.; pinnulae usque 12-jugae, pinnatae usque profunde pinnatifidae, segmentis ultimis denticulatis; laminae membranaceae, sine paleis, utrinque glabrae vel subtus stipitato-glandulosae (praesertim axibus); indusia orbiculari-reniformia, ca. 1 mm. diametro, persistentia, fulva, ad marginem stipitato-glandulosa; 1 = 4).
Hototype. Mexico. Chiapas: municipio El Porvenir, 3-4 km. W. of El Porvenir along road from Huixtla to Siltepec, 2800 m., Breedlove & Smith 31772 (DS):
PaRATYPE. Mexico. Chiapas: municipio San Cristobal Las Casas, Cerro Huitepec (Muk’ta vits), 2700 m., Breedlove & Smith 32058 (DS).
Remarks. This species can be compared only to D. nubigena Maxon & Morton, from which it differs by the more densely scaly stipe bases, strongly ascending pinnae, the larger, more persistent indusia, and the less glandular, thinner-textured blades.
The paratype cited showed 2” = 41 II at meiotic metaphase.
Dryopteris munchii A. Reid Smith, sp. nov. (Figure 3, A-B.)
Rhizomata ignota (probabiliter erecta); stipites usque 35 cm. longi, 5 mm. diametro, brunneoli, basin versus squamati, paleis anguste lanceolatis, usque ca. 10 mm. longis, 1(2) mm. latis, saepe bicoloribus, apice fulvis integris, basin versus denigratis dentatis, non lustratis; rhachides fulvae, eglandulosae, epilosae, paleis dispersis; laminae infra subtripinnatae, sursum_ bipinnato- pinnatifidae, ca. 45 cm. longae, 27 cm. latae; pinnae basi suboppositae, sursum alternae, ca. 16-jugae, ad angulum 60-80° ascendentes, infimae deltoideae, usque 15 cm. longae, 8.5 cm. latae, petiolulatae usque 6 mm., basiscopicae, ana- dromae, ceterae gradatim aequilaterae; pinnae secondariae oppositae usque alternae, patentes (ca. 90°), lanceolatae, fere aequilaterae, sessiles usque petiolulatae ca. 1 mm.; pinnulae usque ca. 10-jugae, apicem versus spinulosae; laminae chartaceae, in costis costulisque infra paleis minutis fulvis ovatis usque linearibus praeditae, alibi glabrae, eglandulosae; indusia orbiculari-reniformia, ca. 0.8 mm. diametro, persistentia, porphyrea, glabra.
Ho.otype. Mexico. Chiapas: Baduitz, Munch 116 (DS).
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Ficure 4. A. Elaphoglossum albomarginatum, plant, Breedlove & Smith 31629, DS. B-D. Elaphoglossum chiapense, Breedlove ¢& Smith 32649, DS: B, plant; C, stipe base scale; D, laminar scale.
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PaRATYPE. Mexico. Chiapas: municipio Tenejapa, paraje Banabil, 2800 m., Breedlove & Smith 22027 (DS).
RemMArRKS. Related to Dryopteris rossi C. Christensen from western Mexico (later treated by Christensen as a variety of D. patula), from which it differs by its larger, narrower, eglandular blades. From D. patula (Swartz) Under- wood it differs in the smaller, bicolorous stipe scales, spreading, nearly equi- lateral secondary pinnae, and eglandular blades.
Elaphoglossum albomarginatum A. Reid Smith, sp. nov. (Figure 4, A.)
Rhizomata breve repentia, crassa, caudices ca. 1 cm. diametro, dense paleacei; stipites ca. 25 cm. longi, 2.5 mm. diametro, straminei, non nisi basi squamati, paleis lanceolatis, usque 2.5 cm. longis, 2 mm. latis, integris, fulvis, glabris, ascendentibus et flexilibus; laminae steriles ovato-lanceolatae, ca. 30 cm. longae, 7 cm. latae, basi rotundatae, apice acutae usque acuminatae, mar- gine scarioso, albido, 0.8 mm. lato, chartaceae, utrinque glabrae; venae 1-2- furcatae plerumque liberae, prope apicem saepe bifurcatae, secus rhachim 3—4 mm. inter se distantes, prope marginem 1.3 mm. inter se distantes; laminae fertiles 13 cm. longae, 3.5 cm. latae, ambitu laminis sterilibus similes, margine albido-hyalino ca. 1.5 mm. lato.
Horotype. Mexico. Chiapas: municipio Union Juarez, SE. side of Vol. Tacana, above Talquian, 2100 m., Breedlove & Smith 31629 (DS).
Remarks. Related to E. latifolium (Swartz) J. Smith, but with a very broad scarious margin, wider-spaced veins, totally glabrous laminae, and very long, twisted stipe base and rhizome scales.
Elaphoglossum chiapense A. Reid Smith, sp. nov. (Figure 4, B—D.)
Rhizomata longe repentia (ca. 1 cm. inter bases stipitum), 2-3 mm. di- ametro, dense squamosa, paleis lanceolatis, 3-4 mm. longis, brunneis, lustratis, denticulatis, plerumque patentibus; stipites usque 15 cm. longi, 2 mm. diametro, straminei usque basin versus fuscati, paleis ca. 2 mm. longis, brunneis, paten- tibus; laminae steriles usque ca. 20 cm. longae, 3.5 cm. latae, apice acutae, basi cuneatae, chartaceae, utrinque in laminis, venis, rhachidibusque paleis ap- pressis, lanceolatis, 0.7-1.5 mm. longis e basi circulari, minute denticulatis, fulvis; laminae fertiles 3.5 cm. longae, 1.2 cm. latae, oblongae, margine re- volutae; venae simplices vel prope basin 1-furcatae, 1.5-2.0 mm. inter se dis- tantes, hydathodis ca. 1 mm. intra marginem terminantes.
Ho.iotype. Mexico. Chiapas: municipio Rayon, 10 km. above Rayon Mezcalapa along road to Jitotal, 1700 m., Breedlove & Smith 32649 (DS).
PARATYPE. Same locality, Breedlove & Smith 32429 (DS).
Remarks. A very distinct species, seemingly related to the smaller E. mathewsii (Fée) Moore, of high elevations; also related to E. alfrediit Rosen-
iss) _
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stock (type from Costa Rica), but differing from that species in the longer, narrower, darker, and more prominently toothed rhizome scales, the longer stipes of the sterile fronds, and the more persistently scaly blade surface and stipes.
Grammitis margaritata A. Reid Smith, sp. nov. (Figure 5, D-E.)
Rhizomata breve repentia, crassa (caudices ca. 0.7 cm. diametro), squamata, paleis numerosis rigidis castaneis opacis anguste lanceolatis, usque 5 mm. longis, basi ca. 0.5 mm. latis, margine evidenter ciliatis, ciliis 0.3 mm. longis, albidis, patentibus; stipites usque 14 cm. longi, 1 mm. diametro, brunnei, teretes, non alati, dense setosi, setis patentibus rigidis usque 3.5 mm. longis brunneis et pilis articulatis brevioribus; rhachides fuscae, setosae supra et infra; laminae subcoriaceae lanceolatae, basi pinnatisectae, ca. 11 cm. longae, 2.5 cm. latae; segmenta ca. 20-juga, usque 1.5 cm. longa, 4.5 mm. lata, margine leviter revoluta setosa, apice rotundata vel vix acuta, basi adnata, supra atro- viridia, infra pallide viridia, utrinque glabra, supra sine hydathodis calcareis; venae ca. 7-jugae, simplices; sori uniseriati mediales usque submarginales ro- tundi, glandulis stipitatis numerosis sphaericis albis mixti; ” = 37.
HototyPe. Mexico. Chiapas: municipio El Porvenir, 3-4 km. W. of El Porvenir along road from Huixtla to Siltepec, 2800 m., Breedlove & Smith 31801 (DS).
REMARKS. Known only from a single collection, growing on rocky ledge, in evergreen cloud forest (with Quercus, Drimys, Clethra, Symplocos). Most closely related to Polypodium (Grammitis) meridensis Klotzsch, but differing in the narrower, less deltoid laminae, the rounded (rather than acute) seg- ments, and the darker, more rigid and stiffly ciliate rhizome scales. The holo- type showed 2n = 37 II at meiotic metaphase.
The specific epithet recalls the pearly glands that adorn and nearly ob- scure the young sori.
Hypolepis melanochlaena A. Reid Smith, sp. nov. (Figure 7, D-E.)
Rhizomata et stipites ignoti; rhachides infra fulvae usque brunneolae, supra secus sulcum pubescentes, pilis rufis pluricellularibus, minute et perparce aculeatae, aculeis 0.2 (0.5) mm. longis; laminae 3-pinnatopinnatifidae, plus quam 60 cm. longae (specimina incompleta), dissectae similes H. repenti (Linnaeus) Presl; pinnae alternae, maximae plus quam 35 cm. longae, 10 cm. diametro; segmenta ultima oblonga, apice rotundata, integra usque lobata ca. 0.5; laminae chartaceae, infra omnino glabrae in statu sicco viridi-brunneae, supra secus sulcos costarum costularumque pubescentes, in statu sicco nigre- scentes; sori 1(2)-jugi per segmentum, indusiis ca. 1 mm. X 0.5 mm. semi- circularibus integris denigratis instructi.
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Horotyre. Mexico. Chiapas: without further locality, Munch 35 (DS); probable isotype: Minch s.n. (DS).
REMARKS. This species differs from all other species of Hypolepis known to me by the glabrous, blackish indusia; additionally it differs from all other Mexican species of Hvpolepis by the glabrous laminae below.
Polypodium surcurrens A. Reid Smith, sp. nov. (Figure 5, A-C.)
Rhizomata repentia, usque 6 mm. diametro, ca. 7 mm. inter bases stipitum, paleis appressis castaneis lustratis comosis vestita; stipites usque 30 cm. longi, 3 mm. diametro, laminis ca. 0.7 plo breviores, fulvi vel brunneoli; laminae usque 45 cm. longae, 25 cm. latae, basi pinnatisectae, apicem versus pinnati- partitae, denique segmento terminali usque 7 cm. longo, 1 cm. lato; pinnae usque 27-jugae, usque 14 cm. longae, 1.4 cm. latae, saepe arcuatae, apice acutae, medio latissimae, valde sursumcurrentes, infra pinnis infimis excisis, deflexis, marginibus undulatis pinnarum; venae discretae, 3—4-furcatae, venis primariis ca. 8 per 3 cm.; costae infra pilis ca. 0.2 mm. longis (pilis similaribus secus margines pinnarum), supra pilis densioribus Ctenitidis; paginae glabrae utrinque laminarum; sori mediales, elliptici, usque 2 mm. longi, 1.3 mm. latae, venula prima acroscopica venarum lateralium locati; sporangia glabra.
Hototype. Mexico. Chiapas: municipio Cintalapa, 16 km. NW. of Rizo de Oro, SE. of Cerro Baul on border with State of Oaxaca, 1600 m., Breedlove & Smith 31311 (DS).
ParaAtyPeEs. Mexico. Chiapas: Cerro del Ocote, 1500 m., Breedlove 28918 (DS); between Bochil and Simojovel, 1400 m., Breedlove & Smith 32493 (DS); Ocotal Grande, 750 m., Breedlove 33118 (DS); Cerro Baul, 1600 m., Breedlove & Smith 21795 (DS). Veracruz: Mirador, Galeotti 6414 (BR, photo US); Jalapa, CL. Smith 2207 (UC).
Remarks. Apparently a relatively common species at middle elevations in Chiapas. Polypodium surcurrens is closely related to P. longepinnulatum Fournier, from which it differs in the strongly surcurrent pinnae.
Polystichum bicolor A. Reid Smith, sp. nov. (Figure 6, E-G.)
Rhizomata erecta, caudices ca. 2 cm. diametro; stipites ca. 25 cm. longi, 5(8) mm. diametro, dense squamosi, paleis ascendentibus, anguste lanceolatis,
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Ficure 5. A-C. Polypodium surcurrens: A, rhizome and stipe base, Breedlove 33118, DS; B, portion of pinna, Breedlove & Smith 31311, DS; C, lamina, Breedlove & Smith 31311, DS. D-E. Grammitis margaritata, Breedlove & Smith 31801, DS: D, rhizome scale; E, plant.
CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
CUCU | yw PTI CTC ot wa cass A
Vor. XL] SMITH: CHIAPAS FERNS
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ca. 1.5 cm. longis, 1-2 mm. latis, integris, distincte bicoloribus, margine stra- mineis, in centro brunneis; rhachides non proliferae, minute squamosae, paleis plerumque 1-3 mm. longis, stramineis, integris sed basi dilatatis et minute fimbriatis; laminae chartaceae, bipinnato-pinnatifidae, usque 90 cm. longae, 28 cm. latae; pinnae usque 36-jugae, leviter inaequilaterae, non imbricatae, lanceolatae, usque 15 cm. longae, 2.4 cm. latae, inferiores 5—8-jugae gradatim reductae (infimae xX 0.3—0.6); pinnulae usque ca. 25-jugae, non congestae, maximae (basales) usque 1.5 cm. longae, 6 mm. latae, in latere acroscopico profunde lobatae usque crenatae, apice spinulosae; costae infra paleis dispersis stramineis similibus illis rhachidum vestitae, costulae venaeque glabrae: sori indusiis peltatis 0.5-0.8 mm. diametro, fulvis, brunnescentibus, margine irre- gularibus.
HootyPe. Mexico. Chiapas: municipio Tenejapa, Colonia ’Ach’lum, Ton 1987 (DS).
PaRATYPES. Mexico. Chiapas: municipio Tenejapa, Colonia ’Ach’lum, Ton 890 (DS); Vol. Tacana, 2100 m., Matuda 2971 (NY); municipio San Andres Larrainzar, summit of Chuchil Ton, NE. of Bochil, Breedlove 34686 (DS).
REMARKS. Distinguished from P. ordinatum (Kunze) Liebmann and P. drepanoides Fournier by the narrowly lanceolate, adpressed, distinctly bicolorous scales on the stipe bases, and by the reduced basal pinnae.
Polystichum erythrosorum A. Reid Smith, sp. nov.
Rhizomata ignota, probabiliter erecta, crassa; stipites usque 60 cm. longi, 0.8 cm. diametro, dense paleacei, aliquot paleis ovato-lanceolatis, usque 2 cm. longis, 0.8 cm. latis, nigricantibus et lustratis, vel paleis bicoloribus margine angusto fulvo, sursum paleis concoloribus, fulvis, parvioribus; rhachides non proliferae, moderate squamosae, paleis plerumque lineari-lanceolatis, usque 0.7 cm. longis, 0.1 cm. latis, fulvis, parce denticulatis vel basin versus fimbriatis; laminae subcoriaceae, bipinnatae, usque 70 cm. longae, 40 cm. latae; pinnae usque 26-jugae, non imbricatae, lineari-lanceolatae, usque 26 cm. longae, 3.2 cm. latae, infimae deflexae vix reductae; pinnulae usque 30-jugae per pinnam, non congestae, maximae 1.6 cm. longae, 0.7 cm. latae, lobo parvo acroscopico praeditae, aliter integrae vel crenatae, apice spinulosae, margine basiscopico arcuato; costae costulae venaeque infra paleis numerosis tortis fulvis capil- laceis vestitae; sori indusiis peltatis 0.6—-0.8 mm. diametro rubiginosis planis margine irregularibus.
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Ficure 6. A-D. Polystichum furfuraceum: A, lower pinna, Miinch 113, DS; B, stipe base, Breedlove & Smith 31807, DS; C—D, stipe scales, Breedlove & Smith 31807, DS. E-G. Polystichum bicolor, Breedlove 34686, DS: E, lower pinna; F, stipe base scales; G, stipe base. H-J. Pteris chiapensis, Breedlove & Smith 22492, DS: H, portion of ultimate seg- ment; I, ultimate segment; J, lamina.
226 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEk.
Horotype. Mexico. Chiapas: municipio Motozintla de Mendoza, road from Huixtla to El] Porvenir and Siltepec, 3000 m., Breedlove & Smith 22709 (DS).
ParAtyPEs. Mexico. Chiapas: Zontehuitz, Miinch 113 (US); Zontehuitz, Mickel 1250 (US); Zontehuitz, 2800 m., Breedlove & Smith 22047 (DS); municipio E] Porvenir, 3-4 km. W. of El Porvenir along road from Huixtla to Siltepec, 2800 m., Breedlove & Smith 31769 (DS).
Remarks. Similar to P. furfuraceum A. Reid Smith in size and dissection and growing in similar habitats, but with numerous black stipe base scales, pinnae and pinnules less crowded, never imbricate, scales not setiform along the margin, and reddish indusia.
Polystichum furfuraceum A. Reid Smith, sp. nov. (Figure 6, A—D.)
Rhizomata ignota, probabiliter crassa, erecta; stipites ca. 30 cm. longi, 0.7 cm. diametro, perdense paleacei, aliquot paleis ovatis, usque 1.8 cm. longis, 0.9 cm. latis, ceteris anguste lanceolatis, ca. 0.3 cm. longis, paleis omnibus ferrugineis usque brunneis, concoloribus vel basin versus leviter fus- catis, margine manifeste setiformis, dentibus usque 0.4 mm. longis; rhachides non proliferae, perdense paleaceae, paleis lanceolatis usque ca. 1 cm. longis, 0.2 cm. latis, similibus illis stipitum; laminae subcoriaceae, bipinnatae, ca. 60 cm. longae, 24-38 cm. latae; pinnae usque 45-jugae, approximatae vel etiam imbricatae, lineari-lanceolatae, usque 20 cm. longae, 2 cm. latae, infimae de- flexae et aliquantum abbreviatae (x 0.7); pinnulae usque 40-jugae per pinnam, approximatae vel imbricatae, maximae 1 cm. longae, 0.4 cm. latae, indivisae sed lobo parvo acroscopico praeditae, apice spinulosae, margine basiscopico arcuatae integrae vel apicem versus parce spinulosae; costae costulae venaeque infra paleis numerosis fulvis tortis capillaceis praeditae; sori indusiis peltatis 0.40.7 mm. diametro fulvis in centro depressis praediti.
Horotypre. Mexico. Chiapas: Zonehuitz |Zontehuitz|, Miinch 113 (DS 267622); isotype: DS 267621.
PARATYPE. Mexico. Chiapas: municipio El Porvenir, 3-4 km. W. of El Porvenir, 2800 m., Breedlove & Smith 31807 (DS).
ReMarRKS. Distinguished from all Mexican and Central American species by the very densely scaly stipe and rachis, the approximate or imbricate entire pinnules, and by the scales’ being prominently setiform. Perhaps most closely related to P. erythrosorum A. Reid Smith.
Pteris chiapensis A. Reid Smith, sp. nov. (Figure 6, H—J.)
Rhizomata erecta, caudices ca. 3 mm. diametro; stipites plus quam 30 cm. longi, ca. 8 mm. diametro, straminei, glabri; laminae chartaceae, ca. 60 cm. longae, basi bipinnatae usque tripinnatae, sursum bipinnatae, ultimo pin-
VoL. XL] SMITH: CHIAPAS FERNS 227
natae; pinnae infimae maximae, ca. 50 cm. longae, 35 cm. latae; segmenta ultima et pinnae distales usque 25 cm. longae, 1.5 cm. latae, basi anguste cuneatae et petiolatae usque 3 cm., integrae, apice sterili acute serratae; venatio areolata, areolis 2—3-seriatis, areolis costalis 2.0-2.5 mm. latis, brevis- simis, margine areolis ca. 1 mm. latis; sori marginales continui (apice excepto).
Hototyre. Mexico. Chiapas: municipio Ocozocoautla de Espinosa, 26-28 km. N. of Ocozocoautla, 700 ft. [213 m.|, Breedlove & Smith 22492 (DS— 2 sheets).
Remarks. Most closely allied to P. mexicana (Fée) Fournier, but differ- ing from that species in the sharply serrate apices of pinnae and pinnules, the narrower and more elongate pinnae and pinnules, fewer rows of areoles, and marginal areoles mostly 1 mm. or more broad (rather than ca. 0.5 mm. broad).
Thelypteris blepharis A. Reid Smith, sp. nov. (Figure 7, F—H.)
Rhizomata repentia, 0.8 mm. diametro; stipites 20-40 cm. longi, 4-6 mm. diametro, brunneoli vel dilute purpurei, paleis numerosis patentibus lineari- lanceolatis usque 8 mm. longis, 1 mm. latis praediti, his glabris vel margine minute glandulosis, atrocastaneis, lustratis; rhachides fulvae, paleis paucis similaribus; laminae chartaceae usque subcoriaceae, atrovirides, 50-70 cm. longae, 30-36 cm. latae, apice gradatim decrescentes; pinnae 20-35 jugae, usque 19 cm. longae, 2.1 cm. latae, basi suboppositae, sursum alternae, fere costis incisae; segmenta obliqua, subfalcata, apice acuta vel obtusa, segmentis basalibus pinnarum infimarum leviter amplificatis, auriformibus; venae promi- nentes, usque 13-jugae, infimae marginem supra sinum attingentes; rhachides costae venaeque infra epilosae vel sparsim pubescentes, glandulibus minutis dispersis stipitatis luteolis, supra pilis usque ca. 0.5 mm. longis; paginae laminarum utrinque glabrae; sori mediales; indusia porphyrea, glandulifera, dense pilosa, pilis usque ca. 1 mm. longis.
Hototypr. Mexico. Chiapas: municipio La Independencia, road from Las Margaritas to Campo Alegre, 2300 m., Breedlove 33605 (DS).
Paratype. Guatemala. Huehuetenango: slope above San Juan Ixcoy, Sierra de los Cuchumantanes, travertine limestone, 2400 m., Steyermark 49997 (F, GH).
Remarks. This is one of the most distinct species in section Cyclosorus, differing from its relatives [7. puberula (Baker) Morton, 7. ovata R. St. John, 7. tuerckheimii (Donnell-Smith) Reed] by the narrow, dark, bristlelike scales on the stipes and by the densely long-hairy indusia but otherwise glabrous laminae. Its discovery reinforces my belief that southern Mexico- Guatemala is the center of evolution of section Cyclosorus in the New World (Univ. Calif. Publ. Bot. Vol. 59, pp. 1-136, 1971).
228
' [Proc. 4TH SER.
Vou. XL] SMITH: CHIAPAS FERNS 229
Thelypteris nubigena A. Reid Smith, sp. nov. (Figure 7, C.)
Rhizomata suberecta; stipites usque 15 cm. longi, 3 mm. lati, puberuli, basi fuscati et parce squamulosi, paleis ovatis, fulvis, margine sparsim ciliatis; frondes usque 65 cm. longae, rhachidibus stramineis; pinnae ca. 25-jugae, in- fimae deinceps 1 mm., 1.5 cm., 4.0 cm., 7.0 cm. longae; pinnae maximae lanceo- latae, ca. 7 cm. longae, 1.8 cm. latae, basi latissimae, ca. 0.9 costam versus lobatae; segmenta obliqua, subfalcata, usque 10 mm. longa, 2 mm. lata, apice subacuta; venae usque 10-jugae, marginem supra sinum attingentes; aerophora usque 1 mm. longa basi pinnarum inferiorum; rhachides costae costulaeque utrinque sine paleis, pilis dispersis interdum subfasciculatis ca. 0.2 mm. longis; glandes infra in venis costulis laminisque numerosa, aurantiaca, sessiles, glutinosa; sori mediales; indusia minuta, glandibus aurantiacis abscondita.
Hototyrr. Mexico. Chiapas: municipio San Cristobal Las Casas, E. side of Zontehuitz near summit, 2800 m., Breedlove & Smith 22057 (DS).
ParAtyPes. Mexico. Oaxaca: Distrito Ixtlan, 19 km. N. of Ixtlan on Rte. 175, 2600 m., Mickel 5533 (NY). Guatemala. Quezaltenango: Vol. Santo Tomas, Steyermark 34709 (F).
Remarks. This species can be compared only to 7. thomsonii (Jenman) Proctor, but the hairs of 7. nubigena are less obviously fasciculate, and the fronds are much smaller, the segments narrower and more acute. It occurs at perhaps a higher elevation than any other Thelypteris in Mexico.
Ctenitis chiapasensis (Christ) A. Reid Smith, comb. nov.
Aspidium chiapasense Christ, Bull. Herb. Boissier II, 5:727. 1905. Type: Mexico, Chiapas, Baduitz, Miinch 117 (isotype DS!).
Ctenitis lanceolata (Baker) A. Reid Smith, comb. nov.
Nephrodium lanceolatum Baker, Syn. Fil, second edition. 498. 1874. Type: Guatemala [Alta Verapaz], Coban, Salvin & Godman s.n.
Ctenitis lindenii (Kuhn) A. Reid Smith, comb. nov. Aspidium lindenii Kuhn, Linnaea 36:116. 1869. Type: Mexico, Tabasco, Teapa, Linden 1489.
Grammitis blepharodes (Maxon) A. Reid Smith, comb. nov.
Polypodium blepharodes Maxon, Contr. U.S. Natl. Herb. 17:407. 1914. Type: Costa Rica, La Palma, Maxon 406.
—
Ficure 7. A-B. Asplenium munchii, Miinch 114, DS: A, frond; B, pinna. C, Thelyp- teris nubigena, lowermost five pinnae, Breedlove & Smith 22057, DS. D-E. Hypolepis mel- anochlaena, Miinch 35: D, ultimate segment; E, pinna. F-H. Thelypteris blepharis, Breed- love 33605, DS: F, lower pinna; G, stipe base; H, portion of pinna.
230 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Grammitis harrisii (Jenman) A. Reid Smith, comb. nov.
Polypodium harrisii Jenman, Gard. Chron. III, 27:241. 1900. Type: Jamaica, Harris.
Grammitis leptostoma (Fée) A. Reid Smith, comb. nov.
Polypodium leptostomum Fée, Mem. Foug. 7:58. 1857. Type: Mexico [Veracruz], near Orizaba, Schaffner 210.
Microgramma nitida (J. Smith) A. Reid Smith, comb. nov.
Phlebodium nitidum J. Smith, Bot. Mag. 72. Comp. 13. 1846. Type: cultivated specimen, originally from Honduras, introduced to Kew in 1844. Maxon (Proc. Biol. Soc. Wash. Vol. 51, p. 38, 1938) discussed the identity of J. Smith’s species. A synonym is Polypodium palmeri Maxon (Contr. US. Natl. Herb. Vol. 17, p. 600, 1917).
Microgramma reptans (Cavanilles) A. Reid Smith, comb. nov.
Acrostichum reptans Cavanilles, Anales Hist. Nat. 1:104. 1799. Type: Ecuador, Guayaquil, Née sn. The type was seen and commented upon by Christensen [Dansk. Bot. Ark. vol. 9, no. 3, p. 9, 1937].
Polypodium ciliatum Willdenow |= Microgramma ciliata (Willdenow)
Alston], the earliest name for this species in Polypodium, becomes a taxonomic
synonym when Microgramma is recognized.
Pleopeltis munchii (Christ) A. Reid Smith, comb. nov. Polypodium munchii Christ, Bull. Herb. Boissier II, 3:147. 1903. Type: Mexico, Chiapas, San Cristobal Las Casas, Miinch 90 (isotype DS!).
Pleopeltis revoluta (Sprengel ex Willdenow) A. Reid Smith, comb. nov.
Grammitis revoluta Sprengel ex Willdenow, Sp. Pl. 5:139. 1810. Type: locality not stated.
This species has gone under a great many names, most notably Polypodium astrolepis Liebmann, which is the earliest name for this species in Polypodium (see Weatherby, Contr. Gray Herb. No. 65, pp. 3-14, 1922, for additional synonymy). However, the epithet revoluta is prior and available in Pleopeltis.
| Warne Biolog PROCEEDINGS
OF THE | N |
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 9, pp. 231-242; 2 figs. August 8, 1975
A NEW GENUS AND SPECIES OF EUCERINE BEE FROM NORTH AMERICA (HYMENOPTERA: ANTHOPHORIDAE)
By Thomas J. Zavortink
Department of Entomology, California Academy of Sciences San Francisco, California 94118
Apstract: A new genus and species of eucerine bee, Simanthedon linsleyi, from the southwestern United States and northern Mexico is described and figured. In southeastern Arizona, this aestival bee is primarily a matinal forager on the flowers of Menodora scabra Gray (Oleaceae).
Shortly after daybreak on a morning in late August 1972, I collected a few eucerine bees that appeared to be Martinapis luteicornis (Cockerell). They were taken from a plant with yellow, star-shaped flowers found growing near the town of Bisbee in southeastern Arizona. Subsequent examination of the bees disclosed that they were not M. luteicornis, but were instead an undescribed species that is sufficiently distinct from all other New World eucerines to be placed into a new genus. Subsequent examination of the plant showed that it was Menodora scabra Gray (Oleaceae). During the summers of 1973 and 1974, I returned to southeastern Arizona in order to study the pollination of M. scabra and collected more of the unusual eucerine bees on it. This new genus and species of bee is described in the present paper.
The generic description includes the characters that have been used to define the numerous New World genera of Eucerini by Moure and Michener (1955) and LaBerge (1957), and numerous additional characters that I believe are also of generic value. Characteristics common to both sexes are included in the generic and specific descriptions of the male.
The abbreviations used in the list of specimens examined and the museums
[231]
232 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
they represent are: AMNH, American Museum of Natural History; ASU, Arizona State University; CAS, California Academy of Sciences; UCB, Cali- fornia Insect Survey Collection, University of California at Berkeley; UCR, P. H. Timberlake Collection, University of California at Riverside.
Simanthedon Zavortink, new genus
Mate. Head: Face narrow, minimum distance between eyes 0.70—0.75 length of eye. Inner margins of eyes parallel to slightly convergent ventrally. Vertex weakly elevated behind ocelli, median ocellus separated from apex of head by 0.35—0.45 of its width in frontal view. Ocelli slightly enlarged, distance from median ocellus to lateral ocellus 0.29-0.44 diameter of median ocellus. Distance between lateral ocelli 1.6—2.1 times distance from lateral ocellus to eye. Lower paraocular carina absent. Minimum oculoclypeal distance 0.3—0.7 mini- mum width of first flagellar segment. Clypeus approximate to eye for distance subequal to basal width of mandible; strongly protuberant, in lateral view extended beyond anterior margin of eye by 0.70—0.85 width of eye; moderately produced, in facial view extended beyond lower end of eye by 0.7—1.1 width of median ocellus; lateral part not reflexed mesad of lateroclypeal carina; upper part narrow, width at level of anterior tentorial pit 0.85—0.95 width of labrum; ‘pug-nosed’, with surface concave in upper 0.65—0.75 medially, strongly reflexed and flat in lower 0.25—0.35. Malar area short, 0.10—0.13 as long as wide. An- terior mandibular articulation slightly farther from eye than posterior one. Antenna: Long, extended to pterostigma in repose. Scape thick, short, slightly thicker than width of median ocellus, as long as to longer than interantennal distance. Flagellum slender, greatest diameter less than width of median ocellus, slightly flattened, tapered distally; segment 1 long, 1.20-1.35 length of scape, 0.75—0.85 length of segment 2; segments 3-10 progressively shorter; segment 11 long, as long as segments 5, 6, or 7, curved, tapered to a blunt point. Mouth- parts: Labrum 0.60—0.65 as long as wide; distal margin convex with shallow median emargination. Mandible short, 0.50—0.60 length of eye; with weak subapical tooth on inner margin. Proboscis moderately long, distal part of galea 1.0—-1.2 length of eye. Maxillary palpus 5-segmented; 0.32—0.37 length of distal part of galea; lengths of segments in ratio of 1.00:0.94:0.82:0.53:0.41. Segment 1 of labial palpus 1.8—2.3 length of segment 2.
Mesosoma: Scutellum 0.22—0.28 length of scutum; strongly convex; with moderately long, median, longitudinal, impunctate ridge or line anteriorly. Metanotum strongly convex. Propodeum with basal part steeply declivous; posterior surface without vertical carina laterally. Without spatuloplumose hairs. Tegula oval. Wings: Forewing with prestigma short. Pterostigma shorter than prestigma, with inner margin barely extended distad of base of vein r. Marginal cell subequal in length to distance from its apex to apex of wing or slightly longer; 3.7—4.3 times as long as wide; basal part subequal in length to
Vot. XL] ZAVORTINK: EUCERINE BEE
ine) W WwW
FicureE 1. Simanthedon linsleyi, new species, head. A, facial view of male; B, lateral view of male; C, facial view of female; D, lateral view of female.
free part. Three submarginal cells; first 1.4-1.7 length of second, 1.0—1.2 length of third; second strongly rhomboid. Cell 1M petiolate, 0.84—0.94 length of marginal cell. Vein C not strongly expanded at base. Vein 2nd r-m sharply angled, usually appendiculate. Vein Ist m-cu ending 0.70—0.82 distance from base of second submarginal cell. Vein 2nd m-cu joined to Cu; in nearly even curve. Hindwing with 13-18 hamuli. Vein cu-v 0.53-0.67 length of 2nd abscissa of vein M + Cu. Jugal lobe 0.87—0.95 length of cell Cu. Legs: Femora slender; femur I 1.0-1.1 length of femur II, flattened dorsoventrally, broadest near base; femur IIT unusually slim, 4.3—-4.8 times longer than maximum width. Tibiae slender; tibia III long, 1.5—1.7 length of tibia II. Tibia I spur long, longer than spurs of tibiae IT and III, 0.6-0.7 length of basitarsus I, with post-
234 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
velar portion of malus long, sinuous; spur II moderately long, 0.43—0.49 distance from its base to anterior tibiofemoral articulation, curved at apex; outer spur III short, inner spur III moderately long, subequal in length to spur II, both gently curved toward each other, so their distal portions are subparallel. Tarsi short; tarsus I 1.0—1.1 length of tibia I; tarsus II 1.5—1.6 of tibia IT; tarsus III subequal in length to tarsus II and tibia III. Basitarsus I 0.6—-0.7 length of tarsus I; basitarsus II 0.9-1.0 length of tibia II, 0.6 length of tarsus IT; basitarsus III 0.6—-0.7 length of tarsus III, 5.3-5.6 times longer than wide, with outer surface sparsely pubescent, flattened or concave apically. Claws small, symmetrical on all legs. Arolia present.
Metasoma: Anterior face of tergum I subequal in length to dorsal face; concave. Terga VI, VII without gradular teeth. Pygidial plate wider than long, not narrowed basally, truncate, with notched lateral margin; marginal carina progressively strengthened posteriorly basad of notch, weak distad of notch; surface sparsely pubescent. Sternum I without conspicuous eminence. Sterna II-IV not emarginate. Terga II-IV with basal and narrower apical bands of pale pubescence; without spatuloplumose hairs. Distal sterna and genitalia: Sternum V weakly trilobed, with mesal edge of lateral lobe bearing clump of posteriorly directed, strong, long, curved, terete, barbed setae. Sternum VI with oblique lateral carina progressively weakened distad, curved laterally proximad; with median longitudinal sulcus bordered by setae; with pair of lateral subquadrate areas delimited by weak carinae on dorsal surface of disk. Sternum VII with lateral plate large, strongly sclerotized, strongly pigmented, with very deep, narrow lateral emargination; clear membranous area long, nar- row, not produced; median plate long, without basal tubercle, with distal portion narrow, straplike, reflexed basally, directed laterad, glabrate; without unpaired median projection; basal apodeme broad, curved, pointed. Sternum VIII narrowed distad of apodeme, with rounded, slightly emarginate apex bear- ing a few short setae; tubercles far basad of apex; dorsal tubercle moderately broad, with rounded apex; ventral tubercle narrow, elevated distad. Gonocoxite with strong, moderately long dorsoapical process; with moderately long ventral process extended over base of penis valve; without ventroapical process; either without or with very few ventroapical setae. Gonostylus with base broad, sparsely pubescent with simple and inconspicuously plumose hairs; distal 0.6 very slender, gently curved inward. Spatha 1.7—1.8 times wider than long; basolateral angle slightly produced; distal margin with pair of deep, oblique emarginations that receive dorsobasal edges of penis valves, and shallow median emargination; without median longitudinal apodeme. Penis valve with dorso- basal edge strongly oblique, set into emargination of spatha basally, barely extended beyond dorsolateral tooth distally; dorsolateral tooth short, not extended to dorsoapical process of gonocoxite; ventromesal margin not ex-
Vor. XL] ZAVORTINK: EUCERINE BEE
ise) WwW wal
eq dima, a w VAS /
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Ficure 2. Simanthedon linsleyi, new species, distal sterna and genitalia of male. A, genitalia, dorsal surface on left; B, sternum VIII, ventral surface on left; C, sternum VII, ventral surface on left; D, sternum VI, left half of ventral surface; E, sternum V, left half of ventral surface. The scale lines represent 1.0 mm.
panded, nearly straight; with broad apodeme extended distad to base of apical process. Penis broad, extended far beyond spatha.
FreMALE. Head: Minimum distance between eyes 0.84—0.91 length of eye. Inner margins of eyes parallel. Median ocellus separated from apex of head by 0.27-0.40 of its width. Distance between lateral ocelli 1.3-1.5 distance from lateral ocellus to eye. Minimum oculoclypeal distance 0.5—-0.8 minimum
236 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
width of first flagellar segment. Clypeus moderately protuberant, extended beyond anterior margin of eye by 0.55—0.65 width of eye; width at level of an- terior tentorial pit 0.95—1.00 width of labrum; surface normal, convex through- out; disk without median longitudinal impunctate ridge or line. Antenna: Scape slender, obconic, longer than interantennal distance. Flagellum slender, greatest diameter less than width of median ocellus; segment 1 0.85—0.95 length of scape, 2.1—2.5 length of segment 2; segment 2 obconic, 0.75—0.95 length of segment 3; segments 3-9 much longer than wide; segment 10 1.8-2.2 times longer than wide. Mouthparts: Labrum 0.55—0.65 as long as wide; with small tuft of dense pubescence around apical emargination. Mandible 0.54—0.66 length of eye; simple; outer lower carina less salient than inferolateral carina. Stipes without hooked hairs.
Legs: Anterior coxa with short apical spine. Femora and tibiae not unusually slender or long. Basitibial plate hidden or defined posteriorly. Scopal hair dense, simple. Tibia I spur short, shorter than spurs of tibiae II and III, 0.45—0.55 length of basitarsus I; spur II long, 0.56—0.66 distance from its base to anterior tibiofemoral articulation, curved at apex; inner and outer spurs III subequal in length, longer than spur II. Tarsi not unusually short. Basitarsus III with inner surface densely hairy. Claws small, symmetrical on all legs.
Metasoma: Gradulus of tergum VI without lateral parts. Pygidial plate more or less V- or Y- shaped, narrow and pointed apically, with distal lateral margin concave; length 1.2 times basal width. Gradulus of sternum IT moderately strongly biconvex. Sternum VI emarginate at apex.
Type species. Simanthedon linsleyi Zavortink, new species.
Etymo.ocy. Simanthedon (feminine) from the Greek simos, pugnosed, and anthedon, bee.
Simanthedon linsleyi Zavortink, new species. (Figures 1, 2.)
Mate. Measurements: Length, exclusive of antenna, 12.0-15.0 mm. Length of forewing 9.0-10.1 mm. Width of head 3.7—4.1 mm. Color: Integument non- metallic. Dark, black or black suffused with red, with clypeus yellow; scape and labrum cream-colored to yellow; flagellum largely yellowish in life but usually more reddish or brownish when dried; mandible yellow basally, dark amber medially, red apically; tegula amber, darker anteriorly; wing membrane weakly infuscated; veins and pterostigma brown and black; legs reddish brown to brown distad; tibial spurs testaceous; apical margin of terga amber; pygidial plate largely red. Vestiture: Face, vertex, posterior surface of head, entire mesosoma, and tergum 1 with long, dense whitish to ochraceous, light fulvous, or light ferruginous pubescence, this densest and darkest on pronotum and scutum; legs with pubescence largely ochraceous to fulvous, but white or whitish
Vor. XL] ZAVORTINK: EUCERINE BEE 237
on posterior surface of trochanter II, femur IJ, and tibiae II and III; hind femur and tibia sparsely pubescent; tergum I with long pubescence extended nearly to apical margin, without an apical band of appressed pale pubescence, with an apicolateral patch of short black pubescence; pubescence of other terga black or brownish black, with basal and/or apical pale bands as follows: basal white pubescent band moderately broad to broad laterally and narrow medially on tergum IT, uniformly broad on tergum III, uniformly broad and fused with apical pale band laterally on tergum IV; apical pubescent band very narrow and whitish on terga II and III, moderately broad and white on tergum IV, moderately broad and white or dingy white laterally and strongly tinged with black or blackish brown medially on tergum V; terga II-VI with scattered, long, erect, black and ochraceous to fulvous hair. Sterna with pubescence short medially on II-V, white to fulvous; strong specialized setae of sternum V orange-brown to brown. Sculpture: Clypeus smooth, shiny, very finely and sparsely punctate medially, tessellate and more coarsely punctate laterally; face roughened, moderately coarsely and moderately closely punctate below, smoother and more finely and closely punctate above; thorax shiny, moderately coarsely and closely to very closely punctate, the punctures more widely separated on posterior center of scutum, anterior part of scutellum, and pleuron; anterior and lower parts of hypoepimeral area impunctate; sides of propodeum roughened, moderately coarsely and closely punctate above, smoother and more sparsely punctate below; basal part of propodeum indistinctly to distinctly tessellate, coarsely and moderately closely punctate; lower part of propodeal triangle smooth, impunctate; dorsal face of tergum I roughened basally, tessellate apically, moderately coarsely and closely to moderately closely punctate; inter- band zones of terga II-IV tessellate, finely and closely to moderately closely punctate.
FEMALE. Measurements: Length, exclusive of antenna, 13.0-15.0 mm. Length of forewing 9.0-10.3 mm. Width of head 4.0-4.5 mm. Color: Without light integumental areas; apical part of clypeus, underside of flagellum, labrum, and base or middle of mandible usually reddish or reddish brown; pygidial plate black. Vestiture: Legs with some pubescence whitish or white on coxae, trochanters, femora, and posterior surface of tibia II; scopal hair stramineous to ochraceous on outer surface of hind tibia and basitarsus, orange to ferruginous on inner surface of basitarsus; basal white pubescent band moderately broad laterally and narrow medially on tergum II, uniformly moderately broad to broad or narrowed medially on tergum III, uniformly moderately broad to broad on tergum IV; apical white pubescent band narrow on tergum IT, narrow to moderately broad on tergum III, moderately broad on tergum IV; tergum V with some white hair apicolaterally; terga V, VI with apical hair ferruginous; terga III, IV with scattered, long, erect, black and ochraceous to fulvous hair.
238 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Pubescence of sterna white and orange to orange-brown. Sculpture: Clypeus tessellate, often diagonally wrinkled, coarsely and moderately closely punctate; interband zones of terga II, III tessellate, finely and closely to moderately closely punctate.
TYPE MATERIAL. Holotype: male, 6.0 mi. SW. Bisbee, Cochise Co., Arizona, United States, 4800 ft. elevation, 26 July 1973, on Menodora scabra, 0615— 0630 Mountain Standard Time (MST), T. J. Zavortink [CAS type No. 12254]. Allotype: female, same data as holotype except collected 0530-0545 MST [CAS]. Paratypes (21 males, 88 females): UNITED STATES. ARIZONA. Cochise Co.: Bisbee (6.0 mi. SW.), 4800 ft., 26 Aug. 1972, on Menodora scabra, 0545— 0615 MST, 14,32, 0615-0645 MST, 12; 26 July 1973, on Menodora scabra, 0530-0545 MST, 12, 0545-0600 MST, 14, 12, 0600-0615 MST, 12; 27 July 1973, on Menodora scabra, 0545-0600 MST, 12, 0600-0615 MST, 22, 0615- 0630 MST, 22, 0700-0715 MST, 12, 0715-0730 MST, 14; 28 July 1973, on Menodora scabra, 0530-0545 MST, 12, 0600-0615 MST, 12, 0615-0630 MST, 1d, 12, 0630-0645 MST, 12; 29 July 1973, on Menodora scabra, 0600-0615 MST, 12, 0615-0630 MST, 22, 0630-0645 MST, 22, on Polygala racemosa, 0545-0600 MST, 12; 30 July 1973, on Menodora scabra, 0545-0600 MST, 12, 0600-0615 MST, 14, 12; 16 Aug. 1974, on Menodora scabra, 0515-0530 MST, 12, 0530-0545 MST, 12, 0545-0600 MST, 22, 0600-0615 MST, 42, 0615-0630 MST, 42, 0630-0645 MST, 22, 0645-0700 MST, 12, 0700-0715 MST, 12, 0715-0730 MST, 22; 26 Aug. 1974, on Menodora scabra, 0515-0530 MST, 32, 0530-0545 MST, 62, 0545-0600 MST, 14, 2%, 0600-0615 MST, 24, 0615-0630 MST, 14, 0630-0645 MST, 12, 0645-0700 MST, 12, 0700- 0715 MST, 12, on /pomoea hirsutula, 0700-0715 MST, 12, T. J. Zavortink [CAS]. Douglas (1.0 mi. E.), 14 July 1962, on Conyza species, 14, M. A. Cazier [ASU]; 26 July 1962, on Mentzelia pumila, 1800-1900 hours, 14, M. A. Cazier [UCB]. Douglas (4.1 mi. E.), 4400 ft., 27 Aug. 1974, on Menodora scabra, 0530-0545 MST, 24, 8%, 0545-0600 MST, 62, 0600-0615 MST, 32, 0615-0630 MST, 14,32, 0630-0645 MST, 14,12, on Desmanthus cooleyi, 0545-0600 MST, 12, on Salvia reflexa, 0700-0715 MST, 14, T. J. Zavortink [CAS]. Portal (1.0 mi. S.), 3 Aug. 1969, on Hoffmannseggia densiflora, 0550— 0559 MST, 32 [UCB], 12 [UCR], flying around Solanum elaeagnifolium, 0530-0544 MST, 12 [UCB], E. G. Linsley. Portal (2.0 mi. NW.), 5100 ft., 21 Aug. 1973, on Menodora scabra, 0600-0615 MST, 12, 0615-0630 MST, 1°; 31 Aug. 1974, on Menodora scabra, 0515-0530 MST, 14, 0545-0600 MST, 1¢, 0600-0615 MST, 14, 0630-0645 MST, 12, T. J. Zavortink [CAS]. New Mexico. Eddy Co.: Whites City, 8 July 1954, 16, M. A. Cazier and W. J. Gertsch [AMNH]. Mexico. Duranco. La Zarca (26 mi. S.), 16 July 1964, on Cevallia sinuata, 1¢, J. A. Chemsak [UCB].
Etymo.ocy. This species is dedicated to E. Gorton Linsley in recognition of his contributions to knowledge of the systematics and biology of bees.
Vor. XL] ZAVORTINK: EUCERINE BEE 239
DISCUSSION
The monotypic genus Simanthedon may be distinguished from all other North American genera of Eucerini as follows: The male by (1) the strongly protuberant, ‘pug-nosed’ clypeus; (2) the long, slender, lightly pigmented antenna, with flagellar segment 1 long and segment 11 long, curved, and tapered to a blunt point (the antenna is similar to that of Martinapis luteicornis, but longer); (3) the combination of the slender femora I and III, the long and slender tibia III, the short tarsal segments 2—5 on I and III, and the sparsely pubescent and apically flattened or concave outer surface of basitarsus IIT; (4) the combination of the long spur on tibia I and the unequally shortened spurs on tibia III; (5) the paired patches of posteriorly directed, strong, long, curved, terete, barbed setae on sternum V; (6) the strongly sclerotized, strongly pigmented, and deeply laterally emarginate lateral plate and the long, narrow, reflexed, glabrate median plate of sternum VII; and (7) the pair of deep, oblique emarginations in the distal margin of the spatha. The female is distinguished by the combination of (1) the parallel inner margins of the eyes; (2) the moderately protuberant clypeus; (3) the short malar area; (4) the 5-segmented maxillary palpus; (5) the absence of spatuloplumose hairs on the mesosoma or metasoma; (6) the simple scopal hair; (7) the small, symmetrical claws on all legs; (8) the narrow, pointed, V- or Y- shaped pygidial plate; and (9) the absence of lateral parts on the gradulus of tergum VI.
Both sexes of Simanthedon linsleyi bear a strong but only superficial re- semblance to Martinapis luteicornis. Simanthedon linsleyi may be distinguished from M. luteicornis by all the features of the male and features 1, 6, 7, and 8 of the female mentioned in the preceding paragraph, and by the following addi- tional significant characters: (1) the absence of a median longitudinal, im- punctate ridge or line on the clypeus of the female; (2) the simple, apically narrowed mandible of the female; (3) the oval tegula (the lateral margin of the tegula of M. luteicornis is stated to be convex in the anterior half by LaBerge (1957), but it is actually distinctly concave); (4) the short pterostigma (the pterostigma of M. luteicornis is unusual in being longer than the prestigma and extended far beyond the base of vein r); (5) the absence of a strong expansion at the base of the costal vein; (6) the absence of a gradular tooth on tergum VI of the male; (7) the more weakly biconvex gradulus of sternum II of the female; (8) the presence of a ventral process on the gonocoxite of the male; and (9) the simple, sparsely pubescent gonostylus of the male.
In LaBerge’s (1957) key to the North and Central American genera of Eucerini, males of Simanthedon linsleyi run to Martinapis and females run to the couplet separating Synhalonia (as Tetralonia) and Xenoglossodes. The numerous characteristics that distinguish Simanthedon from the North American Martinapis (M.) luteicornis have been enumerated above; females of Siman- thedon may be separated from those of Synhalonia by the five segmented
240 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
maxillary palpus and the narrow, pointed, V- or Y- shaped pygidial plate, and from those of Xenoglossodes by the moderately protuberant clypeus and the longer galea. In Moure and Michener’s (1955) key to the South American genera of Eucerini, both sexes of S. linsleyi run to Martinapis if the reflexed median plate of sternum VII of the male and the simple apex of the mandible of the female are ignored. Unfortunately, specimens of the Argentine Martinapis (Svastropsis) bipunctata (Friese) have not been available for study and I cannot give a detailed list of the differences between S. linsleyi and that species. However, on the basis of the descriptions and illustrations in Moure and Michener (1955), males of S. linsleyi probably differ from those of /. bipunctata (female unknown to Moure and Michener) by all seven of the distinguishing features given in the first paragraph of this section.
The affinities of S¢manthedon are unknown; it does not appear to be closely related to any North or South American genus of Eucerini. It does belong, however, in the “central group” of eucerine genera, as defined by Moure and Michener (1955). Sternum VII of the male of S. linsleyi is remarkably similar to that of the Chilean Svastrides melanura (Spinola), but beyond that, there is little resemblance between these species.
I have not observed any significant variation in the specimens of S. linsleyi available for study; even the males from New Mexico and Durango, Mexico appear to be indistinguishable from those from Arizona.
The majority of the known specimens of Simanthedon linsleyi have been collected from the flowers of Menodora scabra. The species is apparently not oligolectic on this plant however, as many females, including some collected on Menodora, bear a heavy load of pollen from other flowers in their scopae. The non-oleaceous pollen that has been collected in significant amounts appears to be of six different kinds, three of which I have been able to determine as Agave palmeri Engelmann, Datura meteloides A. P. de Candolle, and Polygala racemosa Blake. This identification was done by comparing, in temporary glycerol mounts, pollen removed from the scopae of females with that removed from anthers of herbarium specimens of plants that were common at the type locality of S. linsleyi in 1973. Numerous other females of S. linsleyi collected on Menodora bear what I consider to be insignificant amounts of non-oleaceous pollen in their general vestiture; this pollen was undoubtedly obtained inci- dentally while these individuals were foraging for nectar only, and I have made no attempt to identify it. A complete list of flower records, along with an indication of the type of pollen borne in significant amounts by the females, for 21 males and 89 females is: Cevallia sinuata Lagasca y Segura, 14; Conyza species, 1 6 ; Desmanthus cooleyi (Eaton) Trelease, 12 (with Menodora pollen) ; Hoffmannseggia densiflora Benthan ex Gray, 42 (none with Hoffmannseggia pollen, 1 with Datura pollen, 1 with Agave and an unknown large spherical pollen, 2 with the unknown large spherical pollen); /pomoea hirsutula J.
Vou. XL] ZAVORTINK: EUCERINE BEE 241
Jacquin, 12 (with 7pomoea and Menodora pollen); Menodora scabra, 174, 812 (452 with pure or nearly pure Menodora pollen, 12 with Menodora and Datura pollen, 12 with Menodora, Datura, and Polygala pollen, 12. with Menodora and Agave pollen, 12 with Menodora and [pomoea pollen, 12 with Menodora and an unknown small spherical pollen, 5° with Datura pollen, 1° with an unknown moderately large oval pollen, 252 without pollen); Mentzelia pumila (Nuttall) Torrey & Gray, 14; Salvia reflexa Hornemann, 1 4 ; Polygala racemosa Blake, 12 (with Datura pollen) ; flying around Solanum elaeagnifolium Cavanilles, 12 (without Solanum pollen, but with Agave and an unknown large spherical pollen).
All but one of the 19 males and 89 females of S. dinsleyi for which the time of collection has been recorded were taken in the early morning, between 0515 and 0730 MST. The single exception is a male taken in the evening, between 1800 and 1900 hours, on Mentzelia pumila near Douglas, Arizona. On the basis of my collections on Menodora scabra, the main flight period, during which 75 percent of the specimens were taken, of S. linsleyi extends from 0.50 hour before sunrise to 0.75 hour after sunrise, the females usually arriving at the flowers slightly earlier than males. However, since the flowers of Menodora scabra are closed at daybreak and do not start opening until 0.50—0.25 hour before sunrise, bees active at dawn cannot forage on this plant, and the daily flight period of S. linsleyi may start earlier than my data show. This hypothesis is supported by the fact that most of the females of S. dinsleyi which bore non- oleaceous pollen and were collected on Menodora scabra were among the first bees to arrive at the flowers of this plant each morning.
The seasonal flight period of S. dinsleyi is during the summer rainy season typical of the region it inhabits. The earliest and latest recorded dates are 8 July and 31 August.
S. linsleyi is presently known from three areas near the edge of the Chi- huahuan Desert: southeastern Arizona and southeastern New Mexico, United States, and northeastern Durango, Mexico. All the specimens collected by me in Arizona have been taken in areas of calcareous soil, where Menodora scabra is particularly abundant. The single specimens from New Mexico and Durango are also from regions characterized by abundant limestone and, incidentally, within or near the known range of Menodora scabra (Steyermark, 1932). All of the specimens of S. linsleyi examined for this study (22 males, 89 females) are part of the type series, cited above.
ACKNOWLEDGMENTS
I am greatly indebted to C. D. Michener and R. R. Snelling for the loan or gift of specimens representing nearly every genus of Eucerini found in the Western Hemisphere. I also thank the following persons for allowing me to study additional specimens in their care: H. V. Daly, F. F. Hasbrouck, P. D. Hurd,
242 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
iiveeeGaLinsley. J. 'G. Rozen, Jr.; P: H. Timberlake, and P. F. Torchio;el thank W. E. LaBerge for his advice on the relationships and rank of Simanthedon linsleyi, P. H. Arnaud, Jr., and M. C. Giles for preparing the photographs, C. L. Mullinex for preparing the drawings, and R. Berlin for typing the manuscript. P. H. Arnaud, Jr., D. H. Kavanaugh, E. G. Linsley, and R. R. Snelling kindly read and criticized the manuscript.
REFERENCES LABeErcE, W. E. 1957. The genera of bees of the tribe Eucerini in North and Central America (Hymenop- tera, Apoidea). American Museum Novitates, no. 1837, pp. 1-44. Mouvre, J. S., and C. D. MICHENER 1955. A contribution toward the classification of Neotropical Eucerini (Hymenoptera, Apoidea). Dusenia, vol. 6, pp. 239-331. STEYERMARK, J. A. 1932. A revision of the genus Menodora. Annals of the Missouri Botanical Garden, vol. 19, pp. 87-176.
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES.
FOURTH SERIES
Vol. XL, No. 10, pp. 243-263; 5 figs.; 4 tables August 8, 1975
THE TAXONOMIC STRUCTURE OF SIX GOLDEN TROUT (SALMO AGUABONITA) POPULATIONS FROM THE SIERRA NEVADA, CALIFORNIA (PISCES: SALMONIDAE)
by J. R. Gold and G. A. E. Gall
Fisheries Biology Research Facility, Department of Animal Science, University of California, Davis, California 95616
ApstraAct: Two hundred and eighty-eight specimens representing six populations of the golden trout (Salmo aguabonita Jordan) from the Sierra Nevada, California, Were examined for similarities in 11 meristic characters. On the basis of mean similarity and phenetic relationships estimated from Euclidian distances, the six populations were divided into three distinct taxonomic groups. Two populations sampled from the eastern Kern River basin, and one from the Owens River drain- age, were identified as the subspecies S. a. aguabonita. Two populations, sampled from the Little Kern River basin, displayed characteristics which tended toward those reported for S. gairdneri Richardson, and were suspected of having a rela- tively recent hybrid origin. The final population, sampled from the headwaters of a stream tributary to the Little Kern River, was tentatively classified as the threatened Little Kern golden trout, S. a. whitei Evermann. The latter classifica- tion is in contrast to an earlier one that held the Little Kern golden trout to be synonymous with the Kern River rainbow trout, S. g. gilberti Jordan.
INTRODUCTION
The historic distribution and zoogeographic relationships among many salmonine fishes are confounded both by the ‘coffee pot’ transfers of fish by the early settlers of the late 1800’s and by the introduction of nonnative, hatchery-reared trout for recreational purposes. Moreover, the absence of
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244 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
complete biological isolating mechanisms among salmonine species, resulting in numerous instances of interspecific hybridization, has further confounded attempts to discern the historic distributions of individual species.
The systematic status and distribution of the golden trout of the Sierra Nevada has been in dispute since the first taxonomic descriptions (Evermann, 1905; Ellis & Bryant, 1920). Currently, the golden trout are classified as one species, Salmo aguabonita Jordan, comprised of two subspecies; S. a. agua- bonita of the Golden Trout Creek, Cottonwood Creek, and South Fork of the Kern River drainages (Curtis, 1935), and S. a. whitei Evermann of the Little Kern River basin (Miller, 1950; Shapovalov, Dill, & Cordone, 1959). Populations of S. a. aguabonita are distinguished from those of S. a. whitei on the basis of less intense spotting, greater brilliance in life colors, and geographic isolation (Evermann, 1905).
Recently, Schreck and Behnke (1971) and Legendre, Schreck, and Behnke (1972) have elaborated not only on the above differences but reported sharp distinctions for a number of meristic characters. Based on their observations, Schreck and Behnke (1971) suggested synonymy of S. a. whitei with the Kern River rainbow trout, S. gairdneri gilberti Jordan, and thus reclassified S. a. whitei to S. a. gilberti. However, their revision was based almost entirely on similarities in the number of lateral scale rows between specimens sampled from the Little Kern River basin during 1967-1969 and a few specimens collected from the Little Kern River and the main Kern River in 1893 and 1904. In addition, observations made during a helicopter flight over the Little Kern River basin led them to the erroneous conclusion that no significant barriers to fish migration existed between the main Kern River and the Little Kern River.
Until recently, a complete knowledge of the general topography of the Little Kern River drainage was not available, and an intimate understanding of the locations of natural barriers restricting directional fish migration was lacking. A thorough survey in 1973 (Evans, Smith, & Bell, 1973) has re- vealed the existence of several natural barriers throughout the watershed, not only near the confluence of the Little Kern River and the Kern River but in most streams tributary to the Little Kern River. The latter findings have two important consequences. First, the presence of barriers restricting fish migration into the Little Kern River basin from the Kern River raises a serious question regarding Schreck and Behnke’s contention of unrestricted gene flow between Little Kern and Kern River trout, and hence to their proposed reclassification of S. a. whiteiz. Secondly, the demarcation of tributary streams throughout the basin into several discrete regions suggests the existence of population subdivisions which would require definitive sampling to ascertain population status and distribution.
Vor. XL] GOLD AND GALL: GOLDEN TROUT POPULATIONS 24
Un
A second source of confusion regarding the status of the Little Kern golden trout stems from the possible hybridization between endemic golden and rainbow trout, S. gairdneri Richardson, introduced for recreational pur- poses. From 1931-1941, almost 100,000 rainbow fingerlings were planted yearly in various streams in the Little Kern River basin (Dill, 1945). Dill (1945 & 1950) and D. P. Christenson (personal communication) have sug- gested that the extensive phenotypic heterogeneity which they observed among the Little Kern trout was due to hybridization and subsequent backcrossing of planted rainbow to endemic golden trout. Although no critical evidence sup- porting successful golden X rainbow hybridization is available, it is generally assumed that extensive crossing occurs (Dill, 1945 & 1950; Needham & Gard, 1959; Schreck, 1969; Schreck & Behnke, 1971). Furthermore, the success of other salmonid hybridizations (Buss & Wright, 1956; Gould, 1966) suggests that isolating mechanisms among salmonids are far from complete.
The purpose of the present investigation was to examine the trout in the Little Kern River basin to discern whether the presumed hybridization between endemic golden and introduced rainbow trout had resulted in sig- nificant alterations in or the loss of S. a. whitei from the basin. This report presents the results of an analysis based on meristic characteristics; an analysis of the chromosome karyotypes has been presented elsewhere (Gold & Gall, 1975).
MATERIALS AND METHODS
Mopet. Studies of hybridization in teleosts have traditionally relied on estimates such as hybrid indices (Hubbs, 1955) and discriminant functions (Smith, 1973) to detect hybrid individuals. These methods require reliable estimates of the parametric values for all characters in each parental popula- tion. Since reliable taxonomic data were not available for either S. a. whitei or the rainbow trout planted in the Little Kern River basin during 1931— 1941, these methods seemed untenable. Furthermore, it was questionable whether these approaches would be valid if taxonomic data were available since about 10 generations had passed since the last rainbow introductions, and backcrossing of hybrid individuals to endemic goldens would surely have occurred.
As an alternative approach, an operational model of population diversity was derived which could be tested through appropriate sampling. The model was based on two observations. First, California Department of Fish and Game personnel and the members of the 1973 Little Kern River basin survey team indicated that while most of the Little Kern trout were pheno- typically heterogeneous, several small isolated populations which might repre- sent ‘pure’ S. a. whitei existed in the headwaters of various streams tributary
246 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
to the Little Kern River. Secondly, Department of Fish and Game records showed that waters inhabited by the subspecies S. a. aguabonita had not re- ceived plantings of rainbow trout; therefore populations of this subspecies could be assumed to represent S. aguabonita.
The model had four premises: 1) If golden by rainbow crossing occurred in the majority of the waters of the Little Kern River basin, as proposed by Dill (1945), then endemic S. a. whitei should be represented only by populations into which individuals from introgressed populations could not migrate; 2) A comparison of rainbow-trout free, geographically isolated populations of S. a. aguabonita should define the degree of naturally occurring diversity to be expected among golden trout; 3) The diversity among S. a. aguabonita populations should be less in relative degree than that expected between S. a. whitei and introgressed populations; 4) Isolated S. a. whitei populations should be more closely related to their sister subspecies, S. a. aguabonita, than to introgressed trout from adjoining waters. It should be noted that this latter premise is in disagreement with that of Schreck and Behnke (1971) who reasoned that S. a. aguabonita and the Little Kern golden trouts ‘‘. . . represent two independent invasions by already divergent forms of the golden trout complex.” Most of their 1967-1969 collections, however, came from waters accessible to planted rainbow trout.
SAMPLING. The locations of the collection sites are shown in figures 1, 2, and 3, and detailed descriptions are presented in appendix tables 1 and 2. Within the Little Kern River basin (fig. 2), one sampling was made from the Little Kern River (LAR) below the mouth of Soda Springs Creek (Zone 1) and another from lower Soda Springs Creek (LSSC) just above the mouth (Zone 2). A third sampling was made near the headwaters of Soda Springs Creek (USSC) above a natural barrier to upstream migration (Zone 3). The approximate locations of rainbow trout introductions during 1931-1941 are also shown in figure 2.
The presence of a barrier restricting upstream migration of trout from sites where rainbows were introduced identified the USSC sample as a po- tential population of S. a. whitei. The barrier at the mouth of Soda Springs Creek, separating LSSC from LKR, was constructed by the U.S. Forest Service in 1970. Consequently, trout from both LSSC and LKR represent the descendants of endemic golden trout and introduced rainbow trout except that LSSC might have received more immigrants from USSC if downstream migration occurs.
Three populations of S. a. aguabonita also were sampled. One was ob- tained from Golden Trout Creek (GTC) at a point adjacent to the South Fork of the Kern River where a second sample (SFA) was obtained. These sites are shown in figures 1 and 3 as Zones 4 and 6, respectively. The third
Vor. XL] GOLD AND GALL: GOLDEN TROUT POPULATIONS 247
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Ficure 1. A map showing the relative locations of the six populations of golden trout. The collection sites are identified as Zone 1-6.
sample was obtained from Cottonwood Creek (CWC) of the Owens River drainage, downstream from the Cottonwood Lakes (Zone 5). Although these three populations are isolated geographically, all share an interrelated history. The low alluvial ridge separating Golden Trout Creek from the South Fork of the Kern River was tunneled in the late 1800’s allowing a short period of exchange between the two drainages (Evermann, 1905). Furthermore, at one time the upper South Fork of the Kern was part of the Golden Trout Creek drainage (Lawson, 1904) indicating that trout from these waters may have a common origin. The CWC trout are ancestrally related to those of GTC and SFK since they are descendents of an 1870’s ‘coffee pot’ trans- plant of 12-13 trout from Mulkey Creek, a tributary of the South Fork of the Kern (Evermann, 1905; Ellis & Bryant, 1920). Since the CWC and
248 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
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Ficure 2. A map of the Little Kern River basin showing the locations of three collec- tion sites (Zone 1-3), the locations of natural barriers to upstream migration, and the approximate sites of rainbow trout introductions during 1931-41.
Cottonwood Lakes waters were barren of fish prior to the transplant, the CWC population was apparently founded by a small sample of fish.
All samples were collected in June, 1973, by electroshock and angling. The specimens were brought live to the trout hatchery at the Fisheries Biology Research Facility at Davis and held until sacrificed. An additional sampling was made at the USSC site in August, 1973. All but a few of the specimens were found to be sexually mature.
Vor. XL] GOLD AND GALL: GOLDEN TROUT POPULATIONS 249
N S COTTONWOOD LAKES OHN MUIR SEQUOIA ie .
NATIONA o GOLDEN TROUT WILDERNESS Sea gS CAMP - ZONE 5 conean Ww E ea ~7T0.000' Vk, i ne—- \ { ) > 5 \ \ iG ] \ 7 Cw ‘“ > q \ w ~ — a \ \ = y 1 S ( ~ ‘ 1 q mae ) S fe ] ) 1900 a 892— 1906 Nee = z S SEQUOIA inva NATIONAL - NATIONAL FOREST FOREST
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1— MAN-MADE TUNNEL CONSTRUCTED BETWEEN GI.C. AND S.FKR. IN 1892; TUNNEL CLOSED IN 1900
2— CWC. ORIGINALLY STOCKED WITH 13 TROUT TRANSPLANTED FROM MULKEY CR., IN 1876
FicureE 3. A map of the upper Kern River basin and the Cottonwood Lakes area showing the location of three collection sites (Zone 4-6), the location of an 1890 tunnel, and Mulkey Creek.
MeEtTHoD oF ANALysIs. Following sacrifice, specimens were tagged for identification, preserved in 10% formalin for one week, and then transferred to 40% isopropyl alcohol. Measurements and counts of meristic characters were taken from the left side. All specimens were examined in a random sequence and identified only by tag number. The twelve characters analyzed
250 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
and the methods of scoring were: Fork length—the distance in millimeters from the tip of the snout to the fork in the caudal fin; Pyloric caeca—each complete tip counted as a single caecum; BO rays—all branchiostegal rays including the most anterior (short) rays; Fin rays—the principal rays of the ventral (pelvic), dorsal, anal, and pectoral fins, counted under a dissecting microscope; Vertebrae—all ossified centra (using radiographs) ; Gill rakers—all gill rakers, including rudiments, on the first gill arch; Scales along LL—the number of oblique scale rows, 2—4 rows above the lateral line, from the anterior- most scale touching the shoulder girdle to the last scale at the structural caudal base; Scales above LL—the number of scales above the lateral line counted obliquely down from the origin of the dorsal fin to, but not including, the lateral line scale; Parr marks—the number of well-defined marks extending both above and below the lateral line.
All data were subjected initially to frequency distribution analysis using the mean, variance, and Fisher’s third and fourth moment statistics. Homo- geneity of variances among the six samples was tested using Bartlett’s method, and homogeneity of means was tested using single classification analysis of variance. If significant heterogeneity among means was detected, mean separa- tion was accomplished using Duncan’s multiple range test weighing the least significant ranges for unequal sample sizes (Sokal & Rohlf, 1969). All statistical analyses were carried out by computer using modifications of pro- grams found in Sokal and Rohlf (1969).
RESULTS
DISTRIBUTION OF THE CHARACTERS. Evaluation of the distributions of all 12 characters in each sample indicated that only the number of parr marks was non-normally distributed. Right skewness and leptokurtosis in four of the six samples suggested that the underlying biological phenomenon in parr mark genesis may not follow the assumptions of the normal probability density function. However, the deviations from normality were generally slight, and in the case of LAR and LSSC resulted from the inclusion of speci- mens with no visible parr marks. This absence of parr marks on some LKR and LSSC specimens may indicate the presence of S. gairdneri influence since parr marks are not retained into adulthood in that species.
VARIABILITY OF THE CHARACTERS. Estimates of the within sample variances of each of the 12 characters are presented in table 1. The variances among samples were homogeneous for fork length, BO rays, dorsal and_ pectoral fin rays, gill rakers, and scales above LL. The variances of pyloric caeca, vertebrae, ventral fin rays, and scales along LL were significantly hetero- geneous at the 5% level; whereas anal fin rays and parr marks were hetero- geneous at the 1% level. Since Bartlett’s test is unduly sensitive to even slight
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252 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
departures from normality, the significance of these results may have resulted from either small sample sizes or from sampling error. In the case of parr marks, the heterogeneity was undoubtedly due, in part, to the non-normality of the parr mark distribution and raises a serious doubt regarding the validity of number of parr marks as a basis for population separation. Examination of the sample variances for anal fin rays suggested differences in variance between the Little Kern River basin samples and those of S. a. aguabonita. Since separation into these two sets correlates with the geographical separation of S. a. aguabonita from S. a. whitei, this observation may reflect a difference in variability at the subspecies level. Why this should be evident in anal fin ray number alone is unclear.
For the remaining characters with heterogeneous variances, sample variances tended to be higher for LAR and LSSC than for the other four samples with the exception of the high variance in ventral fin rays in USSC and the low variance in pyloric caeca in LSSC. The high variability observed for LKR and LSSC would be expected if the trout from LAR and LSSC represented ‘hybrid’ or introgressed populations (Anderson, 1949; Hubbs, 1955).
One notable exception to the above was the high variance in along LL scale row number in the SFK sample relative to that observed for USSC, GTC, and CWC. Two seemingly unrelated observations provide a_ possible explanation for this result. First, during an examination of individual scales, it was noted that the SFA specimens possessed an unusually high number of regenerated scales. Secondly, E. P. Pister (personal communication) has informed us that extensive electroshocking has been carried out in the past few years to remove brown trout (S. trutta) from the general area of the SFK collection site. It is possible that a regenerative response to replace damaged or lost scales caused by heavy electroshocking or excessive handling may have produced the increased variability.
One further result requiring explanation was the reduced variance in fork length observed for the CWC sample. Since the Cottonwood Creek region is the most accessible of the six regions sampled and probably the most in- tensely angled by fishermen, the lower variability and smaller average size (see below) was likely a function of the legal size limit (6 in. or 15 cm.) in effect prior to our sampling.
MEAN VALUES OF THE CHARACTERS. The observed means for the 12 char- acters in each of the six samples are given in table 2 along with the error mean square obtained in the analysis of variance. The levels of heteroscedasticity observed were considered to be within the limits of the robustness of the analysis of variance for all the characters except number of parr marks. Homogeneity of the parr mark distributions of the six samples was tested by the non-parametric Kruskal-Wallis one-way analysis of variance (Siegel, 1956).
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254 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.
Taste 3. Mean similarity matrix describing the distributions of means among the six samples of golden trout for 10 meristic characters.* Upper values in each comparison refer to the number of characters with similar means; lower values refer to the number of char- acters with significantly different means (P < 0.05).
Sample LSSC USSC GTC cwc SFK LKR 9 3 3 4 33 1 i 7 6 7 LSSC 5 4 5 4 5 6 5 6 USSC 6 2 4 4 8 6 GTC 8 9 2 1 CWC 8 2
* Fork length and number of parr marks were not included.
The corrected H statistic was highly significant (H = 257, X? = 11.1 at P = .01 with 5 df) demonstrating that the population distribution for number of parr marks was not the same for all six regions sampled. The heterogeneity was probably the result of both greater variability within the LAR and LSSC samples, and the higher number of parr marks on the USSC fish.
The mean values for 9 of the remaining 11 characters were found to be significantly heterogeneous (P < .05); differences were not observed in the numbers of anal fin rays or gill rakers. The significant heterogeneity found for fork length was due solely to the small mean size of the CWC fish. Al- though the differences are not completely consistent for all four characters, the data suggested that trout from the Little Kern River basin, i.e., LAR, LSSC, and USSC, tended to have a larger number of BO rays and a larger number of principal rays in the ventral, dorsal, and pectoral fins than did specimens of S. a. aguabonita.
The remaining four characters, pyloric caeca, number of vertebrae, scales along LL and scales above LL, consistently discriminated between the LKR and LSSC samples and that from USSC. The latter had significantly lower numbers of pyloric caeca and vertebrae and a larger number of scales both along LL and above LL. In addition, the pyloric caeca and number of scales observed for the USSC sample were very similar to that for GTC and SFK. A large number of lateral scale rows was a distinctive character of the CWC sample, and the USSC sample appeared to have a unique number of vertebrae.
PHENETIC DISTANCE BETWEEN GRoupS. Mean similarity and Euclidian distance matrices were generated from the data in table 2 in an effort to ex-
Vout. XL] GOLD AND GALL: GOLDEN TROUT POPULATIONS 255
LKR
LSSC
USSC
GTC
SFK
CWC
10 5 0 Number of similar means Ficure 4. Dendrogram from UPGMA average linkage clustering showing the relation-
ship among the six samples of golden trout based on the number of characters with similar mean values. The cophenetic correlation coefficient rcs is 0.911.
press more clearly the collective distributions of the characters among the six samples. Fork length and number of parr marks were omitted as characters in both matrices.
The mean similarity matrix (table 3) provided a qualitative expression of the degree of similarity among the samples. The paired numbers in the matrix represent, for each sample by sample comparison, the number of characters for which the sample means were not found to differ (upper number) and the number of characters for which the sample means did differ (lower number), based on a 5% level of significance. A dendrogram (fig. 4) was derived from the matrix, using the unweighted pair-group method using arithmetic averages (UPGMA) average linkage algorithm outlined in Sneath and Sokal (1973), to pictoralize the degree of association among samples based on the number of characters with similar means.
The method identified two closely aligned groupings or clusters. The LKR and LSSC samples were similar in mean value for 9 of the 10 characters; the GTC, SFK, and CWC samples shared 8 of 10 character means in common, although GTC and SFK were more similar to each other than to CWC. These two main groupings differed markedly from each other, being similar for only 3—5 of the 10 characters. The USSC sample, on the other hand, appeared to be as different from LAR and LSSC as from GTC, SFK, and CWC, being similar to each of the two main groupings for an average of only 4 of the 10 characters.
256 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Taste 4. Euclidian distance matrix of 10 meristic characters* for six samples of golden trout.
Sample LSSC USSC GTC cwc SFK LKR 4.4 22.4 24.7 38.5 De LSSC 22.2 24.0 38.1 22.0 USSC 10.2 18.3 11.5 (CANe 15.6 4.7 CWC 19.0
* Fork length and number of parr marks were not included.
Euclidian distance estimates were obtained for each comparison to provide a quantitative evaluation of the similarities among the samples. This estima- tion of phenetic distance considers differences in the magnitude of the means as well as the number of means involved. It is important to note that this method does not give equal consideration to all 10 characters but rather gives greatest weight to those characters which show the greatest differences.
The distance estimates were calculated using only those differences which were found statistically significant (P < .05) as the remaining differences were attributed to sampling variability. When sample means for a given character were not found to differ (table 2), the best estimate of the mean for each sam- ple involved was calculated as the average of the actual observed means. The sample means were then standardized to remove scaling effects by dividing the deviation of each sample mean from the grand mean of all samples by the standard deviation of a mean. The latter was estimated from the harmonic mean number of observations per sample and the error mean square from the analysis of variance. Finally, the distances shown in table 4 were calculated
from the formula: k=10 VY, Dy = ee Gan] k=1
where Dj; = the distance between the i'" and j'" sample Zi; — the value of the k character in the i™ sample Z;x = the value of the k‘" character in the j'® sample.
The values of D represent the phenetic distance in standard deviation units between any two samples in a 10 dimensional hyperspace (Sokal, 1961; Good- man, 1972).
The resulting dendrogram, obtained using the UPGMA average linkage algorithm, is shown in figure 5. The Euclidian distance estimates identified a close phenetic relationship between LAR and LSSC (4.4 units), and between
Vor. XL] GOLD AND GALL: GOLDEN TROUT POPULATIONS 257
LKR
ESSC
USSC
GTC
IK ———_—— |
CWC
Euclidian Distance
Ficure 5. Dendrogram from UPGMA average linkage clustering showing the relation- ship among the six samples of golden trout based on Euclidian distance. The cophenetic correlation coefficient rcs is 0.860.
GTC and SFK (4.7 units). USSC was found to be more closely related to the GTC-SFK group (10.8 units) than was CWC (17.3 units). The latter clustered with the GT'C-SFK-USSC group at 17.6 units. This is in contrast to the results obtained from the mean similarity matrix which demonstrated CWC joining the GTC-SFK cluster before USSC. This disparity apparently stemmed from the large difference in lateral scale rows between the CWC and the GTC and SFK samples in contrast to their high degree of similarity to the USSC sample. Finally, the LAR-LSSC group clustered with the GT'C-SFK-USSC- CWC group at 26.8 units, indicating a marked distinction between the two groups.
DISCUSSION
Sharp distinctions were observed among the six populations of golden trout. Significant differences were found in either the variance, the mean, or both for all meristic characters examined except number of gill rakers. The data indi- cated discrete groupings from which inference regarding taxonomic classifica- tion can be drawn. However, classification in the salmonine fishes, particularly among the subgenus Parasalmo, is at best partially subjective. Hubbs (1943)
258 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
considered that the practical consideration of degree of difference may be the most efficient method. Since there is little or no known genetic incompatibility among most western North American Salmo (the subgenus Parasalmo), the classifications discussed below are based on “degree of difference” and not on any previous evidence of isolating mechanisms creating genetic incompatibility.
The most evident subgroup comprised trout from GTC, SFK, and CWC which shared common means for 83.3% of the characters studied, but an average of only 38.9% with trout from LAR, LSSC, and USSC. Based on a comparison of these data with the meristic data provided by Schreck (1969) and Schreck and Behnke (1971), the GTC, SFK, and CWC trout were identified as the golden trout subspecies, S. a. aguabonita. Also, these trout displayed the brilliant coloration and sparse spotting characteristic of S. a. aguabonita (Ever- mann, 1905; Curtis, 1935).
One notable exception to the classification of CWC with GTC and SFK was the significantly greater number of lateral scale rows on CWC trout as compared to both GTC and SFK trout. Although the number of lateral scale rows for S. a. aguabonita is reported to range as high as 210 (Schreck & Behnke, 1971), the large mean value for CWC coupled with the low variance raises the question of whether these trout should be given a separate classification. The effect of these differences was evident in the Euclidian distance estimates. The CWC trout differed from those of GTC and SFK in number of scales both along LL and above LL; whereas the latter two differed only in the number above LL. However, separation on this basis alone does not seem warranted. Possible explanations for the large observed difference include: 1) non-random sampling combined with small sample size as only 25 trout were sampled from CWC; 2) a ‘founder’ effect since the Cottonwood Creek was initially founded with only 12-13 trout (Evermann, 1905; Ellis & Bryant, 1920); or 3) environmental modification due to a lower ambient temperature (Garside, 1966; Wallace, 1973) since CWC is over 1,000 feet higher in ele- vation than the GTC and SFK sites.
It is of interest that despite complete geographic isolation for over 80 years (about 30-40 generations) a high degree of similarity exists among the S. a. aguabonita populations. This suggests that the various selection pressures in each region are sufficiently similar to maintain a high degree of homogeneity in meristic characters. The limited distribution and narrow range of S. a. aguabonita support this hypothesis.
The situation involving the three samples from the Little Kern River basin is more difficult to interpret. The LAR and LSSC trout were indistin- guishable for 9 of the 10 characters, differing only slightly in number of ventral fin rays, and comprised a second major subgroup. They differed from trout in the other four samples for an average of 61.2% of the characters studied and were found to be only distantly related to them phenetically.
Vor. XL] GOLD AND GALL: GOLDEN TROUT POPULATIONS 259
The final subgroup, trout from USSC, differed markedly from the other two major subgroups in mean similarity (60% of the characters), but was found to be closely related to S. a. aguabonita from GTC, SFK, and CWC in terms of Euclidian distance. It is, therefore, tentatively proposed that the USSC trout represent the endemic Little Kern golden trout, S. @. whitei Ever- mann, a conclusion based largely on the close phenetic relationship to S. a. aguabonita. However, there also was a remarkable similarity in the coloration and spotting of the USSC specimens to the original color plate of S. whitei shown in Evermann (1905).
What about the trout from LAR and LSSC? Originally, the Little Kern River basin was thought to include only golden trout of the subspecies S. a. whitei, having differentiated from S. a. aguabonita primarily in coloration and spotting (Evermann, 1905). Clearly, the present fish sampled from LAR and LSSC were only distantly related phenetically to those from USSC. More- over, the means for many of the characters, particularly pyloric caeca, number of vertebrae, and scales along LL and above LL, tended to be intermediate between those observed for S. a. whitei and those reported for rainbow trout, S. gairdneri (see Needham & Gard, 1959; Schreck & Behnke, 1971). This intermediateness suggested that trout in LAR and LSSC may have had a hybrid origin.
Schreck and Behnke (1971), following a study of trout from the Little Kern River basin, considered that S. a. whitei and the Kern River rainbow trout, S. gairdneri gilberti (Jordan & Henshaw, 1878; Evermann, 1905), were synonyms, and thus proposed the classification S. a. gilberti. To support this revision they noted that the ranges and means for certain meristic characters, principally oblique lateral scale rows, of trout collected in the Little Kern River in 1893 and the Kern River in 1904 were not apparently different from a limited sample collected from the Little Kern River basin in 1967-69. They further noted that there was no evidence that trout from the Kern River and the Little Kern River were isolated from each other.
For those meristic characters reported, the mean values for the LAR and LSSC samples were similar to those described for S. a. gilberti by Schreck and Behnke (1971). They acknowledged the possibility that these fish were of hybrid origin but dismissed it since some early specimens of S. a. gilberti were found to have basibranchial teeth, a character they felt demonstrated a primitive golden trout-like state.
The present finding in upper Soda Springs Creek of an exceptional golden trout population phenetically less similar to the proposed “S. a. gilberti”’, resident only a few miles downstream, than to the geographically distant S. a. aguabonita, raises a serious question as to whether the LAR and LSSC trout are in fact an integral part of the golden trout complex referred to by Legendre, et al. (1972). The recent survey of the Little Kern River basin (Evans, e¢ al.,
260 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
1973) has revealed the existence of significant barriers near the mouth of the Little Kern River which restrict the migration of trout from the Kern River into the Little Kern River. Therefore, there has not been a free exchange of genes between Kern River and Little Kern River populations, at least for an indefinite period of time. Based on these considerations, it is unlikely that the Kern River rainbow, S. g. gilberti, is synonymous with S. a. whitei.
Further, one of us (JRG) has examined a few of the specimens from the early Kern River and Little Kern River collections, now maintained at the California Academy of Sciences. On one specimen, IU 1113 from the 1904 Little Kern River collection, a lateral scale row count showed in excess of 175 scale rows, a count in sharp disagreement with those of Schreck and Behnke (1971) who reported a mean of 159 and a maximum of 169 lateral scale rows from these trout. However, the condition of most specimens precluded an accurate count and comparison of early specimens with fresh collections seems a dubious prospect.
Finally, no evidence of basibranchial dentition was found on any of the specimens in our collections. Since the same was stated for S. a. aguabonita by Schreck and Behnke (1971), it appears that basibranchial dentition may not be a golden trout characteristic.
In summary, the presence of the unique upper Soda Springs Creek golden trout population suggests that a ‘pure’ Little Kern golden trout still persists in the Little Kern River basin. The origin of the LAR and LSSC trout is speculative. Their intermediateness between S. a. whitei and S. gairdneri is strongly suggestive of a hybrid origin. Since numerous S. gairdneri were introduced throughout waters of the Little Kern River basin during 1931— 1941, hybridization between endemic S. a. whitei and the introduced rainbows could have produced the trout now present in the Little Kern River. If the foregoing hypothesis is true, then the impassable barrier separating the upper Soda Springs Creek trout from those downstream has prevented any possible genetic contamination and preserved a vestige population of the original Little Kern golden trout, S. a. whitei. Further samplings of other isolated headwater populations throughout the Little Kern River basin should test this hypothesis.
ACKNOWLEDGMENTS
The authors wish to express their gratitude to D. P. Christenson, S. J. Nicola, E. P. Pister, T. Mills, and R. C. Smith for their assistance in pro- curing the specimens for this study. Miss Julie Kent is also to be thanked for the preparation of the maps in figures 1, 2 and 3. Mr. L. Shapovalov and Drs. W. E. Eschmeyer and R. R. Miller have read the manuscript and pro- vided many illuminating comments which were appreciated and incorporated into the manuscript. The project was partially supported by the Department of Fish and Game, The Resources Agency of California, Contract No. S—813.
VoL. XL] GOLD AND GALL: GOLDEN TROUT POPULATIONS 261 APPENDIX TABLE 1. Geographic location* of collection sites.*
Collection Longitude Latitude Altitude ; ; site W N (in feet) County Area ZONE 1 118°30’43” BI UH 6,160 Tulare Sequoia
(Little Kern National River) Forest ZONE 2 IWS 3427 SOmelt5 eS 6,400 Tulare Sequoia (Lower Soda National Springs Creek) Forest ZONE 3 118°35’47” 36° 1849” 8,800 Tulare Sequoia (Upper Soda National Springs Creek) Park ZONE 4 Sel Oe Ome 54 8,800 Tulare Inyo (Golden Trout National Creek) Forest ZONE 5 AS S03 277 8 OmZieA Ge 10,000 Inyo Inyo (Cottonwood National Creek) Forest ZONE 6 LTO? 36°21'49” 8,800 Tulare Inyo (South Fork of National the Kern River) Forest * Geographic locations were taken from U.S. Geological Survey, 15 minute series topographic maps, scale 1:62,500. + Fish collections in Zones 1, 3, and 6 were made in about 0.25 miles of stream; collections
from Zone 2 were made in about 0.7
APPENDIX TABLE 2.
Ss miles of stream.
The linear, cross-country distances (miles) between the collection
sites. The three values below the diagonal refer to stream distances (miles).
ZONE 1 ZONE 2 ZONE 3 ZONE 4 ZONE 5 ZONE 6 Location (LKR) (LSSC) (USSC) (GTC) (CWC) (SFR) Kernville, California 34 36 40 43 51 43 ZONE 1 (LKR) 2.4 Hell 15.5 25 US) ZONE 2 (LSSC) 2.9 4.9 15.0 24 14.9 ZONE 3 (USSC) 9.6 6.7 WA fo) 25 7/all ZONE 4 (GTC) 9.7 0.4 ZONE 5
(CWC) 9.7
262 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
LITERATURE CITED ANDERSON, E. 1949. Introgressive hybridization. John Wiley and Sons, Inc., New York; Chapman and Hall, Limited, London. 109 pp. Buss, K., and J. E. Wricut 1956. Results of species hybridization within the family Salmonidae. The Progressive Fish-Culturist, vol. 18, no. 4, pp. 149-158.
Curtis, B. 1935. The golden trout of Cottonwood Lakes. California Fish and Game, vol. 21, no. 2, pp. 102-109. Dr, W. A.
1945. The Little Kern River drainage, Tulare County. Progress Report No. 2, Cali- fornia Department of Fish and Game, Inland Fisheries Administrative Report 45-29, 19 pp. (Mimeo.) 1950. A preliminary report on the status of the golden trout fishery of California. California Department of Fish and Game, Inland Fisheries Administrative Report 50-44, 28 pp. (Mimeo.) Eviis, S. L. N., and H. C. Bryant 1920. Distribution of the golden trout in California. California Fish and Game, vol. 6, no. 4, pp. 142-152. Evans, W. A., R. C. SmitH, and M. BELi 1973. A reconnaissance survey of the fish resources of the Little Kern River drainage, California. United States Department of Agriculture Forest Service, Region 5, and California Department of Fish and Game, Draft (Preliminary Report), 41 pp. EVERMANN, B. W. 1905. The golden trout of the southern High Sierras. Bulletin of the United States Bureau of Fisheries, vol. 25, pp. 1-51. Garsipg, E. T. 1966. Developmental rate and vertebral number of Salmonids. Journal of the Fisheries Research Board of Canada, vol. 23, no. 10, pp. 1537-1551. GoLp, J. R., and G. A. E. Garr 1975. Chromosome cytology and polymorphism in the California High Sierra golden trout (Salmo aguabonita). Canadian Journal of Genetics and Cytology, vol. 17, pp. 41-53. GoopMAN, M. M. 1972. Distance analysis in biology. Systematic Zoology, vol. 21, pp. 174-186. Goutp, W. R. 1966. Cutthroat trout (Salmo clarkii Richardson) x golden trout (Salmo aguabonita Jordan) hybrids. Copeia, no. 3, pp. 599-600. lations, (C, We 1943. Criteria for subspecies, species, and genera, as determined by researches on fishes. Annals of the New York Academy of Sciences, vol. XLIV, art. 2, pp. 109-121. 1955. Hybridization between fish species in nature. Systematic Zoology, vol. 4, pp. 1-20. Jorpan, D. S., and H. W. HensHaw 1878. Report upon the fishes collected during the years 1875, 1876 and 1877, in California and Nevada. Annual Report of the United States Chief of Engi- neering, 1878, pt. 3, pp. 1609-1622.
Vor. XL] GOLD AND GALL: GOLDEN TROUT POPULATIONS 263
Lawson, A. C. 1904. The geomorphogeny of the upper Kern basin. University of California Publica- tions, Bulletin of the Department of Geology, vol. 3, no. 15, pp. 291-376. LEGENDRE, P., C. B. ScHreck, and R. J. BEHNKE 1972. Taximetric analysis of selected groups of western North American Salmo with respect to phylogenetic divergences. Systematic Zoology, vol. 21, pp. 292-307. Mirter, R. R. 1950. Notes on the cutthroat and rainbow trouts with the description of a new species from the Gila River, New Mexico. Occasional Papers of the Museum of Zoology, University of Michigan, no. 529, pp. 1-42. NEEDHAM, P. R., and R. Garp 1959. Rainbow trout in Mexico and California: With notes on the cutthroat series. University of California Publications in Zoology, vol. 67, pp. 1-124. ScHReEck, C. B. 1969. Trouts of the upper Kern River basin, California. M.S. Thesis, Colorado State University, Ft. Collins, Colorado, 120 pp. ScHRECK, C. B., and R. J. BEHNKE 1971. Trouts of the upper Kern River basin, California, with reference to systematics and evolution of Western North American Salmo. Journal of the Fisheries Research Board of Canada, vol. 28, pp. 987-998. SHAPOVALOV, L., W. A. Diz, and A. J. CoRDONE 1959. A revised checklist of the freshwater and anadromous fishes of California. California Fish and Game, vol. 45, pp. 159-180. SIEGEL, S. 1956. Non-parametric statistics. McGraw-Hill Book Company, New York-Toronto- London, 312 pp. SMITH, G. R. 1973. Analysis of several cyprinid fishes from Western North America. Copeia, no. 3, pp. 395-410. SNEATH, P. H. A., and R. R. Soka 1973. Numerical taxonomy. W. H. Freeman and Company, San Francisco, 573 pp. Sokal, R. R. 1961. Distance as a measure of taxonomic similarity. Systematic Zoology, vol. 10, pp. 70-79. Socal, R. R., and F. J. Rouir 1969. Biometry. W. H. Freeman and Company, San Francisco, 776 pp. WALLACE, C. R. 1973. Effects of temperature on developing meristic structures of smallmouth bass, Micropterus dolomieui WLacepede. Transactions of the American Fisheries Society, vol. 102, no. 1, pp. 142-144.
PROCEEDINGS OF THE | al
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 11, pp. 265-334; 25 figs.; 4 tables. October 3, 1975
THE SCORPAENID FISHES OF THE HAWAIIAN
ISLANDS, INCLUDING NEW SPECIES AND NEW RECORDS (PISCES: SCORPAENIDAE)
Hy William N. Eschmeyer
California Academy of Sciences, Golden Gate Park, San Francisco, California 94118
and
John E. Randall Bernice P. Bishop Museum, Honolulu, Hawaii 96818
Agstract: The number of species of the fish family Scorpaenidae now known from the Hawaiian Islands totals 26. This includes two new species (Scorpaenopsis brevifrons and Scorpaena pele), and four new records (Phenacoscorpius nebris, Scorpaenodes corallinus, Scorpaenodes hirsutus, and Scorpaenodes littoralis). In addition, the following are treated as valid Hawaiian species: Dendrochirus barbert, Ectreposebastes imus, Iracundus signifer, Neomerinthe rufescens, Plectrogenium nanum, Pontinus macrocephalus, Pterois sphex, Rhinopias xenops, Scorpaena ballieui, S. colorata, S. coniorta, S. galactacma, Scorpaenodes kelloggi, S. parvipinnis, Scor- paenopsis altirostris, S. cacopsis, S. diabolus, S. fowleri, Setarches guentheri, and Taenianotus triacanthus. All species are at least briefly described, all are figured, and a key is provided. Synonymies based on Hawaiian references are included for all species; non-Hawaiian references and new Indo-Pacific records are given for a few species.
TABLE OF CONTENTS Included in brackets are junior synonyms with Hawaiian type localities
Tirantbyeray ervey: ca ei SR Sie oe a a eee EN Tbr anal SG RTOS EGS 9 <a sen I a ee ee Net Oc Seen ater ee tO ee wa pad 9 tn Dn Dna iE eg Rie ah drt es Nt = Keyatonscorpiontishessotather awaltanw slams) sss ee ne
[265]
266 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Cenusmeteros Oken ee een, Se ae aes ESE 2 2 ee 271 micnOiasp ies lordanvandusvermann = eee 272 (CErnism Der OnOGHIMUSMOWAINSOM 22.222 2 ee OS Dendrochirus barberi (Steindachner) |D. hudsoni Jordan and Evermann, D. GHVONE US MCNIKANS || ee a8 ee ha ee eee DiS (Genusms con aetodess Bleecker) 20 eer 6 6 ee ee eee eee Dili] Scorpaenodes hirsutus (Smith) — ge i Bt ee Se 277 Scorpaenodes kelloggi (Jenkins) — Le eee tee pv ae 279 SGonpaeno nes parvipinnises((Ganrett)) see ee 280 SGOMPUCIVOGMES MITE GON CITS ian (MANN Kel) eee Sg 282 SOA DARIO (COPA Syoovhl\ e e 283 Genuswlectrocenvin (Gilbert =e ee TM a a er 284 Rlectrozenium nanum (Gilbert) =e = ee ee eee 284 Genus) *iaenanotussitacépede» 2. 2 = 2 2 ee ee ee a ee eee eee 285 Taenianotus triacanthus Lacépéde |7. garretti Giinther, T. citrinellus Gilbert] —— 285 (Conus Mnacrra@s \\@rmcben eingel IDxierAene vo 287 Inicio SaeaaP Wola gincl IB\yemeneiy 287 (Cems Searelses OnmeOm — pt eS 289 Setarches guentheri Johnson [S. remigera (Gilbert and Cramer) ] — 289 Genuswe cine p ose bas tess Garin cn eee ee 291 Ectreposebastes imus Garman __...___ 2 in Ws ARIE SE os er 291 GenussBhenacoscorpius, Rowler 22 == ss ee ee ee eee 293 PUN CTHOACD SDE DLIS GCM DIS NOONE 293 Genus.;Pontinus Roby =2-5-- 08. 2 ew kd EE ee ee 294 Pontinus macrocephalus (Sauvage) [P. spilistius Gilbert] 294 Cons Saornirnopys Isieekell aint. Se 296 Scorpaenopsis fowleri (Pietschmann) — noo ge ee 297 Scorpaenopsis brevifrons Eschmeyer and Randall, new species 299 Scorpaeno psis diabolus Cuvier [S. catacola Jordan and Evermann| — 305 SCOAVUCHIODSIS CUCOWSOS Seman 308 SCO; MEMO NSOS Chiacostids AChE 2 309 Genus Rhinopias Gill — So ac ee a 310 Rhinopias xenops (Gilbert) etic Se ee ee 311 Genus Neomerinthe Fowler ____ a eee ee es 312 Weomermtihesraurescens. ((Gillyert) yee 313 Genus’ Scorpaenasinnaeus: == ee 314 SCO NC HO (Monn Haner ae! (A\EMRNNS)) a 314 MEO ANU Conmorads (V\ermllsinS) . 316 Scorpaena ballieui Sauvage [S. corallicola Jenkins, S. peruana Hildebrand, ? S. iy NAA on lexeiaveve(iel|| eee I ee Dee ee wae oe We 317 Scorpaena pele Eschmeyer and Randall, new species _.....___ = 320 Scorpaena colorata (Gilbert) — ogres eon be Se ee 326 Iiteratunes Cited) Sasa sees ae ot ee eee 327 INTRODUCTION
The shallow-water fish fauna of the Hawaiian Islands is of particular interest because of the high level of endemism associated with its isolated geographic location. The family Scorpaenidae is representative, for nine of a
VoL. XL] ESCHMEYVER AND RANDALL: HAWAIIAN SCORPAENIDAE 267
total of 26 species presently are known only from Hawaiian waters. Two species are known only from Hawaii and Japan. The remaining species of scorpaenids occur elsewhere in the Indo-Pacific region and two of these also occur in the Atlantic. Certain groups of scorpionfishes are absent from the Hawaiian Islands but are present in the central and western Pacific; for example, no stonefishes (Synanceiinae) occur in Hawaii.
The scorpionfishes of Oceania are not well known except at the Hawaiian Islands. This is particularly true of those occurring in depths below about 30 meters. We suspect that at least some of the deeper-living species known only from Hawaii will be found at other localities in the Pacific as appropriate depths are sampled.
Many of the scorpaenids which live at depths below about 50 meters were described from specimens collected by the United States Fish Commission steamer Albatross in 1901-1902. In the last few years, new material has been taken by the National Marine Fisheries Service vessel Townsend Cromwell, and these specimens have been made available to us. Extensive recent collect- ing in inshore waters to depths of about 200 feet using scuba gear has re- sulted in additional valuable material. Rather than report solely on the new species and new records of Hawaiian scorpionfishes, we have provided a synopsis of all Hawaiian species. Also, for some species, we have presented new records from other Indo-Pacific localities, as well as comments on possible synonyms or close relatives.
ACKNOWLEDGMENTS
Persons aiding the study were many. We especially thank Paul J. Struh- saker, National Marine Fisheries Service (NMFS), Honolulu, for making available specimens collected by the Townsend Cromwell. John Fowler (NMFS) provided fresh specimens of 2 offshore species collected by the Townsend Cromwell. William D. Madden provided information on /racundus signifer and provided the specimens of Phenacoscorpius megalops. We wish to thank the following persons for assistance during visits to museums: Alwyne C. Wheeler, British Museum of Natural History (BM (NH)); James E. Bohlke, Academy of Natural Sciences of Philadelphia (ANSP); M. L. Bauchot, Museum National d’Histoire Naturelle (MNHN), Paris; the staff of the United States National Museum of Natural History (USNM); William A. Gosline and George Losey, University of Hawaii (UH); Paul Kausbauer, Naturhistorisches Museum, Wien (NMW); and John R. Paxton, Australian Mu- seum, Sydney (AMS). R. J. McKay, Queensland Museum, Brisbane (QMB), generously loaned the type of Scorpacnopsis macrochir. Margaret M. Smith, Rhodes University (RU), Grahamstown, loaned a specimen of Scorpaenodes littoralis. Richard Rosenblatt, Scripps Institution (SIO), provided a_speci- men of Ectreposebastes niger. Bruce B. Collette, Systematics Laboratory,
268 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
National Marine Fisheries Service, arranged for a loan of a specimen of E£. niger and supplied data on the holotype of that species. We are indebted to various persons for aiding in scuba collections with Randall, particularly Gerald R. Allen, Deetsie Neil Chave, and Paul M. Allen. David T. Anderson assisted in analysis of the humpbacked species of the genus Scorpaenopsis. Melissa Barbour, Lillian Dempster, W. I. Follett, Warren Freihofer, Maurice Giles, James Gordon, Tomio Iwamoto, John McCosker, Cherryl Pape, Stuart Poss, Katherine Smith, Pearl Sonoda, and Beverly Wesemann of the Cali- fornia Academy of Sciences (CAS) aided the study as did Helen Randall from the Bernice P. Bishop Museum (BPBM). W. I. Follett and Lillian Dempster provided advice on nomenclatural problems. Kaza V. Rama Rao, Zoological Survey of India, provided comments on the manuscript. We es- pecially thank Lillian Dempster for assistance with literature and for re- viewing the manuscript.
Funds from a National Science Foundation grant (NSF 15811) allowed Eschmeyer to visit museums. Many of the shallow-water specimens were collected with grant support from the National Geographic Society and Na- tional Science Foundation to Randall for studies on other fishes.
METHODS
Measurements, counts, and terminology of head spines follow Eschmeyer (1969b) with a few modifications. Preorbital spines are here termed lachrymal spines. The last soft ray in the dorsal and anal fins in all Hawaiian species is a double ray, often appearing as two close-set rays. The fraction %, as in anal rays 5%, is used to draw attention to the fact that the last ray is double and should not be counted as 2 rays.
In MATERIAL EXAMINED sections we list representative specimens from typical habitats and more briefly indicate additional material. The number of specimens followed by the range in standard length is given in parentheses. If only the number of specimens is given, then no fin-ray counts were made on any of the specimens in that lot.
Abbreviations of depositories of specimens are given in the INTRODUCTION with the following exceptions: SU (Stanford University) and GVF (George Vanderbilt Foundation), these specimens now being housed at the California Academy of Sciences. Specimens listed as NMFS uncat. are now at the National Marine Fisheries Service Laboratory, Honolulu, but most of this material even- tually will be deposited at the United States National Museum of Natural History. Much of the University of Hawaii collection has been transferred to the Bernice P. Bishop Museum; the specimens reported with both UH and BPBM numbers were transferred, while those with only a UH number remain at the University of Hawaii.
VoL. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 269
— — Le Sy CORONAL “los = SPINES
\ = 2 NN SN Cos ~ 7
he 5 OS
la. 1b.
2a. 2b.
3a.
3b.
4a.
4b.
Sa.
6a.
6b.
Kry TO SCORPIONFISHES OF THE HAWAIIAN ISLANDS
Note. Consult figure 1 for diagrams accompanying key. ‘‘Deepwater’’ means that the species occurs deeper than about 50 meters. The coloration for most species is diagnostic and the reader should consult the appropriate figures.
DD OTSalespINeSwell Sia .cnssses seneerseeescte tee BR ea ST oe RSE REA ene ee eT ee eee 2 TD OTSA SPINES wales weeks. sree ede Reews snes a cea oe tek. eet ascehen oes i ee a Steen eae Ee Serre: Pectoral rays unbranched; 61% anal soft rays. ....... : .....Pterois sphex (figure 2a) Somempectoraleraysepnranched 52/0 alaaleSOlity may Syren see eecreereeeeas or eeeee serene eee es:
Dorsal spines long, longest nearly as long as or longer than depth of body............ Re Ni oes ok Pecan ee cat sdcbacosoespononseteneaneeuagesoconeeonoccened DA BOIFOCUEROS (xno? (Grune 7)s))) Dorsal soft rays short, the longest less than 4% depth of body.........000.0........ eee
Vertical scale rows (counted 2 or 3 rows above lateral line from above first lateral line scale to end of hypural) fewer than 32; dorsal soft rays usually 84.0.0... 5
Vertical scale rows more than 40; dorsal soft rays usually 914 (except Scorpaenodes GORGIITUS mawitlhe S/o) eee ee ae eee ee ee ener ata ea chhdic ta earn De ee pe teG
Pectoral rays 17-18; suborbital ridge with 4 spines plus extra spine below main rvabege {(GiteqbbRayy LIT)) cb yncboendseneatebeneecrec cece becca ee csescacee ..... Scorpaenodes hirsutus (figure 3) Pectoral rays 18-20, usually 18-19; suborbital ridge usually with 3 spines (fig- Line Mall) gare ena ete a ee a ences ike Tie ee -....Scorpaenodes kelloggi (figure 4) Suborbital ridge with more than 5 spinous points (frequently 10 or even 20 or IMOLE)) eae esac ecsesoret Renisk he Ne 6 tie pak na Scorpaenodes parvipinnis (figure 5a) Suborbital ridge with 3 or fewer spinous points... 2 a
7b.
8a.
8b.
9a.
9b.
10a.
10b.
lla.
11b.
12a.
l3a.
CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
. Dorsal soft rays 914; dusky spot on subopercle behind preopercular spines (fig-
WHA TIC) ~ scosooesoad oe Secon EDR PRES ERDEe co Ree PRen ntact oanireeee cena Scorpaenodes littoralis (figure 5b) Dorsal soft rays 84%; dark spot not present on subopercle behind preopercular SS TD TITLES Ne I ee Sect cn nt bw cleo cesar cadne sate ara erate Scorpaenodes corallinus (figure 5c)
Pectoral rays 22-23; dorsal soft rays 742; tenth dorsal spine very short, nearly separated from ninth and eleventh; a deepwater species ............00..000ccceeeee sosscceihcode See ie e R Ere OR are as es ma oe Oe Plectrogenium nanum (figure 6)
Pectoral rays fewer than 20; dorsal soft rays 9% or more; tenth dorsal spine joined by membrane to ninth and eleventh spines............... J hinlighee eet See eee 9
Pectoral rays 14-15, usually 14; body strongly compressed, width usually about one-fourth of body depth; body covered with rough papillae instead of normal
SOA CS te ee ee eee en ie Ded Taenianotus triacanthus (figure 7) Pectoral rays 16 or more; body not greatly compressed; body covered with nor- mal scales ............. sett dangdeas oh cece ceie. ete eaiabiok dedi e heme aalsuchdoncdersninalet x atk ee ee 10
Fourth dorsal spine especially elongate in specimens over about 60 mm. S.L. (smaller specimens may be identified solely by the next character); black pig- ment between spines 1-3 or 2-3 (figure 1d)................... Tracundus signifer (figure 8)
Fourth dorsal spine not especially elongate; no black spot between spines 1-3.......... 11
Lateral line a more or less continuous trough (covered by thin membranous scales which are usually lost on capture); head cavernous, with ossification weak;
Scales tinyspanrd “Cycloide tex cout se. cr esctan cocenstcncetseaiepesaiee eee eee eee 12 Lateral line normal, scales tubed; head not cavernous, ossification normal; scales Cy. ClOId OR MCLEM OTE Fee ce nieces stees goss east RTs ees va eae ee 13
Orbit diameter subequal to interorbital width; pectoral rays 20-22; anal soft rays
usually 544; a deepwater species.......0....0.00.0.0ccceeee Setarches guentheri (figure 9) Orbit diameter about one-half of interorbital width; pectoral rays 19-20; anal soft rays usually 6144; a deepwater specieS.........0.....0.00..... Ectreposebastes imus (figure 10)
Enlarged, black melanophores on caudal peduncle as in figure 11; lateral line incomplete, only 4 or 5 tubed scales present anteriorly; a deepwater species ....... eee ei ie See ae ee vooveettetvsteveetssitttssessessssssssseseee Phenacoscorpius megalops (figure 11)
. No enlarged black melanophores on caudal peduncle; lateral line complete................ 14
. Pectoral fin rays unbranched; a deepwater SpeCieS ...........0.......cccccccc cece ceeeteeeeteeees
NAD oe Re CORSE RE ay AEE hat Clr EIN Feel Pontinus macrocephalus (figure 12)
Upper pectoral fin rays branched (look carefully at tips) ....0..00.0000ccccce cee 15 Palatine teeth rabbsent!tnerisecncist osc ees Bee eee genus Scorpaenopsis............ 16 Palatine. teeth: presents ..:5202hicac. ho sdaecbavsaceautnetyss cose he es ee ee 20
. Pectoral rays 16 (15 or 17 should be expected occasionally); (a tiny species not
Exceeding sa boutesOlmimies Salas) eeeeeeeeeeeeeeeee sae Scorpaenopsis fowleri (figure 13) Pectoral srays, 7=19) smiostiys Sito Oi sey cee ee eee ee 17
Pectoral rays usually 19; dark spot frequently present at midheight of spinous
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 271
portion of dorsal fin between spines 3 or 4 to 6 or 7..
rs ech ene nen aoe ee Scorpaenopsis brevifrons new species Gicties 14-15) 17b. Pectoral rays usually 17 or 18, rarely 19; no dark spot present on spinous portion
ObRGOTSANE Tine. sees er eee hake tees Sees eh neces As eee ea Ade 18
18a. A hump behind head (as in figure 16); (see also figure 17b for coloration inside Otvpectoral fini) psc eee Meee es ee ... Scorpaenopsis diabolus (figures 16-17) TUES Dy. DNL SV aU a 0) ope cl heey 0 OS See ar a eee a i Re ere oe ee EAMES eel
19a. Eye diameter smaller than snout length; numerous tentacles present on lower jaw afagdecelthaoag soe RSanactac eet aadRtecch iS A RAEE PEEDE rene te oe tte Reena td Scorpaenopsis cacopsis (figure 18) 19b. Eye large, nearly 144 times snout; 1 or 2 tentacles present on lower jaw; a
GeepwatersspecieSs ee set eee oe Scorpaenopsis altirostris (figure 19) 20a. Head compressed and orbit elevated; vertical scale rows about 70; a deepwater
SDC CLES ieee tere eM etch Fs fanned nth ee ee cate Se tare Rhinopias xenops (figure 20) 20b. Head normal; vertical scale rows usually fewer than 50 ........0..........c.cccecceseeeecereeeeeee 21 21a. Pectoral rays 18-19; a deepwater species...................... Neomerinthe rufescens (figure 21) 21b. Pectoral rays 17 or fewer, very rarely 18; most species in shallow water................
REN a cnn oath teat tee es po ak ye eR A ee genus Scorpaena............ 22 UoteeS CAlESwOMNSIGES, GY, CLOIG. ... ...25--2=2.222s.-0ecs.0- sees beekoscesseetecesoy Scorpaena galactacma (figure 22a) DES CAlesmOnNe SiGeSmCtCNOLG! Mer... Sete ogertrhs eect ee eee he en ates See 2S 2B echoralnaysmusualliys WGsmsmallowy) Water) oss. ceeeeeeces ce cee ness eee estes ee eee 24 opm eectoralenaysp usually. W7i (deepwater 22. cecc-ceece-ese ee eee eee ese ee Fe es 5)
24a. Body and fins with conspicuous dark spots; coronal spines absent (not as in ITPA OTR “TYS)). > ae: Aa Re OR eee ps ae re Oa Scorpaena coniorta (figure 22b)
24b. Body and fins without conspicuous dark spots; coronal spines present (figure le) ee nS one CORSE Tae ted oh eis SOW ccestecet Scorpaena ballieui (figure 23)
25a. Scales on breast buried (area appears unscaled) ; 4 spines on suborbital ridge, the hinstmOnee lateral meta cen Otewla clirayare) lity O11 erecta tee eee ee Pe ear WIN Rit meee aoe oe Scorpaena pele new species (figures 24, 25a)
25b. Breast with obvious scales; 3 spines on suborbital ridge.......000...0..00..0cccccccecteeeeeeee ee aR ees ee a ao Ade Me 9 hit Scorpaena colorata (figure 25b)
Genus Pterois Oken
Pterois OKEN, 1817, p. 1182 [misprinted as p. 1782] (type-species Scorpaena volitans Lin- naeus by monotypy; based on Cuvier’s ‘Les Pterois’).
REMARKS. Generic synonyms are not included. The genus Pterois is very closely related to the genus Dendrochirus; the 2 genera are separated on the basis of long unbranched pectoral rays (a juvenile feature) in adults of the genus Pterois; some rays are branched in adults of the genus Dendrochirus. Juveniles are very similar in all features, and separate genera possibly may not be warranted. We do not recognize the genus Pferopterus Swainson as
22. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
distinct from Pterois as does Smith (1957b). Only one species occurs in Hawaiian waters.
Pterois sphex Jordan and Evermann.
(Figure 2a.)
Pterois sphex JorRDAN & EVERMANN, 1903a, p. 201 (original description; type locality Ha- waiian Islands, one specimen from Honolulu; holotype USNM 50650); Jorpan & EVERMANN, 1905, pp. 464-465, fig. 203 (description from Jordan and Evermann, 1903a; figure of holotype); JORDAN & SEALE, 1906, p. 379 (name only; Hawaiian Islands) ; JORDAN & JORDAN, 1922, p. 56 (listed) ; Fowler, 1925, p. 27 (listed); JorDAN & EVER- MANN, 1926, p. 10 (listed); Fowler, 1928, p. 294, pl. 35A (synonymy; description; photograph of colored cast); Tinker, 1944, pp. 270-271, fig. (brief diagnosis; figure from Jordan and Evermann, 1905); SmirH, 1957b, pp. 78, 81 (compared with P. russelli and P. mombasae); GOSLINE & Brock, 1960, pp. 284-286, 341, fig. 262 (brief mention; in key; line drawing); GosLInr, 1965, p. 825 (depth distribution).
MATERIAL EXAMINED. USNM 50650 (1, 112 mm. S.L., holotype of Pterois sphex), Oahu, Honolulu, D. S. Jordan and B. W. Evermann, 19 Jan. 1904. BPBM 7812 (1, 36.8), Hawaii, Kona Coast, off Puako, coral bottom in 15 meters, J. E. Randall, 7 Aug. 1969. BPBM 7813 (4, 37.1-68.5), Oahu, Pupakea and Waimea Bay, in 15 meters, rock ledge, J. E. Randall et al., 24 Aug. 1969. BPBM 7814 (1, 148), Oahu, off Makua, at base of ledge in 30 meters, G. R. Allen, 20 March 1969. BPBM 7882 (3, 16.8-27.7), Oahu, Waimea Bay, in 3.5—5.5 meters, G. R. Allen, 30 May 1968. BPBM 7883 (1, 30.0), Oahu, Kaneohe Bay, ledge, J. E. Randall and E. Chave, 10 Oct. 1969. BPBM 9771 (1, 70.0), Oahu, off Makaha Shores Condominium, small caves in reef in 14 meters, J. E. Randall and A. R. Emery, 26 Apr. 1970. BPBM 10625 (1, 44.8), Oahu, Waimea Bay, G. R. Allen, Sept. 1966. CAS 15721 (1, 23.5), Oahu, Waimea Bay, W. P. Davis, 15 July 1967. Additional mate- rial is available in the BPBM and CAS collections, and specimens collected by the Townsend Cromwell are present in the NMFS collection.
DISTINGUISHING FEATURES. Dorsal fin rays normally XIII, 10% (some- times XIII, 11%). Anal fin rays III, 6%. Pectoral fin rays 15-16, usually 16. Pectoral rays all unbranched, very long, and free from membrane distally. Dorsal spines very long, some about as long as body depth. Scales ctenoid, about 50-55 vertical scale rows. Coronal spines present. Most head spines become multiple with growth. Lachrymal and_ suborbital bones densely covered with spines at specimen length of 100 mm. S. L. Supraocular tentacles banded with black, frequently tentacles absent in large specimens. Coloration (fig. 2a) is also diagnostic (small specimens have fewer bars on pectoral and pelvic fins).
DistRIBUTION. This species is known only from the Hawaiian Islands. It has been collected by scuba diving in depths from about 3 to 30 meters. These collections were made during daytime hours, and the specimens were found
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE
274 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
TABLE 1. Counts of dorsal and anal fin rays in Hawaiian scorpionfishes (based only on Hawaiian specimens).
Dorsal Anal Anal spines Dorsal soft rays spines soft rays
12 143 7% 8% 9% 10% 11% 3 5% 6% 7%
Pterois sphex — 13 —- —- — 12 1 12 — 30 Dendrochirus barberi — 23 — 1 21 1 — 23 23 —- — Scorpaenodes hirsutus — 12 a 11 1 —-— — 12 12 —_- — Scorpaenodes kelloggi — 15 1 14 — — — 15 56 — — Scorpaenodes parvipinnis* 1 15 —- — 16 —- — 16 15 —- — Scorpaenodes littoralis —_ 10 —- — 10 —- — 10 9 1 —_ Scorpaenodes corallinus -- 4 — 4 —- — — 4 4 — — Plectrogenium nanum* 14 —_ 13 —- —- => — 14 4 —- — Taenianotus triacanthus*® 12 —- = 1 1 100 — 7 1 100 — Tracundus signifer 13 —- — — 12 1 — 13 13 —- — Setarches guentheri 32 —- => — 1 31 — 32 31 1 — Ectreposebastes imus 19 —- —- — — 18 1 19 —_ 17 2 Phenacoscorpius megalops 2 —- — — 2 —- — ZB z, —- — Pontinus macrocephalus 15 —- — — 2 13 -— 15 6 — — Scorpaenopsis diabolus* 24 — — 1 20 es 2S 1 — Scorpaenopsis cacopsis 11 — — 1 10 — — 11 110 — — Scorpaenopsis brevifrons’ 12 — — — 13 — — 13 130 —- — Scorpaenopsis fowleri 4 0—-— —~— — 4 — — 4 4 —- — Scorpaenopsis altirostris 40 — — — 4 — — 4 4 — — Rhinopias xenops 4 - —- — — 4 — — 4 4 —- — Neomerinthe rufescens 15 1 = 1 14 1 = 16 16 —- — Scorpaena coniorta 11 — — — 10 1 a 11 ia — — Scorpaena ballieui 10 — — — 10 — — 10 10 — — Scorpaena galactacma' 14 — — — 12 1 — 14 14 — — Scorpaena pele 19 — — 1 138 — — 19 19 — — Scorpaena colorata 17 —- —- — Ly —_- — 17 17 —- —
1One with 5 soft anal rays.
2 Holotype with 6% soft dorsal rays.
3 One with 6 soft anal rays.
4 One with 9 soft dorsal rays.
° One with 6 + scaled area + 1 dorsal spines.
under ledges and in or near caves. If the behavior of this species is sim- ilar to that of the other members of the genus Pterois as observed by G. R. Allen and by us, then the species tends to stay in concealed areas during day- light hours but ranges over adjacent areas while feeding at night. The Townsend Cromwell trawled 6 specimens from 5 stations off Molokai in 119-124 meters. These specimens, with one exception, were all taken at night.
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 275
Genus Dendrochirus Swainson
Brachirus SwWAINson, 1839, p. 71 (see Remarks).
Dendrochirus Swainson, 1839, p. 180 (see Remarks).
Brachyrus SwWAINson, 1839, p. 264 (see Remarks; type-species Pterois zebra Cuvier by subsequent designation of Swain, 1883, p. 277).
Nemapterois Fow er, 1938a, p. 73 (type-species Nemapterois biocellatus Fowler by original designation, monotypic).
REMARKS. Swainson (1839) first used the spelling Brachirus on p. 71; then on p. 180 he substituted the name Dendrochirus, but on p. 264, where species were included, he used the name and spelling Brachyrus. Moreover, on p. 303 he used the name Brachirus for a genus of sole. We treat Brachirus (of p. 71) as an inadvertent spelling error. Bleeker (1876b, p. 42) serves as the first revisor selecting Dendrochirus over Brachyrus.
Dendrochirus barberi (Steindachner). (Figure 2b.)
Pterois barberi STEINDACHNER, 1900a, p. 175 (original description; type locality South Pacific [one small specimen found by Captain Barber in the plankton during the trip from Honolulu to Cape Horn, 1896-1897]); STEINDACHNER, 1900b, pp. 491-492, pl. 3, fig. 2 (more complete description including the figure).
Dendrochirus hudsoni JorDAN & EVERMANN, 1903a [April 11], pp. 202-203 (original de- scription; type locality Hawaiian Islands, Oahu, Waikiki; holotype USNM 50652); SNYDER, 1904, p. 536 (listed, Honolulu); BOnLKE, 1953, p. 120 (location of types).
Dendrochirus chloreus JENKINS, 1903 [July 23], pp. 498-499 fig. 41 (original description ; type locality Hawaiian Islands, Honolulu; holotype USNM 50701); JorpAN & SNYDER, 1904b, p. 126 (one from Honolulu market); JorpAN & EVERMANN, 1905, pp. 465-466, fig. 204 (mostly compiled from Jenkins, 1903; figure of type of D. chloreus from Jenkins, 1903); JorpDAN & SEALE, 1906, p. 379 (name only, Hawaiian Islands) ; JORDAN & JORDAN, 1922, p. 55 (listed) ; BOHLKE, 1953, p. 120 (location of types).
Dendrochirus barberi: JoRDAN & SNYDER, 1904b, p. 126 (hudsoni a synonym of barber?) ; JorpAN & EVERMANN, 1905, p. 465, color pl. 73 (description mostly from Jordan and Evermann, 1903; hudsoni as a synonym of barberi) ; JoRDAN & SEALE, 1906, p. 379 (listed, Hawaii; hudsoni as a synonym); JORDAN & JORDAN, 1922, p. 56 (listed).
Brachirus chloreus: Fow rr, 1925, p. 27 (listed, Hawaiian Islands).
Dendrochirus zebra (not of Quoy and Gaimard): Fowter, 1928, pp. 294-295 (in part; barberi wrongly as young of zebra); WaAHLERT, 1955, p. 326 (type of barberi in Ubersee Museum, Bremen).
Dendrochirus brachypterus (not of Cuvier): Fowter, 1928, p. 295 (in part; hudsoni and chloreus included in synonymy); PIETSCHMANN, 1930, p. 18 (Kaneohe Bay, Oahu) ; Fow ter, 1931, p. 349 (in part; two specimens from Kona, Hawaii); Fowler, 1934, p. 430 (in part; one specimen from Kewalo Bay, Oahu); PIETSCHMANN, 1938, pp. 5, 30-31 (brief description; color like chloreus; one from French Frigate Shoal) ; TrvKer, 1944, p. 271, fig. 9 on pl. 6, text fig. (brief description; figure of type of chloreus from Jenkins, 1903); Gostine &« Brock, 1960, p. 341 (wrongly included barberi in synonymy).
Remarks. The type specimen of D. barberi is in the Ubersee Museum in Bremen, West Germany (Wahlert, 1955). It has not been examined by us.
276 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
TasLe 2. Counts of left pectoral fin rays in Hawaiian scorpionfishes (specimens from Hawai only; footnotes are used when the left or right fin-ray count differed such that the count for one side was unusual for the species).
14 15 16 17 18 19 20 21 22 23 24
Pterois sphex — 1 12 — — = == Ss —_ ss =e Dendrochirus barberi — — — 2 21 == — = aa el Scorpaenodes hirsutus —- —- — 4 §s —- —- =—- —- ~— =— Scorpaenodes kelloggi —- —- —> — 5 9 vr—-— — — — Scorpaenodes parvipinnis 1330" 3 0 See Scorpaenodes littoralis _ —- — 3 4 3 = F- FO Scorpaenodes corallinus —- —- — 1 Po eS eae SS Plectrogenium nanum —- =| | | S| Se Se 4 9 1 Taenianotus triacanthus 11 1 — — — — — Tracundus signifer = => = 4c (Cr oe ee Setarches guentheri —- —- FOO Eee tL BB ae Ectreposebastes imus — — o — -= 10 8 1° — — — Phenacoscorpius megalops — — — 20 — eee ee a Pontinus macrocephalus — — 2 13 — — == = _— = —— Scorpaenopsis diabolus —- — — Oe Scorpaenopsis cacopsis — — == 1@ @P2 — ss an Bm = — Scorpaenopsis brevifrons — — a — 1 10 a a5 2 nies Scorpaenopsis fowleri —- — | a ee ae Te Scorpaenopsis altirostris —-— — — 1 3 a Rhino pias xenops — a a 3 1 Ben OS Eee Neomerinthe rufescens — — — — 1S 4¢ a — ee —! ea Scorpaena coniorta — 1 10 —_ — = = ae = pi _ Scorpaena ballieui = = 9 1 = —_ ea oe a = Scorpaena galactacma = i aa ee eS es Scorpaena pele —>) 1 TS oe ee eS EE EES Scorpaena colorata’ _—- — {Pe ie DD es ee eer
® One with 19 on right.
> One with 17 on right.
© One with 18 on right.
4 23 on right in one specimen.
e 20 on right.
f 16 on right.
£ Two with 18 on right.
1One abnormal specimen with 15+ 16.
The type was reported as 31 mm. in standard length and was reportedly taken in the plankton during the trip from Hawaii to Cape Horn. A 61l-mm. S. L. specimen came to a nightlight station and was captured by G. R. Allen at Hanalei Bay, Kauai, so we know that small specimens can be found in surface waters. Because of this and because the species is otherwise known only from the Hawaiian Islands, we suspect the small holotype was collected very soon after leaving port in the Hawaiian Islands.
MATERIAL EXAMINED. USNM 50652 (1, 33.4, holotype of Dendrochirus hudsoni), Oahu, Waikiki Beach, U. S. Bureau of Fisheries, 1901. USNM 50701 (1, 97.4, holotype of Dendrochirus chloreus), Oahu, Honolulu, O. P. Jenkins,
VoL. XL] ESCHMEVER AND RANDALL: HAWAIIAN SCORPAENIDAE 277
1889. SU 23294 (5, about 35—55.3, paratypes of D. chloreus) and SU 23315 (1, 72.8, paratype of D. chloreus), Oahu, Honolulu, O. P. Jenkins, 1889. USNM 126089 (1, 43.5), Oahu, Honolulu, Albatross, 1902. SU 7467 (2, 33.8- 37.5, paratypes of D. hudsoni), Oahu, Honolulu, U. S. Fish Commission, 1901. ANSP 87600 (1, 62.9), Honolulu, J. W. Thompson, 1910. ANSP 104663 (1, 72.7), Honolulu, C. M. Cooke, 16 Oct. 1923. BPBM 6540 (1, 50.3), Oahu, Moku Manu, north side in 49 meters, base of vertical dropoff, J. E. Randall et al., 9 Oct. 1968. BPBM 7975 (1, 66.7), Oahu, Pokai Bay, in 9 meters, J. E. Randall et al., 29 July 1969. CAS 15690 (2), Oahu, west side of Waimea Bay, in 6-9 meters, J. E. Randall e¢ al., 25 Aug. 1969. CAS 15693 (9, 36.8-54.2), Oahu, off rocky islet at SW. end of Waimea Bay, in 1—-10.5 meters, J. E. and L. A. Randall and P. M. Allen, 27 July 1970. CAS 15714 (1, 80.8), Hawaiian Leeward Islands, reef on NW. side of Laysan, 25°46’27’N., 171°44°37’W., GVi Kes. no. 26, V. E. Brock, R. R. Harry ¢é @:, 3 July 1951. SU 7846 (2, 40.0-114), Oahu, Honolulu, U. S. Fish Commission, 1901. SU 8420 (2, 98.5— 116), Honolulu, E. L. Berndt, 1902. Additional material is available in the ANSP, BPBM, CAS, and USNM collections.
DISTINGUISHING FEATURES. Dorsal fin rays normally XIII, 9%. Anal fin rays normally III, 5%. Pectoral fin rays 17-18, mostly 18. Pectoral fin large, upper rays branched distally. Dorsal fin spines longer than '2 body depth, membranes deeply incised. Scales ctenoid, about 50-55 vertical scale rows. Coronal spines present. Some branching of head spines in large specimens. Suborbital ridge a single row of spines, not as broad patch of tiny spinules. Supraocular tentacle when present short, less than orbit diameter, and usually absent; not banded with black.
DistrisuTion. Dendrochirus barberi is known only from the Hawaiian Islands. It has been collected in depths from near shore to about 50 meters.
Genus Scorpaenodes Bleeker Scorpaenodes BLEEKER, 1857, p. 371 (type-species by monotypy, Scorpaena polylepis Bleeker, 1851). Generic synonyms and a discussion of the limits of the genus Scorpaenodes are given by Eschmeyer (1969a, pp. 2-3). Five species of Scorpaenodes occur in Hawaiian waters; all are widespread Indo-Pacific species.
Scorpaenodes hirsutus (Smith). (Figure 3.)
Parascorpaenodes hirsutus SmirH, 1957a, p. 63, fig. 5, pl. 1E (original description; as type of new genus; type locality western Indian Ocean, Bazaruto, 21°30’S., 35°30’E.; four paratypes, from Pinda, Bazaruto, and Aldabra; comparisons with S. kellogg?).
No Hawaiian references apply to this species.
278 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
MATERIAL EXAMINED. BPBM 13742 (12, 30.6-41.8), Oahu, Kahe Point, E. Chave, 9 March 1968. CAS 13473, formerly UH 1644 (1, 33.3), Oahu, Hauula Park, W. A. Gosline e¢ al., 11 May 1952.
DISTINGUISHING FEATURES. Dorsal fin rays normally XIII, 8%. Anal fin rays III, 5%. Pectoral rays 17-18, usually 17. Vertical scale rows about 30 (as in S. kelloggi; see key couplet 5 to separate Hirsutus and kelloggi). Sub- orbital ridge usually with 4 spines, first on lateral face of lachrymal bone; usually 1 additional spine below suborbital ridge at level between second and third spines. Interorbital spines present. No small spines at midline between tympanic spines.
DisTRIBUTION. Until now this species has been known only from the type specimens from the western Indian Ocean and a recent listing from Tahiti (Randall, 1973), but it appears that Scorpaenodes hirsutus is a widespread Indo-Pacific species. We can report it from the following localities besides Hawaii: Johnston Island (BPBM 13740), Palmyra (BPBM 7787) and Fan- ning (BPBM 7548) in the Line Islands, Tahiti (BPBM 7809), the Ryukyu Islands (BPBM 8703), Rarotonga (BPBM 13913), and Mauritius (BPBM 16384). CAS specimens are from Ifaluk and Kapingamarangi in the Caroline Islands, Comoro Islands, Chagos Archipelago, Red Sea, and Taiwan. Depths of capture where available are from near shore to 40 meters, and the habitat
of this species appears to be coral reef areas.
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE
Scorpaenodes kelloggi (Jenkins). (Figure 4.)
(Synonymy based only on Hawaiian references. )
Sebastopsis kelloggi JENKINS, 1903, pp. 492-493, fig. 37 (original description; type locality coral rocks on reef at Honolulu, Hawaii; holotype USNM 50694); Snyper, 1904, p. 535 (listed, Honolulu) ; Jorpan « EVERMANN, 1905, pp. 462-463, fig. 202 (description; addi- tional Hawaiian specimens; figure trom Jenkins, 1903); JoRDAN & SEALE, 1906, p. 374 (abundant at Hawaii; compared with S. guamensis and S. scaber); BOHLKE, 1953, p. 124 (location of types).
Scorpaenodes kelloggi: JORDAN & JORDAN, 1922, p. 54 (listed; common on reefs); JORDAN & EVERMANN, 1926, p. 10 (listed) ; Boroptn, 1930, p. 59 (listed, Pearl Harbor) ; Fow Ler, 1928, p. 290 (synonymy; brief description) ; Fowler, 1934, p. 430 (listed); Prerscu- MANN, 1938, pp. 5, 30, pl. 18 (brief description; one specimen from Kaneohe Bay [second specimen and figure possibly S. hirsutus|); Tinxer, 1944, pp. 266-267, fig. (brief diagnosis; figure of type from Jenkins, 1903); FowLrr, 1949, p. 107 (compiled synonymy) ; SMITH, 1957a, p. 63 (compared with S. hirsutus) ; ScHuLTz, 1966, pp. 35- 36, fig. 142 (figure of type from Jenkins, 1903; remainder of account deals with Marshall and Marianas Islands specimens).
Scorpaenodes guamensis (not of Quoy and Gaimard): Tanaka, 1928, p. 826 (S. kelloggi in synonymy).
Scorpaenodes parvipinnis (not of Garrett): GosLINE & Brock, 1960, p. 340 (S. kelloggi in synonymy).
MATERIAL EXAMINED. USNM 50694 (1, 37.7, holotype of Sebastopsis kel- loggi) and SU 23306 (1, about 24, paratype of S. kelloggi), Oahu, coral rocks at Honolulu, O. P. Jenkins, 1889. BPBM 8593 (9, 21.0-37.5), Oahu, Waimea Bay, in 6-9 meters, J. E. Randall e¢ al., 25 Aug. 1969. BPBM 8594 (1, 32.9), Oahu, Kahe Point, E. Chave, 9 Mar. 1968. BPBM 9787 (5, 19.2-31.9),
280 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Oahu, off Makaha Shores Condominium, in 14 meters, small caves in reef, J. E. Randall and A. R. Emery, 26 Apr. 1970. BPBM 10967 (5), Oahu, off Waikiki, in 7.5 meters, sand and small coral head and coral rubble, J. E. Randall, E. Chave, and students, 30 Mar. 1969. BPBM 12288 (4), Oahu, off Pokai Bay in 24 meters, J. E. Randall et al., 29 July 1969. CAS 15689 (3), Oahu, artificial reef off Aina Haina, in 23 meters, J. E. Randall e¢ al., 12 Sept. 1969. Additional Hawaiian material is present in the BPBM and CAS collections.
DISTINGUISHING FEATURES. Dorsal fin rays normally XIII, 8%. Anal fin rays III, 5%. Pectoral fin rays usually 18-19, rarely 20. Vertical scale rows about 30 (as in S. hirsutus; see key couplet 5). Suborbital ridge with 3 or 4 spines in a single row. Interorbital spines present. No small spines at midline between tympanic spines.
DistTrIBuTION. Scorpaenodes kelloggi is a small species inhabiting coralline areas in depths from near shore to at least 24 meters. It appears to be wide- spread in the Indo-Pacific. We can report this species from Tahiti (BPBM 8650, 8368), and localities represented by CAS specimens include Palmyra, Raiatea, Palau Islands, Gilbert Islands, Caroline Islands, and Taiwan.
Scorpaenodes parvipinnis (Garrett). (Figure 5a.)
(Synonymy based only on Hawaiian references.)
Scorpaena parvipinnis GARRETT, 1864, pp. 105-106 (original description; type locality Sandwich Islands [Hawaii]); GUNTHER, 1873, p. 75, fig. D on pl. 52 (description; Hawaii and Raiatea).
Sebastopsis guamensis (not of Quoy and Gaimard): Fowter, 1900, p. 535 (one specimen, ANSP 12207).
Sebastopsis parvipennis: SEALE, 1902, p. 20 (specific name misspelled; listed).
Sebastopsis parvipinnis: SNYDER, 1904, p. 535 (listed, Honolulu); JorpAN & EvERMANN, 1905, pp. 462-463 (description; brief synonymy; one specimen taken by the Albatross) ; JoRDAN & SEALE, 1906, p. 374 (compiled).
Scorpaenodes parvipinnis: JORDAN & JORDAN, 1922, p. 54 (listed; very rare); JORDAN & EVERMANN, 1926, p. 10 (listed) ; Fowler, 1928, p. 290 (synonymy; description; Ha- waiian specimens) ; Fowler, 1941, p. 257 (one from Hawaii); Tinker, 1944, p. 266 (compiled) ; Fow er, 1949, p. 107 (synonymy); GOSLINE & Brock, 1960, pp. 284, 287, 341, fig. 264 (brief description; in key; line drawing; S. kelloggi included in synonymy) ; GOSLINE, 1965, p. 825 (depth distribution).
MATERIAL EXAMINED. BPBM 7818 (1, 50.8), Oahu, off Waikiki, in 24 meters in coral, J. E. Randall and S. Swerdloff, 3 Sept. 1969. BPBM 7822 (1, 45.0), Oahu, Moku Manu, in 26 meters, J. E. Randall and W. J. Baldwin, 6
=>
FicurEe 5. a. Scorpaenodes parvipinnis, BPBM 7823, 69 mm. S. L., Hawaii. b. Scor- paenodes littoralis, BPBM 13732, 80 mm. S. L., Oahu. c. Scorpaenodes corallinus, BPBM 13730, 36 mm. S. L., Oahu. ( Photographs of fresh specimens.)
282 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Oct. 1969. BPBM 7823 (1, 66.5), Hawaii, Kona Coast, off point at north end of Honaunau Bay, in 43-49 meters, J. E. Randall e¢ al., 16 Aug. 1969. BPBM 7892 (1, 91.3), Oahu, Diamond Head Park, in 2 meters, G. R. Allen, 6 Mar. 1966. BPBM 9859 (3, 48.8-63.6), Oahu, off Lahilahi Point, in 27 meters, J. E. Randall and P. M. Allen, 11 July 1970. BPBM 13741 (1, 94.3), Hawaii, Mauna Loa, lava flow kill, H. Moore on O’Malley, 3 June 1950. CAS 13478, formerly UH 1644 (8, 50.4—92.5), Oahu, Hauula Park, W. A. Gosline e¢ al., 11 May 1952. CAS 15696 (2), Oahu, off rocky islet at SW. end of Waimea Bay, 1-10.5 meters J. E. and L. A. Randall and P. M. Allen, 27 July 1970. CAS 15712 (2), Oahu, Waikiki, % mi. off Niumalu Hotel, in 7.5—-9 meters, W. A. Gosline et al., GVF Reg. no. 54, 7 Sept. 1951. Additional Hawaiian material is available in the BPBM collection.
DISTINGUISHING FEATURES. Dorsal fin rays normally XIII, 9%. Anal fin rays III, 5%. Pectoral fin rays 17-19, usually 18. Scales very strongly ctenoid; vertical scale rows 45-55; interorbital area and snout scaled. Small spines at midline between tympanic spines frequently present. Interorbital spines present. Extra spines usually present on upper rear margin of eye after supraocular spine. Dorsal spines short, usually none longer than orbit diameter. Body often covered with small skin flaps (as in fig. 5a).
This species is distinguished from other species of Scorpaenodes by having the suborbital ridge with more than 5 spinous points, usually with 10 or more in adults.
DISTRIBUTION. Scorpaenodes parvipinnis has been taken in depths from near shore to about 45 meters in Hawaii. This species occurs widely in the Indo- Pacific from Africa and the Red Sea to the central Pacific.
Scorpaenodes littoralis (Tanaka).
(Figure 5b.)
Sebastella littoralis TANAKA, 1917, p. 10 (original description; type locality Misaki, Japan). Hawaiian reference:
?Scorpaenodes guamensis (not of Quoy and Gaimard): GosLINE & Brock, 1960, pp. 284,
287, 341 (brief description; Hawaii).
Scorpaenodes guamensis was reported from Hawaii by Gosline and Brock (1960), but this species does not occur in Hawaiian waters. At least some specimens in the University of Hawaii collection available to Gosline and Brock and labeled S. guamensis are specimens which appear to be referable to S. littoralis (Tanaka), the type locality of which is Japan.
MATERIAL EXAMINED. BPBM 10059 (2, 41.8-69.0), Oahu, off Pupukea on north shore, in 21 meters, rock and sand bottom, from small cave, J. E. Randall, 9 Aug. 1970. BPBM 10174 (4, 50.0-73.6), Oahu, off Kahana Bay, west side, “4 mi. out, cave in 26 meters, J. E. Randall, 27 Sept. 1970. BPBM 13731 (2, 55.8-60.9), Oahu, artificial reef off Pokai Bay, in 26 meters, J. E. Randall e¢
Vot. XL] ESCHMEVER AND RANDALL: HAWAIIAN SCORPAENIDAE 283
al., 21 June 1969. BPBM 13732 (2, 67.7-76.5), Oahu, Waimea Bay, west side, large boulders and some sand, J. E. Randall e¢ al., 25 Aug. 1969. CAS 13476, formerly UH 360 (2, 61.3-63.9), Oahu, Hauula Park, W. A. Gosline and class, 28 June 1949. CAS 13477 (1, 73.0), Oahu, Kaena Point, W. A. Gosline and class, 4 Mar. 1950.
DISTINGUISHING FEATURES. Dorsal fin rays normally XIII, 9%. Anal fin rays normally III, 5%. Pectoral rays 17-19, usually 18-19. Vertical scale rows about 45. Suborbital ridge with single row of 3 spines. Interorbital spines usually present, sometimes not well marked; small spines at midline between tympanic spines sometimes present.
This species is distinguished from other Hawaiian species of Scorpaenodes by the dark spot on the subopercle behind the preopercular spines. General body coloration also is diagnostic.
DISTRIBUTION. Scorpaenodes littoralis has been collected at Hawaii in depths to 26 meters. Habitat appears to be rocky or coral areas and caves. S. englerti Eschmeyer and Allen, 1971 (Easter Island) is closely related. We also can report Jittoralis-like specimens from a variety of localities in the Indo-Pacific: Rapa (BPBM 11242, 11247, 11249, 11251, 11255) in depths from about 3 to 27 meters, the Marquesas (BPBM 11111, 11134, 11147) in depths from 4.5—9 to 35-41 meters, Taiwan (USNM uncat.), One Tree Island off Australia (CAS 13855), and Africa (RU 970-142). This complex needs additional study.
Scorpaenodes corallinus Smith.
(Figure 5c.)
Scorpaenodes corallinus Smitrn, 1957a, pp. 64-65, 68, fig. 5, pl. 3E (original description ; type locality western Indian Ocean, Baixo Pinda; paratypes from Mozambique, Matemo, Tekomazi, Zanzibar, Pemba, Kenya, Aldabra, and Assumption).
No Hawaiian references apply to this species.
MATERIAL EXAMINED. BPBM 6451 (2, 38.2-53.6), Oahu, Moku Manu, west side, cave in 18 meters, rocky bottom, J. E. Randall, E. Reese, G. S. Losey, and L. Harris, 30 Sept. 1968. BPBM 13730 (1, 34.5), Oahu, Kahe Point, Waianae Coast, in 12 meters, G. R. Allen, 7 Sept. 1969. CAS 15724 (1, 34.8), Hawaii, Keahuolu Point, N. of Kailua, 19°38’45”N., 156°01’30’W., in 7.5-12 meters, Te-Vega cruise 8, station 366, R. L. Bolin et al., 15 Aug. 1965. Additional BPBM lots from Oahu are available.
DISTINGUISHING FEATURES. Dorsal fin rays XIII, 8%. Anal fin rays III, 5%. Pectoral fin rays 17-18. Vertical scale rows in low 40’s. Suborbital ridge with 3 spinous points in a single row. Interorbital spines absent or present as a lump; small spines at midline between tympanic spines present.
Body coloration (see figure 5c and Smith, 1957a, pl. 3E) is sufficient to distinguish this species from all other species of Scorpaenodes.
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DISTRIBUTION. Scorpaenodes corallinus was previously known only from the type specimens from the western Indian Ocean and a recent listing from Tahiti (Randall, 1973). Besides the Hawaiian specimens reported above we also can report this species from Moorea in the Society Islands (CAS 15804) at a depth of 9-12 meters, and from the Mentawei Islands in Indonesia (CAS 15805) in .5—2 meters. Depths of capture for the Hawaiian examples are 7.5— 12 to 18 meters. Little is known of its habitat or habits. Smith (1957a, p. 64) reported that the species was rather rare, occurred only in coral, and normally was found well below the low tide mark, usually in 1-5 fathoms (2—9 meters).
Genus Plectrogenium Gilbert
Plectrogenium GtLBerT, 1905, p. 634 (type-species by original designation, Plectrogenium nanum Gilbert, 1905).
Plectrogenium is a monotypic genus which Matsubara (1934) treats in a separate subfamily, Plectrogeniinae.
Plectrogenium nanum Gilbert. (Figure 6.)
Synonymy based only on Hawaiian references:
Plectrogenium nanum GILBERT, 1905, pp. 634-635, fig. 248 (original description; type locality Hawaiian Islands, off north coast of Maui, Albatross station 4082, in 220-238 fathoms [435 meters]; holotype USNM 51598; paratypes from Albatross stations 3952, 4079-82, 4132); JorDAN & SEALE, 1906, p. 378 (listed); JORDAN & JORDAN, 1922, p. 55 (listed) ; JorDAN & EVERMANN, 1926, p. 10 (listed); Fowter, 1928, p. 291 (compiled from Gilbert, 1905); TinKer, 1944, p. 269, fig. (compiled; figure of type from Gilbert, 1905); Fowter, 1949, p. 107 (compiled); BonrKeE, 1953, pp. 120-121 (location of types) ; GosLIne & Brock, 1960, pp. 285, 288, 341 (in key; compiled) ; CLARKE, 1972, p. 313 (submarine observation in 380 meters, off Barbers Point, Oahu).
MATERIAL EXAMINED. USNM 51598 (1, 56.3, holotype of Plectrogenium nanum), Maui, north coast off Puniawa Point, in 402-445 meters, bottom of gray sand, 10-ft. Blake trawl, Albatross station 4082, 21 July 1902. SU 8652 (4, paratypes of P. nanum), Hawaiian Islands, Albatross, no other data. BPBM 13738 (10, 46.3-55.4), CAS 15704 (18) and CAS 31300 (2, cleared and stained), Lanai, Kealaikahiki Channel, 20°38.1’—-41.3’N., 155°41.1—41.0’;W., in 292 meters, shrimp trawl, Townsend Cromwell cruise 33, station 38, 9 Nov. 1967. CAS 15700 (3, 49.1-52.3) and CAS 15702 (1, cleared and stained), Hamakua, off coast of Hawaii, 19°54’—57’N., 155°03.17—04.8’W, in 280 meters, shrimp trawl, Townsend Cromwell cruise 35, station 8, 29 Mar. 1968. Ad- ditional paratypes are present in the USNM collection. Additional Townsend Cromwell specimens are in the NMFS collection.
DISTINGUISHING FEATURES. Dorsal fin rays normally XII, 7%. Anal fin rays III, 5%. Pectoral fin rays 22-24. Scales ctenoid, about 30-35 vertical scale rows. Suborbital ridge with very well developed flat spines (somewhat
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 26
UL
NSS Ficure 6. Plectrogenium nanum, USNM 51598, off north coast of Maui. (Figure from Gilbert, 1905, fig. 248, but modified by addition of more dusky pigment.)
as in platycephalids). A number of extra spines on head, particularly on lachrymal bone and above orbits. Dorsal fin divided into 2 fins, second fin headed by 2 spines. Mouth somewhat ventral in location.
Body shape is diagnostic for this species. The normal presence of 7% soft dorsal rays together with the high pectoral ray count characterizes this species.
DistTRIBUTION. Plectrogenium nanum is a small species and apparently is very common in the Hawaiian Islands in the depth range of 274 to 640 meters; Struhsaker (1973) indicates a peak in abundance at depths of 300 to 450 meters based on Townsend Cromwell trawling operations. This species is also known from Japan (Matsubara, 1943, pp. 330-335).
Genus Taenianotus Lacépéde
Taenianotus LACEPEDE, 1802, p. 304 (type-species Taenianotus triacanthus Lacépéde, by subsequent designation of Cuvier in Cuvier & Valenciennes, 1829, p. 371).
Taenianotus triacanthus Lacépéde. (Figure 7.)
Taenianotus triacanthus LACEPEDE, 1802, pp. 305, 308 (original description; no type locality). Hawaiian references:
Taenianotus garretti GUNTHER, 1873, p. 83, fig. C on pl. 57 (original description; type locality Hawaiian Islands) ; JorpAN «& EvERMANN, 1905, p. 471 (compiled from Giinther, 1873); JORDAN & SEALE, 1906, p. 378 (name only); JORDAN & JORDAN, 1922, p. 55 (listed) ; ScHuLtz, 1938, p. 206 (suggested garretti may be distinct from triacanthus if Giinther’s figure were accurate).
Taenianotus citrinellus GILBERT, 1905, pp. 636-637, pl. 81 (original description; type locality Hawaiian Islands, south of Molokai, in 43-73 fathoms, Albatross station 3849; holotype USNM 51634); JorpAN & SEALE, 1906, p. 378 (name only); JoRDAN & JORDAN, 1922, p. 55 (listed; compiled locality data); Boroprn, 1930, p. 59 (Pearl Harbor record; reference to Fowler).
286 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.
Ficure 7. Taenianotus triacanthus, BPBM 6205, 47 mm. S. L., Oahu. (Photograph of fresh specimen.)
Taenianotus triacanthus: Fow.rr, 1923, p. 387 (listed; Honolulu); Fowter, 1925, p. 27 (listed) ; Jorpan, 1925, p. 21 (listed) ; FowLer, 1928, pp. 296-297 (synonymy; garretti and citrinellus in synonymy; Hawaiian specimens) ; FOWLER, 1934, p. 431 (reference to Borodin, 1930); ScHuLtz, 1938, p. 206 (brief description; color phases; citrinellus a synonym); TINKER, 1944, p. 272, fig. (brief description; figure from Gilbert, 1905) ; GosLINE & Brock, 1960, pp. 284, 285, 341 (in key; brief description; synonymy) ; GosLINE, 1965, p. 825 (depth distribution).
The type localities of garretti and citrinellus are Hawaii. They have been recognized as color phases of Taenianotus triacanthus for a number of years.
MATERIAL EXAMINED. USNM 51634 (1, 35.8, holotype of Taenianotus citrinellus), south coast of Molokai, N. 71° and W. 21.9’ from Lae-o Ka Laau Light, 10-ft. Blake trawl in 134-79 meters, bottom of coarse sand, broken shells and coral, Albatross station 3849, 8 April 1902. BPBM 4405 (1, 60.5), Oahu, Kahala, C. M. Cooke, Jr., 17 March 1919. BPBM 4406 (2, 51.2—56.6), Oahu, C. M. Cooke, Jr., 3 Dec. 1923. BPBM 4891 (1, 59.4), Oahu, Laie, C. M. Cooke III, 15 Jan. 1939. BPBM 6203 (1, 45.3), Oahu, Ala Moana Reef, 200 feet from shore in .5 meters, S. M. Trefz, 17 April 1966. BPBM 6984 (1, 38.2), Oahu, Diamond Head Park, in 2 meters, G. R. Allen, March 1968. BPBM 8821 (1, 37.9), Oahu, Makua, in 12 meters, G. R. Allen, March 1968. BPBM 9777 (1, 41.0), Oahu, off Makaha Shores Condominium, small caves
Vox. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 287
in reef, in 14 meters, J. E. Randall and A. R. Emery, 26 April 1970. BPBM 10628 (2, 56.5-59.7), Oahu, Hanauma Bay, G. R. Allen, June 1965. BPBM 10950 (1, 29.0), Oahu, Waimea Bay, in 3-6 meters, G. R. Allen, July 1970. CAS 15691 (2, 35.7-52.8), Oahu, Waimea Bay, among large boulders with some sand, at west side of Waimea Bay, in 6—9 meters, J. E. Randall e¢ al., 25 Aug. 1969. Additional Hawaiian material is available in the BPBM and NMFS collections.
DISTINGUISHING FEATURES. Dorsal fin rays usually XII, 10%. Anal fin rays III, 6’2 (sometimes III, 512). Pectoral rays usually 14 (sometimes 15), all rays unbranched. Dorsal fin high, 3rd or 4th spine longest. Scales as small spiny papillae. Suborbital ridge without spines, or with lump at end of ridge. Preopercle with 2 indistinct spines only.
This species is characterized by having the body extremely compressed, the soft dorsal fin attached to the caudal fin, and the fewest pectoral rays (usually 14) of any Hawaiian scorpionfish.
DIsTRIBUTION. Taenianotus triancanthus is a widespread, shallow water Indo-Pacific species occurring from Africa to the central Pacific. Depths of capture in Hawaii ranged from .5 to 14 meters, with one trawled from 79-134 meters.
Remarks. The coloration of this species is variable, from nearly all yellow, to red, brown, or nearly black, and variously mottled with darker pigment. Coloration evidently is related partly to shedding of a cuticular layer con- taining algae (see Wickler and Nowak, 1969).
Genus Iracundus Jordan and Evermann
Tracundus JORDAN & EVERMANN, 1903b, pp. 209-210 (type-species Jracundus signifer Jordan and Evermann, 1903b, by original designation, monotypic).
Tracundus signifer Jordan and Evermann. (Figure 8.)
Tracundus signifer JORDAN & EVERMANN, 1903b, p. 210 (original description; type locality Hawaiian Islands; one specimen from coral reef at Honolulu; holotype USNM 50886) ; JorDAN & SNYDER, 1904b, p. 126 (one specimen from Honolulu market); JORDAN & EVERMANN, 1905, pp. 470-471, fig. 207 (mostly compiled from Jordan and Evermann, 1903b, additional Hawaiian specimens; figure of holotype); JoRDAN & SEALE, 1906, p. 374 (name only; Hawaii); JorDAN & JORDAN, 1922, p. 55 (listed) ; Fow irr, 1923, p. 387 (listed; Honolulu) ; Fowrer, 1925, p. 27 (listed); JorpDAN, 1925, p. 20 (listed) ; JorDAN & EveRMANN, 1926, p. 10 (listed); Fowtrerr, 1928, p. 292 (synonymy; description; Hawaiian specimens) ; TrvKER, 1944, p. 269, fig. (compiled; figure of type from Jordan and Evermann, 1905); Fowrer, 1949, p. 107 (compiled) ; GosLine & Brock, 1960, pp. 284, 286, 341 (in key; compiled) ; Gostrne, 1965, p. 825 (listed) ; Mappen, 1973, p. 145 (from salvaged vessel; luring behavior) ; SHALLENBERGER & MAppDEN, 1973, pp. 33-47, figs. 1-6 (luring behavior; aspects of biology).
Non-Hawaiian reference:
Tracundus signifer rarotongae WHITLEY, 1965, pp. 113-114 (original subspecies description ; type locality Cook Islands, Rarotonga; holotype AMS IA5226).
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Remarks. Whitley (1965, pp. 113-114) described Jracundus signifer varotongae from Rarotonga Island in the Cook Islands. The type (AMS IA5226) was examined by Eschmeyer. The differences reported by Whit- ley between J. s. rarotongae and /. s. signifer do not appear to be valid ones. Whitley reported that his subspecies had one more pectoral ray (18), but we find many Hawalian examples with 18 pectoral rays. Whitley re- ported 39 scale rows above the lateral line but Eschmeyer counts about 70 vertical scale rows for Whitley’s type. Whitley gives the gill rakers as 6 + 10, but the type has 6 or 7 + 11, including rudiments. Other differences reported by Whitley seem to result from incorrect interpretation of Jordan and Ever- mann’s account of this species. At the time of Whitley’s description of /. signifer rarotongae, I. signifer was known only in Hawaiian waters; we now know that the species is widespread in the Indo-Pacific (see below). We feel that the Cook Island population is not subspecifically distinct.
MATERIAL EXAMINED. USNM 50886 (1, 77.8, holotype of /racundus sig- nifer), Oahu, reef at Honolulu, U. S. Fish Commission. BPBM 6354 (3, 86.9- 97.4), Oahu, Kahe Point Beach State Park, reef in 10.5 meters, J. E. Randall, G. R. Allen, ef al., 30 March 1968. BPBM 7319 (2, 45.2-53.4), Oahu, off Lahilahi Point, near cave in 27-30 meters, coral rubble, J. E. Randall and University of Hawaii students, 10 Aug. 1968. BPBM 7924 (6, 54.8-78.3),
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Oahu, Moku Manu, in 49 meters, J. E. Randall, W. J. Baldwin, and A. Stark, 9 Oct. 1968. CAS 24990 (18), Oahu, SE. of Poka Bay, Waianae coast, caves adjacent to coral rubble and sand in 21-30 meters, J. E. Randall et a/., 20 July 1969. SU 8602 (1), Oahu, Honolulu, E. L. Berndt. Additional Hawaiian material is available in the BPBM collection.
DISTINGUISHING FEATURES. Dorsal fin rays usually XII, 9%. Anal fin rays III, 542. Pectoral rays usually 18 (sometimes 17; 19 should be expected). Palatine teeth absent. Scales ctenoid; about 65-75 vertical scale rows. Lachrymal bone with 2 spines over maxillary; first points forward; second broad, pointing out and to rear. Suborbital ridge without spines except one at rear before preopercle. Preopercular spines short, usually only 3 developed; no supplemental preopercular spine at base of first spine.
This species is characterized by a dark spot on the spinous dorsal fin be- tween spines 1 or 2 and 3, and vertical scale rows of about 70. The fourth dorsal spine is characteristically elongate in specimens over about 50 or 60 mm. S. LL:
DistriBuTION. Jracundus signifer occurs in Hawaiian waters in coral rubble areas, particularly in or near concealed locations beneath ledges or in caves, in depths from about 10.5 to 61 meters. William D. Madden (1973, p. 141, and pers. comm.) reports capture of specimens inside a sunken hull raised from 67 meters. Although this species is known in the literature only from the Hawaiian Islands and from the Cook Islands (type of /. s. rarotongae), we suspect it is widely distributed in the Indo-Pacific. Randall has taken speci- mens from several localities in French Polynesia and from Mauritius.
Genus Setarches Johnson
Setarches Jounson, 1862, p. 177 (type-species Setarches giintheri Johnson, 1862, by monotypy).
This genus has been treated on a world basis by Eschmeyer and Collette (1966, pp. 355-356).
Setarches guentheri Johnson. (Figure 9.)
Setarches giintheri JouNSoN, 1862, pp. 177-179, pl. 23 (original description; type locality Madeira).
Synonymy of Hawaiian references only:
Scorpaena remigera GILBERT & CRAMER, 1897, pp. 404, 418-419, pl. 40 (original description ; type locality off Hawaiian Islands, in 298 fathoms, Albatross station 3476, holotype USNM 47726); BouLKE, 1953, p. 121 (location of types).
Setarches remiger: G1LBERT, 1905, p. 634 (brief description; new combination; additional specimens from Hawaii); JoRDAN & SEALE, 1906, p. 377 (name only, Hawaii); JORDAN & JorDAN, 1922, p. 55 (listed; common in deepwater); Fowrer, 1928, pp. 291-292 (synonymy; description from Gilbert and Cramer); TInKER, 1944, Pp. 270, text fig. (compiled; figure from Gilbert and Cramer, 1897) ; FowLer, 1949, p. 107 (compiled) ;
290 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.
Ficure 9. Setarches guentheri, BPBM 29322, 109 mm. S. L., north of Maui. (Photo- graph of preserved specimen.)
GosLINE & Brock, 1960, pp. 285, 341 (in key; compiled); Ciarkr, 1972, p. 313 (submarine observation in 310 meters, off Barbers Point, Oahu).
Setarches guentheri: ESCHMEYER & COLLETTE, 1966, pp. 358-360 (remigera in synonymy; description; distribution; remarks).
The reader is referred to Eschmeyer and Collette (1966) for a thorough treatment of this species.
MATERIAL EXAMINED. USNM 47726 (1, 83.2, holotype of Scorpaena remigera), south of Oahu, 21°09’N., 157°53’W., in 545 meters Albatross station 3476, 6 Dec. 1891. BPBM 14109 (16, 66.3-88.0) and CAS 29322 (16, 68.6-92.5), north of Maui, 21°10.1’-10.6’N., 156°25.8’-33.2’°W., 41-ft. shrimp trawl in 494-512 meters, Townsend Cromwell cruise 35, station 17, 1 April 1968. Additional specimens are available in the BPBM, USNM, and NMFS collections; paratypes of S. remigera are present in the USNM and SU collec- tions. (Counts in tables 1 and 2 based on BPBM 14109 and CAS 29322, not from Eschmeyer and Collette, 1966.)
DISTINGUISHING FEATURES. Dorsal fin rays normally XII, 10’. Anal fin rays III, 5%. Pectoral fin rays in Hawaiian specimens 20-22, mostly 21. Body covered by tiny cycloid scales. Lateral line more or less a continuous trough covered by thin membranous scales (these covering scales usually lost during capture) as in EF. imus. Body grayish or pinkish in coloration. Distin- guishable from E. imus by having 5% rather than 6% anal soft rays, higher average number of pectoral rays, and orbit diameter about equal to interorbital width rather than about '% interorbital width.
Vot. XL] ESCHMEYVER AND RANDALL: HAWAIIAN SCORPAENIDAE 291
DISTRIBUTION. Setarches guentheri occurs in offshore areas from Hawaii circumtropically west to the western Atlantic (Eschmeyer and Collette, 1966). In Hawaiian waters the Albatross took specimens at 14 stations (see Gilbert, 1905) between about 350 and 550 meters (with one station in 177-364 and one in 351-644 meters). The Townsend Cromwell captured a total of 184 specimens from 21 stations in the depth range 238-686 meters (Struhsaker, 1973). Setarches guentheri is a bottom or near-bottom species which may feed only on pelagic organisms (Eschmeyer and Collette, 1966).
Genus Ectreposebastes Garman
Ectreposebastes GARMAN, 1899, p. 53 (type-species Ectreposebastes imus Garman, 1899, by monotypy).
This genus has been treated on a world basis by Eschmeyer and Collette (1966, pp. 366-367).
Ectreposebastes imus Garman. (Figure 10.)
Ectreposebastes imus GARMAN, 1899, pp. 53-55, pls. 7, 9, fig. 1 on pl. 71 (original descrip- tion; type locality Galapagos Islands, Albatross station 3403, 702 meters).
Hawaiian references:
Ectreposebastes imus: ESCHMEYER, 1969b, p. 104 (new record for Hawaii based on per- sonal communication from P. Struhsaker) ; CoLLETTE & UYENO, 1972, pp. 26-28, fig. 1 (Pontinus niger Fourmanoir, 1970, as a synonym; range including Hawaii; first record for Japan; various comments; good figure).
The reader is referred to Eschmeyer and Collette (1966) for a more thorough treatment of this species, and to Collette and Uyeno (1972) for re- cent range extensions and references.
Pontinus niger Fourmanoir, type locality near the Marquesas Islands, was placed in the genus Ectreposebastes and synonymized with E. imus by Collette and Uyeno (1972). Examination by Eschmeyer of a specimen from the south central Pacific at 10°51~29’S., 123°28’-48’W. (SIO 73-43), the specimen reported by Collette and Uyeno from near New Caledonia at 22°02’S., 167°57’E. (USNM 206500), along with notes made by Collette on the holotype of P. niger, indicate that E. niger possibly is a valid species differing most notably in having scales noticeably larger than in specimens of EF. imus from other areas. There also may be differences in depth of the caudal peduncle, pectoral fin length, and length of lachrymal and preopercular spines. A detailed com- parison of specimens from different populations is needed.
MATERIAL EXAMINED. BPBM 14110 (18, 82.5-140) and CAS 29321 (33, 76.3-135), Kaiwi Channel between Oahu and Molokai, 21°08.8—07.4’N., 157° 42.2’-49.6'W., in 585-640 meters, 41-ft. shrimp trawl, Townsend Cromwell
292 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 10. Ectreposebastes imus, CAS 29321, 88.7 mm. S. L., between Oahu and Molokai. (Photograph of preserved specimen.)
cruise 36, station 31, 6 May 1968. Additional Hawaiian specimens are avail- able in the NMFS collection.
DISTINGUISHING FEATURES. Dorsal fin rays usually XII, 10%. Anal fin rays III, 6%. Pectoral fin rays usually 19 or 20. Body delicate, flabby, with weak fin spines and poor ossification. Scales tiny and cycloid. Lateral line more or less a continuous trough covered by thin membranous scales (these covering scales usually lost during capture) as in S. guentheri. Juveniles black, adults maroon and black.
This species may be distinguished from all species except Setarches guentheri on the basis of its characteristic lateral line. It differs from Setarches guentheri in having 6% rather than 5’ anal rays, 3 rather than 2 prominent lachrymal spines, and interorbital width about twice orbit rather than about equal to orbit.
DisTRIBUTION. Ectreposebastes imus is known from the eastern and western Atlantic, the southeastern Pacific off the Galapagos Islands and Peru, Hawaii, and Japan.
It is an offshore, near-bottom species occurring in depths of about 150 to about 850 meters, excluding those depths deeper than 850 meters where the specimens may have been captured during retrieval of nets. Some captures have been made by midwater trawl hauls well off the bottom, and Eschmeyer and Collette (1966) suggest that this fish is the most modified of scorpionfishes for life in midwater. Struhsaker (1973) states that the species may occur near bottom during the day and undertake vertical feeding migrations into mid-
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 293
water areas at night. In Hawaiian waters the Townsend Cromwell took speci- mens at five bottom trawl stations in depths between 567 and 686 meters. Two hundred twenty-one specimens were taken on cruise 36 at station 31 (see Material examined) in 622 meters during daytime hours while a repeat tow in the same area at night resulted in only 2 specimens (Struhsaker, 1973).
Genus Phenacoscorpius Fowler
Phenacoscorpius FOWLER, 1938a, pp. 69-70 (type-species Phenacoscorpius megalops Fowler, 1938a, by original designation).
This genus has been discussed by Eschmeyer (1965b, pp. 522-523).
Phenacoscorpius megalops Fowler. (Figure 11.)
No Hawaiian literature applies to this species.
Phenacoscorpius megalops Fow er, 1938a, pp. 70-71, fig. 30 (original description; type locality Philippine Islands, Albatross station 5387, in 209 fathoms, 11 March 1909; plus a series of paratypes from the Philippines and East Indies) ; Bohlke, 1953, p. 120 (location of types); EscHMEyER, 1965b, pp. 522-523 (compared with an Atlantic species) .
MATERIAL EXAMINED. BPBM 13761 (1, 29.3), Oahu, 6 miles off Makapuu Point in direction of Molokai, depth of 366 meters, collected with pink coral, W. D. Madden, 5 May 1972. BPBM 16416 (1, about 51, poor condition), Oahu, Kaiwi Channel off Makapuu Pt., depth 366 meters, collected with pink coral from submarine, presented by W. D. Madden, 11 Feb. 1974. (The holotype, USNM 98903, and some paratypes, SU 40198-40200, of Phenacoscor- pius megalops were examined. )
DISTINGUISHING FEATURES. (Based also on specimens from the Philippines. ) Dorsal fin rays XII, 9%. Anal fin rays III, 5%. Pectoral fin rays usually 17. Palatine teeth absent. Scales on sides weakly ctenoid; about 55 (?) vertical scale rows. Lachrymal bone with 2 spines over maxillary; first as a broad lobe, second as a broad spine pointing down. Suborbital ridge well marked, with 5 or 6 or more spines. Second preopercle spine small or absent.
This species is easily distinguished from other Hawaiian scorpionfishes by having only the anterior 4 or 5 tubed lateral line scales present, with the remainder of the lateral line absent. The large melanophores on the caudal peduncle are also characteristic.
Coloration in life is mostly red.
DistTRIBUTION. Phenacoscorpius megalops is known from the Philippine Islands and the East Indies and now from Hawaii in reported depths of 68—622 meters. A wider distribution in the central Pacific Ocean is expected when these depths are sampled more thoroughly.
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Genus Pontinus Poey
Pontinus Pory, 1860, p. 172 (type-species Pontinus castor by inference from text (see Eschmeyer, 1965b, p. 527) or by subsequent designation of Jordan and Gilbert, 1883, p. 669).
Generic synonyms and nomenclatural problems are discussed by Eschmeyer (1965b, pp. 526-528).
Pontinus macrocephalus (Sauvage). (Figure 12.)
Sebastes macrocephalus SAUVAGE, 1882, pp. 169-170 (original description; type locality Hawaiian Islands, collected by Ballieu).
Merinthe macrocephala: SNyprER, 1904, p. 535 (brief description; 2 specimens from Hono- lulu); JorpDAN & EVvERMANN, 1905, p. 461, pl. 55 (description; two specimens from Hawaii) ; JoRDAN & SNYDER, 1907, p. 217 (common in winter market; brilliant orange with sparse dots and mottlings); JoRDAN & JoRDAN, 1922, p. 55 (name; size; deep- water) ; JoRDAN, 1925, p. 20 (variation in skin appendages; color description) ; FOWLER, 1925, p. 27 (listed); JorDAN & EVERMANN, 1926, p. 10 (listed); FowLer, 1928, p. 291 (synonymy; description); Fowrerr, 1931, p. 349 (description of 2 specimens from Honolulu market); Tinker, 1944, p. 267, fig. (compiled brief description; figure from Jordan and Evermann, 1905); FowLer, 1949, p. 107 (compiled literature reference) ; GOSLINE & Brock, 1960, pp. 285, 288, 341, fig. 266 (in key; brief characterization; figure from Jordan and Evermann, 1905); Gostine, 1965, p. 825 (depth distribution).
Pontinus spilistius GILBERT, 1905, pp. 633-634, fig. 247 (original description; type locality Hawaiian Islands, off Maui, in 174-238 meters, Albatross station 4077; Holotype USNM 51644) ; JorDAN & SEALE, 1906, p. 377 (listed) ; JorDAN & JORDAN, 1922, p. 55 (listed) ; Jorpan & EvERMANN, 1926, p. 10 (listed); FowLer, 1928, p. 291 (compiled from Gilbert, 1905); Boroprn, 1930, p. 59 (listed, Oahu); TinKer, 1944, p. 268, fig. (brief
Vot. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 295
Ficure 12. Pontinus macrocephalus, BPBM 10573, 285 mm. S. L., Honolulu fish market. (Photograph of fresh specimen.)
description compiled; figure of type from Gilbert, 1905); FowLer, 1949, p. 107 (compiled literature reference); BOHLKE, 1953, p. 121 (location of types); GosLINE & Brock, 1960, pp. 285, 289, 342 (in key; compiled).
Pontinus macrocephalus: ESCHMEYER, 1969b, p. 24 (spilistius a synonym; macrocephalus restricted to Hawaii); Ciarker, 1972, p. 313 (submarine observations and _ gill-net capture, off Oahu, in 120-300 meters).
Remarks. Pontinus spilistius appears to be based on juveniles of Pontinus macrocephalus. Specimens under about 100 mm. S. L. have a dark spot on the spinous dorsal fin but this is absent or not well marked in large specimens. Small specimens have the eye almost as large as the snout while the eye is proportionally smaller in large specimens. The supraocular tentacle often is absent; when present it may be two or more times the size of the eye.
MATERIAL EXAMINED. MNHN A. 4165 (1, about 365, holotype of Sebastes macrocephalus), Hawaii, collected by Ballieu, no other data. USNM 51644 (1, 67.7, holotype of Pontinus spilistius), Maui, off north coast, Puniawa Point, 5. 45°45’, E. 6.1’, in 181-194 meters, bottom of fine coral sand and foraminifera, 10-ft. Blake trawl, Albatross station 4077, 21 July 1902. USNM 51663 (1, 47.1, paratype of P. spilistius) and SU 8638 (1, 44.3, paratype of P. spilistius), Maui, off north coast, Puniawa Point, S. 52°30’, E. 6.5’, in 174-238 meters bottom of coral, sand and foraminifera and rock, 10-ft. Blake Trawl, Albatross station 4098. ANSP 66366—-67 (2, 192-210), Oahu, Honolulu, R. Wehrl, 1929. BPBM 4386 (2, 270-360), Oahu, Honolulu, J. W. Thompson. BPBM 7944 (1, 340), Oahu, Campbell Park, Waianae, in 183 meters, gill net, T. A.
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Clarke, and P. Struhsaker, 11 Dec. 1968. BPBM 8150 (1, 127), Oahu, off Campbell Industrial Park, Barber’s Point, in 183-219 meters, T. A. Clarke, 14-15 Dec. 1968. BPBM 10573 (1, 285), Oahu, Honolulu fish market, J. E. andettiA, Randall, 7 Oct. 1969. BPBM 13736 (2, 67.2-99.5), CAS 15703 (2, 67.3-103) and USNM 214048 (3, 60.0—-82.5), off Hawaii, Hamakua coast, 20°07.2-08.3’N., 155°24.7-28.2’W., in 238-252 meters, shrimp trawl, Townsend Cromwell cruise 35, station 12, 30 Mar. 1968. USNM 88259 (2, 205-206), Oahu, Honolulu, H. W. Fowler. USNM 151639 (1, 196), Oahu, Honolulu, E. K. Jordan.
DISTINGUISHING FEATURES. Dorsal fin rays normally XII, 10’2, some- times XII, 9%. Anal fin rays III, 5%. Pectoral fin rays 16-17, usually 17. Scales ctenoid; about 45—50 vertical scale rows. Lachrymal bone with 2 spines over maxillary, first points out and back, second points mostly back. Sub- orbital ridge with 3 or 4 spines. Second preopercular spine usually absent in large specimens. Mostly red in life.
The pectoral rays are unbranched in adults of this species (as in Taenianotus and Pterois). Pterois differs in having 13 dorsal spines and Taenianotus in having 15 or fewer pectoral rays, among other features. Most difficulty comes in separating small specimens of Pontinus from Scorpaena or Scorpaena-like specimens, such as ones of Neomerinthe rufescens, when the specimens are at a size at which their pectoral rays are still unbranched (usually under 30 or 40 mm. S. L.). Useful characters for separating small specimens include counts (tables 1-2), presence or absence of a dark spot on the spinous dorsal fin (present in juveniles of P. macrocephalus and a few other species), as well as general coloration, depth of capture, and lachrymal and suborbital spination. See also the account of Neomerinthe rufescens.
DisTRIBUTION. Pontinus macrocephalus is an upper slope species. Depths of capture range from about 180 to 250 meters, with one collection from the range 174-278 meters. We know this species only from the Hawaiian Islands (see Eschmeyer, 1969b, p. 24).
Genus Scorpaenopsis Heckel
Scorpaenopsis HrCKEL, 1837, p. 159 (type-species Scorpaena gibbosa Schneider in Bloch and Schneider, 1801, by subsequent designation of Bleeker, 1876, p. 4; not an originally included species but Bleeker synonymized it with mnesogallica Cuvier in Cuvier and Valenciennes on p. 28 of the same work; see Article 69(a) (iv), International Code of Zoological Nomenclature, 1964).
Scorpaenichthys BLEEKER, 1856, pp. 388, 402 (type-species Scorpaena gibbosa Schneider in Bloch and Schneider, 1801, by subsequent designation of Jordan, 1919, p. 267; pre- occupied by Scorpaenichthys Girard, 1854, p. 161, a genus of cottid fishes).
Dendroscorpaena SmiTH, 1957a, pp. 51, 60 (replacement name for Scorpaenichthys Bleeker, 1856; therefore taking the same type-species as Scorpaenichthys Bleeker, despite the statement by Smith on p. 60 “Genotype Perca cirrhosa Thunberg, 1793”; see Article 67(i), International Code of Zoological Nomenclature, 1964).
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 297
The genus Scorpaenopsis consists of two groups of species: humpbacked ones and non-humpbacked ones. Smith (1957a) recognized each group as a separate genus, Scorpaenopsis for the former and Dendroscorpaena for the latter. As discussed in the synonymy above, Smith wrongly thought the type- species of Dendroscorpaena was the non-humpbacked species cirrhosa when in fact the type-species was the humpbacked species gibbosa. We agree, how- ever, with Matsubara (1943) that all species should be placed in one genus. Of the four humpbacked species, one has only a slight hump (S. neglecta).
The differences between Scorpaenopsis and Scorpaena are few, the major difference being that species of Scorpaena have palatine teeth while species of Scorpaenopsis lack palatine teeth. We suspect that palatine teeth have been lost more than once in different Scorpaena-like species. Most of the species usually referred to Scorpaenopsis seem more closely related to each other than to the species in various subgroups of Scorpaena, but there are a few species which stand apart. They are usually placed in Scorpaenopsis because they lack palatine teeth. Among these are S. fowleri and S. altirostris from Hawaii. On the other hand, there are a few species of the genus Scorpaena which seem perhaps more closely related to species of Scorpaenopsis than to subgroups of Scorpaena, for example, Scorpaena orgila from Easter Island. So the limits of Scorpaenopsis and Scorpaena are unclear, as are the differences between them. We have allocated the Hawaiian species for now solely on the basis of presence or absence of palatine teeth.
The non-humpbacked species are poorly known. For example, there are several species which are usually confused under the name Scorpaenopsis cirrhosa. Among the humpbacked ones, there are four species as discussed under the account of Scorpaenopsis diabolus.
Scorpaenopsis fowleri (Pietschmann).
(Figure 13.)
Scorpaenodes fowleri PirTSCHMANN, 1934, pp. 99-100 (original description; type locality Hawaiian Islands, Makaua, Oahu [see lectotype designation below]); PirtscHMANN, 1938, pp. 5-6, 30, pl. 9 (redescription; figure of one type); FowLer, 1949, p. 107 (compiled synonymy; no specimens; questioned Pietschmann’s generic placement, thought related to asperella).
Scorpaena ballieui (not of Sauvage): Gostine & Brock, 1960, p. 342 (in part; questionably included fowler in synonymy).
7
Remarks. Pietschmann described this species from 3 small specimens (NMW 6341-6343); these specimens were examined briefly by Eschmeyer and one (NMW 6341) was loaned to him for more detailed study. We designate NMW 6341 as the lectotype of S. fowleri. We are unable to determine if this was the specimen figured by Pietschmann (1938, pl. 9).
Placement of this species in the genus Scorpaenopsis, rather than in Scorpaena, is based on the absence of palatine teeth, but the limits of the
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genera Scorpaenopsis and Scorpaena are uncertain as was discussed under the account of Scorpaenopsis. (This species does not belong in Scorpaenodes as placed by Pietschmann.)
Except for a recent listing from Tahiti (Randall, 1973), Scorpaenopsis fowlert (Pietschmann) has not been recognized as a valid species since its description in 1938. The three types and subsequent specimens reported here are all less than 30 millimeters in standard length. Attempts by us to identify this species with the young stages of other species failed. Dissection showed that two of our specimens were egg-laiden mature females, and this left no doubt that the species is a valid one. This species almost certainly matures at the smallest size (about 25 mm. S. L.) of any scorpaenid now known. As discussed in the distribution section, collections by Randall have resulted in capture of specimens from areas outside the Hawaiian Islands.
MATERIAL EXAMINED. NMW 6341 (1, 28.1, lectotype of Scorpaenodes fow- leri), Oahu, Makaua, collected by T. T. Dranga, Dec. 1927. NMW 6342-43 (2, paralectotypes of S. fowler’), same data as lectotype (not examined in detail). BPBM 7853 (1, 24.6), Oahu, Moku Manu, off north side in 23 meters at entrance to cave, bottom of sand and coral rubble, J. E. Randall, W. J. Bald- win, and G. S. Losey, 3 Oct. 1968. BPBM 7854 (1, 25.8), Oahu, Moku Manu, in 26 meters, bottom of sand, coral rubble, and reef, J. E. Randall and W. J. Baldwin, 6 Oct. 1968. CAS 30738 (1, 27.0), Hawaii, Kona coast at
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 299
south end of Kailua Airport, between Keahole and Puhili Points, in 14 meters, coral and rocky bottom, J. E. Randall, 7 Nov. 1972.
DISTINGUISHING FEATURES. A very small species, probably not exceeding 30 or perhaps 35 mm. in standard length (about 40-45 mm. total length). Dorsal fin rays XII, 9%. Anal fin rays III, 5%. Pectoral fin rays 16, unbranched in available specimens. Vertical scale rows about 35; scales on sides ctenoid. Palatine teeth absent. Lachrymal bone with 2 spines over maxillary, first points forward, second points down and slightly forward. Suborbital ridge without spines or with a low spine at rear end. Nasal spine small, sometimes absent.
This species differs from other species of Scorpaenopsis by having only 16 pectoral rays rather than 17 or more. It differs from species of Scorpaena by lacking palatine teeth. The posterior lachrymal spine over the maxillary points forward, a feature characteristic for the species and rare in scorpion- fishes generally.
Preserved specimens are mostly pallid; frequently one or two small dark bars radiate back from rear of orbit. In life, body and fins mottled with red and white.
DistrrBuTion. No data on depth of capture are available for Pietschmann’s types. Subsequent collections of this species in Hawaii have been in depths from 14 to 26 meters. We also can report this species from the Tuamotu Archipelago at Mangareva Island (BPBM 13576) in 12 meters and Takaroa Island (BPBM 11159) in 9 to 15 meters, Tetiaroa in the Society Islands (BPBM 14968), from American Samoa at Tutuila (BPBM 17246) in 27 meters, and from Enewetak (formerly Eniwetok) in the Marshall Islands (CAS 31807, 31808, collected by R. S. Nolan). A wider distribution in the central Pacific is expected.
Scorpaenopsis brevifrons Eschmeyer and Randall, new species.
(Figures 14-15.)
Sebastapistes asperella (not of Bennett): PretscHMANN, 1938, pp. 27-28 (five specimens from French Frigate Shoal in the Hawaiian Islands; brief description) ; Fowler, 1949, p. 106 (in part; reference to Pietschmann, 1938).
Scorpaenopsis cacopsis (not of Jenkins): Epmonpson, 1946, pp. 344-345, fig. 211b (color notes; fairly good figure).
MATERIAL EXAMINED. Holotype: BPBM 10958 (103 mm. S. L.), Oahu, reef in Kaneohe Bay, in 1 meter, J. E. Randall, A. H. and D. Banner, R. E. and J. Brock, 14 July 1971.
Paratypes: ANSP 84921 (1, 98.6), Honolulu, collected by J. W. Thomp- son. BPBM 4350 (1, 119), Honolulu, J. W. Thompson. BPBM 7816 (3, 27.4-53.8), Oahu, Waimea Bay, in 6-9 meters, rock with some sand, J. E. Randall et al., 25 Aug. 1969. BPBM 10182 (1, 108), Oahu, off channel to Kaneohe Bay, in 30 meters, cave at base of small dropoff, J. E. Randall and
300 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 14. Scorpaenopsis brevifrons. a. BPBM 6491, paratype, 95.5 mm. S. L., lacking dark spot on spinous dorsal fin, Oahu. b. BPBM 10958, holotype, with dark spot on dorsal fin, Oahu. (Photographs of fresh specimens.)
EK. Chave, 29 Sept. 1970. BPBM 11985 (1, 81.3), Oahu, % mile off Niumalu Hotel, Waikiki, E. S. Herald, R. R. Rofen, V. E. Brock, W. A. Gosline, eé al., 7 Sept. 1951. BPBM 13350 (1, 116), Oahu, off Makua, in 18 meters, bottom mostly of coral rock and rubble, J. E. Randall, 9 Sept. 1972. BPBM 13351 (1, 57.8), Oahu, Makua, rocky shore in 6 meters, J. E. Randall, G. S. Losey
Vot. XL] ESCHMEVER AND RANDALL: HAWAIIAN SCORPAENIDAE 301
Figure 15. Scorpaenopsis brevifrons, diagram of head spines, based mostly on BPBM 6491.
and class, 9 Sept. 1972. CAS 30231, formerly BPBM 6491 (2, 54.4-95.5), Oahu, off Lahilahi Point, near cave in 27-30 meters, coral rubble and rock, J. E. Randall and University of Hawaii students, 10 Aug. 1968. USNM 214047 (1, 89.5), Pokai Bay, 60 meters seaward from deep dropsite, ledge in 24 meters, W. A. Gosline ef al., Aug. 1969.
ADDITIONAL MATERIAL. NMW 4996-5000 (5, 87.5—-99.5), French Frigate Shoal [no other data; these specimens reported as Sebastapistes asperella by Pietschmann, 1938].
Dracnosts. A species of Scorpaenopsis with pectoral fin rays usually 19 (18-20), a blunt head profile, long jaw (21-27% S. L.), no hump _ behind head, longest dorsal spines 4-6, and normally 5 spines on the suborbital ridge and 3 on the lachrymal bone overlying the maxillary. (See also Comparisons below. )
Description. Measurements and counts summarized in table 3; body shape and coloration as in figures 14-15.
Dorsal fin rays XII, 9%, longest spines the third through sixth. Anal fin rays III, 5’, third anal spine extends beyond second when depressed. Pectoral fin rays usually 19 (18-20), rays 1 or 2 through 6 or 7 branched in larger specimens, branching begins at specimen length of just under 30 mm S. L. Gill rakers, including rudiments, on outside of first arch 14-16, 4—6 on upper arch, 8 plus 2 or 3 indistinct rudiments on lower arch.
Head spination as in figure 15. Lachrymal bone usually with 3 spines over maxillary, first points forward, posterior two close set and point mostly down (specimens under about 50 mm. S. L. usually with only 2 spines). Suborbital ridge usually with 5 spines, first 2 on lachrymal. Preopercle with supplemental and 5 preopercular spines. Other spines include nasal, pre-,
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304 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
supra-, and postocular, tympanic, nuchal, parietal, upper and lower post- temporal, opercular (2, sometimes double), sphenotic (multiple), pterotic, supracleithral, and cleithral. Also small spines (postorbital) behind eye. Occiput with a shallow pit.
Scales on sides ctenoid; head before eye unscaled, otherwise with cycloid scales. Vertical scale rows about 45; lateral-line scales 23 + 1 on caudal fin. Vertebrae 24. Head and body with skin appendages. Supraocular tentacle variable, about equal to eye as maximum, frequently absent. Small flaps present on most head spines, pectoral fin, some body scales, a few on eye, and on the lower jaw and underside of head.
Measurements are summarized in table 3. Orbit smaller than snout, orbit into snout 1.1—-1.6, large specimens with proportionally smaller orbits; orbit into head 3.9—5.6, large specimens with lower values.
Color in alcohol similar to that in figure 14 taken of fresh specimen. Body and head marked with dark patches on a pale background, darkest usually below middle of spinous dorsal fin, below soft dorsal fin, and at base of caudal fin. Fins usually with some dark pigment, especially at center of anal fin and as a broad bar across the caudal fin. Dark spot on spinous dorsal fin present or absent, when present usually between spines 4 through 7 at about midheight of fin. Color in life variable. Upper part of body and most of head brownish to gray with bluish-green to green areas and some yellow, mottled with white. Lower parts with more red and orange. Pelvic fin red, streaked with white, mostly white distally. Anal fin with prominent white bar across anterior base, greenish in middle, otherwise red streaked with white. Dorsal fin mostly pale, mottled with brown, orange, and green.
ComPARISONS. This species is rather typical for a Scorpaenopsis: of moderate size, robust, well colored, with a large mouth and strong spination. It lacks the humpback found in some species (see account of S. diabolus). It has a smaller eye, a longer maxillary, and a blunter head profile than most species of Scorpaenopsis of the Indo-Pacific.
In Hawaiian waters, S. brevifrons most likely would be confused with S. cacopsis and S. diabolus—all brightly colored, moderate to large species. S. diabolus has a characteristic color pattern on the inside of the pectoral fin (fig. 17b), is humpbacked, and has the interorbital width greater than the eye diameter. S. cacopsis has a more pointed snout and usually 18 rather than 19 pectoral rays; the longest dorsal spine in S. cacopsis usually is the third while the longest dorsal spines in S. brevifrons are near the middle of the fin (4-6).
DIsTRIBUTION. This species is known only from Hawaii in depths from 1 to 30 meters, usually in coral or rocks.
Name. The species name is a noun in apposition formed by the combina- tion of the Latin words ‘brevis’ (short) plus ‘frons’ (brow, forehead), re-
Vot. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 305
ferring to the steep head profile in this species as compared to the more pointed, longer snout found in most species of Scorpaenopsis.
Scorpaenopsis diabolus Cuvier. (Figures 16-17.)
Scorpaenopsis diabolus Cuvier in Cuvier and Valenciennes, 1829, p. 312 (original descrip- tion; type locality “le grand Océan oriental”).
Hawaiian references:
Scorpaenenopsis diabolus: Fowtrer, 1900, p. 515 (generic name misspelled; dried skin from Hawaii [could be cacopsis]).
Scorpaena gibbosa (not of Schneider): STEINDACHNER, 1900b, p. 491 (one from Honolulu).
Scorpaenopsus gibbosa (not of Schneider): SEALE, 1902, p. 18 (generic name misspelled; Honolulu).
Scorpaenopsis catocala JORDAN & EVERMANN, 1903a, pp. 201-202, pl. 56 (original description; type locality Hawaiian Islands, holotype (USNM 50651) from Honolulu, paratypes from Honolulu and Hilo); Snyper, 1904, p. 536 (listed; Hawaii); JoRDAN & SNYDER, 1904a, p. 946 (coloration; Honolulu); JorpAN & EvERMANN, 1926, p. 10 (listed; Hawaii).
Scorpaenopsis gibbosa (not of Schneider): JorDAN « EVERMANN, 1905, pp. 468-470, pl. 56, fig. 206 (description; good figures, pl. 56 wrongly with catocala as caption; synonymy) ; JoRDAN & SEALE, 1906, pp. 178, 375 (in part; catocala in synonymy; common about Hawaii); JorpAN & JORDAN, 1922, p. 55 (synonymy; abundant); Fowter, 1925, p. 27 (listed) ; GosLINE & Brock, 1960, pp. 284, 287, 341 (in key; brief description) ; GosLInr, 1965, p. 825 (depth distribution) .
Scorpaenopsis gibbosus: Fow rr, 1925, p. 27 (listed; Hawaii); Fowler, 1928, pp. 286-287 (in part; diabolus and catocala in synonymy; Hawaiian specimens listed); TINKEr, 1944, p. 264, fig. (brief description; figure from Jordan & Evermann, 1905); EpmMonp- SON, 1946, p. 344 (brief description; coloration; reefs of Hawaiian Islands); Fow rr, 1949, p. 106 (in part; reference to Fowler, 1938; record from Hawaii).
Remarks. The species of Scorpaenopsis which have a humpback have at times been recognized as a separate genus as discussed by us under the genus Scorpaenopsis. We follow a broader interpretation of the genus. Within the humpbacked subgroup, there are four species: Scorpaenopsis diabolus, S. gibbosa (Schneider in Bloch and Schneider, 1801), S. macrochir Ogilby, 1910, and S. neglecta Heckel, 1837. Scorpaenopsis diabolus and S. gibbosa fre- quently are confused, and S. macrochir, type locality northeastern Australia, is widespread in the central Pacific based on CAS and BPBM specimens, but has not been reported from there (so far only reported from Australia). Scorpaenopsis neglecta is an Oriental species characterized by serrated ocular spines and the inside of the pectoral fin marked about as in figure 17d. Scorpaenopsis macrochir also has the pectoral fin markings as in figure 17d, but macrochir has unserrated ocular spines among other differences. Scor- paenopsis gibbosa (with S. nesogallica (Cuvier) a synonym) apparently is restricted to the Indian Ocean and S. diabolus, the only humpbacked species occurring in Hawaii, is widespread in the Indo-Pacific faunal region. Scor- paenopsis diabolus and S. gibbosa may be separated easily on the basis of
306 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
pectoral fin coloration (see figure 17a-c); S. diabolus also has a longer snout and wider interorbit than S. gibbosa, and usually 18 rather than 17 pectoral rays.
MATERIAL EXAMINED. USNM 50651, formerly SU 7754 in part (1, 185, holotype of Scorpaenopsis catocala), Oahu, Honolulu, U. S. Fish Commission. SU 7754 (5, 56.3-152) and SU 7466 (5, 115-181), paratypes of S. catocala, Oahu, U. S. Fish Commission. CAS 17488 (1, 58), Leeward Is., Laysan Is., 25-4627 N., 171°4437°W., im depths to 3:5 meters, SU 19344 (Gao), Oahu, Honolulu, D. S. Jordan. SU 23393 (2, 95.2-101), Oahu, Honolulu, E. K. Jordan. Counts are also included for the following Hawaiian speci- mens: CAS 389 (1), CAS 996 (1), CAS 11073 (); CAS 1l0ssG)Raexs 17485 (1), CAS 17486 (1), CAS 17487 (1), CAS 31393 (1) and SU 8406 (1). Much additional Hawaiian material is available in the BPBM collection.
Norte. The type specimens of S. catocala were to be distributed to a number of museums (Jordan and Evermann, 1903a, p. 202) but they were not sent and are now found in SU 7466 and 7754. The specimens bear separate field numbers so it will be possible to distribute them as originally intended, and this will be done at the time of publication of this paper.
DISTINGUISHING FEATURES. Dorsal fin rays XII, 9%. Anal fin rays III, 5%. Pectoral fin rays usually 18. Back arched (see figure 16). Scales ctenoid; about 45 vertical scale rows. Lachrymal bone with 2 or 3 spines over maxillary, first points forward, followed by 1 or by 2 closeset spines which point down and back. Suborbital ridge with 4 or more spinous points, usually
Vor. XL] ESCHMEVER AND RANDALL: HAWAIIAN SCORPAENIDAE 307
Ficure 17. Coloration on inside of pectoral fin in humpbacked species of the genus Scorpaenopsis. a. S. diabolus (restricted dark patch distally); b. S. diabolus, Hawaiian island populations (dark patch distally, note dark pigment at base of fin); c. S. gibbosa; d. S. macrochir and S. neglecta (band complete, no black patch as in c).
more than 8 or 10 points, not in a row and of various sizes. A shallow pit below front corner of eye.
This species is most likely to be confused with Scorpaenopsis cacopsis and S. brevifrons. The arched back distinguishes S. diabolus (fig. 16) along with the characteristic coloration on the inside of the pectoral fin (fig. 17b). Scorpaenopsis brevifrons usually has 19 instead of 18 pectoral rays, while S. diabolus and S. cacopsis usually have 18 pectoral rays. The interorbital width is greater than the orbit diameter in S. diabolus but smaller than the orbit diameter in S. brevifrons and S. cacopsis (in very large specimens, over 200 mm S. L., of cacopsis the interorbital width is about equal to the orbit diameter ).
DistRIBUTION. Scorpaenopsis diabolus is the most widespread species in the genus, occurring from the Red Sea and eastern Africa to the central Pacific as far east as the Society Islands. Specimens from Hawaii seem to
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represent a distinct and recognizable population based on the pectoral fin coloration (fig. 17c). In Hawaii, specimens of the species have been taken in depths from .5 to 10.5 meters.
Scorpaenopsis cacopsis Jenkins.
(Figure 18.)
Scorpaena cookii GUNTHER, 1873, pl. LV (in part; plate only; plate based on a Hawaiian specimen).
Scorpaenopsis cacopsis JENKINS, 1901, pp. 400-402, figs. 13-14 (original description; type locality Hawaiian Islands, Honolulu; holotype USNM 49690); SEare, 1901, p. 11, fig. 5 (description; good figure; two specimens from Honolulu); JENKINS, 1903, p. 497 (one specimen from Hawaii); JoRDAN & EVERMANN, 1905, pp. 467-468, figs. 205, 205a, color pl. 71 (description; 14 specimens from Hawaii; figs. 205 and 205a copied from Jenkins, 1901); JorDAN & SEALE, 1906, p. 375 (listed, Hawaii and Tahiti); JorpAN & JoRDAN, 1922, p. 55 (listed) ; JorpDAN & EVERMANN, 1926, p. 10 (listed) ; Fow Ler, 1928, p. 286, pl. 34B (synonymy; Hawaiian specimens; altirostris wrongly in synonymy); TINKErR, 1944, pp. 263-264, fig. (brief description; compiled range as Society Islands and Hawaii; figure copied from Jenkins, 1901) ; Fow Ler, 1949, p. 106 (reference) ; GOSLINE & BRocK, 1960, pp. 284, 288, 341 (brief description and synonymy) ; GosLInE, 1965, p. 825 (depth distribution) .
Scorpaenopus cacopsis: SEALE, 1902, p. 22 (generic name misspelled; listed).
MATERIAL EXAMINED. USNM 49690 (1, 330, holotype of Scorpaenopsis cacopsis), Hawaiian Islands, collected by T. D. Wood, no other data. BPBM 4387 (1, about 330), Oahu, Haleiwa Bay, Waialua, in 3.5 meters, E. Y. Hosaka, 2 Aug. 1932. BPBM 7852 (2, about 300-360), Oahu, Honolulu, J. W. Thompson, no other data. BPBM 10527 (1, 222), Oahu, Moku Manu, in 26 meters, J. E. Randall and W. J. Baldwin, 6 Oct. 1969. BPBM 13352 (2, 74.3-143), Oahu, Makua, rocky shore in 6 meters, J. E. Randall, G. S. Losey and class, 9 Sept. 1972. BPBM 13733 (1, 41.1), Oahu, Waimea Bay, in 4.5 meters, G. R. Allen, 2 July 1967. BPBM 13739, formerly UH 2095 (1, 22.5), Maui, Baldwin Packer’s property, about 3 miles W. of Lahaina, W. A. Gosline and E. Hunter, 5 Aug. 1955. BPBM 13744 (1, 227), Oahu, off Pakai Bay, in 9 meters, J. E. Randall, S. N. Swerdloff and D. Chave, 29 July 1969. SU 23263 (1, 220), Honolulu, Albatross.
DISTINGUISHING FEATURES. Dorsal fin rays XII, 9%. Anal fin rays III, 5%. Pectoral fin rays normally 18 (17-19). Scales ctenoid; about 50-55 vertical scale rows. Lachrymal bone with 2 spines over maxillary; first points down and forward, second points down and back, sometimes second split distally into 2 points. Suborbital ridge usually with 5 spines, sometimes some spines split in larger specimens. See also the distinguishing features sections of Scorpaenopsis brevifrons and S. diabolus.
DisTRIBUTION. This species is confined to the Hawaiian Islands. Available depths of capture are from near shore to 26 meters.
Vor. XL] ESCHMEYVER AND RANDALL: HAWAIIAN SCORPAENIDAE 309
Scorpaenopsis altirostris Gilbert.
(Figure 19.)
Scorpaenopsis altirostris GiLBerT, 1905, pp. 628-630, fig. 244 (original description; type locality Hawaiian Islands, off Molokai, Albatross station 3849; holotype USNM 51636); JORDAN & SEALE, 1906, p. 376 (name only); JoRDAN & JORDAN, 1922, p. 55 (name; suggested may belong in a genus separate from Scorpaenopsis); JORDAN & EVERMANN, 1926, p. 10 (listed); BoHrKeE, 1953, p. 121 (location of types); GOSLINE & Brock, 1960, pp. 285, 289, 342 (compiled); Gostinr, 1965, p. 825 (compiled depth distribution).
Remarks: See Remarks under the genus Scorpaenopsis regarding generic placement of this species.
MATERIAL EXAMINED. USNM 51636 (1, 45.6, holotype of Scorpaenopsis altirostris), USNM 51671 (3, 32.0-39.9, paratypes of S. altirostris, poor con- dition) and SU 8620 (2, 33.0-39.2, paratypes of S. altirostris), off south coast of Molokai, N. at 71°, W. 21.9’ from Lae-o Ka Laau Light, in 134— 79 meters, 10-ft. Blake trawl, bottom of coarse sand, broken shells, and coral, Albatross station 3849, 8 April 1902.
DISTINGUISHING FEATURES. Dorsal fin rays XII, 9%. Anal fin rays III, 5%. Pectoral fin rays 17-18, mostly 18. Scales ctenoid; about 45 vertical scale rows. Lachrymal bone with 2 spines over maxillary; anterior spine points forward and down, posterior spine points down and back. Suborbital ridge usually with 4 spines, first on lachrymal below main suborbital ridge,
310 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
second under eye and below main ridge, third and fourth to rear. A slight occipital pit present.
This species and S. fowleri resemble species of Scorpaena more than species of Scorpaenopsis but they lack palatine teeth (see Remarks under the genus Scorpaenopsis). Scorpaenopsis fowleri has only 16 pectoral rays as opposed to 17-18 in altirostris, and S. fowleri is a near-shore species while S. alti- rostris occurs in deeper water.
DIsTRIBUTION. Scorpaenopsis altirostris is an upper slope species known only from the type specimens listed in the material examined section. Depth of capture was between 79-134 meters.
REMARKS. Scorpaenopsis cotticeps Fowler, 1938, from the Philippines might prove to be the same species. Fowler wrongly described it as having 14 pec- toral fin rays but the type (USNM 98891) has 17. The type specimen is only 27.4 mm. S. L.
Genus Rhinopias Gill
Rhinopias GiL, 1905, p. 225 (type-species Scorpaena frondosa Giinther, 1891, by original designation). Peloropsis GILBERT, 1905, p. 630 (type-species Peloropsis xenops Gilbert, 1905, by original designation) . ’
The genus Rhinopias has been treated in detail by Eschmeyer, Hirosaki, and Abe (1973).
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 311
Rhinopias xenops (Gilbert). (Figure 20.)
Synonymy based only on Hawaiian references:
Peloropsis xenops GILBERT, 1905, pp. 630-631, fig. 245 (original description; type locality Hawaiian Islands, between Maui and Lanai, Albatross station 3872; holotype USNM 51604); JorDAN & SEALE, 1906, p. 379 (listed); JoRDAN & JORDAN, 1922, p. 55 (listed; compiled locality data); JorpAN & EvERMANN, 1926, p. 10 (listed); FOWLrr, 1928, p. 287 (compiled) ; TinKEr, 1944, pp. 262-263, fig. (compiled; figure of type copied from Gilbert, 1905) ; Gostine & Brock, 1960, pp. 284, 286, 341 (in key; compiled) ; GosLInE, 1965, p. 825 (depth distribution compiled).
Rhinopias xenops: EESCHMEYER, Hirosaki, & ABE, 1973, pp. 288, 292-295, figs. 3-4 (synonymy; additional Hawaiian specimen and Japanese material; description; com- parisons; shedding of cuticle; figure of holotype copied from Gilbert, 1905; second Hawaiian specimen figured).
MATERIAL EXAMINED. USNM 51604 (1, 110 mm. S. L., holotype), Auau Channel, between Maui and Lanai islands, in 59 to 79 meters, Albatross station 3872, 12 Apr. 1902. BPBM 13988 (1, 115), Oahu, off Haleiwa, in 110 meters, shrimp trawl, Robert May, Oct. 1972 [kept alive in an aquarium for 2 months]. UH 3102 (1, 153), Oahu, off Waikiki, otter trawl in 73 meters, P. S. Lobel, 9 Feb. 1974. USNM 209415 (1, 107), Hawaii, Haleiwa,
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21°39.6—42’N., 158°07.3’-05’W., shrimp trawl, in 95-110 meters, Townsend Cromwell cruise 36, station 19, 3 May 1968.
DISTINGUISHING FEATURES. Dorsal fin rays XII, 9%. Anal fin rays III, 5%. Pectoral fin rays 18. Scales cycloid; vertical scale rows about 70-75. Palatine teeth absent. Lachrymal bone with 2 rounded lobes over maxillary. Suborbital ridge with 3 or 4 spines, not well marked and sometimes split, first spine on lateral face of lachrymal bone.
This species is distinguished from other Hawaiian species by its com- pressed head and body, somewhat elevated rostrum, long snout, and colora- tion. A complete description is given by Gilbert (1905) and by Eschmeyer, Hirosaki, and Abe (1973).
The specimen figured differs from previously known specimens in having a dark spot on the spinous dorsal fin and it lacks dusky pigment at the base of the soft dorsal fin. When fresh it was reddish with large areas of light purple. In other features it agrees with other specimens of R. xenops. UH 3102 differs from previously known specimens by having 19 pectoral rays and the third dorsal spine is not elongate.
DiIsTRIBUTION. We know this offshore species at Hawaii only from the holotype and 3 additional specimens as given in our “Materials examined” section. One additional specimen was taken off Molokai in 124 meters by the Townsend Cromwell (Paul Struhsaker, pers. comm.) but this specimen was lost. Depths of capture were 59-79, 73, 95-110, and 124 meters. Rhinopias xenops is known also from Japan (see Eschmeyer, Hirosaki, and Abe, 1973).
Remarks. William D. Madden and Randall observed the feeding of the specimen shown in figure 20 when it was maintained in an aquarium. Live Gambusia were introduced into the tank as food. The Rhinopias moved very slowly and intermittently towards its prey by “creeping” on its lower pectoral rays which were in contact with the bottom. The dorsal fin was held fully erect. Occasionally the RAinopias moved a little to and fro, reminiscent of the rocking of Taenianotus. When it came to within about 2.5 cm. of the prey, it engulfed the small fish with incredible rapidity. Just prior to striking the prey, the dorsal fin was slowly folded back.
Genus Neomerinthe Fowler Neomerinthe Fow rr, 1935, pp. 41-42 (type-species Neomerinthe hemingwayi Fowler, 1935, by original designation).
Eschmeyer (1969b, p. 93) remarked that the Hawaiian species described as Helicolenus rufescens Gilbert did not belong in the genus Helicolenus of the subfamily Sebastinae but should be classified instead in the subfamily Scorpaeninae. The species rufescens seems to be referable to the genus Neomerinthe Fowler. Neomerinthe is a genus which contains Pontinus-like species, but unlike species of Pontinus those of Neomerinthe have some
Vot. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 313
branched pectoral rays. Previously, only two Atlantic species have composed the genus Neomerinthe.
Neomerinthe rufescens (Gilbert).
(Figure 21.)
Helicolenus rufescens GILBERT, 1905, pp. 631-633, fig. 246 (original description; type locality Hawaiian Islands, holotype (USNM 51628) from near Kauai at Albatross station 4133, paratype from off Maui at Albatross station 4074); JorDAN & SEALE, 1906, p. 378 (listed) ; JoRDAN & JORDAN, 1922, p. 54 (name; locality data compiled); JoRDAN «& EVERMANN, 1926, p. 10 (listed); Fowtrer, 1928, p. 291 (synonymy; brief description compiled from Gilbert, 1905); Tinker, 1944, pp. 267-268, fig. (compiled; figure of type from Gilbert, 1905); BouriKr, 1953, p. 120 (location of types) ; GosLINE & Brock, 1960, pp. 285, 289, 342 (in key; compiled) ; Gosting, 1965, p. 825 (depth distribution).
MATERIAL EXAMINED. USNM 51628 (1, 83.4, holotype of Helicolenus rufescens), off Kauai, S. at 40° and W. 4.4’ from Hanamaulu warehouse, in 75-302 meters, bottom of fine gray sand and rubble, 8-ft. Albatross-Blake trawl, station D. 4133, 1 Aug. 1902. BPBM 13737 (5, 59.1-90.8), CAS 15697 (9, 61.3-92.4) and CAS 15689 (3, 75.0-100, cleared and stained), all off Molokai, 21°14.4-15.6’N., 157°07.8"-13.5’°W., shrimp trawl in 124 meters, Townsend Cromwell cruise 40, station 106, 28 Nov. 1968. SU 8619 (1, 59.5, paratype of Helicolenus rufescens), north coast of Maui, S. at 70° and E. 8.5’ from Puniawa Point, in 143-155 meters, bottom of coral, sand, and forami- nifera, 8-ft. Tanner beam trawl, staton D. 4074, 19 July 1902. Additional Hawaiian material is available in the NMFS collection.
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DISTINGUISHING FEATURES. Dorsal fin rays XII, 9%. Anal fin rays III, 5%. Pectoral fin rays usually 18, sometimes 19. Palatine teeth present. Scales ctenoid; about 50 vertical scale rows. Lachrymal bone with 2 spines over maxillary, both point back and down. Suborbital ridge with 2—4, usually 3, spines; one on lateral face of lachrymal bone usually absent. No occipital pit.
This species is most likely to be confused with species of Scorpaena or Pontinus. Neomerinthe rufescens has 18 or 19 pectoral rays while all Hawaiian species of Scorpaena and Pontinus usually have 17 or fewer pectoral rays (S. colorata sometimes with 18). Upper pectoral rays in N. rufescens branch when specimens reach about 70 mm. S. L., but in Pontinus macrocephalus they never branch; specimens of rufescens usually have 9% soft dorsal rays and macrocephalus usually 10'2. Species of Scorpaenopsis differ in lacking palatine teeth.
DistRIBUTION. Neomerinthe rufescens is an offshore species known only from the Hawaiian Islands. Townsend Cromwell operations captured a total of 63 specimens from 18 stations in depths between 108 and 124 meters (Struhsaker, 1973). The holotype was taken somewhere between 75 and 302 meters and the paratype between 143 and 155 meters by the Albatross.
Genus Scorpaena Linnaeus
Scorpaena LINNAEUS, 1758, p. 266 (type-species Scorpaena porcus Linnaeus, 1758, by subse- quent designation of Bleeker, 1876; see Opinion 77 and “Official List of Generic Names’’).
Problems of the limits of this genus and generic synonyms have been mentioned by Eschmeyer (1965a, p. 89; 1969b, p. 54) and Eschmeyer and Allen (1971, p. 521). Some of the Hawaiian species are referable to Sebastapistes, but we follow Matsubara (1943) in treating Sebastapistes as a synonym of Scorpaena. It is likely that the genus Scorpaena is a ‘catch- basket’ genus, and a more thorough treatment on a world basis is needed. See also our remarks under the genus Scorpaenopsis.
Scorpaena galactacma (Jenkins). (Figure 22a.)
Sebastapistes galactacma JENKINS, 1903, pp. 496-497, fig. 40 (original description; type locality Hawaiian Islands, coral reef at Honolulu; holotype USNM 50692); JorpAaN & EVERMANN, 1905, pp. 455, 459-460, fig. 201 (description mostly from Jenkins, 1903; figure of type from Jenkins, 1903); JorpAN & JORDAN, 1922, p. 54 (listed; common on reefs).
Sebastapistes galactacme: JORDAN & SEALE, 1906, p. 377 (species name misspelled; listed).
Sebastapistes albobrunneus (not of Gtinther): Fowter, 1928, p. 287 (in part; galactacma in synonymy).
?Sebastopsis galactma: Boropin, 1930, p. 59 (species name misspelled; specimens from Oahu; compared with type).
Scorpaena balliewt (not of Sauvage): GosLInE & Brock, 1960, p. 342 (in part; galactacma in synonymy).
FicurE 22. a. Scorpaena galactacma, BPBM 7925, 33 mm. S. L., Oahu (Photograph of a preserved specimen). b. Scorpaena coniorta, BPBM 6464, 47 mm. S. L., Oahu. (Photo- graph of fresh specimen.)
MATERIAL EXAMINED. USNM 50692 (1, 49.3 mm. S. L., holotype of Sebastapistes galactacma), and USNM uncat. (2, about 36-40, paratypes of S. galactacma), Oahu, Honolulu, O. P. Jenkins, 1889. BPBM 7835 (7, 33.4— 47.8), Oahu, off Waikiki, in 7.5 meters, J. E. Randall, E. Niel, and students, 30 Mar. 1969. BPBM 7925 (1, 32.8), Oahu, Waimea Bay, in 6-9 meters, J. E. Randall, W. F. Smith-Vaniz, e¢ al., 25 Aug. 1969. BPBM 13470 (3,
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33.5-36.4), Oahu, off Waikiki in 29 meters, from vicinity of small ledge, jE. Randall, 30 Sept. 1972. CAS 13471, formerly UH 2484 (10), Kauai, Port Allen, in 24-29 meters, Miss Honolulu, 10 Sept. 1959. CAS 13472, formerly UH 2481 (40), Kauai, Port Allen, in 10.5—24.5 meters, Miss Honolulu, 12 Sept. 1959. CAS 15725 (2), Maui, dredged off Lahaina, in 18-64 meters, E. M. Ehrhorn. SU 7615 (6), Oahu, Honolulu, U. S. Fish Commission, 1901. Additional material is present in the BPBM collection.
DISTINGUISHING FEATURES. Dorsal fin rays XII, 9%. Anal fin rays III, 5’. Pectoral fin rays 15-17, usually 16. Coronal spines absent. Virtually no occipital pit. Vertical scale rows about 45. Lachrymal bone with 2 widely diverging spines over maxillary, first points forward, second back. Suborbital ridge with 2 or 3 poorly marked spines, usually one under eye below main ridge, one at end, and sometimes one on lachrymal bone below main ridge. A dark smudge sometimes present between dorsal spines 6—7 to 9.
This species differs from other Hawaiian species of the genus Scorpaena by having cycloid to emarginate scales; the other species have ctenoid scales at least on their flanks.
DIsTRIBUTION. This small species appears to inhabit coral and rubble areas in depths from near shore to 29 meters (one collection from 18 to 64 meters).
Scorpaena coniorta (Jenkins). (Figure 22b.)
Sebastapistes strongia (not of Cuvier): STREETS, 1877, pp. 62-63 (description; Honolulu) ; Fow er, 1900, p. 515 (one from Hawaii; fin formula).
Sebastapistes coniorta JENKINS, 1903, pp. 495-496, fig. 39 (original description; type locality Hawaiian Islands, coral rocks on reef at Honolulu; holotype USNM 50693); Snyper, 1904, p. 536 (listed; Honolulu) ; Jorpan « EverMANN, 1905, pp. 455, 458-459, fig. 200 (description; brief synonymy; figure from Jenkins, 1903); JorDAN & SEALE, 1906, p. 376 (listed) ; JoRDAN & JORDAN, 1922, p. 54 (listed; common on reefs) ; FowrLer & Batt, 1925, p. 20 (Pearl and Hermes Reef); JorDAN & EvERMANN, 1926, p. 10 (listed) ; BOHLKE, 1953, p. 122 (location of types) ; ScHuLTz, 1966, p. 28 (coniorta distinct from albobrunneus) .
Sebastapistes albobrunneus (not of Gunther): Fow rrr & Batt, 1925, p. 20 (Pearl and Hermes Reef); Fow ter, 1928, p. 287 (in part; S. coniorta in synonymy; listed some Hawaiian specimens); PIrTSCHMANN, 1938, pp. 28-29, pl. 3 (description; specimens from Pearl and Hermes Reef [some misidentified]; photograph of 7 specimens; year- groups by length frequencies [based on misidentifications]) ; Tinker, 1944, pp. 264-265, fig. (brief description; figure of type of coniorta from Jenkins, 1903); Epmonpson, 1946, p. 344, fig. 21la (brief description).
Sebastapistes albo-brunneus: Fowtrr, 1941, p. 257 (one specimen from Mogua Reef).
Scorpaena coniorta: GosLiInrE, 1955, pp. 461-462 (described in key, Johnston Island) ; GosLINE & Brock, 1960, pp. 285, 288, 341 (brief diagnosis; in key; brief synonymy) ; GosLINnE, 1965, p. 825 (vertical zonation).
MATERIAL EXAMINED. USNM 50693 (1, 49.8 mm. S. L., holotype of Se- bastapistes coniorta), Oahu, off Honolulu, reefs, O. P. Jenkins, 1889. SU 23589
VoL. XL] ESCHMEYVYER AND RANDALL: HAWAIIAN SCORPAENIDAE 317
(57, paratypes of S. coniorta) and SU 23330 (122, paratypes of S. coniorta), Oahu, reefs off Honolulu, O. P. Jenkins. BPBM 6464 (4, 40.5-45.8), Oahu, western end of Waimea Bay, in 6 meters, all from head of Pocillopora mean- drina, J. E. Randall, 28 Sept. 1968. BPBM 9987 (1, 35.3) and CAS 15695 (4), Oahu, off rocky islet at SW. end of Waimea Bay, in 1—10.5 meters, J. E. Randall, L. A. Randall and P. M. Allen, 27 July 1970. CAS 13480, for- merly UH 809 (10), Oahu, Diamond Head, W. A. Gosline and class, 16 May 1950. CAS 15710 (16) and CAS 15711 (3, 28.5-47.8, cleared and stained), Oahu, Waikiki Beach, about % mi. off Niumalu Hotel, in 7.5—9 meters, W. A. Gosline e¢ al., GVF Reg. no. 54, 7 Sept. 1971. UH uncat. (5, 30.2-53.7), Wake Island, near wreck, 0-6 meters, W. A. Gosline and J. E. Randall, 9 June 1953. Additional material is available in the BPBM and CAS collections.
DISTINGUISHING FEATURES. Dorsal fin rays normally XII, 9'2. Anal fin rays III, 5%. Pectoral fin rays usually 16 (15-17). Coronal spines absent. No occipital pit. Scales ctenoid; about 50-55 vertical scale rows. Lachrymal bone with 4 spines; two spines lie over the maxillary, first points forward, second points back; a spine above each of these. A small spine sometimes present at base of first spine over maxillary in large specimens. Suborbital ridge more or less a double ridge; a spine at end; sometimes a spine under eye; fourth spine on lachrymal bone is far below level of suborbital ridge and not counted as part of suborbital ridge. Supplemental preopercular spine well above first preopercular spine. Well-marked spines at upper rear of orbit in area of sphenotic.
The coloration of this species is diagnostic: body and fins with scattered small dark spots, no dark spot on spinous portion of dorsal fin; see figure 22b.
DIsTRIBUTION. This species is known from the Hawaiian Islands and from Johnston Island and the Line Islands (e.g., CAS 13479). Other coniorta-like species occur in the Indo-West Pacific area as discussed below. In Hawaii the species occurs in shallow water in depths from near shore to about 24.5 meters.
REMARKS. Scorpaena coniorta belongs in the subgroup Sebastapistes within Scorpaena. Within this subgroup, it appears to be related to the widespread Indo-Pacific Scorpaena albobrunnea Ginther. Scorpaena coniorta also seems closely related to the little-known Asian S. tinkhami Fowler (1946), and _ to BPBM specimens of a similar species taken by Randall in French Oceania. A more thorough study is needed of this complex.
Scorpaena ballieui Sauvage. (Figure 23.)
Scorpoena ballieui SAuvacE in Vaillant and Sauvage, 1875, pp. 278-279 (original description ; type locality Iles Sandwich [Hawaiian Islands]; generic name misspelled).
Scorpaena ballieui: SAuvacE, 1878, pp. 123-124, fig. 4 on pl. 2 (description; good figure) ; JorpAN & SEALE, 1906, p. 376 (listed); GostINE & Brocx, 1960, pp. 285, 289, 341-342
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(in key; brief description; galactacma and fowleri included in synonymy); GosLInE, 1965, p. 825 (depth distribution).
Sebastapistes corallicola JENKINS, 1903, pp. 493-495, fig. 38 (original description; type locality Hawaiian Islands, Honolulu; holotype USNM 50691); Snyper, 1904, p. 535 (listed from Honolulu and Hilo); Jorpan « EverRMANN, 1905, pp. 455-458, fig. 199 (mostly after Jenkins, 1903; figure from Jenkins, 1903); JorpAN « SEALE, 1906, p. 376 (thought might be same as asperella); JORDAN & JORDAN, 1922, p. 54 (compiled) ; FowLer & Batt, 1925, p. 19 (Pearl and Hermes Reef, Ocean Island, and Laysan) ; PIETSCHMANN, 1938, pp. 5, 29-30 (brief description; Pearl and Hermes Reef); BOHLKE, 1953, p. 122 (location of types) ; ScHuLtz, 1966, pp. 29-32, figs. 138b, 140 (name used for Marshall and Marianas specimens; figure of holotype of corallicola from Jenkins, 1903).
Sebastapistes ballieut: JORDAN & EVERMANN, 1905, pp. 455-456, color pl. 72 (lengthy de- scription; Honolulu, Waikiki, Hilo); Jorpan «& JorDAN, 1922, p. 54 (name; rather common); FowLer & Barr, 1925, p. 19 (Pearl and Hermes Reef).
Sebastapistes albobrunneus (not of Gtinther): Fowrer, 1928, p. 287 (in part; coniorta and galactacma wrongly in synonymy; Hawaiian specimens among others listed).
Sebastapistes asperella (not of Bennett): Fowrer, 1931, pp. 348-349 (color description of Honolulu specimens) ; Fowrer, 1949, p. 106 (in part; references).
Sebastapistes nuchalus (not of Giinther): ScHurtz, 1943, p. 174 (in part; type of coralli- cola only) ; Fow er, 1949, p. 107 (in part; corallicola wrongly included in synonymy).
Scorpaena peruana HILDEBRAND, 1946, pp. 445-448, fig. 86 (original description; type locality Peru [in error, see remarks below]).
MATERIAL EXAMINED. MNHN 6883 (2, about 32 and 86.0), MNHN 8993 (1, 79.5), and MNHN 9557 (2, 62.4-74.3), all syntypes of Scorpaena ballieui, Hawaii, collected by Ballieu. USNM 50691 (1, about 95, holotype of Se- bastapistes corallicola), Oahu, Honolulu. SU 7729 (1, paratype of Sebastapistes corallicola), Oahu, Honolulu, O. P. Jenkins. BPBM 4369 (1, 64.5), Oahu, C. M. Cooke, Jr., 5 May 1923. BPBM 7817 (2, 47.3-61.5), Oahu, off Waikiki, in 7.5 meters, J. E. Randall, E. Niel, and students, 30 Mar. 1969. BPBM 9338 (1, about 67), Oahu, off Kewalo Basin, in about 1 meter, P. M. Allen, 20 Mar. 1970. CAS 13470, formerly UH 264 (15), Oahu, Waianae coast, W. A. Gosline and class, 22 Feb. 1949. CAS 15692 (8), Oahu, large boulders and some sand at west side of Waimea Bay, in 6-9 meters, J. E. Randall et al., 25 Aug. 1969. CAS 15694 (8), Oahu, off rocky islet at SW. end of Waimea Bay, 1-10.5 meters, J. E. and L. A. Randall and P. M. Allen, 27 July 1970. CAS 15709 (2), Oahu, % mi. off Waikiki Beach, in 7.5—-9 meters, GVF Reg. no. 54, W. A. Gosline e¢ al., 7 Sept. 1951. CAS 22841 (44) and CAS 15688 (3, 40.9-79.8, cleared and stained), Kauai, Kapaa, P. R. Needham and J. P. Walsh, 8 Aug. 1949. Additional material is present in the BPBM and CAS collections.
DISTINGUISHING FEATURES. Dorsal fin rays normally XII, 9%. Anal fin rays III, 5%. Pectoral fin rays 16. Coronal spines present. Virtually no occipital pit. Scales ctenoid; about 40-45 vertical scale rows. Lachrymal bone with 3 spines over maxillary, posterior 2 closeset and pointing down
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 319
and to rear, first points mostly forward. Suborbital ridge with one spine at posterior end of ridge.
This is the only Hawaiian species which has coronal spines (fig. le). A dark spot frequently is present on the spinous dorsal fin between about spines 7—10.
DISTRIBUTION. Scorpaena ballieui occurs in or near coral at depths from near shore to about 10.5 meters. We know it definitely only from Hawaii but similar species occur in the Pacific and need study.
REMARKS. Scorpaena asperella Bennett (1829) was the first scorpaenid described from the Hawaiian Islands, but it still remains unidentifiable in our opinion. The type specimen is lost. This problem has been discussed by Gosline (1955) and mentioned by Springer (1967). No Hawaiian species fits Bennett’s description well. The capture site of a blenny described by Bennett from Hawaii actually was off South America (see Springer, 1967), and this adds doubt to the provenance of the specimen described as asperella by Bennett. The collections used by Bennett originated from a cruise of the Blonde, which stopped at Madeira, several areas on the Pacific coast of South America, Hawaii, and other ports in the Pacific. We have been unable to link Bennett’s brief description of asperella with any scorpaenid from the localities visited by the Blonde and we retain it for now as an unidentifiable species.
Scorpaena peruana Hildebrand (1946) is a synonym of S. balliewi; the
320 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 47TH SER.
type specimens of peruana had incorrect locality data accompanying them (see Greenfield, MS).
Scorpaena pele Eschmeyer and Randall, new species. (Figures 24, 25a.)
No literature applies to this species.
MATERIAL EXAMINED. Except for the specimen from the Honolulu market, all specimens were collected by the Townsend Cromwell using shrimp trawls.
Holotvpe: USNM 214046 (124 mm. S. L.), north coast of Oahu, 21°40'N., 158°08’W., 176-202 meters, cruise 59, station 3, 7 July 1972.
Paratypes: ANSP 130800 (1, 116), off Maui, Pailolo Channel, 21°02.3’— 20°58.9'N., 156°44.4’-49.3’W., in 229-238 meters, cruise 40, station 56, 18 Nov. 1968. BPBM 4350 (1, about 134, soft), Honolulu market, J. W. Thompson, no other data. BPBM 13635 (1, 76.5), off Molokai, Penguin Bank, 21°09.7’N., 157°25’-29.8’W., in 177-188 meters, cruise 35, station 33, 7 April 1968. BPBM 17247 (2, 88.5-99.0), off Maui, 21°02.1—20°59.0N., 156°44.4’-44.0’°W., in 238 meters, cruise 40, station 61, 18 Nov. 1968. CAS 15707 (1, 81.4), off Maui, Pailolo Channel, 21°01.2’-20°57.2’N., 156°44.1— 47.1°W., in 210 meters, cruise 40, station 47, 16 Nov. 1968. CAS 30236 (1, 88.7), Pailolo Channel, in 219 meters, no other data. CAS 30237 (2, 110- 112), off Maui, 21°01.7—20°58.8’N., 156°43.1-45.1’W., in 185-232 meters, cruise 35, station 4, 28 Mar. 1968.~CAS 30238 (3, 84.8, 98.9; dij)eeott Maui, 21°01.7-20°57.3’N., 156°43.1—47.4’W., in 221 meters, cruise 40, station 51, 17 Nov. 1968. CAS 30239 (1, 118), taken with the holotype. USNM 214045 (1, 90.8), off Maui, Pailolo Channel, 21°01.6’—20°57.3’N., 156°43.0— 47.4°W., in 223 meters, cruise 40, station 54, 18 Nov. 1968. USNM 214043 (1, 87.1), off Maui, 21°02.5’-20°58.9'N., 156°44.0-49.0’°W., in 229-243 meters, cruise 40, station 55, 18 Nov. 1968. USNM 214044 (2, 75.9-110), off Maui, 21°03.0’-01.2’N., 156°45.4’-50.2’W., in 199-230 meters, cruise 40, station 6/7, 19 Nov. 1968. USNM 214042 (1, 135), off Maui, 21°03.1—01.4'N., 156°45.5— 50.8’°W., in 198-223 meters, cruise 40, station 68, 20 Nov. 1968.
Additional material is available in the NMFS collections, but was not used in the description of the species.
Diacnosis. A species of Scorpaena with 9% dorsal soft rays, 16-17 pec- toral rays, and 15-17 gill rakers. Scales on sides ctenoid. Occiput with a shallow pit. Lachrymal bone with 3 spines over maxillary; suborbital ridge with 4 spines. Coronal spines absent. Color mostly red, marbled and spotted with white.
DescripTIon. Measurements and counts summarized in table 4; body shape and coloration as in figures 24, 25a.
Dorsal fin rays normally XII, 9’, third spine usually the longest. Anal fin rays III, 5%, second spine usually extending beyond third when depressed.
321
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE
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CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH Ser.
Scorpaena pele, CAS 30236, paratype, 88.7 mm. S. L., Pailolo Channel. b: Scorpaena colorata, BPBM 14130, 63.0 mm. S. L., Pailolo Channel (second dorsal spine abnormally short). (Photographs of fresh specimens.)
Pectoral fin rays 16-17, almost always 17, rays 2 through 7 or 8 branched in adults. Gill rakers, on outside of first arch including rudiments total 15-17, 4—6 on upper arch, 10—12 on lower arch, usually 5 + 11.
Head spination as in figure 24. Lachrymal bone with 3 spines over
VoLt. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 323
maxillary, first points forward, second near base of or on first and points forward, third points down and slightly to rear. Suborbital ridge with four spines, first on lateral face of lachrymal bone. Preopercle with supplemental and 5 preopercular spines, fifth preopercular spine sometimes tiny. Other spines include nasal, pre-, supra-, and postocular, tympanic, nuchal, parietal, upper and lower posttemporal, opercular, sphenotic (multiple), pterotic, supra- cleithral, and cleithral. Interorbital ridges moderate, ending at or near base of tympanic spines. Occiput with a shallow to moderate pit, usually poorly defined.
Scales on sides ctenoid; scales on belly and pectoral fin base cycloid. Head before eyes unscaled. Buried scales present behind eye and on cheek. Breast with buried scales. Vertical scale rows about 45; lateral-line scales usually 23 plus 1 on caudal fin. Vertebrae 24. Head and body with small skin appendages. Supraocular tentacle somewhat frilly distally, flattened, its length usually about one-half of orbit diameter. Lachrymal, preopercular, and preorbital spines with small skin flaps; small flaps on eye, on anterior nostril, and a few small ones at other locations on head.
Measurements are summarized in table 4. Orbit diameter about equal to snout length (orbit into snout .9-1.1); orbit into head 3.3-3.8; larger speci- mens tend to have proportionally smaller orbit diameters.
Color in alcohol as in figure 24. Body and head marbled and _ spotted with brown on a pale background. Dark spot on dorsal fin between spines 6-10, sometimes this spot reduced or absent. Large dark patches on flanks under scales (these prominent, purplish in some specimens). Smaller, scat- tered black spots present on body and on caudal fin and usually also on other fins. Broad lines radiate from eye, two ventral ones most noticeable. Color in life mostly red, marbled and spotted with white. All dark areas in figure 25a red in life.
COMPARISONS. Scorpaena pele is very similar to Scorpaena colorata in shape, live coloration, and spination. Both are upper slope species. They differ in that S. colorata lacks roundish black spots on the caudal fin, and usually on other fins, which are present in S. pele. The tooth patch on the palatine of S. colorata is wider and the palatine teeth are scattered, while in S. pele the teeth are in a narrow band of not more than two rows. The scales on the breast of S. pele are buried so the area appears unscaled, while they are readily apparent in S. colorata. Scorpaena pele invariably has a spine on the lateral face of the lachrymal bone, yielding a total of 4 on the “suborbital ridge”; S. colorata lacks a spine on the lateral face of the lachrymal bone and has 3 spines on the suborbital ridge. These two species are distinguished from other species by the characters presented in the key.
DISTRIBUTION. Known only from the Hawaiian Islands; all specimens were trawled in depths from about 180-240 meters.
[Proc. 47TH Ser.
CALIFORNIA ACADEMY OF SCIENCES
324
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326 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SEr.
Name. The specific name is based on Pele, the Hawaiian goddess of vol- canoes and volcanic fires, alluding to the red coloration of the species. It is to be treated as a noun in apposition.
Scorpaena colorata (Gilbert).
(Figure 25b.)
Sebastapistes coloratus GILBERT, 1905, pp. 627-628, fig. 243 (original description; type locality Hawaiian Islands, off Molokai, Albatross station 3849, holotype USNM 51631, plus paratypes from Albatross stations 3849 and 3850); JoRDAN & SEALE, 1906, p. 376 (listed from Hawaii) ; JorpAN & JoRDAN, 1922, p. 54 (compiled); JorpAN « EVERMANN, 1926, p. 10 (listed) ; BOHLKE, 1953, p. 122 (location of types).
Sebastapistes bynoensis (not of Richardson): TINKER, 1944, p. 266 (compiled; figure of type of colorata copied from Gilbert, 1905).
Scorpaena coloratus: GOsLINE & Brock, 1960, pp. 285, 288, 289, 341 (in key; compiled brief description; synonymy); GosLINE, 1965, p. 825 (compiled depth distribution).
MATERIAL EXAMINED. USNM 51631 (1, 57.2 mm. S. L., holotype of Sebastapistes coloratus), and USNM 51667 (2, 44.0-50.8, paratypes of Se- bastapistes coloratus), south coast of Molokai, N. at 71°, W. 21.9’ from Lae-o Ka Laau Light, in 133.5—-78.5 meters, coarse sand, broken shells and coral, 10-ft. Blake trawl, Albatross station 3849, 8 Apr. 1902. SU 8618 (1, para- type of Sebastapistes coloratus), south coast of Molokai, N. at 74°15’, W. 22.2’ from Lae-o Ka Laau Light, in 78.5-121 meters, coarse sand, broken shells, and coral, 10-ft. Blake trawl, Albatross station 3850, 8 Apr. 1902. BPBM 8808 (2, 43.8-63.3), Oahu, off Haleiwa, 21°40’N., 158°07’W., Town- send Cromwell, 3 Mar. 1968. BPBM 14130 (1), off Pailolo Channel, in 219 meters, Townsend Cromwell (no other data). BPBM 13734 (4, 61.2—70.3), CAS 15705 (8, 43.9-88.0), and CAS 15706 (3, 75-95, cleared and stained), off Oahu, Haleiwa, 21°39.4’-42.5’N., 158°07.1’W., shrimp trawl in 102 meters, Townsend Cromwell cruise 40, station 112, 30 Nov. 1968. CAS 15699 (1), off Molokai, 21°14.4-15.7’N., 157°08.2’-14.5’'W., shrimp trawl in 119 meters, Townsend Cromwell cruise 40, station 41, 13 Nov. 1968. Additional material is present in the NMFS and CAS collections.
DISTINGUISHING FEATURES. Dorsal fin rays XII, 9%. Anal fin rays III, 5%. Pectoral fin rays usually 17 (16-18). Coronal spines absent. Shallow to moderate occipital pit. Scales on sides ctenoid; about 45 vertical scale rows. Lachrymal bone usually with 3 spines, second small and at base of first and both of these directed mostly forward, posterior spine points out and slightly back; middle spine develops when fish is about 40 mm. S. L. Suborbital ridge usually with 3 spines, first under eye, second under rear of eye, third at end of ridge. Often a dark blotch on distal part of spinous dorsal between about spines 7-9,
Scorpaena colorata closely resembles Scorpaena pele in live coloration and in most features, but S. colorata has a scaled breast, while the area before
Vout. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 327
the pelvic fins is unscaled or with only a few buried scales in S. pele (see also the account of S. pele).
DISTRIBUTION. Scorpaena colorata is known only from the Hawaiian Islands in depths of about 100-219 meters, with some from uncertain depths between 79-133 meters.
LITERATURE CITED
BENNETT, E. T. 1829. Observations on the fishes contained in the collection of the Zoological Society. On some fishes from the Sandwich Islands. The Zoological Journal, London, vol. 4, art. 3, pp. 31-42. BLEEKER, PIETER 1856. Bijdrage tot de kennis der ichthyologische fauna van het eiland Boeroe. Natuur- kundig Tijdschrift voor Nederlandsch-Indié, vol. 11, pp. 383-414. 1857. Bijdrage tot de kennis der ichthyologische fauna van de Sangi-eilanden. Natu- urkundig Tijdschrift voor Nederlandsch-Indié, vol. 13, pp. 369-380. 1876a. Genera familiae scorpaenoideorum conspectus analyticus. Verslagen en Medede- elingen der Koninklijke Akademie van Wetenschappen, Amsterdam, Afdeeling Natuurkunde, series 2, vol. 9, part 3, pp. 294-300. 1876b. Mémoire sur les espéces insulindiennes de la famille des scorpénoides. Ver- handelingen der Koninklijke van Wetenschappen. Afdeeling Natuurkunde, vol. XVI, pp. 1-100, col. pls. I-V. BouHLKE, JAMES E. 1953. A catalogue of the type specimens of recent fishes in the Natural History Museum of Stanford University. Stanford Ichthyological Bulletin, vol. 5, pp. 1-168. Boropin, N. A. 1930. Scientific results of the yacht “Ara” Expedition during the years 1926 to 1930, while in command of William K. Vanderbilt. Fishes (collected in 1929). Bulletin of the Vanderbilt Marine Museum, vol. 1, art. 2, pp. 39-64, pls. 1-2. CLARKE, THoMAs A. 1972. Collections and submarine observations of deep benthic fishes and decapod crustacea in Hawaii. Pacific Science, vol. 26, no. 3, pp. 310-317. CoLLETTE, Bruce B., and Teruya Uyeno 1972. Pontinus niger, a synonym of the scorpionfish Ectreposebastes imus, with exten- sion of its range to Japan. Japanese Journal of Ichthyology, vol. 19, no. 1, pp. 26-28, 1 fig. Cuvier, GrorGes, and ACHILLE VALENCIENNES 1829. Histoire naturelle des poissons. F. G. Levrault, Paris, vol. 4, xxvi + 518 pp. EDMONDSON, CHARLES HowArD 1946. Reef and shore fauna of Hawaii. Bernice P. Bishop Museum, Special Publication 22, pp. 232-381, 223 figs. EsCHMEYER, WILLIAM N. 1965a. Western Atlantic scorpionfishes of the genus Scorpaena, including four new species. Bulletin of Marine Science, vol. 15, no. 1, pp. 84-164, 12 figs. 1965b. Three new scorpionfishes of the genera Pontinus, Phenacoscorpius and Idiastion from the western Atlantic Ocean. Bulletin of Marine Science, vol. 15, no. 3, pp. 521-534.
328 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
1969a. A new scorpionfish of the genus Scorpaenodes and S. muciparus (Alcock) from the Indian Ocean, with comments on the limits of the genus. Occasional Papers of the California Academy of Sciences, no. 76, 11 pp., 1 fig.
1969b. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scorpaenidae). Occasional Papers of the California Academy of Sciences, no. 79, 130 pp., 13 figs.
EscHMEYER, WILLIAM N., and Greratp R. ALLEN
1971. Three new species of scorpionfishes (family Scorpaenidae) from Easter Island. Proceedings of the California Academy of Sciences, vol. 27, no. 19, pp. 515— SYA, Ab sale
EscCHMEYER, WILLIAM N., and Bruce B. CoLLeTTE
1966. The scorpionfish subfamily Setarchinae, including the genus Ectreposebastes.
Bulletin of Marine Science, vol. 16, no. 2, pp. 349-375. EscHMEYER, WILLIAM N., YosuHitsucu Hirosaki, and TokrHAru ABE
1973. Two new species of the scorpionfish genus Rhinopias, with comments on related genera and species. Proceedings of the California Academy of Sciences, vol. 27, no. 16, pp. 285-310, 10 figs.
Fow Ler, Henry W.
1900. Contributions to the ichthyology of the tropical Pacific. Proceedings of the Academy of Natural Sciences of Philadelphia, for year 1900, pp. 493-528, pls. 18-20.
1923. New or little-known Hawaiian fishes. Occasional Papers of the Bernice Pauahi Bishop Museum of Polynesian Ethnology and Natural History, vol. 8, no. 7, pp. 375-392.
1925. Fishes of Guam, Hawaii, Samoa, and Tahiti. Bernice P. Bishop Museum, Bulletin 22, 38 pp.
1928. The fishes of Oceania. Memoirs of the Bernice P. Bishop Museum, vol. 10, iii + 540 pp., 82 figs., pls. 1-49.
1931. The fishes of Oceania—Supplement 1. Memoirs of the Bernice P. Bishop Mu- seum, vol. 11, no. 5, pp. 313-381, 7 figs.
1934. The fishes of Oceania—Supplement 2. Memoirs of the Bernice P. Bishop Mu- seum, vol. 11, no. 6, pp. 385-466, 4 figs.
1935. Description of a new scorpaenid fish (Neomerinthe hemingwayi) from off New Jersey. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 87, pp. 41-43.
1938a. Descriptions of new fishes obtained by the United States Bureau of Fisheries Steamer “Albatross,” chiefly in Philippine Seas and adjacent waters. Proceed- ings of the United States National Museum, vol. 85, no. 3032, pp. 31-135, figs. 1-61.
1938b. The fishes of the George Vanderbilt South Pacific Expedition, 1937. The Academy of Natural Sciences of Philadelphia, Monographs, no. 2, 349 pp., 12 pls.
1941. The George Vanderbilt Oahu Survey—the fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 93, pp. 247-279, 29 figs.
1946. A collection of fishes obtained in the Riu Kiu Islands by Captain Ernest R. Tinkham A.U.S. Proceedings of the Academy of Natural Sciences of Philadel- phia, vol. 98, pp. 123-218, 76 figs.
1949. The fishes of Oceania—Supplement 3. Memoirs of the Bernice P. Bishop Mu- seum, vol. 12, no. 2, pp. 37-186.
VoL. XL] ESCHMEYVER AND RANDALL: HAWAIIAN SCORPAENIDAE 329
FowLer, Henry W., and STANLEY C. BAL 1925. Fishes of Hawaii, Johnston Island, and Wake Island. Bernice P. Bishop Museum, Bulletin 26, 31 pp. GARMAN, SAMUEL 1899. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission steamer “Albatross,” during 1891, Lieut. Com- mander Z. L. Tanner, U.S.N., commanding. XXVI. The fishes. Memoirs of the Museum of Comparative Zoology at Harvard College, vol. 24, text: 431 pp., pls. 1-98. GARRETT, ANDREW 1864. Descriptions of new species of fishes—no. II. Proceedings of the California Academy of Natural Sciences, vol. 3, pp. 103-107. GILBERT, CHARLES HENRY 1905. The deep-sea fishes of the Hawaiian Islands. Jn the aquatic resources of the Hawaiian Islands. Bulletin of the U. S. Fish Commission, vol. 23, part 2, pp. 575-713, figs. 230-276, pls. 66-101. GILBERT, CHARLES Henry, and FRANK CRAMER 1897. Report on the fishes dredged in deep water near the Hawaiian Islands, with descriptions and figures of twenty-three new species. Proceedings of the U. S. National Museum, vol. 19, no. 1114, pp. 403-435, pls. 36-48. GILL, THEODORE 1905. Note on the genera of synanceine and pelorine fishes. Proceedings of the United States National Museum, vol. 28, no. 1394, pp. 221-225, 1 fig. GosLINE, WILLIAM A. 1955. The inshore fish fauna of Johnston Island, a central Pacific atoll. Pacific Science, vol. 9, pp. 442-480, 4 figs. 1965. Vertical zonation of inshore fishes in the water layers of the Hawaiian Islands. Ecology, vol. 46, no. 6, pp. 823-831. GosLINE, WILLIAM A., and VERNON E. Brock 1960. Handbook of Hawaiian fishes. Hawaii Press, Honolulu, 372 pp. GREENFIELD, TERESA ARAMBULA MS. Systematics and zoogeography of eastern Pacific scorpionfishes of the genus Scorpaena (Pisces: Scorpaenidae). Thesis, California State University, San Francisco. GUNTHER, ALBERT C. L. G. 1873. Andrew Garrett’s Die Fische der Siidsee. Heft I. Journal des Museum Godeffroy, vol. 2, Heft III, pp. 1-128, pls. 41-60. HECKEL, JOHANN JAKOB 1837. Ichtyologische Beitrage zu den Familien der Cottoiden, Scorpaenoiden, Gobioiden, und Cyprinoiden. Annalen des Wiener Museums der Naturgeschichte, vol. 2, fasc. 1, pp. 143-164. HILDEBRAND, SAMUEL F. 1946. A descriptive catalog of the shore fishes of Peru. Bulletin of the United States National Museum, Bulletin 189, 530 pp., 95 figs. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 1922. Opinion 77. Thirty-five generic names in Protozoa, Coelenterata, Trematoda, Cestoda, Cirripedia, Tunicata, and Pisces placed in the official list of generic names. Jn Opinions rendered by the International Commission on Zoological Nomenclature. Smithsonian Miscellaneous Collections, vol. 73, no. 71-74.
330 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
1964. International Code of Zoological Nomenclature adopted by the XV International Congress of Zoology. International Trust for Zoological Nomenclature, Lon- don, 176 pp.
JENKINS, OLIVER P.
1901. Description of fifteen new species of fishes from the Hawaiian Islands. Bulletin of the U. S. Fish Commission for 1899, pp. 387-404, 16 figs.
1903. Report on collections of fishes made in the Hawaiian Islands, with descriptions of new species. Bulletin of the United States Fish Commission, vol. 22, pp. 417-511, 50 figs., 4 pls.
JoHNson, JAMES YATE
1862. Descriptions of some new genera and species of fishes obtained at Madeira. Proceedings of the Zoological Society of London for 1862, pp. 167-180, pls. 22-23.
Jorpan, Davin Starr, and BARTON WarRREN EVERMANN
1903a. Descriptions of new genera and species of fishes from the Hawaiian Islands. Bulletin of the United States Fish Commission, vol. 22 (for 1902), pp. 161-208.
1903b. Descriptions of a new genus and two new species of fishes from the Hawaiian Islands. Bulletin of the United States Fish Commission, vol. 22 (for 1902), pp. 209-210.
1905. The shore fishes of the Hawaiian Islands, with a general account of the fish fauna. In The Aquatic Resources of the Hawaiian Islands, Bulletin of the United States Fish Commission, vol. 23, part 1, xxvii + 574 pp., 229 figs., pls. 1-65 + col. pls. I-LXIII.
1926. A check-list of the fishes of Hawaii. Bulletin of the Pan-Pacific Research Insti- tution, vol. 1, no. 1, pp. 3-15.
Jorpan, Davip Starr, and CHARLES HENRY GILBERT
1883. Synopsis of the fishes of North America. Bulletin of the United States National
Museum, no. 16, lvi + 1018 pp. Jorpan, Davin Starr, and Eric KNIGHT JORDAN
1922. A list of the fishes of Hawaii, with notes and descriptions of new species.
Memoirs of the Carnegie Museum, vol. 10, no. 1, pp. 1-92, 7 figs., pls. 1-4. Jorpan, Davip Starr, and ALVIN SEALE
1906. The fishes of Samoa. Description of the species found in the archipelago, with a provisional check-list of the fishes of Oceania. Bulletin of the Bureau of Fisheries, vol. 25, pp. 173-455, 111 figs., pls. 38-53.
Jorpan, Davin Starr, and JOHN OTTERBEIN SNYDER
1904a. Notes on collections of fishes from Oahu Island and Laysan Island, Hawaii, with descriptions of four new species. Proceedings of the United States National Museum, vol. 27, no. 1377, pp. 939-948.
1904b. Description of a new species of fish (Apogon evermanni) from the Hawaiian Islands, with notes on other species. Proceedings of the United States Na- tional Museum, vol. 28, no. 1386, pp. 123-126.
1907. Notes on fishes of Hawaii, with descriptions of new species. Bulletin of the Bureau of Fisheries, vol. 26, pp. 205-218, pls. XII—-XIII.
Jorpan, Eric KNIGHT
1925. Notes on the fishes of Hawaii, with descriptions of six new species. Proceedings
of the United States National Museum, vol. 66, no. 2570, pp. 1-43, pls. 1-2. LacrepEDE, B. G. E. de 1802. Histoire naturelle des poissons. Chez Plasson, Paris, vol. 3, 558 pp., 34 pls.
Vor. XL] ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 331
LINNAEUS, CAROLUS
1758. Systema naturae per regna tria naturae, edition 10. Laurentii Salvii Holmiae,
824 pp. Mappen, WitiiAm D.
1973. The collection of live fishes from a salvaged vessel. Copeia, 1973, no. 1, pp.
144-145. Matsusara, KryoMaTsu
1943. Studies on the scorpaenoid fishes of Japan. The Transactions of the Sigenkagaku
Kenkyusyo, no. 1, 486 pp., 4 pls. OXKEN, LORENZ
1817. Cuviers und Okens Zoologien neben einander gestellt. Isis, oder Encyclopadische Zeitung von Oken, vol. 8, nos. 144-148 [28 pp. inconsistently numbered by columns 1145-1184, including unnumbered columns; 1780-1782 misprint for 1180-1182. ]
PIETSCHMANN, VICTOR
1930. Remarks on Pacific fishes. Bernice P. Bishop Museum, Bulletin 73, 24 pp., pls. 1-4.
1934. Drei neue Fische aus den Hawaiischen Kiistengewassern. Akademie der Wissen- schaften in Wien, Mathematisch-Naturwissenchaftliche Klass, Anzeiger, 71. Jahrgang, pp. 99-100.
1938. Hawaiian shore fishes. Bernice P. Bishop Museum Bulletin 156, 56 pp., 18 pls.
Pory, FELIPE
1860. Poissons de Cuba. Memorias sobre la historia natural de la isla de Cuba, vol. 2,
pp. 115-356. RANDALL, JOHN E.
1973. Tahitian fish names and a preliminary checklist of the fishes of the Society Islands. Occasional Papers of Bernice P. Bishop Museum, vol. 24, no. 11, pp. 167-214.
SauvAGE, Henri EMILE
1878. Description de poissons nouveaux ou imparfaitement connus de la collection du Muséum d’Histoire Naturelle. Famille des scorpénidées, des platycéphalidées et des triglidées. Nouvelles Archives du Muséum d’Histoire Naturelle, 2nd ser., vol. 1, pp. 109-159, pls. 1-2.
1882. Description de quelques poissons de la collection du Muséum d’Histoire Naturelle. Bulletin de la Société Philomathique de Paris, ser. 7, vol. 6, pp. 168-171.
ScHULTz, LEoNaARD P.
1938. Notes on the scorpaenid fish, Taenianotus triacanthus, from the Hawaiian Islands. Copeia, 1938, no. 4, p. 206.
1943. Fishes of the Phoenix and Samoan islands collected in 1939 during the expedition of the U.S.S. “Bushnell.” Smithsonian Institution, United States National Museum, Bulletin 180, 316 pp., 27 text figs., 9 pls.
1966. Fishes of the Marshall and Marianas Islands, vol. 3. United States National Museum, Bulletin 202, vii + 176 pp., 155 figs., pls. 124-148. [With collab- orators. |
SEALE, ALVIN
1901. New Hawaiian fishes. Occasional Papers of the Bernice Pauahi Bishop Museum of Polynesian Ethnology and Natural History, vol. 1, no. 4, pp. 1-15, 7 figs. (= 7 lel
1902. Ichthyology, In Director’s report for 1903. Occasional Papers of the Bernice
332 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Pauahi Bishop Museum of Polynesian Ethnology and Natural History, vol. 2, no. 2, pp. 18-21. (Authorship inferred.) SHALLENBERGER, Ropert J., and Witt1am D. MaApprEn 1973. Luring behavior in the scorpionfish, Jracundus signifer. Behavior, vol. 47, pp. 33-47, 2 pls. SmiTH, J. L. B. 1957a. The fishes of the family Scorpaenidae in the western Indian Ocean. Part I. The sub-family Scorpaeninae. Ichthyological Bulletin, Rhodes University, no. 4, pp. 49-72, pls. 1-4. 1957b. The fishes of the family Scorpaenidae in the western Indian Ocean. Part II. The subfamilies Pteroinae, Apistinae, Setarchinae and Sebastinae. Ichthyo- logical Bulletin, Rhodes University, no. 5, pp. 75-88, 9 figs., pls. 1-6. SNYDER, JOHN OTTERBEIN 1904. A catalogue of the shore fishes collected by the steamer ALBATROSS about the Hawaiian Islands in 1902. Bulletin of the United States Fish Commission, vol. 22 (for 1902), pp. 513-538. SPRINGER, VICTOR G. 1967. The Pacific South American blenniid fish, Hypsoblennius sordidus. Copeia, 1967, no. 2, pp. 461-465, 1 fig. STEINDACHNER, FRANZ 1900a. Fische aus dem Stillen Ocean. Ergebnisse einer Reise nach dem _ Pacific (Schauinsland, 1896-1897). Anzeiger der Kaiserlichen Akademie der Wissen- schaften in Wien. Mathematisch-Naturwissenschaftliche Klasse, Jahrgang 1900, no. 16, pp. 174-178 [Abstract of 1900b.] 1900b. Fische aus dem Stillen Ocean, Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896-97). Denkschriften der Kaiserlichen Akademie der Wis- senschaften. Mathematisch-Naturwissenschaftliche Classe, vol. 70, pp. 483- 521, pls. I-VI. [Separate published in 1900, volume in 1901.] STREETS, THOMAS HALE 1877. Contributions to the natural history of the Hawaiian and Fanning Islands and Lower California, made in connection with the United States North Pacific Surveying Expedition, 1873-75. Bulletin of the United States National Mu- seum, vol. 7, pp. 1-172. STRUHSAKER, PAuL J. 1973. Contribution to the systematics and ecology of Hawaiian bathyal fishes. Ph.D. Thesis. University of Hawaii, 482 pp. SWAIN, JOSEPH 1883. A review of Swainson’s genera of fishes. Proceedings of the Academy of Nat- tural Sciences of Philadelphia, for the year 1882, pp. 272-284. SWAINSON, WILLIAM 1839. The natural history of fishes, amphibians, and reptiles, or monocardian animals. London, vol. 2, vi + 448 pp., 135 text figs. TANAKA, S. 1917. “Eleven new species of fish from Japan.” Dobutsugaku Zasshi (Zoological Magazine), Tokyo, vol. 29, pp. 7-12. [In Japanese. ] 1928. Figures and descriptions of the fishes of Japan, including Riukiu Islands, Bonin Islands, Formosa, Kurile Islands, Korea, and Southern Sakhalin. vol. 42, pp. 809-830, pls. 172-174. TINKER, SPENCER WILKIE 1944. Hawaiian fishes, a handbook of the fishes found among the islands of the
VoL. XL]
VAILLANT, 1875.
WAHLERT,
1955.
WHITLEY, 1965.
WICKLER, 1969.
ESCHMEYER AND RANDALL: HAWAIIAN SCORPAENIDAE 333
Central Pacific Ocean. Tongg Publishing Co., Honolulu, 404 pp., numerous figs., 8 col. pls.
M. M., and H. E. Sauvacre
Note sur quelques espéces nouvelles de poissons des iles Sandwich. Revue et Magasin de Zoologie pure et appliquée . . . 3rd ser., vol. 3, pp. 278-287.
GERD VON
Die Typen und Typoide des Ubersee-Museums Bremen, 2: Pisces. Veréffentlich- ungen aus dem Uberseemuseum Bremen, ser. A, vol. 2, Heft 5, pp. 323-326, figs. 1-2.
GILBERT P.
Illustrations and records of fishes. The Australian Zoologist, vol. 13, no. 2, pp. 103-120, 13 figs.
Wo trcanc, and CuHristeL Nowak
Hautung und andere Verhaltensweisen von Taenianotus triacanthus, einem Verwandten der Skorpionfische. Natur und Museum, vol. 99, no. 10, pp. 441-456, 9 figs.
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 12, pp. 335-416; 59 figs. October 3, 1975
MUSCULAR ANATOMY OF THE HIND LIMB OF THE SEA OTTER (ENHYDRA LUTRIS)
By L. D. Howard
FOREWORD
In December 1973 the California Academy of Sciences published Dr. Lot D. Howard’s work on the muscular anatomy of the forelimb of the sea otter, Enhydra lutris (Howard, 1973). This paper represented the results of several years of detailed dissection and anatomical drawing of the front limb muscu- lature of a unique carnivore that has become specialized for life in a marine environment. As Dr. Howard noted in his introduction, one would expect a mammal that lives in the sea and uses its forelimbs for securing food from the bottom, for breaking shellfish on a rock placed on its abdomen, for elaborate grooming of fur, and for holding the young, to have a high degree of muscular specialization. This did not prove to be so. The anatomical features of the limb show a marked resemblance to those of related land mammals. Its functional specialization is basically a result of bimanual use.
Dr. Howard, prior to his retirement to Pebble Beach on the Monterey Peninsula of California, was a world-renowned hand surgeon who had spent years on the staff of Stanford University Medical School. Much of his time in his later years was devoted to watching sea otters in the ocean in front of his home as well as to the anatomical dissection of these animals. Following the completion of his manuscript on the front limb of the sea otter, Dr. How- ard began work on the muscular anatomy of the hind limb. Knowing of his own fatal illness, he worked against time to try to complete the dissections and drawings which are presented in this paper and at the time of his death the work was nearly finished. Mr. Judson E. Vandevere of Monterey, Cali-
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fornia, an authority on sea otters, and Dr. and Mrs. Donald T. Abbott of Stanford University’s Hopkins Marine Station at Pacific Grove contributed to the completion of the manuscript.
Robert T. Orr Associate Director, California Academy of Sciences.
INTRODUCTION
The following study is a presentation of the musculature of the hind limb of the sea otter. Having published the muscular anatomy of the forelimb of the sea otter (Howard, 1973) it seems only logical that a similar study of the hind limb be undertaken.
The hind limb is grossly different from the forelimb both anatomically and functionally, being used almost exclusively for swimming and showing a high degree of specialization for this purpose. The limb is relatively short and heavily muscled. The hip and knee are semi-flexed and for the most part the foot is in the plantar flexed position. Detailed studies of the use of the hind limbs in swimming have been made by Tarasoff et al. (1972).
The hind foot is large compared to the front foot, its length equaling that of the lower leg. Both metacarpals and phalanges are elongated and a gen- erous skin web exists between the digits. The 5th digit is the largest and the longest, and the Ist digit the smallest. The foot becomes twice as wide when the toes are spread. In the hind foot the elongated digits and generous webbing make for individual digit mobility far in excess of that possible in the front foot.
DESCRIPTION OF THE LOWER EXTREMITY OF THE SEA OTTER
Like the upper extremity, the lower extremity is relatively short in com- parison with body length and is heavily muscled in the upper and lower leg.
The foot is highly developed for aquatic propulsion and is completely covered with fur. The skin of the foot is soft and mobile with the exception of the terminal digital pads. Broad, soft webbing of skin is present between the toes extending in slightly scalloped fashion to the very tips of the digits. Unlike the forefoot or hand and in spite of the heavy fur cover, the individual digits are grossly identifiable on both volar and dorsal surfaces.
Even in the preserved specimen, the individual digital joints are highly mobile. Three phalanges are present in digits five, four, three, and two, with but two phalanges in the ‘great toe’ which in this case is the smallest digit. The proximal or metacarpal phalangeal joints hyperextend approxi- mately 40° beyond the straight line. Flexion is to 110°. The middle toe joints or proximal interphalangeals extend to 55° beyond the straight line
Vor. XL] HOWARD: SEA OTTER HIND LIMB 337
and flex to 115°. The distal joints or distal interphalangeal joints extend to 45° above the straight line and flex to 65°. For digit one, only a single interphalangeal joint is present. This single interphalangeal joint extends to approximately 45° beyond the straight line and flexes to approximately 75°
When spread, the distance between the tips of toes one and five is twice, or a little more than twice, the distance when the toes are together.
An arched and curved claw is present for each digit. Each is about equal in size, the one on digit four being perhaps slightly heavier. The distal pads cover the plantar aspect of the terminal phalanges covering approximately the distal one-third or one-fourth of the middle phalanx. A small pad also occurs at the base the ‘great toe.’
Due to the extreme mobility of the small joints of the foot, when the metacarpal, phalangeal, and interphalangeal joints are flexed to their passive limits the tips of the toes actually touch the sole of the foot, curving around an ample area to permit actual grasp of objects (two or three fingers in a grip-like manner). Vandevere (pers. commun.) has seen on numerous occasions food, a tool, and a pup held against the body but never grasped by the toes.
Also because of the interdigital webs, the individual digits themselves are more highly mobile and thus the pad of the fifth can touch the pads of the fourth and the third in the manner of apposition, and can touch the pads of the second and the first in the configuration of opposition.
Thus it would appear that the foot has the potential of being rather highly prehensile in contrast to the hand, yet in spite of this the foot serves mainly for locomotion in water and to a degree for grooming. The foot appears to be more suitable than the hand as an organ for grasping food and the hand is superior to the foot as an organ for locomotion on land.
Thus the extremities present a kind of a paradox for a mammal returning to the sea. The hind legs and feet have made ready adaption for aquatic locomotion, the forefoot or hand seeming to persist in a form for ambulation on land. An amusing conclusion would be that the animal entered the sea backwards! Perhaps the forelimbs are becoming functionally obsolete as organs of locomotion in swimming.
An outstanding feature is the development of the outer side of the foot, with the fifth digit, the shortest in land mammals, becoming more massive and longer than all of the remaining digits. In seals the fifth digit has also become massive and long, but not noticeably more massive or longer than the great toe. When sea otters are swimming on their backs, this development of their fifth digit gives maximum skulling surface and the largest excursion of motion. In underwater swimming, accomplished by undulations of the body in the AP diameter, the trailing feet can take on the general configuration of the flukes of a whale.
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Vor. XL] HOWARD: SEA OTTER HIND LIMB 339
MuSscLES OF THE HINDLIMB OF THE SEA OTTER ALPHABETICAL LISTING
Muscle No. Muscle No. Abductor digiti quinti 40 Interossei (interosseus) 46 Abductor hallucis 43 Lumbricales 48 Adductor digiti quinti 41 Obturator externus 21 Adductor femoris 5s = superficial Obturator internus 23
5d = deep Opponens digiti quinti 42 Adductor hallucis 44 Panniculus carnosis A Adductor longus 5-A Pectineus 6 Biceps femoris 2 Peroneus brevis 30 Caudofemoralis ia Peroneus digiti quinti 29 Calcaneometatarsalis 39 Peroneus longus 28 Extensor digitorum brevis 47 Plantaris 31 Extensor digitorum longus 27 Popliteus 32 Extensor hallucis proprius (longus) 26 Presemimembranosus 17 Flexor digitorum brevis 37 Piriformis 19 Flexor digitorum longus 36 Quadratus plantae 38 Flexor hallucis longus 35 Rectus femoris 7 Gastrocnemius 24 Sartorius 1 Gemelli (gemellus) 20 Semimembranosus 4 Gluteus maximus 9 Semitendinosus 3 Gluteus medius 16 Tenuissimus 13 Gluteus minimus 18 Tibialis anterior 25 Gracilis 2 Tibialis posterior 34 Ihocapsularis (quadratus femoris) 22 Vastus medialis 14 Iliopsoas 8 Vastus lateralis 10
DESCRIPTIONS OF MUSCLES A. Panniculus carnosis.
This voluntary muscle for moving the skin is well developed in the area of the hind limb of the sea otter. The skin and subcutaneous tissues are very mobile over the underlying deep fascia. The panniculus carnosis muscle is a broad thin layer of rather coarse longitudinal muscle fibers closely attached to the undersurface of the skin, so closely that rather meticulous sharp dis- section is required to separate them.
OricGIN. For the hind limb, this muscle layer is a continuation of that present in the trunk area. The fibers paralleling the middorsal line cover most of the lateral aspect of the hind limb and wrap about the knee to reach the medial surface of the limb to some degree in this area.
INSERTION. Insertion of this muscle is in the deep layers of the skin as far posteriorly as the skin web between the tail and the ankle.
Action. This muscle serves to voluntarily mobilize the skin over the hind limb from the trunk to the heel area.
340 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 2. The plantar view of the right hind foot. Note the relatively small pads over- lying the terminal phalanges of each digit. A smaller pad at the metatarsophalangeal joint of digit 1 is covered with hair and therefore is out of view. The pads have a rough texture similar to the pads of the forefoot. Hair coverage on the plantar surface of the foot is extensive.
1. Sartorius.
This muscle is fleshy, somewhat flat, and superficial in position, overlying the anterior aspect of the hind limb connecting the pelvis to the knee area.
Oricin. The muscle arises by fleshy and short tendinous fibers from the iliac crest. It is the most superficial muscle overlying in part the origins of
Vor. XL] HOWARD: SEA OTTER HIND LIMB 341
the glutei musculature. The muscle passes distally from its origin over the anterior thigh, narrowing somewhat and shifting slightly medial. INSERTION. The insertion is along the full length of the patella tendon with some short tendinous fibers entering the medial side of the tibial tubercle. Action. This muscle assists in extension of the knee through its patellar tendon attachment and also assists in flexion of the hip.
2. Gracilis.
This broad, flat, superficial muscle is on the medial aspect of the lower limb and connects the pelvis with the tibia.
OriciIn. The muscular fibers of origin come from the full width of the pubis in the area of the symphysis. From its origin, the muscle maintains a constant width and courses like a muscular strap toward the upper tibia.
INSERTION. The insertion is by fleshy and short tendinous fibers into the tibial crest beginning at and extending distal to the tibial tubercle. The in- sertion overlies the insertion of the semitendinosus (3).
Action. This muscle serves to adduct the lower limb and assists in flexion of the knee and internal rotation of the extremity.
3. Semitendinosus.
This muscle is a flat, somewhat triangular superficial muscle joining the caudal area of the axial skeleton to the tibia. This muscle participates with the biceps femoris (12) to form a sort of muscular web between the caudal area and the knee.
OricIN. This muscle arises as fleshy and short tendinous fibers from the spinous processes and associated fascia of caudal vertebrae 1-7. The more superior area of origin overlies the origin of the caudofemoralis (11) at the caudal 1—2 level. The muscle now crosses toward the medial side of the tibia and while so doing underlies and gives off a group of muscle fibers to the biceps femoris (12).
INSERTION. The muscle narrows as it approaches the tibia and terminates as a short flat tendon which inserts on the crest of the tibia partly under the insertion of the gracilis (2).
Action. This muscle assists in flexion of the knee and internal rotation of the lower limb.
4. Semimembranosus.
This is a somewhat flat muscle connecting the pelvis with the upper tibia.
Oricin. Muscular fibers arise superficially along the ischial tuberosity overlying and almost in common with a portion of the superficial division of the adductor femoris (5). The muscle then extends obliquely toward the knee underlying the gracilis (2) and paralleling the adductor femoris (5).
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Ficure 3. Tracing from an x-ray of the fore limb and hind limb of the same animal to show relationship of length of foot to length of limb and also relative length of digits compared one to the other. Key: F, fibula; G, digit 5; R, radius; T, tibia; U, ulna.
VoL. XL] HOWARD: SEA OTTER HIND LIMB 343
Ficure 4. Lateral view of right lower extremity and caudal area. The skin has been carefully dissected from the panniculus carnosis muscle and reflected to show this most superficial muscle (A). The rather coarse parallel muscle fibers terminate in the deep layer of the skin in an irregular manner along the web-like fold of skin between the tail and the heel. Key: C, spinous process of the first caudal vertebra; D, reflected skin and sub- cutaneous tissue; F, foot; S, spinous process of the first sacral vertebra; A, panniculus carnosis; 3, semitendinosis; 12, biceps femoris; 13, tenuissimus.
INSERTION. Nearing its insertion, the muscle narrows to form a_ short flat tendon which passes under the medial collateral ligament of the knee and inserts on the tibia just medial to the tuberosity at the level of the junction of the insertion of the gracilis (2) and sartorius (1).
AcTIon. This muscle serves as a flexor of the knee and an internal rotator of the lower limb.
5. Adductor femoris.
The adductor femoris is a large muscle joining the pelvis with the femur. It is divided into two: the superficial and deep portions.
344 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
fapon yy al 1X
Ficure 5. Lateral view of right lower extremity and hip area. The panniculus carnosis muscle (A) has been removed to show the disposition of the superficial musculature beneath. Key: C, spinous process of first caudal vertebra; F, foot; I, iliac crest; S, spinous process
of first sacral vertebra; 1, sartorius; 3, semitendinosus; 9, gluteus maximus; 10, vastus lateralis; 11, caudofemoralis; -12, biceps femoris; 13, tenuissimus.
SUPERFICIAL PORTION.
OrIcIN. The superficial portion arises from the most distal area of the external surface of the pelvis adjacent to the origin of the semimembranosus and in close association with the deep portion of the adductor femoris. From the semitriangular area of origin the muscle broadens and flattens as it ap- proaches the knee area where it underlies the broad sartorius muscle (1).
INSERTION. The insertion by muscular and short tendinous fibers is over a fairly broad area about the knee. The more distal fibers insert into the medial epicondyle area of the femur and the remainder insert into the deep fascia on the medial side of the knee joint.
Action. Adduction of the femur.
Vou. XL] HOWARD: SEA OTTER HIND LIMB 345
Ficure 6. Lateral view of right lower extremity and hip area. The sartorius (1) and the gluteus maximus (9) have been reflected superiorly from their origins, and the semi- tendinosus (3) has been reflected dorsally. Origins of the biceps femoris (12) and the caudofemoralis (11) are more clearly visualized. Seen in the deeper layer of muscle are the gluteus medius (16) and the rectus femoris (7). Key: C, spinous process of the first caudal vertebra; D, ischial tuberosity; E, knee joint level; F, foot; G, caudal musculature; I, iliac crest; W, femur (greater trochanter); 1, sartorius (reflected); 3, semitendinosus (reflected) ; 7, rectus femoris; 9, gluteus maximus (reflected); 10, vastus lateralis; 11, caudofemoralis; 12, biceps femoris; 16, gluteus medius.
DEEP PORTION.
OricIn. The deep portion of this muscle arises from the external surface of the ischium and pubis immediately proximal to and almost in common with the superficial portion. The greater area of the crescent shaped origin is from the pubic bone, where it borders the gracilis muscle (2) origin medially and the origin of the adductor longus (5A) superiorly. From its origin, the muscle parallels the superficial portion as it passes toward the femur.
INSERTION. Distally, the muscle spreads fan-like to insert by tendinous fibers into the posteromedial aspect of the femur. The more distal portion underlies the insertion of the superficial portion of the adductor femoris in
346 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 7. Lateral view of the right lower extremity and hip area, similar to figure 6 but with the caudofemoralis (11) detached at its origin and reflected anteriorly. The deeper muscles are thus exposed, and the area of origin of the biceps femoris (12) and gluteus medius (16) are clearly seen. Key: C, spinous process of first caudal vertebra; D, ischial tuberosity; F, foot; G, caudal musculature; I, iliac crest; W, femur (greater trochanter); 1, sartorius (reflected); 3, semitendinosus (reflected); 9, gluteus maximus (reflected) ; 10, vastus lateralis; 11, caudofemoralis (reflected); 12, biceps femoris; 13, tenuissimus; 16, gluteus medius; 17, presemimembranosus; 19, piriformis.
the area of the medial epicondyle of the femur. Insertion extends proximally along the medial ridge to the level of the lesser trochanter. Action. The muscle is an adductor of the femur.
5A. Adductor longus.'
This muscle is the most superior of the adductor group and is smaller than the others.
OricIN. The origin of this muscle is by short tendinous and muscular fibers from the external surface of the superior ramus of the pubis. At the symphysis the origin is adjacent to the gracilis muscle (2) and superiorly
1 There is some indication in Dr. Howard’s notes that he intended to renumber this muscle as ‘15.’ Editor.
Vor. XL] HOWARD: SEA OTTER HIND LIMB 347
Ficure 8. Lateral view of right lower extremity and hip area. The large biceps femoris (12) has been detached from its origin leaving a short stump and reflected ante- riorly at its insertion. As a result, the musculature of the lower leg is exposed and the tenuissimus (13) and presemimembranosus (17) are seen from their origin to their insertion. Key: F, foot; G, caudal musculature; J, iliac crest; W, femur (greater trochanter); 3, semitendinosus; 10, vastus lateralis; 11, caudofemoralis (reflected); 12, biceps femoris (reflected from the tibia and also the stub from the pelvis); 13, tenuissimus; 16, gluteus medius; 17, presemimembranosus; 19, piriformis; 24, gastrocnemius; 25, tibialis anterior; 28, peroneus longus; 30, peroneus brevis.
the origin extends to that of the pectineus (6). On the pubis, the origin of the deep portion of the adductor femoris (5) is immediately adjacent distally.
INSERTION. This muscle parallels the other adductors to a long insertion the full length of the medial ridge of the femur adjacent and anterior to the deep portion of the adductor femoris (5) and just posterior to the pectineus (6), thus inserting between these two muscles.
348 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr.
Action. Adduction and internal rotation of the femur would result from contraction of this muscle.
6. Pectineus.
This is a smaller muscle, as far as total mass ‘s concerned, but is clearly associated with the adductor group connecting the pelvis to the femur.
Oricin. The muscle arises in an elliptical configuration from the full width of the anterior surface of the pelvic bone directly opposite the acetabulum and just proximal to the adductor longus (5A).
INSERTION. From its origin, the muscle fans out paralleling the adductors
VoL. XL] HOWARD: SEA OTTER HIND LIMB 349
Ficure 10. An enlarged and localized lateral view of the right hip area. The sartorius (1) and gluteus maximus (9) have been reflected from the iliac crest. The gluteus medius (16) has been peeled off the wing of the ilium and, with the piriformis (19), rolled upward to disclose the gluteus minimus (18). Also visualized are the origins of the rectus femoris (7) and vastus lateralis (10). Key: I, iliac crest and wing of ilium; W, femur (greater trochanter) ; 1, sartorius (reflected); 7, rectus femoris; 9, gluteus maximus (reflected) ; 10, vastus lateralis; 16, gluteus medius (reflected); 18, gluteus minimus; 19, piriformis (re- flected).
to insert along the full length of the anterior ridge of the femur just anterior
to the adductor longus (5A). Action. This muscle would give adduction and internal rotation to the
femur.
7. Rectus femoris. This muscle is round in cross-section proximally and laterally compressed distally. It is a large fusiform muscle overlying the anterior surface of the
femur joining the pelvis to the tibia. Oricin. A short strong tendon of origin arises from the pelvic bone
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Ficure 11. Medial view of the right lower extremity. The limb has been abducted approximately 90° so that the pelvic area presents an anterior view. With the skin and subcutaneous tissues removed, the superficial musculature is shown. Key: F, foot; I, iliac crest; L, patellar ligament; P, symphysis pubis; R, pelvic rim; T, tibial tuberosity; U, perineal muscles; V, penis; 1, sartorius; 2, gracilis; 3, semitendinosus; 4, semimembranosus; 5, adductor femoris; 5A, adductor longus; 6, pectineus; 7, rectus femoris; 8, iliopsoas.
(ilium) just above the acetabulum in the anterolateral area. At the site of origin, the bone has two shallow crater-like depressions joined by a slight ridge of bone. From its tendinous origin the muscle enlarges in a spindle-like form passing distally along the anterior femur overlying and being bordered by the vastus medialis (14) and vastus lateralis (10).
INSERTION. Tendinous fibers of insertion begin developing in the distal one-third of the muscle, particularly on the medial side. A strong tendon is present at the insertion into the proximal end of the patella. Laterally, the tendon of insertion of the vastus lateralis (10) joins in to make a common insertion. Medially, the vastus medialis (14) with fleshy fibers underlies the rectus femoris tendon but inserts into the patella to the medial side of the rectus femoris tendon. The strong patellar tendon then continues the insertion of all three muscles to the tibial tubercle.
AcTIon. Strong extension of the knee.
Vor. XL] HOWARD: SEA OTTER HIND LIMB 351
Ficure 12. Medial view of abducted right lower leg. The sartorius (1) and the gracilis (2) have been reflected at their insertions to show relationship of the underlying deeper musculature. The medial collateral ligament of the knee takes its origin from the femur in the area of insertion of the superficial portion of the adductor femoris (5). Key: F, foot; J, tibial plateau; K, medial collateral ligament of the knee; O, patella; P, symphysis pubis; R, pelvic rim; U, perineal muscle; V, penis; 1, sartorius (reflected) ; 2, gracilis (reflected) ; 3, semitendinosus; 4, semimembranosus; 5, adductor femoris; 5A, adductor longus; 6, pec- tineus; 7, rectus femoris; 8, iliopsoas; 14, vastus medialis; 16, gluteus medius; 24, gastroc- nemius; 34, tibialis posterior.
8. Iliopsoas. This is a large heavy muscle connecting the lumbar spine and ilium with the femur. OricIN. Superior portion of this large muscle arises from the volar surfaces of the transverse processes of the last three lumbar vertebrae and adjacent fascia. As the muscle forms and passes caudally into the pelvis, an additional
352 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 13. Medial view of abducted right lower extremity, similar to figure 12. The semimembranosus (4) and the superficial portion of the adductor femoris (5) have now been removed at their origin from the pubis and reflected at their insertions. Further detail of the adductor muscles is made visible, and other deep muscles shown in relationship to one another. Note the contribution of muscle fibers from the semitendinosus (3) to the biceps femoris (12). That portion of the sartorius (1) which arises from the iliac crest is shown reflected at that site. Key: F, foot; H, femur (medial epicondyle); P, symphysis pubis; R, pelvic rim; T, tibia (medial malleolus) ; U, perineal muscles; V, penis; 1, sartorius (reflected) ; 2, gracilis (reflected) ; 3, semitendinosus, origin and reflected muscle; 4, semi- membranosus, origin and reflected muscle; 5, adductor femoris (origin of superficial por- tion); 5S, adductor femoris (superficial portion reflected); 5D, adductor femoris (deep portion); 5A, adductor longus; 6, pectineus; 7, rectus femoris; 8, iliopsoas; 11, caudo- femoralis; 12, biceps femoris; 13, tenuissimus; 14, vastus medialis; 16, gluteus medius; 17, presemimembranosus; 24, gastrocnemius; 34, tibialis posterior.
area of origin develops from the anterior surface of the ilium immediately adjacent to the sacroiliac joint.
INSERTION. The muscle becomes elliptical in transverse section and tapers into a heavy tendon which dips deeply along the side of the vastus medialis (14) to insert on the medial side of the lesser trochanter of the femur.
Action. Flexion and external rotation of the femur.
Vou. XL] HOWARD: SEA OTTER HIND LIMB 353
9. Gluteus maximus.
This is a flat triangular muscle lying over the extensor surface of the hip area connecting the pelvis with the femur and lateral fascia of the tibia.
OricIn. This muscle arises as a broad thin tendon from the outer side of the iliac crest. The origin is more or less fused with the tendon of origin of the gluteus medius (16). The anterior one-half of the origin underlies the origin of the muscle sartorius (1). The posterior one-half arises from the lumbar fascia overlying the spinal musculature in the area of the last lumbar vertebra. The muscle is superficial in position and from its origin triangulates toward the greater trochanter of the femur.
INSERTION. Near the upper femur, a short flat tendon develops from the greater portion of the muscle and inserts into the distal end of the greater trochanter, more or less in common with the proximal end of the caudofemoralis (11) insertion. Some of the more lateral fibers remain superficial and appear to insert into the deep fascia on the lateral aspect of the hip with continuation toward the knee. This portion may represent a tensor fascia lata muscle, although such a muscle is not specifically present in the sea otter.
10. Vastus lateralis.
This is the larger of the two vasti muscles which join the femur with the tibia through the patella and patella ligament.
OricIn. Fleshy fibers take origin from the proximal and lateral one-half of the femur. The origin begins on the anterior surface of the greater trochanter adjacent to the insertion of the gluteus minimus (18) and medius (16). The muscle then passes distally toward the knee joint where it joins with the rectus femoris (7).
INSERTION. The more lateral muscle fibers continue along the patella laterally and insert distal to but in the same plane as the tendon of the rectus femoris (7). The more medial fibers, which join in part with the rectus femoris (7), also form a strong short flat tendon which inserts into the lateral one-half of the proximal pole of the patella just under the rectus femoris (7) tendon.
AcTIoN. Extension of the knee through the patella and patella tendon.
11. Caudofemoralis.
This heavy superficial triangular muscle on the lateral aspect of the hip area connects the spine to the femur.
OricIn. The muscle arises from the spinous processes and deep fascia over the spinal muscles from the level of the last lumbar vertebra, the sacrum, and the first two caudal vertebrae. Inferiorly, the origin overlies two-thirds of the origin of the biceps femoris (12) and also the origin of the tenuissimus (13). Superiorly, the origin borders out of the gluteus maximus (9).
354 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
INSERTION. From its origin, the muscle triangulates somewhat and, passing over the greater trochanter, spreads to its insertion along the lateral femoral ridge of the inferior border of the greater trochanter to the upper border of the lateral femoral condyle.
Action. Abduction and external rotation of the femur.
12. Biceps femoris.
This is a very heavy and strong triangular muscle appearing superficially as the most prominent muscle on the lateral aspect of the lower limb. The muscle connects the pelvis to the tibia.
OrIGIN. The origin is by muscular tendinous fibers arising from a double bony exostosis-like protuberance along the posterior margin of the ischium just above the ischial tuberosity.
INSERTION. From its origin, the muscle spreads out fan-like to encompass the full length of the lower leg. Terminally, a thin flat tendon forms which inserts along the full length of the tibial crest laterally. Proximally, the insertion begins on the midlateral area of the tibia just below the joint surface. Distally, the tendinous fibers spread out into the deep fascia covering the entire lateral aspect of the ankle.
Action. This muscle acts as a flexor of the knee and external rotator of the lower limb.
13. Tenuissimus.
This is a long, uniformly thin and flat strap-type muscle bridging the sacrum to the ankle.
OricIN. This muscle arises as mostly muscular fibers from the transverse process of the third sacral vertebra and adjacent spinal muscle fascia. From here, the muscle courses diagonally toward the ankle, overlying the presemi- membranosus (17) and underlying the caudofemoralis (11) and the biceps femoris (12).
INSERTION. At the ankle, the muscle inserts into a fascia-like structure on the lateral aspect which has attachments to the lateral malleolus and to the calcaneus.
Action. Action of this muscle would be to flex the knee and to plantar flex the ankle.
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Ficure 14. Medial view of abducted right lower extremity, similar to figure 13, but in addition to the other reflected muscles, the deep portion of the adductor femoris (5D) has been removed from its origin and reflected distally. The relationship between the adductor longus (5A) and the pectineus (6) is now seen, and also the origin of the obturator
Vou. XL] HOWARD: SEA OTTER HIND LIMB 355
356 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Ficure 15. Medial view of abducted right lower extremity, similar to figures 13 and 14, but in addition the adductor longus (5A) and the pectineus (6) have been detached from their origins on the pelvis and reflected distally. By so doing, the head of the femur and the hip joint are exposed. Also, the insertion of the iliopsoas (8) can be seen as well as the relationship of the adjacent muscle. Key: D, ischial tuberosity; M, femur (lesser tuberosity) ; N, head of femur; Q, pelvis; R, pelvic rim; 4, semimembranosus; 5A, adductor
VoL. XL] HOWARD: SEA OTTER HIND LIMB 357
14
Ficure 16. Medial view of abducted right lower extremity and ventral view of pelvis. The origin and insertion of the iliopsoas (8) muscle is shown. All of the deep muscles can be identified. Key: B, transverse process, last lumbar vertebra; I, iliac crest; M, femur (lesser trochanter) ; N, head of femur; P, symphysis pubis; Q, pelvis; 4, semimembranosus (reflected) ; 5A, adductor longus (reflected) ; 5S, adductor femoris (superficial portion, re- flected) ; 5D, adductor femoris (deep portion, reflected) ; 6, pectineus (reflected); 7, rectus femoris; 8, iliopsoas; 14, vastus medialis; 16, gluteus medius; 21, obturator externus; 24, gastrocnemius; 34, tibialis posterior.
14. Vastus medialis.
This smaller of the two vasti muscles joins the femur to the tibia as a component of the extensor mechanism of the knee.
OricIn. Fleshy fibers arise from the medial one-half of the anterior surface of the femur. Proximally, the origin tapers to a point on the base of
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longus (reflected); 5S, adductor femoris (superficial portion, reflected); 5D, adductor femoris (deep portion, reflected); 6, pectineus (reflected); 7, rectus femoris; 8, iliopsoas; 14, vastus medialis; 24, gastrocnemius; 34, tibialis posterior.
358 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
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Ficure 17. Medial view of abducted right lower extremity, similar to figure 16. The iliopsoas (8) has been detached from its origin on the lumbar vertebra and ilium and reflected posteriorly. Doing so uncovers the origin of the rectus femoris (7) and, with the rectus femoris displaced anteriorly a bit, the proximal origin of the vastus medialis (14) and vastus lateralis (10) can be seen. Also, the gluteus medius (16) can be seen going toward its insertion on the greater trochanter. Key: I, iliac crest; M, femur (lesser trochanter); N, head of femur; O, patella; Q, pelvis; W, femur (greater trochanter) ; 1, sartorius (reflected); 5A, adductor longus (reflected); 5S, adductor femoris (superficial portion, reflected) ; 5D, adductor femoris (deep portion, reflected) ; 6, pectineus (reflected) ; 7, rectus femoris (displaced) ; 8, iliopsoas (reflected) ; 10, vastus lateralis; 12, biceps femoris; 14, vastus medialis; 16, gluteus medius; 21, obturator externus; 24, gastrocnemius; 34, tibialis posterior.
VoL. XL] HOWARD: SEA OTTER HIND LIMB 359
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Ficure 18. Medial view of right pelvis after disarticulation through the sacroiliac joint. The femur is viewed end-on from its proximal end, and the hip joint is visible. The gluteus medius (16) and piriformis (19) are reflected outward, and the origins and insertions of the short hip muscles can be seen. Key: I, iliac crest; N, head of femur; O, obturator foramen; P, symphysis pubis; Q, pelvis; S, sacroiliac joint; W, femur (greater trochanter) ; X, ischial spine; 16, gluteus medius; 19, piriformis; 20, gemelli; 22, iliocapsularis; 23, obturator internus.
the neck of the femur. Distally, the origin extends to just proximal to the articular surface, at which point it underlies the rectus femoris (7).
INSERTION. A short tendon forms distally and inserts into the medial one-half of the proximal end of the patella. This tendon partially underlies that of the vastus lateralis (10), which in turn underlies in part the rectus femoris (7) tendon.
AcTIon. The action of this muscle is to extend the knee through the patella and its patellar tendon.
15. There is some indication in Dr. Howard’s notes that muscle 5A—the ad- ductor longus—should be numbered ‘15.’ Editor.
16. Gluteus medius.
This large pyramidal-shaped muscle of coarse fibers connects the wing of the ilium to the greater trochanter of the femur. Orictn. The muscle arises as fleshy fibers from most of the lateral surface
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Ficure 19. Lateral view of right pelvis with hip flexed and internally rotated so that the femur presents a posterior view showing both greater and lesser trochanters. Visible also are the head and neck of the femur and the hip joint. The obturator externus (21) is seen throughout its course. The other muscles have been reflected to disclose their in- sertions. Key: M, femur (lesser trochanter); N, head of femur; P, symphysis pubis; Q, pelvis; W, femur (greater trochanter); Y, femur (posterior ridge); 5, adductor femoris; 5A, adductor longus; 6, pectineus; 8, iliopsoas; 16, gluteus medius; 19, piriformis; 20, gemelli; 21, obturator externus.
of the wing of the ilium from the crest area to the lower border of the sacro- iliac joint dorsally. The origin is practically inseparable from that of the gluteus minimus (18), which is immediately anterior and inferior on the iliac wing surface. Posteriorly, the muscle is practically inseparable from
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Ficure 20. Anterior view of right femur and pelvis. The hip joint is exposed. The distal end of the femur is seen. The patella has been reflected laterally between the vastus
Vout. XL] HOWARD: SEA OTTER HIND LIMB 361
Ficure 21. Posterior view of right knee and lower leg to show origin and insertion of the large superficial gastrocnemius (24) muscle. Note muscle insertions on the medial side of the tibia and their relation to the medial collateral ligament of the knee. Key: C, calcaneus; D, femur (distal end); F, foot; K, medial collateral ligament of knee; 2, gracilis (reflected) ; 3, semitendinosus (reflected); 4, semimembranosus (reflected); 5S, adductor femoris (superficial portion, reflected) ; 5D, adductor femoris (deep portion, reflected) ; 13, tenuissimus (reflected) ; 24, gastrocnemius; 31, plantaris; 34, tibialis posterior.
the piriformis (19), which has the bulk of its origin from the axial skeleton, thus marking the division between the two.
INSERTION. The gluteus medius terminates in a short heavy tendon insert- ing on the outer anterior and superior aspect of the greater trochanter of the femur.
Action. This muscle abducts and extends the femur.
17. Presemimembranosus.
A rather thin, flat, roughly triangular muscle connecting the first caudal vertebra with the femur.
Orictn. This muscle arises by a short strong tendon from the superior border of the transverse process of the first caudal vertebra. The muscle passes immediately over the dorsum of the ilium just above the tuberosity directed toward the femur and lies beneath the tenuissimus (13) and the caudofemoralis (11).
INSERTION. The muscle spreads out for a linear insertion on the femur (lateral ridge area) immediately posterior to the caudofemoralis (11). The insertion extends from a short distance below the greater trochanter to the distal femur, swinging somewhat medially at the epicondyle level.
Action. The action of this muscle is to extend the femur.
VoL. XL] HOWARD: SEA OTTER HIND LIMB 363
Ficure 22. Lateral view of right knee and lower leg. The biceps femoris (12) has been reflected anteriorly to show the lower leg musculature. Both heads of the gastroc- nemius (24) have been detached at their origins and the muscle reflected posteriorly and rotated somewhat to show the undersurface. Thus, the plantaris muscle (31) is exposed. Note the relationship of other lower leg muscles in this view. Key: A, fibula (head); B, sesamoid; D, femur (distal end); E, fibula (lateral malleolus); F, foot; O, patella; 5D, adductor femoris (deep portion, reflected) ; 7, rectus femoris (reflected) ; 10, vastus lateralis (reflected) ; 12, biceps femoris (reflected) ; 13, tenuissimus (reflected); 14, vastus medialis (reflected) ; 24L, gastrocnemius, lateral head (reflected) ; 24M, gastrocnemius, medial head (reflected) ; 25, tibialis anterior; 27, extensor digitorum longus; 28, peroneus longus; 29, peroneus digiti quinti; 30, peroneus brevis; 31, plantaris.
18. Gluteus minimus.
This is the lesser of the three glutei muscles and lies in close relation to the gluteus medius (16) in both origin and insertion.
OricIn. Muscular fibers arise from the inferior anterior aspect of the outer surface of the iliac wing, practically inseparable from the gluteus medius (16) and in close proximity to the origin of the rectus femoris (7). Toward its
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Ficure 23. Anterolateral view of right lower extremity to show relationship of the more superficial extensor muscles. The distal femur and knee joint are shown. The medial collateral ligament has been detached from the femur. The extensor musculature of the knee has been reflected anteriorly and rotated to show the undersurface of the patella and the patellar tendon. Note the transverse ligaments at ankle level; these serve as pulleys to prevent the extensor muscles from bowstringing when the foot is dorsiflexed. Key: D, femur (distal end); E, fibula (lateral malleolus); G, tibia (medial malleolus); L, patellar tendon; M, medial collateral ligament of knee; N, lateral collateral ligament of knee; O, patella; T, tibial crest; X, metatarsophalangeal joint level; 1, sartorius (reflected); 7, rectus femoris (reflected) ; 10, vastus lateralis (reflected) ; 14, vastus medialis (reflected) ; 25, tibialis anterior; 26, extensor hallucis longus (proprius); 27, extensor digitorum longus; 28, peroneus longus; 29, peroneus digiti quinti; 30, peroneus brevis.
insertion, the muscle becomes a little more individualized as its short strong tendon of insertion develops.
INSERTION. Insertion by the tendon is on the outer surface of the greater trochanter, just distal and anterior to the gluteus medius (16) tendon with which it is closely associated.
Action. Abduction and internal rotation of the femur.
19. Piriformis.
This large strong muscle of the buttock connects the axial skeleton to the femur.
OricIN. Tendinous and fleshy fibers arise from the fascia overlying the spinal musculature adjacent to the wing of the ilium and from the last lumbar and first sacral vertebrae. A few deeper fleshy fibers arise from the outer surface of the ilium adjacent to the sacroiliac joint. On surface viewing, it appears inseparable from the gluteus medius (16). The muscle mass curves superiorly around the greater trochanter.
Ww oO iat
Ficure 24. Lateral view of right lower leg including knee and ankle area. The tibialis anterior (25) has been removed from its origin and the distal portion reflected. The origin of the extensor digitorum longus (27) is now disclosed. As in figure 23, the patella and its tendon are again shown, reflected in a manner to disclose the undersurface. Key: D, femur (distal end); E, fibula (lateral malleolus) ; F, fibula (head); L, patellar tendon; N, lateral collateral ligament of knee; O, patella; T, tibial tuberosity; 7, rectus femoris (re- flected) ; 10, vastus lateralis (reflected) ; 14, vastus medialis (reflected) ; 25, tibialis anterior (reflected) ; 26, extensor hallucis longus (proprius); 27, extensor digitorum longus; 28, peroneus longus; 29, peroneus digiti quinti; 30, peroneus brevis.
INSERTION. The short heavy tendon inserts on the outer posterior aspect of the greater trochanter appearing as a continuation of the tendon of the gluteus medius (16).
Action. Abduction and distention of the femur.
20. Gemelli.
These two small short muscles designated as superior and inferior join the pelvis to the femur, lying one on either side of the obturator internus tendon (23).
Or1IGIN. Both muscles arise from the rim of the sciatic notch; the inferior one between the obturator internus (23) and sciatic spine, and the superior one adjacent to the obturator internus (23) to the border of the acetabulum.
INSERTION. The two gemelli join with the obturator internus (23) to form a common tendon which inserts on the medial surface of the greater trochanter as described for the obturator internus (23).
Action. The same action as the obturator internus (23).
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Ficure 25. Anterolateral view of the right lower leg including knee and ankle areas. The tibialis anterior (25), the extensor digitorum longus (27), and the peroneus longus (28) have been reflected. The double origin of the peroneus digiti quinti (29) is visible. The origin of the extensor hallucis longus (proprius) (26) from the intermuscular septum is also seen. Key: D, femur (distal end); E, fibula (lateral malleolus); F, fibula (head) ; I, interosseus membrane; L, patellar tendon; M, intermuscular septum; 7, rectus femoris; 10, vastus lateralis; 14, vastus medialis; 25, tibialis anterior; 26, extensor hallucis longus (pro- prius) ; 27, extensor digitorum longus (reflected muscle and stump of origin); 28, peroneus longus; 29, peroneus digiti quinti; 30, peroneus brevis.
21. Obturator externus.
Lying deep to the adductor musculature, this strong pear-shaped muscle connects the distal pelvis with the greater trochanter of the femur.
ORIGIN. Fleshy fibers arise from the entire outer surface of the obturator membrane, which covers the obturator foramen, and from the adjacent bony rim of the ischium and pubis.
INSERTION. The converging fibers form a strong tendon which inserts into the posteromedial aspect of the greater trochanter just inferior to the combined tendons of the obturator internus (23) and gemelli (20).
AcTIoN. External rotation of the femur.
2D Hiocapsularis.
This is a small, flat, triangular muscle arising from the acetabular rim and inserting on the greater trochanter.
OricIN. Fleshy fibers arise from the posterior rim of the acetabulum start- ing immediately adjacent to the superior gemelli (20). The muscle overlies the hip joint capsule as it triangulates toward the greater trochanter.
INSERTION. A short tendon is formed and the insertion is into the medial
VoL. XL] HOWARD: SEA OTTER HIND LIMB 367
Ficure 26. Lateral view of right lower leg and ankle. The peroneus digiti quinti (29), peroneus longus (28), and extensor digitorum longus (27) have been detached at their origins and reflected anteriorly to disclose the origin of the peroneus brevis (30). Note how, in the ankle area, the peroneus brevis (30) invests the peroneus digiti quinti (29) in a sling-like manner. Key: C, calcaneus; E, fibula (lateral malleolus); F, fibula (head) ; I, interosseus membranes; T, tibia; 24, gastrocnemius (reflected); 27, extensor digitorum longus; 28, peroneus longus; 29, peroneus digiti quinti (origin from fibula with muscle belly reflected) ; 30, peroneus brevis; 31, plantaris (reflected).
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Ficure 27. Lateral view of right knee following removal of the overlying musculature. The tendon of origin of the popliteus muscle (32) is seen. The extensor digitorum longus (27) has been divided near its origin and the distal portion reflected anteriorly. The stumps of origin of adjacent muscles are identified. A lateral semilunar cartilage is interposed between the joint surfaces. Key: D, femur (distal end); F, fibula (head); N, collateral ligament (lateral); S, semilunar cartilage; T, tibia; 27, extensor digitorum longus; 28, peroneus longus; 29, peroneus digiti quinti; 32, popliteus.
surface of the greater trochanter adjacent and anterior to the obturator (23) and gemelli (20) common tendon.
Action. This small muscle probably serves to stabilize the hip joint to some degree.
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Ficure 28. Posterior view of right lower leg with knee and ankle areas included. The foot is in plantar flexion. The popliteus muscle (32) has been detached at its tendinous origin and reflected medially. The origins, dispositions, and relationships of the tibialis posterior (34), flexor hallucis longus (35), and flexor digitorum longus (36) are shown. Note tendon arrangement at ankle. Key: C, calcaneus; D, femur (distal end); F, fibula (head) ; G, talus; I, interosseus membrane; QO, fibula (lateral malleolus); P, tibia (medial malleolus) ; T, tibia; X, metatarsal 1 (base); 24, gastrocnemius (reflected); 31, plantaris (reflected) ; 32, popliteus; 34, tibialis posterior; 35, flexor hallucis longus; 36, flexor digi- torum longus.
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23. Obturator internus.
This is a rather small flat muscle arising from the inner surface of the pelvis and extending to the femur.
OrtcIn. This muscle arises by fleshy fibers from the distal and inferior surface of the obturator membrane and from along the ischium in the region of the spine and extends anteriorly along the pubic ramus. A _ heart-shaped indentation seems to separate the ischial from the pubic origin. The muscle tapers abruptly to pass over the rim of the sciatic notch where it is flanked by the superior and inferior gemelli (20).
INSERTION. A common tendon is formed with the gemelli which then in- serts into the medial surface of the greater trochanter beneath the insertion of the piriformis (19) and between the insertion of the iliocapsularis (22) and the obturator externus (21).
Action. Abduction of the femur and also external rotation of the femur.
24. Gastrocnemius.
This large calf muscle arises by two heads on the distal femur and inserts on the calcaneus. The muscle covers much of the posterior aspect of the lower leg.
Oritcin. The medial head arises by heavy tendinous fibers from the pos- terior aspect of the distal femur just proximal to the medial condyle. The lateral head arises from a comparable position just proximal to the lateral condyle. The more medial fibers are fleshy and take origin from a slightly depressed area in the bone. The lateral tendinous fibers arise from a bony ridge along the posterior margin of the epicondyle. The sesamoid bone is in the tendon near its origin. The two heads meet about midway down the lower leg and at this point the tendon of insertion begins forming on the outer surface.
INSERTION. Distally, the tendon inserts at the tip of the calcaneus, sep- arating slightly to let the plantaris (31) tendon pass through the groove in
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Ficure 29. Sole of right foot with skin and subcutaneous tissue removed to show superficial musculature and tendon insertion in the proximal area. Also note fibrous flexor tendon sheaths of the digits. The plantaris (31), surrounded by the insertion of the gastroc- nemius (24), emerges from a groove in the calcaneus to assist in formation of the flexor digitorum brevis (37). Key: V, fibrous flexor tendon sheath of digits; X, base of metacarpal 1; Y, base of metacarpal 5; 24, gastrocnemius; 28, peroneus longus; 30, peroneus brevis; 31, plantaris; 34, tibialis posterior; 35, flexor hallucis longus; 36, flexor digitorum longus; 37, flexor digitorum brevis; 38, quadratus plantae; 40, abductor digiti quinti; 41, adductor digiti quinti; 43, abductor hallucis; 44, adductor hallucis; 48, lum- bricales.
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this bone. Muscular fibers continue a short distance down the medial side of the calcaneus.
Action. Contraction of this muscle gives strong plantar flexion of the foot.
25. Tibialis anterior.
The most anterior of the anterolateral muscle group is this large roughly pyramidal-shaped muscle connecting the tibia to the foot.
OricIn. Fleshy fibers arise from the lateral surface of the proximal one- third of the tibia, plus a small area from the adjacent interosseus membrane. Some posterior fibers of origin span over the tendon of origin of the extensor digitorum longus (27). The muscle then tapers and passes distally in close approximation to the tibia. In the upper two-thirds, the muscle is immediately anterior to the extensor digitorum longus (27), and in the lower one-third to the extensor hallucis proprius (26). At the ankle, a strong tendon has formed which, passing under the restraining transverse tarsal ligament, deviates me- dially to the base of the first metatarsal.
INSERTION. The tendon inserts into the base of the first metatarsal, at- taching to a lateral bony prominence.
Action. The action is dorsoflexion of the foot and inversion of the foot.
26. Extensor hallucis proprius (longus).
This is a relatively small muscle, somewhat triangular in shape and deep in origin, connecting the lower leg with digit 1.
OriciIn. The muscle arises by fleshy fibers from the lower one-half of the anterior surface of the intermuscular septum and from the anterior surface of the fibula in the same area, with a few fibers arising from the very margin of the adjacent intermuscular septum. The fleshy fibers of origin triangulate as they progress toward the medial side of the ankle. The muscle continues distally in front of the ankle to about the tarsal and metatarsal junction where a thin tendon is formed. This tendon, passing under the transverse ligament of the ankle, proceeds directly along the first metatarsal.
INSERTION. On reaching the metatarsal phalangeal joint level, the tendon spreads slightly making some fascial attachments, but continues down the dorsum of the proximal phalanx to insert at the dorsum of the base of the terminal phalanx.
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Ficure 30. Sole of right foot, identical with figure 29 except plantaris (31) has been transected at the calcaneus permitting the flexor digitorum brevis (37) to be reflected distally exposing the flexor digitorum longus (36) and lumbricale (48) musculature. The sketch at the right is a diagrammatic enlarged view of the flexor digitorum longus (36) for
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digits 4 and 5 to show the two superficial lumbrical muscles (48) (unshaded) which arise from these tendons and pass to the ulnar side of each digit respectively. Key: C, calcaneus; X, metatarsal 1 (base); Y, metatarsal 5 (base); 24, gastrocnemius; 28, peroneus longus; 30, peroneus brevis; 31, plantaris; 34, tibialis posterior; 35, flexor hallucis longus; 36, flexor digitorum longus; 37, flexor digitorum brevis (reflected); 38, quadratus plantae; 48, lumbricales.
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Action. Action of this tendon is to extend both the proximal and distal joints of the first digit.
27. Extensor digitorum longus.
This is a spindle-shaped muscle in the lateral aspect of the lower leg, which connects the femur to the digits.
OricIn. Origin of this muscle is by long, strong, rounded tendons from the lateral aspect of the distal femur adjacent to the articular surface of the knee. Crossing the knee joint, the tendon lies in the groove of the upper tibia and under the muscular fibers of origin of the tibialis anterior (25). The spindle- shaped muscle belly then forms and courses the length of the lower leg between the tibialis anterior (25) and the peroneus digiti quinti (29), and overlying the extensor hallucis proprius (26). Just above the ankle, the tendon of in- sertion develops and passes under the transverse ankle ligament where it divides promptly into four separate tendons which pass directly toward digits 2, 3, 4, and 5.
INSERTION. On reaching the metatarsal phalangeal joint level, the tendon enters the extensor aponeurosis of the respective digit. The tendon fibers continue distally in the aponeurosis and most of them insert into the base of the middle phalanx, but some may continue to the distal joint (see details of extensor aponeurosis).
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Ficure 31. Anterolateral view of the fifth digit with the fibrous flexor tendon sheath (V) reflected to show relationship and insertions of flexor digitorum longus (36) and flexor digitorum brevis (37). This is the pattern for digits 2 to 5. Note the concentration of transverse connective tissue fiber strands (W) in flexor tendon sheath (V) to form a pulley mechanism for the flexor tendons. Also note the sling-type structure (S) from flexor digitorum brevis (37) surrounding flexor digitorum longus (36) at metatarsal head level. Digit 1, of course, is without a flexor digitorum brevis tendon. The diagrammatic sketch to the right represents an anterior view at proximal interphalangeal joint level showing the manner of insertion of flexor digitorum brevis (37) at the base of the middle phalanx. Key: J, proximal phalanx; K, middle phalanx; L, distal phalanx; S, sling-like structure of brevis surrounding longus; V, flexor tendon sheath; W, flexor tendon sheath pulleys; X, metatarsal; 36, flexor digitorum longus; 37, flexor digitorum brevis.
Ficure 32. Sole of right foot with flexor digitorum brevis (37) and flexor digitorum longus (36) reflected to include the lumbricales (48), thus exposing the deep layer of intrinsic muscles and the ventral joint level of the tarsus. Key: C, calcaneus; S, sesamoid (base of metatarsal 5); T, capsule of tarsus; X, base of metatarsal 5; Y, base of meta- tarsal 1; 24, gastrocnemius; 28, peroneus longus; 30, peroneus brevis (reflected) ; 31, plan- taris; 34, tibialis posterior; 35, flexor hallucis longus (reflected) ; 36, flexor digitorum longus (reflected) ; 37, flexor digitorum brevis (reflected) ; 38, quadratus plantae (reflected) ; 40, abductor digiti quinti; 41, adductor digiti quinti; 42, opponens digiti quinti; 43, abductor hallucis; 44, adductor hallucis; 46, interossei; 48, lumbricales.
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Plantar view of metatarsal area of right foot. All small muscles have been removed, except the interossei (46). The plantar joint capsules of the metacarpophalangeal joints, except for digit 2, have been detached from the metacarpal and reflected distally, thus exposing the joints and revealing the two sesamoid bones in the capsule. For digit 2, the plantar joint capsule remains in place. Key: C, calcaneus; S, sesamoid bone; T, plantar capsule of tarsus; U, plantar joint capsule of metatarsophalangeal joint, reflected; X, meta-
tarsal 1; Y, metatarsal 5; 28, peroneus longus; 30, peroneus brevis; 34, tibialis posterior ; 46, interossel.
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Ficure 34. Plantar view of metatarsotarsal area. All small intrinsic muscles have been removed. The plantar joint capsule has been incised and reflected proximally to reveal the course of the peroneus longus (28) tendon as it passes in a groove in the cuboid to cross the foot and insert at the base of the first metatarsal. Key: CC, calcaneus; D, cuboid; T, capsule of tarsus (reflected) ; X, base of metatarsal 1; Y, base of metatarsal 5; 28, peroneus longus; 30, peroneus brevis; 34, tibialis posterior.
Action. The extensor action through the aponeurosis is to extend the proximal or metatarsal phalangeal joint, and also the middle and distal joints providing the metatarsal phalangeal joint is not in hyperextension.
28. Peroneus longus.
This spindle-type muscle connects the upper tibia with the foot. OricIN. The muscle arises with mostly tendinous fibers from the anterior,
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lateral, and inferior surfaces of the head of the fibula. Passing distally down the lower leg in the lateral position, the muscle lies between the extensor digitorum longus (27) and the peroneus digiti quinti (29). At the ankle, a strong round tendon develops which passes over the anterior surface of the lateral malleolus, under strong restraining ligament, and at this point the tendon overlies the peroneus digiti quinti (29) tendon and the peroneus brevis (30) tendon. Passing through a groove (perineal groove of the cuboid), the tendon dives deeply into the sole of the foot.
INSERTION. The tendon lies in its own sheet as it crosses the foot, buried deep in the plantar fibrous capsule of the tarsal area. Its actual insertion is into the base of the first metatarsal on the ventral aspect.
Action. The action of this muscle is to plantar flex and pronate the foot.
29. Peroneus digiti quinti.
This is a strong fleshy muscle of double origin, which joins the femur and fibula to the fifth digit.
OrIcIN. Origin of this muscle is by two heads; the larger and most pos- terior head arises by fleshy fibers from the lateral side of the distal femur
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and immediately posterior to the lateral collateral knee ligament and from a very slight bony prominence at the site. The lesser and more anterior head arises from the anterolateral surface of the very proximal portion of the fibular shaft and immediately adjacent to the peroneus longus (28) origin. Two muscle bellies join at about the junction of the upper and middle thirds of the fibula and continue distally between the peroneus brevis (30) and longus (28) muscles. Near the lateral malleolus, an ovoid strong tendon forms which then passes posteriorly and deep in the groove behind the malleolus to con- tinue along the lateral side of the ankle passing under the peroneus longus (28) tendon. The tendon of the peroneus brevis (30) lies immediately posterior and, near its insertion at the base of metatarsal 5, a fibrous-type investment sheath arises and surrounds the tendon of the peroneus digiti quinti as the restraining ligament.
INSERTION. The peroneus digiti quinti tendon continues distally along the lateral border of the fifth metatarsal to the proximal joint level, then on dis- tally to enter the extensor aponeurosis of this digit. The tendon continues to stay well to the lateral side, and at middle joint level it inserts into the base of the middle phalanx.
ActTIon. In view of the location of this tendon, it would serve as an ex- tensor of the metatarsal phalangeal joint of the fifth digit when working with the extensor digitorum longus (27), but when working independently it would
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FIGURE 36. Semidiagrammatic plantar view of fourth and fifth digits of right foot to show Origin and insertion of the two superficial lumbrical muscles (48). The flexor digitorum brevis (37) tendons are reflected just prior to their entering the flexor tendon sheath. Note the small tendons of the more superficial lumbrical muscles (48) joining the flexor digi- torum brevis (37) tendon on the lateral side at this site. Key: V, flexor tendon sheath (muscles) ; 36, flexor digitorum longus; 37, flexor digitorum brevis; 48, lumbricales (both deep and superficial).
FicurE 37. Dorsal view of left foot. All skin and subcutaneous tissue have been re- moved, including the interdigital webbing. Visible is the distribution pattern of the toe extensors, and the extensor aponeuroses over the dorsum of digits 2, 3, 4, and 5. The proximal end of the aponeurosis forms the tendinous hood about the metatarsophalangeal joint. Key: A, extensor aponeurosis; H, extensor hood; J, metatarsophalangeal joint level; K, proximal interphalangeal joint level; L, distal interphalangeal joint level; X, metatarsal 1; Y, metatarsal 5; 25, tibialis anterior; 26, extensor hallucis longus (proprius); 27, ex- tensor digitorum longus; 29, peroneus digiti quinti; 47, extensor digitorum brevis.
Ficure 38. Anterior or dorsal view of left foot. The extensor digitorum longus (27) tendons have been reflected distally to show the origin of the extensor digitorum brevis musculature (47). Note that the extensor digitorum brevis (47) has good representation for digits 2, 3, and 4. The token representation to digits 1 and 5 has tiny tendons which appear to join into the extensor digitorum longus (27) for each digit. Key: 25, tibialis anterior; 26, extensor hallucis longus (proprius) ; 27, extensor digitorum longus (reflected) ; 29, peroneus digiti quinti; 47, extensor digitorum brevis.
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Ficure 39. An enlarged medial view of the second digit of the right foot to show the usual relationship of long extensor (27), interossei (46), and lumbrical (48) tendons. Note the tendinous hood about the metatarsophalangeal joint, and the extensor aponeurosis, which continues distally. The tendon of the interosseus (46) forms the lateral band of the aponeurosis. The small lumbrical (48) tendon inserts into the proximal phalanx. Key: A, extensor aponeurosis; B, lateral band of extensor aponeurosis; H, extensor hood; J, proximal phalanx; M, metatarsophalangeal joint; X, metatarsal; 27, extensor digitorum longus; 46, interossei; 48, lumbricales.
Ficure 40. Medial view of the fifth digit of the right foot. Note that there is no interosseus muscle for this digit, and that the lumbrical (48) tendon, instead of inserting into the proximal phalanx, joins the extensor (27) aponeurosis to form the lateral band on the medial side. Key: A, extensor aponeurosis; B, lateral band of extensor aponeurosis; J, proximal phalanx; M, metatarsophalangeal joint; X, metatarsal 5; 27, extensor digitorum longus; 48, lumbricales.
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Ficure 41. Diagrammatic representation of the extensor mechanism of the digits of the right foot. Note that only for digits 2, 3, and 4 are interossei (46) and extensor digitorum brevis (47) present. For digit 5, the peroneus digiti quinti (29) serves as an extensor digi- torum brevis, and the lumbrical (48) joins the extensor aponeurosis instead of inserting directly into the proximal phalanx. Digit 1 shows the simple arrangement of a single extensor tendon made possible by the presence of only two joints. Key: 26, extensor hal- lucis longus (proprius); 27, extensor digitorum longus; 29, peroneus digiti quinti; 46, in- terossei; 47, extensor digitorum brevis; 48, lumbricales.
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Ficure 42. Anterior view of the sacrum. Key: A, sacroiliac joint; 8, iliopsoas (origin).
serve to abduct digit 5 at the metatarsal phalangeal joint level. Actually, the action would be mostly abduction from the adduction position, since abduction per se of this joint is practically nil past the straight line with the metatarsal.
30. Peroneus brevis.
This is the most posterior of the peroneal group connecting the fibula with the foot.
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Ficure 43. Right lateral view of sacrum. Key: A, sacroiliac joint; B, spinous process of first sacral vertebra; 19, piriformis (origin).
OriciIn. The muscle arises from the distal two-thirds of the shaft of the fibula from the lateral, posterior, and anterior surfaces and from the posterior side of the interosseus membrane in this area. The origin is by fleshy fibers for the most part. Superiorly and posteriorly, the origin is in close association
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Ficure 44. Anterior view of right pelvis. Key: A, ilium; B, ischium; C, pubis; D, acetabulum; E, iliac crest; F, symphysis pubis; G, ischial tuberosity; H, obturator foramen; 1, sartorius (origin) ; 2, gracilis (origin) ; 4, semimembranosus (origin) ; 5A, adductor longus (origin) ; 5D, adductor femoris (deep portion—origin) ; 5S, adductor femoris (superficial portion—origin) ; 6, pectineus (origin); 7, rectus femoris (origin); 8, iliopsoas (origin) ; 21, obturator externus (origin).
VoL. XL] HOWARD: SEA OTTER HIND LIMB 389
with the flexor digitorum longus (36). The muscle then courses distally along the side and posterior to the peroneus digiti quinti (29). Near the ankle, a heavy tendon forms which rounds the lateral malleolus posterior to, but in a common sheath with, the tendon of the peroneus digiti quinti (29). A few muscle fibers on the posterior surface of the tendon continue with the tendon around the lateral malleolus.
INSERTION. It continues distally along the lateral side of the ankle under the peroneus longus (28) tendon to the base of metatarsal 5. Near its inser- tion, the fibrous sheath-like structure envelops the more anterior peroneus digiti quinti (29) tendon for a short distance.
Action. Eversion and plantar flexion of the foot.
31. Plantaris.
This muscle bears a close association with the lateral head of the gastroc- nemius (24) and, for most of its distance in the muscular area, it is insep- arable from this lateral head.
Oricin. Distinct tendinous fibers arise in conjunction with the origin of the lateral head. These fibers are the most lateral ones, and from the origin to midway to the ankle distinct separation of the lateral head does not occur. At midpoint, however, the muscle becomes separate and moves medially to the midline. A tendon of insertion begins to form at this point. By the time the calcaneus is reached, a distinct, somewhat flattened tendon has formed which lies directly under the tendon of the gastrocnemius (24).
INSERTION. The tendon now enters a groove at the posterior end of the calcaneus as the gastrocnemius (24) tendon is inserting to either side. Passing through the tendon of the gastrocnemius (24), the plantaris tendon reaches the sole of the foot where it enters into the flexor digitorum brevis (37) mechanism.
Action. This tendon would serve to plantar flex the foot and to also flex the digits through the flexor digitorum brevis (37) complex.
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Ficure 45. Posterior view of right pelvis. Key: D, acetabulum; E, iliac crest; F, symphysis pubis; G, ischial tuberosity; H, obturator foramen; I, sacroiliac joint; 12, biceps femoris (origin); 20, gemelli (origin); 22, iliocapsularis (origin); 23, obturator internus (origin).
Ficure 46. Lateral view of right pelvis. Key: D, acetabulum; E, iliac crest; F, symphysis pubis; G, ischial tuberosity; H, obturator foramen; J, ischial spine; 1, sartorius (origin) ; 5, adductor femoris (origin); 5A, adductor longus (origin); 7, rectus femoris (origin); 9, gluteus maximus (origin); 12, biceps femoris (origin); 16, gluteus medius (origin) ; 18, gluteus minimus (origin); 19, piriformis (origin); 20, gemelli (origin); 21, obturator externus (origin).
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32. Popliteus.
This is a large posterior muscle of the lower leg connecting the femur with the tibia. The muscle lies beneath the medial head of the gastrocnemius (24).
OricIN. The origin of this muscle is by a very strong tendon in the lateral aspect of the distal femur, adjacent to the lateral condylar articular surface and distal to the tendon of origin of the extensor digitorum longus (27). The tendon lies transverse to the axis of the femur; it passes posteriorly under the lateral collateral ligament of the knee and over the lateral semilunar cartilage and joint capsule. Reaching the posterior aspect of the knee, the tendon spreads rapidly into a large triangular-shaped muscle which immediately starts its insertion into the posteromedial aspect of the upper tibia. The lat- eral side of the muscle parallels and overlaps somewhat the tibialis posterior (34) and the flexor hallucis longus (35).
INSERTION. Muscular fibers insert on the medial side of the posterior aspect of the tibia, starting at joint level and passing diagonally medially from the mid-upper tibia to a point distally and medially a distance one-fourth from the distal end of the tibia.
Action. This would be a strong medial rotator of the tibia and a flexor of the knee.
33. Dr. Howard’s notes state “Not found” after the number ‘33.’ Editor.
THE ORIGIN OF THE DEEP POSTERIOR MUSCULATURE
The tibialis posterior (34), the flexor hallucis longus (35), and the flexor digitorum longus (36) form a more or less common combined origin from the full length of the posterior aspect of the fibula, the interosseus membrane, and the distal posterior surface of the tibia. Intercommunicating fibers exist for all three muscles.
The fibers of origin of the tibialis posterior (34) rise mainly from the head of the fibula, interosseus membrane, and lower tibia.
The fibers of origin of the flexor hallucis longus (35) come mainly from the posterior surface of the head of the fibula, medial to and overlying those of the tibialis posterior (34).
The fibers of origin of the flexor digitorum longus (36) arise mainly from the posterior aspect of the fibula for its full length and the adjacent inter- osseus membrane.
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Ficure 47. Anterior view of the right femur. Key: A, head of femur; B, greater trochanter of femur; C, lesser trochanter of femur; 10, vastus lateralis (origin); 14, vastus medialis (origin); 16, gluteus medius; 18, gluteus minimus.
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The fiber interdigitation of these muscles is mainly in the upper two-thirds. The tendon of each develops in the lower one-third.
34. Tibialis posterior.
This is a deep muscle in the posterior aspect of the lower leg connecting the leg to the foot.
OricIn. Muscular fibers are mainly from the head of the fibula and lower end of the tibia and interosseus membrane. Near the ankle, a tendon forms that is most medial and close along that of the flexor hallucis longus (35). The tendon passes around the medial malleolus with a sheath in common with the flexor hallucis longus (35).
INSERTION. The tendon flattens somewhat as it passes distally to the base of the first metatarsal, inserting into the plantar aspect of the proximal end of this bone.
Action. The action is plantar flexion of the ankle and foot.
35. Flexor hallucis longus.
This is a slender muscle passing from the lower leg to the first toe.
Oricin. The origin is mainly by muscular fibers from the head of the fibula on the medial side of the posterior aspect. The origin is closely asso- ciated with the tibialis posterior (34) until the tendon starts to form. The tendon parallels that of the tibialis posterior (34) and passes around the medial malleolus in the same sheath. Just distal to the talus, the tendon joins with the flexor digitorum longus (36), but shortly thereafter again be- comes a separate tendon passing down the line of metatarsal 1 to enter the fibrous flexor tendon sheath at the metatarsal phalangeal joint level. At this point distally, the tendon passes within the sheath to its insertion on the ventral lip of the distal phalanx adjacent to the articular surface.
Action. Flexion of the proximal and distal joints of digit 1.
36. Flexor digitorum longus.
This is a deep posterior muscle connecting the lower leg to the digits.
OricIN. The origin is mainly by muscular fibers from the posteromedial aspect of the fibula for its full length and adjacent interosseus membrane. The muscle broadens distally and abruptly narrows into a tendon at ankle joint level. The tendon then passes through the groove in the posterior talus
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Ficure 48. Posterior view of right femur. Key: A, head of femur; B, greater tro-
chanter; C, lesser trochanter; 5A, adductor longus; 5D, adductor femoris (deep portion) ;
6, pectineus; 11, caudofemoralis; 17, presemimembranosus; 19, piriformis; 20, gemelli; 21,
obturator externus; 22, iliocapsularis; 23, obturator internus; 24, gastrocnemius (medial and lateral heads—origins) ; 31, plantaris (portion of origin).
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to join the tendon of the flexor hallucis longus (35) as both emerge into the sole of the foot at the distal medial side end of the calcaneus. Here, inter- digitation of fibers occurs for a short distance. The flexor hallucis longus (35) then again separates and passes down the line of the first metatarsal to the proximal or metatarsal interphalangeal joint level. The remaining tendon (flexor digitorum longus) continues into the sole of the foot, underlying the flexor digitorum brevis (37). From the area of interdigitation with the flexor hallucis longus (35) to just beyond the calcaneal-cuboid articulation, the long digital flexors join from the lateral side by the musculotendinous fibers of the short, flat, and broad quadratus plantae (38).
37. Flexor digitorum brevis.
This muscle develops the connection with the plantaris (31) tendon as it appears in the sole of the foot.
OricIN. As the plantaris tendon (31) enters the sole of the foot from the groove in the calcaneus, it expands into a musculotendinous structure (flexor digitorum brevis) (37). This structure is superficial and occupies or covers much of the proximal foot in a manner similar to a plantar fascia. Distally at about the half-way point of the sole, the musculotendinous mass separates into four slightly flattened tendons which proceed fan-like to the metacarpal head areas of digits 2, 3, 4, and 5. At this point, each tendon directly overlies the flexor digitorum longus (36) tendon, and together they enter the flexor tendon sheath which extends for the full length of the digit. This flexor tendon sheath is a fibrous connective tissue tunnel with transverse reinforce- ments (termed pulleys) at proximal and middle joint levels. The pulleys and sheath act to prevent bowstringing of the flexor tendons as the digits are flexed. At proximal joint level, the flexor digitorum brevis forms a sling-like structure which completely surrounds the profundus tendon for a short dis- tance and holds the two tendons in close proximity. Just within the flexor tendon sheath, the tendon splits and assumes a more dorsal position, which permits the flexor digitorum longus (36) to become superficial. Continuing on either side of the flexor digitorum longus (36), the flexor brevis now passes with the longus under the distal flexor tendon sheath pulley at proximal inter- phalangeal joint level. Each half of the tendon then broadens. The two halves join together deep to the flexor digitorum longus, forming a flat tendon which inserts into the ventral lip of the middle phalanx at proximal interphalan- geal joint level. Vincula are present as the tendon approaches its insertion.
>
Ficure 49. Medial view of right femur. Key: A, head of femur; C, lesser trochanter of femur; D, medial epicondyle of femur; 5S, adductor femoris (superficial portion) ; 5D, adductor femoris (deep portion); 6, pectineus; 8, iliopsoas.
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Action. The tendon of the flexor digitorum brevis can serve to flex inde- pendently the proximal interphalangeal joint, but can flex the metatarsal phalangeal joint as well.
38. Quadratus plantae.
This is a short, broad, flat muscle joining the calcaneus to the flexor digitorum longus (36) tendons.
OricIn. This muscle arises from the lateral aspect of the calcaneus for its full length. The muscle triangulates somewhat, passing distally and cov- ering most of the plantar surface of the distal one-half of the calcaneus on its plantar aspect. Mostly fleshy fibers then join the flexor digitorum longus tendon (36) just opposite where the flexor hallucis longus (35) tendon also joins, and just proximal to the point of division of the flexor digitorum longus (36) into the separate tendons which pass to digits 2, 3, 4, and 5.
Action. The exact function of this muscle is not clear, but it probably aids in holding the long digital flexors laterally during the time that they are contracting.
39. Calcaneometatarsalis.
This is an extremely small muscle overlying the quadratus plantae (38), joining the calcaneus with the proximal end of the fifth metatarsal.
OriciIn. This muscle arises from the small area on the posterolateral aspect of the calcaneus immediately adjacent to the point of exit of the plantaris (31) tendon. This small muscle then tapers as it passes obliquely over the quadratus plantae (38) toward the base of metatarsal five. A thin tendon forms at the distal one-third, continues distally, and overlies the peroneus longus tendon (28) where it dips deeply into the sole, then underlies the medial side of the peroneus brevis (30).
INSERTION. This tiny tendon then inserts into the base of the fifth meta- tarsal just proximal to the insertion of the peroneus brevis (30).
AcTION. Just what the action of this muscle is would be difficult to deter- mine. From the standpoint of origin and insertion, the muscle could support the longitudinal arch of the foot.
THE INTRINSIC MUSCLES OF THE FOooT
This group of small muscles has origin and insertion within the foot proper. They serve to balance the action of the longer and more powerful prime movers
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Ficure 50. Lateral view of right femur. Key: A, head of femur; B, greater trochanter of femur; E, attachment of lateral collateral ligament of knee; 9, gluteus maximus; 11, caudofemoralis; 16, gluteus medius; 18, gluteus minimus; 19, piriformis; 27, extensor digi- torum longus (origin) ; 29, peroneus digiti quinti (origin) ; 32, popliteus (origin).
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of the digits, and to give finer and more individualized skill motions to the digits. The broad webbing and elongated digits of the hind limb, as compared to the forelimb, permits more individualized digital motions.
The intrinsic muscles fall into three groups for descriptive purposes: the lumbrical muscles (48) which arise from the flexor digitorum longus (36) in the foot, the small muscles grouped about the first and fifth metatarsals, and the deeply situated interossei musculature (46).
The extensor digitorum brevis (47), although its origin and insertion are within the foot, is not considered an intrinsic muscle, but it is described in this general group for convenience.
INTRINSIC MUSCLES OF THE FiFTH DiciT (40, 41, 42)
The fifth digit in the sea otter is the largest, and therefore the small muscles about this digit have a greater total volume than those about the first digit. The opponens muscle (42) is very small and, in view of its insertion, could hardly be expected to give opposition to this digit.
40. Abductor digiti quinti.
This is one of the three small muscles grouped about the fifth metatarsal.
Oricin. This largest of the small foot muscles is the most lateral in loca- tion. The muscle arises proximally from the heavy ligamentous capsule and small sesamoid bone overlying the plantar aspect of the base of the fifth meta- tarsal on the medial side and from the base of the metatarsal laterally adjacent to the insertion of the peroneus brevis (30). The origin continues distally from the lateral side of the fifth metatarsal for its full length. The muscle passes somewhat obliquely across the plantar surface of the fifth metatarsal, covering most of the ventral surface of the bone. At the metatarsal head, a short broad tendon develops which blends into the joint capsule, making at- tachments to the lateral sesamoid and the base of the proximal phalanx on its lateral side.
Action. This muscle serves to abduct and flex the proximal joint of the fifth digit.
41. Adductor digiti quinti.
This muscle is located opposite the abductor (40) and opposes its action. Oricin. This muscle is considerably smaller than the abductor (40). It takes its origin from the capsular structures just medial to the abductor (40)
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Ficure 51. Anterior view of right tibia. Key: A, tibial plateau; B, tibial tuberosity; C, tibial crest; D, medial malleolus; E, patella tendon (combined tendons of 7, 10, and 14) ; 1, sartorius; 2, gracilis; 3, semitendinosus; 25, tibialis anterior (origin).
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and also from the medial side of the fifth metatarsal shaft in its proximal one-half. The muscle then courses distally to the medial side of the proximal joint where a short flattened tendon develops.
INSERTION. This tendon blends into the joint capsule, attaching to the medial sesamoid to make its insertion into the base of the proximal phalanx on the medial plantar aspect. Distally, the thin tendon of the opponens digiti quinti (42) courses over the tendinous expansion.
Action. This muscle serves to adduct and flex the proximal joint of the fifth digit.
42. Opponens digiti quinti.
This is a small intrinsic muscle and from its position is given the name opponens.
Oricin. The muscle arises from the midcentral area of the fibrous capsule of the tarsus and superficially overlies the proximal origin of all interossei (46) except the most medial one. The muscle tapers promptly into a very thin tendon at the level of the junction of the middle and proximal thirds of the fourth metatarsal. The tendon then continues obliquely toward the medial side of the proximal joint of the fifth digit. Keeping to the medial side, the tendon passes over the tendon of the adductor hallucis (44) and appears to join into the extensor aponeurosis one-third of the way down the proximal phalanx.
AcTIon. This small muscle could assist in flexion of the proximal joint, adduction of the proximal joint, and possibly to the extensor aponeurosis, giving some assistance in the extension of the middle and distal joints. In spite of its name, its insertion would exclude any opposition motion of the fifth digit.
INTRINSIC MUSCLES OF THE First Dicit (43, 44)
The first digit with its two phalanges is the shortest toe in the foot, and the two associated intrinsic muscles are likewise small. There is no opponens muscle for this digit.
43. Abductor hallucis.
This muscle is the most medial of the small foot muscles. OrIcIN. This muscle arises from the fibrous plantar capsule of the tarsus at the base of the first metatarsal. The muscle is fusiform in shape and passes
>
Ficure 52. Posterior view of right tibia. Key: A, tibial plateau; D, tibia (medial malleolus) ; 32, popliteus; 34, tibialis posterior (origin).
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distally overlying the medial one-half of the first metatarsal. A short flat tendon develops just proximal to the proximal joint level and passes medially over the medial sesamoid to which it has fibrous attachment, then continues in an aponeurosis-like structure to the base and side of the proximal phalanx for its insertion.
Action. Flexion of the proximal joint and abduction of the proximal joint of digit one.
44. Adductor hallucis.
OriciIn. This small fusiform muscle and companion of the abductor (43) arises from the fibrous plantar capsule on the lateral side of the base of meta- tarsal one. Proceeding distally and parallel to the abductor, it overlies the lateral one-half of metatarsal one on its ventral surface. Distally, a short flat tendon develops which passes to the lateral side, inserting in a similar manner to the abductor. Fibrous attachments are present to the lateral sesamoid and joint capsular structures, and eventually insertion is into the proximal side of the proximal phalanx.
Action. Flexion and adduction of the proximal joint of digit one.
45. Dr. Howard omitted mention of any muscle numbered ‘45.’ Editor.
THE INTEROSSEI MUSCULATURE
The interossei muscles are six in number. They lie deep in the sole of the foot, being cradled in the transverse metatarsal arch. There are two interossei for each of digits two, three, and four, and they insert respectively on the medial and lateral sides of the digits.
46. Interossel.
Excluding the small muscles about the first and the fifth digits, there are six small interossei muscles: two each for digits two, three, and four. For descriptive purposes, they can be identified from the medial to the lateral side as 46a-f.
Oricin. The interossei muscles arise in sequence from the heavy fascia covering the plantar surface of the tarsus, from the lateral side of the base of metatarsal one to the medial side of the base of metatarsal five. Muscle fibers also arise from the periosteum of the proximal ends of the metatarsal. The origin of 46a and b overlies the proximal end of metatarsal two. The origin
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Ficure 53. Medial view of right tibia. Key: B, tibial tubercle; D, medial malleolus of tibia; F, anterior crest of tibia; G, medial collateral ligament; 1, sartorius; 2, gracilis; 3, semitendinosus; 4, semimembranosus.
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of 46c and d overlies the proximal end of metatarsal three. The origin of metatarsal 46e and f overlies the proximal end of metatarsal four, with some additional overlapping of f onto the proximal end of metatarsal five.
INSERTION. The muscles pass distally to assume their respective positions on either side of metatarsals two, three, and four. Near the metatarsophalan- geal joint, flat tendons form which spread out with attachments to the joint capsule and adjacent sesamoid in a more ventral area, and the more dorsal area of tendon extends distally and dorsally into the extensor hood to join the extensor aponeurosis of the digit.
Action. The action of the interossei muscles is to flex the metatarso- phalangeal joint and to give a degree of lateral motion at this joint. Also, through the lateral bands, these muscles would act to extend the middle and distal interphalangeal joints.
47. Extensor digitorum brevis.
This muscle is an accessory digital extensor which lies deep to the extensor digitorum longus (27) and joins the ankle area with the digits.
OricIN. This muscle arises from the calcaneus and adjacent capsular struc- tures on the dorsolateral aspect of the ankle. The origin lies directly under the extensor digitorum longus (27) tendons where they pass through the sheath in front of the ankle. Three distinct muscle bellies are present, and these extend distally to about the metatarsal junction of digits five, four, and three. The muscle terminates in three long flat tendons Which pass to the ulnar sides of digits four, three, and two where they enter the hood structure and extensor aponeurosis.
INSERTION. These tendons enter the extensor aponeurosis to the ulnar side of the extensor digitorum longus (27) and the tendinous fibers continue dis- tally to form a generally lateral border of the aponeurosis. At the interphalan- geal joint level, the fibers are generally central with those of the longus, and the insertion is into the base of the middle phalanx. Tendinous fibers may continue to form, in part at least, the extensor tendon going to the distal joint. In the specimen dissected, there arose from the medial and lateral muscles tiny
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Ficure 54. Lateral view of right tibia. Key: D, medial malleolus of tibia; F, tibial crest; 12, biceps femoris; 25, tibialis anterior (origin).
Ficure 55. Anterior view of right fibula. Key: A, head of fibula; B, lateral malleolus; C, articulation with tibia; 26, extensor hallucis longus (proprius—origin) ; 28, peroneus longus (origin); 29, peroneus digiti quinti (anterior head—origin); 30, peroneus brevis (origin).
Ficure 56. Posterior view of right fibula. Key: A, head of fibula; B, lateral malleolus; C, articulation with tibia; 30, peroneus brevis (origin) ; 36, flexor digitorum longus (origin).
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tendinous slips which joined respectively with the extensor digitorum longus (27) of the fifth and the extensor hallucis proprius (26) of the first. It is highly probable that individual variations occur.
AcTIon. The tiny tendinous slips that go to the first and fifth digits are so small that the function would be minimal, if any. For the main tendons going to digits two, three, and four the action would be the same as the extensor digitorum longus (27), as both are acting through the extensor apo- neurosis of the digits.
48. Lumbricales.
These small slender muscles arise from the long flexor tendons of digits two, three, four, and five, and are generally included in the intrinsic muscle group.
ORIGIN. Out of a total of six muscles, four follow a basic pattern of arising from the radial side of each flexor tendon (flexor digitorum longus—36) for each of the digits two, three, four, and five. They are thin fusiform muscles, each forming a small thin tendon just proximal to the metatarsophalangeal joint level on the radial side of each digit. The other two more superficial muscles arise more from the ventral surfaces of the flexor digitorum longus (36) tendon of four and five. They are closely associated proximally, but separate distally into two distinct muscles. Each forms a thin tendon, the more medial one passing toward the fourth digit and the more lateral toward the fifth digit.
INSERTION. Of the four lumbricales arising from the basic pattern, each has a thin long tendon which goes respectively to the radial sides of digits two, three, four, and five. For digits two, three, and four, a slender tendon inserts on the radial side of the proximal phalanx a short distance beyond the proximal joint and ventral to the extensor aponeurosis. For digit five, the tendon joins with the extensor digitorum longus (27) to form the radiolateral margin of the extensor aponeurosis.
The more superficial lumbrical muscles arising from the long flexor tendons of four and five, are much smaller than the others. Their thin tendons pass
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Ficure 57. Lateral view of right fibula. Key: A, head of fibula; C, lateral malleolus
of fibula; 28, peroneus longus (origin) ; 29, peroneus digiti quinti (anterior head—origin) ; 30, peroneus brevis (origin).
Ficure 58. Anterior view of right tibia and fibula (articulated). Key: T, tibia; F, fibula; I, interosseus membrane; 2, gracilis; 4, semimembranosus; 25, tibialis anterior (ori- gin); 26, extensor hallucis longus (proprius—origin) ; 28, peroneus longus (origin).
Ficure 59. Posterior view of the right tibia and fibula (articulated). Key: T, tibia; F, fibula; I, interosseus membrane; 32, popliteus; 34, tibialis posterior; 35, flexor hallucis longus; 36, flexor digitorum longus.
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ventrally and join respectively the ulnar sides of the sublimus flexor tendons just prior to their disappearance into the flexor tendon sheath at the meta- tarsal joint level.
Action. The action of the four lumbricales which follow the basic pattern would be to assist in flexion of the metatarsophalangeal joint. They would also give some radial lateral flexion of the joint. The one passing to digit five would also assist in extension of the distal two joints through the extensor aponeurosis. The more superficial lumbrical muscles, which serve to join the profundus and sublimus flexors, undoubtedly have some function but it would be difficult to state just what this function is.
EXTENSOR APONEUROSIS
The extensor mechanism for the digits is complicated and not completely understood. The tendinous flattened structure overlying the dorsum of digits 2 through 5 inclusive provides the mechanical arrangement for the insertion and action of the extrinsic and intrinsic musculature. By extrinsic is meant the extensor digitorum longus (27), the extensor digitorum brevis (47), the extensor hallucis longus (26), and the peroneus digiti quinti (29). By intrinsic musculature is meant the interossei (46) and for digit 5 the lumbrical (48) muscle.
The general plan of this mechanism is as follows: The long tendons of the extrinsic muscles for digits 2, 3, 4, and 5 join at the metatarsophalangeal joint level in a fibrous hood-like structure which marks the beginning of the aponeurosis and which attaches ventrally to the capsular structures on either side. The long tendons do not insert into the proximal phalanx, but act to extend the metatarsophalangeal joint by virtue of this hood-type structure. The aponeurosis now proceeds distally as a continuation of the tendons with the cross fibers, with the addition of the interrossei tendons medially and laterally for digits 2, 3, and 4, to form lateral bands; for the fifth digit the lumbrical performs this function on the radial side.
At proximal interphalangeal joint level, direct insertion at the base of the middle phalanx occurs for most of the extensor tendon fibers, but some con- tinue on to insert terminally at the base of the distal phalanx. Both insertions are on the dorsal aspects. The fibers continuing beyond the interphalangeal joint level form a thin and rather poorly developed tendon.
In fresh specimens, traction on the extrinsic tendons will extend all three joints of digits 2, 3, 4, and 5, provided the metatarsophalangeal joint is not permitted to hyperextend. If the metatarsophalangeal joint is allowed to hyperextend, the distal two joints drop into slight flexion as the excursion of the long tendons is limited by the hood attachment. This claw-type deformity is corrected, for the most part, when traction is made along the lateral bands of the aponeurosis.
Vout. XL] HOWARD: SEA OTTER HIND LIMB 415
Although passive hyperextension is present for both proximal and distal interphalangeal joints, active hyperextension is not obtained by traction on the aponeurosis. In fact, the tendon action distal to proximal interphalangeal joint level seems limited and weak.
See figure 41 for a diagrammatic representation of the aponeurosis and the various tendon components for the specimen study. Undoubtedly, variations would be noted if many specimens were to be dissected.
ACKNOWLEDGMENTS
The State of California Department of Fish and Game was most coop- erative in providing the specimen for anatomical study. The California Acad- emy of Sciences provided the invaluable aid of a disarticulated sea otter skeleton.
The Anatomical Studies of the California River Otter, by Edna M. Fisher, proved to be an excellent guide during the dissection, and her unpublished notes on sea otter anatomy, which were made available through the courtesy of Mr. Fred Tarasoff, were also of value. Unpublished notes and sketches by Fred Tarasoff were also available and of some assistance.
The dissection and storage of materials took place at the Hopkins Marine Station of Stanford University in Pacific Grove, California, and the personnel of this institution, particularly Dr. D. Abbott, gave encouragement and as- sistance in every way possible.
BIBLIOGRAPHY
BarABASH-NIKIFOROV, I. I. 1962. The sea otter (Enhydra lutris L.). Biology and econemic problems of breeding. Translated from the Russian by the Israeli Program for Scientific Translations. National Science Foundation and U.S. Department of the Interior, Washington, D.C., 227 pages. FisHER, Epna M. 1939. Habits of the southern sea otter. Journal of Mammalogy, vol. 20, no. 1, pp. 21-36. 1942. Osteology and myology of the California river otter. Stanford University Press, Stanford, California, vi + 66 pages. No date. Notes and sketches on the myology of the southern sea otter. (Uupublished.) Courtesy of Fred Tarasoff.
Goup_Er, F. A. 1925. Bering’s voyages. American Geographical Society, Research Series, no. 2, 290 pages.
GRINNELL, JOSEPH, S. DIxon, AND JEAN M. LINSDALE 1937. Fur-bearing mammals of California. University of California Press, Berkeley, California, 2 vols., 777 pages. Bian Es Re 1926. Muscular anatomy of three mustelid mammals, Mephitis, Spilogale, and Martes. University of California Publications in Zoology, vol. 30, no. 2, pp. 7-38.
416 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Jeary I 1G We
1963. Tool-using performances as indicators of behavioral adaptability. Current
Anthropology, vol. 4, Dec., pp. 479-494. Harris, C. J.
1968. Otters: A study of the recent Lutrinae. Weidenfeld and Nicolson, London, xiv ++
397 pages. Howarp, L. D.
1973. Muscular anatomy of the forelimb of the sea otter (Enhydra lutris). Pro- ceedings of the California Academy of Sciences, ser. 4, vol. 39, no. 20, pp. 411-500.
Howe tt, A. B. 1930. Aquatic mammals. Charles C. Thomas, Springfield, Ill., and Baltimore, Mary- land, xii + 338 pages. JoLiie, M. 1962. Chordate morphology. Reinhold, New York, 478 pages. Jacost, A.
1938. Der Seeotter. Monographien der Wildsaugetiere, Band VI, 93 pages. (Tyansla-
tion: Fisheries Research Board of Canada, No. 521.) Kenyon, K. W.
1969. The sea otter in the eastern Pacific Ocean. North American Fauna, no. 68, U.S. Department of the Interior, Bureau of Sport Fisheries and Wildlife, Washington, xiii + 352 pp.
Mourig, O. J.
1940. Notes on the sea otter. Journal of Mammalogy, vol. 21, no. 2, pp. 119-131. Orr, R. T.
1966. Vertebrate biology (second edition). W. B. Saunders Co., Philadelphia, xii +
483 pages. Romer, A. S. 1964. The vertebrate body. W. B. Saunders Co., Philadelphia and London, vii -+ 643 pages.
SoxoLov, A. S., anp I. I. SoxoLov 1970. Some specific features of the locomotor organs of the river otter and sea otter associated with their mode of life. Moscovskoe Obshchestvo Ispytatelel Priorody, Otdel Biologicheskii, vol. 75, no. 5, pp. 5-17. (Translated from the Russian, Hopkins Marine Station, Pacific Grove, California, 1972.) Tavyior, W. P. 1914. The problem of aquatic adaption in the Carnivora, as illustrated in the osteology and evolution of the sea otter. University of California Publications, Bulletin of the Department of Geology, vol. 7, no. 25, pp. 465-495. TarRASOFFE, F. J. No date. Notes and sketches on the myology of the sea otter. (Unpublished.) TarasorF, F. J., A. Bisattton, J. Prerarp, AND A. P. Wur1pPt 1972. Locomotory patterns and external morphology of the river otter, sea otter, and harp seal (Mammalia). Canadian Journal of Zoology, vol. 50, no. 7, pp. 915-929. VANDEVERE, J. E. 1969. Feeding behavior of the southern sea otter. Proceedings of the Sixth Annual Conference on Biological Sonar and Diving Mammals, Stanford Research Institute, pp. 87—94.
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 13, pp. 417-427; 2 figs. October 3, 1975
THE MORAY EELS (PISCES: MURAENIDAE) OF THE GALAPAGOS ISLANDS, WITH NEW RECORDS AND SYNONYMIES OF EXTRALIMITAL SPECIES
By John E. McCosker
Steinhart Aquarium, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118
and
Richard H. Rosenblatt
Scripps Institution of Oceanography, University of California, San Diego, La Jolla, California 92037
ABSTRACT: Sixteen species of muraenid eels are recognized from the Galapagos Islands. These are (new records in boldface): Anarchias galapagensis (Seale), Echidna nocturna (Cope), E. zebra (Shaw), Gymnothorax dovii (Giinther), G. buroensis (Bleeker), G. castaneus Jordan and Gilbert, G. panamensis (Stein- dachner), G. pictus (Ahl), Enchelycore octaviana (Myers & Wade), E. lichenosa (Jordan & Snyder), Muraena lentiginosa Jenyns, M. clepsydra Gilbert, M. argus (Steindachner), Uropterygius polystictus Myers & Wade, U. necturus (Jordan & Gilbert), and an undescribed species of Uropterygius. The following new synonymies are proposed: Rabula Jordan & Davis, 1891 = Gymnothorax Bloch, 1795. Rabula davisi Fowler, 1912 = Gymnothorax mordax (Ayres, 1859). Sideria chlevastes Jordan & Gilbert, 1883 = Gymnothorax rueppelliae (McClelland, 1845). Echidna scabra Garman, 1899; E. chionostigma Fowler, 1912; Muraena acutis Seale, 1917; and Rabula rotchit Clark, 1936 = Echidna nocturna (Cope, 1872). Gymnothorax arae Borodin, 1928 =G. dovii (Giinther, 1870). Gymnothorax thomsoni Borodin, 1928 =G. pictus (Ahl, 1870). Murenophis marmoreus Valenciennes, 1855; Muraena aquae-dulcis Cope, 1872; M. insularum Jordan and Davis, 1891; and Lycodontis xanthospilus Fowler, 1944 = Muraena lentiginosa Jenyns, 1842. Muraena albigutta Hildebrand, 1946 = M. argus (Steindachner, 1870).
[417]
418 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
INTRODUCTION
The Galapagos Islands (Archipiélago de Colon) possess a large and diverse marine eel fauna, many species of which are muraenids. The moray fauna of the islands has not been treated as a whole since Jordan and Evermann (1896). We herein provide a listing of the 16 valid species, the first records of seven additional species, and comments on the validity of several Galapagos and ex- tralimital nominal species of morays. A key is provided for the identification of known Galapagos species.
The following list includes those species whose existence at the Galapagos we have verified: Anarchias galapagensis (Seale), Echidna nocturna (Cope), Gymnothorax dovii (Gunther), G. buroensis (Bleeker), G. castaneus Jordan & Gilbert, G. panamensis (Steindachner), G. pictus (Ahl), Enchelycore octaviana (Myers & Wade), E. lichenosa (Jordan & Snyder), Muraena lentiginosa Jenyns, M. clepsydra Gilbert, M. argus (Steindachner), Uropterygius polystictus Myers & Wade, U. necturus (Jordan & Gilbert), and an undescribed species of Uropterygius.
Fowler’s (1938) record of Lycodontis afer from South Seymour (Baltra) Island is probably based on Herre’s (1936, p. 44) listing of Gymnothorax funebris (James E. Bohlke, personal communication, 13 August 1974). We have reidentified Herre’s specimen (SU 24386) as G. panamensis. Other doubtful records and invalid species have recently been treated. Rosenblatt et al. (1972) corrected Herre’s (1936) erroneous records of Indo-Pacific species said to be from the Galapagos. Herre’s listing of Gymnothorax chilospilus and G. undulatus was based on small specimens of Muraena lentiginosa. Randall and McCosker (in press) placed Lycodontis umbra Fowler (1944) in the synonymy of Gymnothorax panamensis (Steindachner ).
ACKNOWLEDGMENTS
We thank the curators and staffs of the following institutions for permission to examine material in their care: Academy of Natural Sciences of Philadelphia (ANSP), California Academy of Sciences (CAS, SU), Museum of Comparative Zoology, Harvard University (MCZ), National Museum of Natural History (USNM), and University of California, Los Angeles (UCLA). Much of the comparative material utilized is housed at the Scripps Institution of Oceanogra- phy (SIO). We also thank Lillian J. Dempster and W. I. Follett for their critical reading of portions of this manuscript, and Kim McCleneghan for allowing us to publish his vertebral data.
Key TO THE GALAPAGOS MURAENIDS
la Dorsal and anal fins developed as skin-covered ridges originating just before caudal 2 1b Dorsal and anal fins skin-covered but distinct, beginning ahead of gill opening and just’ behind ‘anus; respectively 2-= 52-2) 3 ee eee
VoL.
2a
2b
3b 4a
4b
Sa
7b
8a
8b
9a
9b
10a
10b
XL] McCOSKER & ROSENBLATT: MORAY EELS 419
Tip of tail hard and pointed; posterior nostril closely associated with an enlarged interorbital pore, so that posterior nostril appears double; Gulf of California to
Rain arrvaedry clan Gralll ayy cg 0S eee Anarchias galapagensis (Seale) Tip of tail blunt, with a skin-covered caudal; posterior nostril not closely associated Withwanten terror }tall lip OKC sree ae aeeneare es ee ee eee ee Se 3
Teeth blunt, molariform; body color dark brown to black, encircled by numerous, narrow white rings; Indo-west Pacific, oceanic islands of eastern Pacific and Guli @i Calhionmin, tuo Jenimneicne), - Echidna zebra (Shaw)
Teeth pointed; body color mottled but without definite rings — 4
Head and trunk longer than tail; ground color light with dark mottling; posterior nostril tubular, located anterior to center of eye; Gulf of California (SIO 65-330) andeGalapap 0sjsa= eee ee ae Uropterygius polystictus Myers & Wade
Head and trunk shorter than tail; ground color dark with rusty mottlings; posterior nostril with a raised rim, located behind center of eye; Gulf of California to Panama and Galapagos __......___»_ Uropterygius necturus (Jordan & Gilbert)
Teeth blunt, becoming molariform in adults; body color even brown to black with scattered white spots; anterior nostrils orange in life; Gulf of California to
RerukandeGalapagoste.==) ee eee Echidna nocturna (Cope) Teeth pointed at all ages; coloration various, not as above —..- 6 Rosteniorenostials Guo Uae ee Ee oe ee ee ae ee 7 Posterior nostril not tubular, at most with a raised rim —. 9
Dorsal and anal margins white; black spot at corner of mouth not preceded by white; body with three rows of large irregular yellow blotches and many scattered small white spots; teeth on shaft of vomer not depressible; Alijos Rocks, Baja California, to) Reruland Galapagos = Muraena argus (Steindachner)
Dorsal and anal margin not white, with at most a few white blotches; black spot at corner of mouth preceded by a white area on lower jaw; body either mostly plain or spotted, but without large yellow blotches; all vomerine teeth de- DIESSID ewe ss etek Mich Abel Wok one Ae ee EU eee te a eee 8
A large black spot, equal to 2.5 or more eye diameters, around gill opening; young with 5 or 6 series of small hour-glass shaped spots, adults speckled with numerous very small irregular spots on body and fins; Cape San Lucas, Mexico, tomberucand \Galapaross 22 aa ee Muraena clepsydra Gilbert
Black area surrounding gill opening not conspicuous or ringed with white, its diameter equal to 1.5 eye diameters or less; color not as above, either tan with rows of round to oblong yellow or white spots, or dark and reticulated with a few small white spots; Gulf of California to Peru and Galapagos ait a ints A Oey Te ees ee OL oe ies bes eae Ser cee Renae Muraena lentiginosa Jenyns
Lower jaw curved, so that a gap is present and teeth are visible when mouth is
CLOSC Cee ce eet ne dren sla ee al Ak eee oe SS OE ee 10 Lower jaw nearly straight so that there is no gap and the teeth are hidden when ITO Uta 1S CO Se Cy eee ee eee oe Ss Od Ne eee 11
Body coloration uniform brown to grey; pores along upper lip elongate slits with crenulate margins; Gulf of California (SIO 65-336) to Colombia and Galapagos SE Ee, aT aera eee ee Enchelycore octaviana (Myers & Wade)
Body coloration dark brown, head and throat overlain with numerous light spots,
a series of large light blotches along sides; all head pores round; Galapagos and iapany> 2.2 a eee Enchelycore lichenosa (Jordan & Snyder)
420 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
lla Body coloration white, profusely covered with dark brown speckling; vomerine teeth biserial, maxillary teeth uniserial; Indo-west Pacific and oceanic islands
OlmCadstenimebd Clilceeme ss 2 ee eee oe Gymnothorax pictus (Ahl) 11b Body background color various, if white not profusely speckled; vomerine teeth not in two equal rows, maxillary teeth uniserial or biserial 12
12a_ Dorsal origin about midway between occiput and gill opening; head pores ringed in white, a black ring around eye; outer series of teeth thickened, bent abruptly backward at tips, their posterior margins serrate; Gulf of California to Panama; Galapagos and! Jian) Femandez and Easter 1s] sis ee ne Pane ares. ns Oe EAS G8 bind ee Gymnothorax panamensis (Steindachner) 12b_ Dorsal origin notably anne: to occiput than gill opening; head pores not ringed in white, no dark ring around eye; teeth in jaws all conical, straight or evenly curved) jtheir margins smooth) —] eee 13
13a Teeth on maxillary in two rows, the outer row smaller than the inner; five longi- tudinal rows of teeth in the front of upper jaw; body ground color dark brown to black, overlain with a wavy, irregular mottling; Indo-west Pacific and oceanieuslandsron caster)» ach cyan Gymnothorax buroensis (Bleeker) i3b Maxillary teeth uniserial; three longitudinal rows of teeth in the front of upper jaw; body ground color brown to black or green, not overlain with a dark, irregular mottling, 8 fe Se Be ee 14
14a Body ground color dark brown to black, with numerous white spots; Panama to Coloma, ame! Calajyeos Gymnothorax dovii (Ginther) 14b Body ground color brown to Arowniche green, usually plain although sometimes with a few white or yellow flecks, mostly on posterior half and dorsal fin; Gulf of California to Panama and Galapagos —___ Gymnothorax castaneus Jordan & Gilbert
NEw REcorpDS Muraena clepsydra Gilbert.
This species is known from Cape San Lucas, Baja California, Mexico, to Panama. Galapagos specimens have come from Barrington (Santa Fe) Island (SIO 55-259; 970 mm; UCLA 55-314; 3, 82-305 mm.).
Gymnothorax buroensis (Bleeker).
A single specimen (SIO 74-103; 465 mm.) was collected at Darwin Bay, Tower (Genovesa) Island by M. Anctil. Gymnothorax buroensis is known from the Indo-west Pacific and Hawaii, and in the eastern Pacific from Clipperton and Cocos islands and nearshore localities in Costa Rica and Panama (Rosen- blatt et al., 1972).
Echidna zebra (Shaw).
The Galapagos record is based on an observation of this species by Gerard M. Wellington, Charles Birkeland, and Peter Glynn at Tower (Genovesa) Island during January, 1975 (Wellington, personal communication, 31 Jan. 1975). Although the specimen was not collected, its unmistakable appearance would preclude its misidentification.
re : 4 ‘ bo . Ss We eae aos”
Ficure 1. Muraena argus, photographed at Gordon Rocks, Galapagos Islands, in a cave at approximately 45 m. depth, by Carl Roessler.
Muraena argus (Steindachner).
This species is known from Alijos Rocks, Baja California, Mexico to Peru (see discussion under Muraena albigutta, page 426). A Galapagos specimen was collected at James Bay, James (San Salvador) Island (UCLA 67-33, 900 mm.), and we have a photograph from Gordon Rocks (fig. 1).
Uropterygius necturus (Jordan & Gilbert).
This species is known from the Gulf of California to Panama, the Revil- lagigedo and Clipperton islands, and has been taken at several Galapagos localities including Indefatigable (Santa Cruz) Island (UCLA 64-19, 230 mm.), Tower (Genovesa) Island (UCLA 67-35; 9, 174-232 mm.), and Charles (Santa Maria) Island (UCLA 67-43; 3, 220-280 mm.).
Enchelycore lichenosa (Jordan & Snyder).
Aemasia lichenosa was described by Jordan and Snyder (1901) as the type species of a new genus on the basis of Japanese market specimens from Wakanoura (the holotype, SU 6480, now at CAS, 52 cm.) and Misaki (a paratype, USNM 49976, 55 cm.). Except for Randall and McCosker’s (in press) synonymy of Aemasia with Enchelycore, this species, to our knowledge, has been mentioned only in Japanese faunal works. We were somewhat
422 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
surprised then, to discover four adult specimens of a hook-jawed moray that we have identified, by comparison with the type material, as this species. Galapagos specimens are from Punta Espinosa, Narborough (Fernandina) Island (UG@EA 64-3, 70 cm.; UCLA 64-8, 76 cm:; ANSP 117435, 60 en ANSE 109855, 63 cm.). We are unable to separate the Galapagos specimens from the Japanese type material on the basis of morphological characters, coloration, or dentition. The holotype has 148 vertebrae (56 preanal); Galapagos specimens ANSP 109855 and ANSP 117435 have 136 (57 preanal) and 142 (53 preanal) vertebrae, respectively.
The addition of Enchelycore lichenosa to the Galapagos’ fauna represents the ninth Indo-Pacific muraenid species known from the eastern Pacific (see Rosenblatt et al., 1972). It is curious that E. lichenosa has not been taken elsewhere in the Pacific.
IDENTITY OF INVALID GENERA AND SPECIES Rabula Jordan & Davis.
Jordan and Davis (1891, pp. 589-590) erected the subgenus Rabula to include those species of Gymnothorax with a dorsal fin origin beginning over or behind the gill opening. They designated as type-species Muraena aquae-dulcis Cope 1872, on the basis of a specimen (USNM 6673) from San Diego, Califor- nia or “probably .. . from farther south,’ and Cope’s description of the holotype. Fowler (1912) subsequently found their specimen to differ from Cope’s species and, in a footnote which we quote, described it as a new species, Rabula davist:
I may note that Jordan and Davis identify an eel from San Diego, Cal., with Cope’s species, and as they do not explicitly designate Cope’s fish the former must be taken as the type of their genus Rabula. Therefore, the Gymnothorax aquae- dulcis (nec Muraena aquae-dulcis Cope) Jordan and Davis requires a new specific name.
We examined the holotype of R. davisi and found it to be an aberrant specimen of Gymnothorax mordax (Ayres, 1859). The dorsal fin appears to arise in the mid-trunk region; however a radiograph (fig. 2) of the specimen clearly shows that the dorsal fin arises above the 4th vertebra, disappears above the 11th, then reappears above the 61st. The specimen has 145 total vertebrae with 64 before the anal fin, not differing significantly from combined data for 54 specimens of Gymnothorax mordax (K. McCleneghan, personal communica- tion, and C. Clothier, unpublished data) with a range of 143-152 total vertebrae (X = 147.8) and 61—67 preanal (x = 65). The specimen also agrees with G. mordax in its coloration, dentition, and morphometry. Rabula thus becomes a synonym of Gymnothorax Bloch 1795.
Other morays currently referred to Rabula include R. fuscomaculata
Vor. XL] McCOSKER & ROSENBLATT: MORAY EELS 423
Ficure 2. Radiograph of the head and trunk regions of the holotype of Rabula davisi. Arrows indicate dorsal fin origin above the 4th vertebra and reappearance above the 61st vertebra.
Schultz, R. marshallensis Schultz, and R. acuta (Parr). We have examined and radiographed specimens of ‘fuscomaculata’ (CAS 31206) and ‘marshallensis’ (CAS 31205) and, with the exception of the posterior dorsal fin origin, are unable to find characters which would allow their generic separation from Gymnothorax. We are hesitant to recognize them as generically distinct solely on the basis of their fin location.
Sideria chlevastes Jordan & Gilbert.
Examination of the holotype (USNM 20385) of Sideria chlevastes has shown the taxon to be a junior synonym of Gymnothorax rueppelliae (Mc- Clelland), a wide-ranging Indo-Pacific and Hawaiian species. The two common Indo-Pacific species of Gymnothorax with conspicuous broad bands have been called G. ruppeli (an unjustified emendation of rueppelliae) and G. petelli, respectively, by most modern authors. Randall (1973, p. 174) has shown however, that both names apply to the same species, and that the former has priority. Thus ‘petelli’ of authors becomes ‘rueppelliae’, and ‘rupelli’ of most authors is without a name. Moreover, ‘reticularis Bleeker’ is not available, as suggested by Randall, since that represents a misidentification of Gymnothorax reticularis Bloch, a distinctly different species.
Jordan and Gilbert’s (1883) terse description of Sideria chlevastes merely states “obtained at the Galapagos Islands by Captain Herendeen.” A search at the National Museum of Natural History by the senior author for further information concerning the type specimen revealed only that it was entered into the catalog on 13 August 1877, with a listing only of the collector and
424 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. “Galapagos Islands.” No other specimens accompanied it. In that no subse- quent Galapagos specimens have appeared since Herendeen’s, we suspect that the locality was in error. It was not uncommon at that time for whalers such as Herendeen to stop at Hawaii and other islands, often returning with speci- mens to be deposited at the USNM. In that manner, it is not unlikely that a locality error might have originated. The 220 mm. holotype has 135 vertebrae, 51 before the anus.
Rabula rotchii Clark.
Clark’s (1936) description is based on an abnormal specimen of Echidna nocturna (Cope, 1872) with a posterior dorsal insertion. The holotype (CAS 4964) in other proportions, coloration, and dentition fits H. nocturna well.
Echidna chionostigma Fowler.
While preparing this study we examined the type specimens (ANSP 14519 and 14520) of E. chionostigma Fowler, 1912, “probably from the Gulf of California,” and found them also to be referable to Echidna nocturna (Cope, LSi2ye
Echidna scabra Garman.
Garman’s (1899) type specimen (MCZ 28451) of Echidna scabra, from Cocos Island, is a juvenile of Echidna nocturna (Cope, 1872).
Muraena acutis Seale.
Seale’s (1917) type specimen (MCZ 3960) of Muraena acutis, from the Gulf of Panama, is clearly referable to Echidna nocturna (Cope, 1872).
Gymnothorax thomsoni Borodin.
Borodin’s (1928) feckless description but passable illustration of G. thomsoni clearly relates to an injured specimen of Gymnothorax pictus (Ahl, 1789), not G. dovii as suggested by Fowler (1938, p. 251).
Gymnothorax arae Borodin.
Borodin’s (1928) inadequate description of G. arae from Darwin Bay, Tower (Genovesa) Island does mention scattered white dots, indicating that G. arae is a junior synonym of G. dovii (Giinther, 1870).
Lycodontis xanthospilus Fowler.
Our examination of the holotype (ANSP 70026) of L. xanthos pilus Fowler (1944) from James (San Salvador) Island revealed it to be a small specimen of Muraena lentiginosa Jenyns, 1842.
Vor. XL] McCOSKER & ROSENBLATT: MORAY EELS 425
Muraena insularum Jordan & Davis.
A study of an adequate series of Muraena from the Galapagos Islands leads us to the conclusion that M. insularum Jordan & Davis, 1891, described from a Chatham (San Cristobal) Island specimen, also is identical with M. lentiginosa Jenyns, 1842. Two extreme color forms may be segregated in the Galapagos material. One has a color pattern of light brown with darker reticulations which almost form bars. In this dark sort there are a few light spots, but these are restricted to the throat. The dark color pattern has been described and figured by Morrow (1957, p. 16) and described by Jordan and Davis in the original description of M. insularum. Extremes of this kind are purplish brown, with a few scattered white spots on the throat. In the other sort, the ground color is light brown and there are several series of white spots down the sides and on the fins. These spots tend to line up in rows. The spots are surrounded by dark brown areas which tend to coalesce to form an interlocking network. There is, however, a complete range of variation between these sorts. Further, Jenyns’ description of M. lentiginosa seems to have been based on a specimen of the dark sort.
The difference mentioned by Jordan and Davis concerning the number of rows of teeth in the upper jaw is invalid, since in all the known eastern Pacific species of Muraena the inner row of enlarged teeth is lost with increasing size.
It is of interest that such variability in coloration is not found in mainland populations. All of a large number of mainland specimens are much like the light island types in coloration, except that the dark brown areas surrounding the light spots are less extensive and never coalesce. However, all of our speci- mens are from Panama north, and Morrow (1957) has reported a specimen of the dark type from Peru. This seems to indicate that the Peruvian and Gala- pagos populations are genetically different from the northern populations. It is, however, difficult to evaluate differences in variability, and we do not feel that specific or even subspecific differentiation is indicated.
Muraena aquae-dulcis Cope.
Our examination of the Costa Rican type specimen (ANSP 14925) of M. aquae-dulcis Cope, 1872 indicates that it is a small and damaged specimen of M. lentiginosa Jenyns, 1842.
Murenophis marmoreus Valenciennes.
Although no specimens of this species have been reported since its descrip- tion (Valenciennes, 1846), the name continues to appear as a species of Gymno- thorax (Jordan & Davis, 1891) or Rabula (Jordan & Evermann, 1896; Fowler, 1938). Jordan and Evermann considered it to be “a doubtful species, perhaps based on Muraena lentiginosa.’ The illustration of the holotype leaves little doubt that this surmise is correct. The coloration (see discussion under Muraena
426 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
insularum) is diagnostic. The indication in the figure of a dorsal origin behind the gill opening is likely an error and Valencienne’s description of the posterior nostril as a slit beneath the eye applies to no known muraenid.
Muraena albigutta Hildebrand.
Hildebrand (1946), in describing Muraena albigutta from Peru, was ap- parently unaware of Steindachner’s (1870) description of Muraena argus from Altata, Sinaloa, Mexico. Our material from the Gulf of California (SIO 60-1; UCLA 56-68) agrees well with Steindachner’s description. We have compared the holotype of M. albigutta (USNM 127840) with a specimen from Guaymas, Sonora, Mexico. They agree, particularly in the distinctive color- ation and in that the teeth on the shaft of the vomer are not depressible. Muraena argus is now known from Alijos Rocks, Baja California, Mexico (SIO 74-104) ; the Gulf of California; Isla La Plata, Ecuador (UCLA 53-419); Lobos de Afuera, Peru; and the Galapagos.
LITERATURE CITED
Boropin, N. A.
1928. Scientific results of the yacht “Ara” expedition during the years 1926 to 1928, while in command of William K. Vanderbilt. Fishes. Bulletin of the Vanderbilt Oceanographic Museum, vol. 1, art. 1, 37 pp.
CrarkK, H. WALTON
1936. The Templeton Crocker expedition of the California Academy of Sciences, 1932. No. 29. New and noteworthy fishes. Proceedings of the California Academy of Sciences, 4th ser., vol. 21, no. 29, pp. 383-396.
Cope, Epwarp D.
1872. Report on the Recent reptiles and fishes of the Survey, collected by Campbell Carrington and C. M. Dawes. Pages 467-476 in F. V. Hayden, Preliminary Report of the United States Geological Survey of Montana and Portions of Adjacent Territories; being a Fifth Annual Report of Progress, part 4.
Fow ter, Henry W.
1912. Descriptions of nine new eels, with notes on other species. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 64, pp. 8-33.
1938. The fishes of the George Vanderbilt South Pacific Expedition, 1937. Monographs of the Academy of Natural Sciences of Philadelphia, no. 2, 349 pp.
1944. Results of the Fifth George Vanderbilt Expedition (1941). Monographs of the Academy of Natural Sciences of Philadelphia, no. 6, pp. 57-529.
GARMAN, SAMUEL
1899. Reports on an exploration of the west coast of Mexico, Central and South Amer- ica, and off the Galapagos islands, in charge of Alexander Agassiz, by the U.S. fish commission steamer “Albatross,” during 1891. The fishes. Memoirs of the Museum of Comparative Zoology, Harvard College, vol. 24, 431 pp.
Herre, A. W. C. T.
1936. Fishes of the Crane Pacific Expedition, Field Museum of Natural History,
publication no. 353, Zoological Series, vol. 21, 472 pp. HILDEBRAND, SAMUEL F.
1946. A descriptive catalog of the shore fishes of Peru. Bulletin of the United States
National Museum, no. 189, 530 pp.
Vor. XL] McCOSKER & ROSENBLATT: MORAY EELS 427
JorpAN, Davip Starr, AND BrApLeEY Moore Davis 1891. A preliminary review of the apodal fishes or eels inhabiting the waters of America and Europe. Report of the United States Commission of Fish and Fisheries for 1888, part 16, pp. 581-677. (Separate issued 1891.) Jorpan, Davin Starr, AND BARTON W. EVERMANN 1896. The fishes of North and Middle America: A descriptive catalogue . . . Panama. Bulletin of the United States National Museum, no. 47, part 1, 1240 pp. JorpDAN, Davip STARR, AND CHARLES H. GILBERT 1883. Description of a new muraenoid eel from the Galapagos Islands. Proceedings of the United States National Museum, vol. 6, pp. 208-210. JorDAN, Davip STARR, AND JOHN O. SNYDER 1901. A review of the apodal fishes or eels of Japan, with descriptions of 19 new species. Proceedings of the United States National Museum, vol. 23, no. 1239, pp. 837-890. Morrow, JAMEs E. 1957. Shore and pelagic fishes from Peru, with new records and the description of a new species of Sphoeroides. Bulletin of the Bingham Oceanographic Collection, vol. 16, art. 2, pp. 5-55. RANDALL, JOHN E. 1973. Tahitian fish names and a preliminary checklist of the fishes of the Society Islands. Occasional Papers of Bernice P. Bishop Museum, vol. 24, no. 11, pp. 167-214. RANDALL, JOHN E., anD JoHN E. McCoskeEr In press. The eels of Easter Island with the description of a new moray. Los Angeles County Museum Contributions in Science. ROSENBLATT, RicHarp H., JouHn E. McCosker, AND I. RUBINOFF 1972. Indo-west Pacific fishes from the Gulf of Chiriqui, Panama. Los Angeles County Museum Contributions in Science, no. 234, 18 pp. SEALE, ALVIN 1917. New species of apodal fishes. Bulletin of the Museum of Comparative Zoology at Harvard College, vol. 61, no. 4, pp. 79-94. STEINDACHNER, FRANZ 1870. Ichthyologische Notizen (X.). Sitzungsberichte der K. Akademie der Wissen- schaften, Wien, Mathematisch-Naturwissenschaiftliche Classe, Band 61, Heft 5, Abth. 1, pp. 623-642. 1846. Poissons, plates 1-10. Zn Abel du Petit-Thouars, Voyage autour du monde sur la frégate la Vénus, pendant les annees 1836-1839. Atlas de Zoologie, Paris.
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 14, pp. 429-437. October 3, 1975
THE TAXONOMIC STATUS OF THE SOUTHERN SEA OTTER
By John Davis and W. Z. Lidicker, Jr. Museum of Vertebrate Zoology, University of California, Berkeley 94720
Apstract: A recent review of the taxonomy of the sea otter (Enhydra lutris) by Roest (1973) concluded that all populations of this species extending from the Commander Islands to southern California should be considered to constitute a single subspecies (E. |. lutris). At present a 1700 mile gap (between Alaska and California) exists within this vast range. We re-examined Roest’s data (and the available literature) for evidence which might suggest alternative taxonomic conclusions. In our opinion, the known facts strongly support the conclusion that the southern sea otter should continue to be recognized as a separate subspecies (E. |. nereis). Marked morphological and behavioral differences seem to exist, and at the present time complete genetic isolation as well. If a northwest-southeast cline in morphology existed formerly, the evidence suggests that there was a pronounced step or change in its slope in southwestern Alaska.
INTRODUCTION
Merriam (1904) described the Californian population of the sea otter (Enhydra lutris) as Latax lutris nereis on the basis of morphological differences between a single skull from San Miguel Island, Santa Barbara County, and skulls of typical E. l. lutris from the Aleutian Islands, Alaska. The validity of £. J. nereis has been variously accepted or questioned by subsequent workers (e. g., Grinnell, 1933; Grinnell, Dixon, and Linsdale, 1937; Scheffer and Wilke, 1950; Hall and Kelson, 1959; Kenyon, 1969). Until recently, lack of adequate material has been a severe handicap to critical evaluation of the taxonomic status of the Californian population. However, Roest (1971) compared large samples of adult skulls of known sex from Alaska (mainly from the Aleutians) and California. On the basis of skull differences and differences between the two populations in
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430 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
weight and total length, he recognized EL. /. nereis as valid and as the proper subspecific designation for the southern sea otter. Subsequently additional skulls, from southwestern Alaska, became available to him. These specimens were inter- mediate in characters between those from the Aleutians and California and he proposed (Roest, 1973) that these characters varied clinally. On that basis he synonymized E. /. nereis and included all of the sea otters from the Commander Islands to California in £. /. lutris.
On the basis of Roest (1973), Miller (1974, p. 2), in an official publication of the California Department of Fish and Game, stated: “Now that E. l. nereis is no longer valid, the question of sea otters in California being rare or endangered is void inasmuch as there are over 120,000 sea otters now extant in Alaskan waters, and over 1,600 in California” (italics ours). In other words, according to Miller, the demographic status of the sea otter in California had been decided by the taxonomy proposed by Roest (1973).
Because of the possible importance of the taxonomy of the California sea otter to this animal’s rare and endangered status, and to future management decisions, and because of the inherently challenging taxonomic questions involved, we re-examined Roest’s data for evidence which might suggest alternative taxonomic conclusions to those chosen by him (1973). We found that there are, in fact, several alternative interpretations of the available data on geographic variation in the sea otter. Moreover, Roest’s interpretation does not seem to fit the known facts as well as do other explanations. In the following discussion of geographic variation in Enhydra lutris, the subspecies FE. 1. gracilis of the Kurile Islands and the southern tip of Kamchatka is not reviewed critically.
DISCUSSION
Roest (1971) assembled 214 adult sea otter skulls from Alaska, mainly from the Aleutian Islands, and 50 adult skulls from California. He took 24 skull measurements and, on the basis of stepwise discriminant analysis, selected four characters as being of taxonomic significance. Using a combination of these characters applied to skulls of known sex only, he designated each of 156 Alaskan and 40 Californian skulls as “Alaskan” or “Californian,” with the following results:
Correctly Incorrectly Not Origin N identified identified referable Alaska 156 141 (90.5%) 2 (1.3%) 13 (8.3%) California 40 38 (95.0%) 0 2. (50723)
On the basis of the high degree of separability of Alaskan from Californian skulls, supplemented by differences in total length and in weight between the two populations, Roest recognized E. /. nereis as a valid subspecies and applied this name to the southern sea otter.
Vor. XL] DAVIS & LIDICKER: SEA OTTER 431
In 1972, skulls from southwestern Alaska (the Alaska Peninsula, southern Alaska, and Prince William Sound) became available and proved to be inter- mediate between those from the Aleutians and California (Roest, 1973). On the basis of this sequential shift in skull morphology from west to southeast, Roest proposed that variation in skull characters was clinal and that all of the sea otters from the Commander Islands to California pertain to a single race, E. l. lutris. From this viewpoint E. /. nereis was relegated to the synonymy of E. 1. lutris although Roest (1973, pp. 8, 14) noted that the Californian population can be distinguished from the Aleutian population, but not from the population of southwestern Alaska.
There are, however, alternative interpretations of the variation in skull characters from the Aleutians to California. One of these has been suggested to us by Dr. Carl L. Hubbs (pers. comm.). Noting that Roest did not examine skulls from the area between southwestern Alaska and California, he points out that we do not therefore know the nature of variation over this distance of 1700 nautical miles (all distances given are great circle distances unless otherwise specified). Hubbs suggests that variation in skull characters may not have been gradually changing in the way suggested by Roest (1973), but they “may have been fairly constant throughout most of this vast distance; if so, we would have two well marked subspecies with a band of intergradation in southwestern Alaska.”
Actually, Roest’s own data suggest this. Of ten specimens from near Umnak Island and along the north shore of the Alaska Peninsula, “3 specimens show primarily Alaskan features, 6 are intermediate, and 1 could be considered Californian ....,’ whereas of 16 specimens from Prince William Sound, he considered two Alaskan, two intermediate, and 12 Californian (Roest, 1973, pp. 8-9). These distributions differ significantly (x7 = 10.69; P<.01). This pronounced shift toward the characters of the southern population, so far north in the sea otter’s range, suggests that a population with Californian skull char- acters may well have extended far north into British Columbian waters. This would be quite compatible with Hubbs’ suggestion of two well-marked subspecies intergrading in southwestern Alaska rather than clinal variation occurring within a single, widely distributed population.
If variation in skull characters was indeed clinal, there must have been a pronounced step or shift in slope steepness between the Alaska Peninsula and Prince William Sound. The great majority of specimens from Amchitka and Adak are of the Alaskan skull type. Between Adak and Port Heiden, the north- easternmost locality on the Alaskan Peninsula from which Roest had specimens, skull type has shifted from Alaskan to predominantly intermediate over a distance of 695 nautical miles. Between Port Heiden and Prince William Sound skull type has shifted from intermediate to predominantly Californian over a distance of 435 nautical miles. If the samples previously discussed indicate
432 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
accurately the morphologic variation in the populations from which they were drawn, the slope of the cline over the 1700 nautical miles from Prince William Sound south to Monterey Bay must have been extremely gentle compared to that from Prince William Sound to Adak.
As regards weight differences between the northern and southern populations, the data presented by Roest (1973, p. 2) suggest that Californian males weigh considerably less than both Aleutian and southwestern Alaskan males; the same is true of females but the difference is less. However, some critical samples are small; further, most of his Alaskan specimens were weighed when fresh and most Californian specimens were weighed some time after death. Additional data on body weights are available from the literature (see summary in Harris, 1968; Kenyon, 1969; Wild and Ames, 1974). However, these do not permit critical geo- graphic comparisons because it is not possible to compare comparable age groups. Nevertheless, they suggest in general that northern otters average heavier and reach a larger maximum weight (100 pounds for males, Kenyon, 1969, p. 21), thus supporting the trend suggested by Roest.
Data on total length (Roest, 1973, p. 7) indicate that males from California are considerably smaller than males from both southwestern Alaska and the Aleutians whereas females from both California and southwestern Alaska are smaller than those from the Aleutians. Unfortunately, total lengths of Alaskan specimens are curvilinear and those of Californian specimens are standard lengths (a fact not mentioned by Roest [1971] although means, ranges, and sample sizes indicate that the same samples were used in both papers). Thus, the generally larger size of Alaskan specimens may be accentuated by the difference in measurement techniques.
Student’s ¢ tests applied to Roest’s 1973 data indicate significant differences in mean total length (P < .001) between males from California and males from both Adak and southwestern Alaska. However, males from the last two localities do not differ significantly. The mean length of females from Adak differs significantly from that of females from southwestern Alaska (P < .01) and Cali- fornia (P < .001), but females from California and southwestern Alaska do not differ significantly.
As regards color, “Alaskan otters are...dark, but those from California are most commonly medium brown in color” (Roest, 1973). It is not possible to estimate the degree of separability of the Alaskan and Californian populations on the basis of pelage color from this information, but apparently most southern sea otters can be distinguished by their color. Data summarized by Harris (1968, pp. 263-264) also support this conclusion.
To summarize the morphological data, the sea otter populations of the Aleutians and California can be differentiated at a level of better than 90% on the basis of skull characters. Aleutian otters are also larger and darker than those from California. Between these populations there is a population in southwestern
VoL. XL] DAVIS & LIDICKER: SEA OTTER 433
Alaska with intermediate skull characters; the males are larger than those from California but the females are similar in size. It is impossible, on the basis of available material, to determine with confidence whether the population in southwestern Alaska represents a point on a long west-southeast cline or whether it is a population intergradient between well-marked northern and southern sub- species. If, however, this population is on a cline, there must be a pronounced step or change in slope steepness between the Alaska Peninsula and Prince William Sound.
In addition to morphological differences, there are also behavioral differences between northern and southern sea otters. The use of hard objects placed on the chest as anvils for breaking open hard-shelled food items is rare in the otters of the Commander Islands and Amchitka, but is common in the otters of California. Since the diet of Alaskan otters includes a much higher proportion of fish than does the diet of southern otters, there may be simply less need for tool-using in the northern part of the species’ range.
Hauling out on land is much more frequent in the northern population. In apparent association with this increased use of land is more efficient terrestrial locomotion in Alaskan otters. Kenyon (1969) states that northern otters walking unhurriedly move with a rolling gait, raising one foot at a time. When animals are startled ‘they arch the back and bound or hop, moving both forefeet then both hind feet forward in rapid succession. Speed of movement is somewhat less than the running speed of a normally agile man.” In contrast, Californian otters do not arch the back and do not move their appendages alternately. Rather, “they either drag their hind quarters or repeatedly pivot their bodies up and forward with the help of their feet” (J. Vandevere, paper presented at AAAS meeting, San Francisco, California, 27 February 1974). These differences suggest possible genetic differences in both behavior and anatomy between northern and southern sea otters. In this regard it is of considerable interest that the Asian sea otter (E. l. gracilis) is also relatively clumsy on land (Lydekker, 1895; Barabash-Nikiforov, 1947).
The ability of Alaskan sea otters to move relatively long distances out of water has been documented by Schneider and Faro (1975). In 1971 a number of otters were trapped by rapidly forming pack ice along the north shore of the Alaska Peninsula, and a number died because they could find no open leads in which to forage. Two dead sea otters were found 8 km. inland from Port Heiden Bay and one dead otter was seen “15 km inland, half way to Pavlof Bay on the Pacific Ocean.” These animals may have been trying to reach the Pacific Ocean in an attempt to find open water where foraging was possible.
Vandevere (1970) described a number of differences in reproductive behavior between northern and southern sea otters. Most striking of these were differences in the posture of the female during copulation. The sequence of events leading to copulation also differed, the nose or side of the face of the female being grasped
434 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.
by the northern male rather late in precopulatory struggling whereas this occurs early in this stage of copulation in southern otters.
The present distribution of the southern sea otter in relation to its nearest conspecifics is also of critical importance in assessing the distinctiveness of this population. As of early December 1974, the northernmost established Californian sea otters were located off Sunset State Beach, Santa Cruz County, about 15 airline miles south-southeast of the city of Santa Cruz. A few individuals have been seen north of this point but none has stayed long enough to suggest estab- lishment. The nearest naturally occurring northern sea otters are in Prince William Sound, 1700 nautical miles to the north, although Pedersen and Stout (1963) reported the sighting of what was probably a single otter off Neahkahnie, northern Oregon, between August 1961 and February 1962. A small introduced population is located near Port Orford, Oregon, 365 nautical miles to the north, where a number of otters from Amchitka were released in 1970 and 1971; about 23 are still present.
At this time, the southern sea otter is well removed from its nearest conspe- cifics and there is no exchange of genetic material between this population and any other. Since 1914, the sea otter has extended its range north in California from Point Sur, Monterey County (Wild and Ames, 1974, p. 23, fig. 5) to Sunset State Beach, a great circle distance of only 35 nautical miles in 60 years. The isolation of the Californian population is thus likely to continue for many years if the present natural situation persists.
Although complete disruption of gene flow between Californian and other sea otters may be relatively recent, resulting from the decimation of the species by fur traders in the last two centuries, the severe reduction of the southern popu- lation may well have had profound effects on its genetic constitution. This is strongly suggested by the findings of Bonnell and Selander (1974) on the northern elephant seal (Mirounga angustirostris). Electrophoretic analysis of blood samples from five isolated breeding colonies in California and Mexico indicated no polymorphisms among 21 proteins encoded by 24 loci. The authors concluded that this homozygosity may have resulted from fixation of alleles brought about by the decimation of the species by sealers in the 19th century. Although comparable evidence is not available for the sea otter, it seems highly likely that the severe reduction in numbers suffered by the southern population, followed by strong geographic isolation, has had some effect on that population’s genetic constitution.
CONCLUSIONS
In the application of formal scientific nomenclature to naturally occurring populations, the taxonomist frequently faces the problems inherent in trying to fit a myriad of biological situations into the rigid framework of a fixed nomen-
Vor. XL] DAVIS & LIDICKER: SEA OTTER 435
clatural system. The best that one can do is to use that nomenclature which best expresses the natural situation with which he is dealing.
In the case of the sea otter, at least two interpretations may be made of the morphological variation observed in the populations between the Commander Islands and Monterey Bay. If we regard this variation as clinal, then the nature of the cline must influence our choice of nomenclature. Three characteristics of this presumed cline are of particular importance: 1) the terminal populations are separable at a level of at least 90% on the basis of skull morphology; 2) there is either a step in the cline, or a pronounced change in slope steepness, between the Alaska Peninsula and Prince William Sound; 3) the population at the southern terminus has been much reduced and has been completely isolated for at least 100 years, with a strong possibility that its genetic constitution has under- gone significant change.
Given these characteristics, whether one includes the entire cline in a single subspecies or treats the terminal populations as distinct subspecies becomes a matter of professional judgement. If we are dealing with a case of primary intergradation in which a gradual cline along western North America sharply changes slope in southern Alaska, it may still be appropriate scientifically to recognize the northern and southern sea otters as distinct subspecies. There are many instances in vertebrate taxonomy in which the terminal populations of a cline are regarded as distinct subspecies if they are distinctly separable in a variety of traits, and/or if they are separated by a pronounced step in the cline. If, in fact, variation is actually clinal in this way, it is just as proper taxonom- ically to regard the northern and southern sea otters as distinct subspecies as it is to combine them in E. /. lutris.
However, it is by no means clear that we are dealing with a case of primary intergradation. It is equally possible that there are two distinct subspecies which in recent history intergraded (secondarily) in southwestern Alaska and, indeed, this is strongly suggested by the nature of the variation described by Roest in the samples available to him. Moreover, some taxonomists have advocated that subspecies designations be applied only to portions of species which are not only genetically distinct, but which are at least partially independent (separately evolving) evolutionary units (e.g., Lidicker, 1962). The southern sea otter clearly meets even these stringent criteria by virtue of its genetic isolation, its morphological and behavioral differentiation, and its recent derivation from what must have been an extremely small remnant population. Further, there are impor- tant differences between the habitats occupied by the northern and southern sea otters. Formation of pack ice as described by Schneider and Faro (1975) never occurs within the range of the southern population. Water temperatures are much lower in the north. Daily sea-water surface temperatures at Scotch Cap in the Aleutians over a five-year period ranged from —1.39° C to 12.22° C, and at Monterey, California, over a similar period, they ranged from 10.1° C to 16.7° C
436 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
(Ricketts and Calvin, 1948, p. 263, fig. 117). Many other habitat differences undoubtedly occur as well. All of these factors imply that the two otter popu- lations are subjected to quite different selective regimes.
Thus, whether one views the evidence as indicating that the southern sea otter population is part of a stepped cline, a remnant of two historically well marked subspecies intergrading in southwestern Alaska, or a recently differentiated and unique evolutionary unit, its formal designation as a subspecies is justified on scientific grounds. Given that such designation is scientifically sound, one may still ask what are the advantages and disadvantages to advocating this nomen- clatural arrangement.
There seems to be no particular advantage to including these populations in one subspecies, but there are a number of disadvantages to doing so. This course of action would conceal the high degree of separability between the southern and northern populations; it would conceal the pronounced step or shift in slope in the indicated cline in southern Alaska; it would fail to consider certain behavioral differences, and possible related anatomical differences, between the northern and southern populations; and it would give no idea of the present geographic isolation of the southern population. To us, formal recognition of the southern population as a distinct subspecies, to which the already available name E. J. nereis may be applied, much more accurately reflects the existing natural situ- ation than would any other nomenclatural arrangement. Furthermore, it is an arrangement which is consistent with recognition of the Asian subspecies E. J. gracilis (cf. Barabash-Nikiforov, 1947; Roest, 1973), a form which appears to be of comparable distinctiveness to EF. /. nereis (differing in size, cranial features, pelage color, and behavior), but much less isolated by distance. We therefore propose that three subspecies continue to be recognized for this unique and important mammal.
ACKNOWLEDGMENTS
We would like to thank N. K. Johnson, R. E. Jones, J. L. Patton, and O. P. Pearson for critically reviewing this manuscript.
LITERATURE CITED
BaARABASH-NIKTFOROY, I. I. 1947. The sea otter. Translated from the Russian by Israel Program for Scientific Translations, Jerusalem, 1962, 227 pp. BonnELL, Micuarrt L., AND Ropert K. SELANDER 1974. Elephant seals: genetic variation and near extinction. Science, vol. 184, pp. 908-909. GRINNELL, JOSEPH 1933. Review of the recent mammal fauna of California. University of California Publications in Zoology, vol. 40, pp. 71-234. GRINNELL, JOSEPH, JOSEPH S. Dixon, AND JEAN M. LINSDALE 1937. Fur-bearing mammals of California. Vol. 1. University of California Press (Berkeley), xii + 375 pp.
Vor. XL] DAVIS & LIDICKER: SEA OTTER 437
Hatt, E. RayMonp, AND KeiTH R. KELSON 1959. The mammals of North America. Vol. 2. The Ronald Press Company (New York), viii + pp. 547-1083, + index, 79 pp. Harris, C. J. 1968. Otters/A study of the Recent Lutrinae. Weidenfeld and Nicolson (London), 397 pp. KENYON, Kari W. 1969. The sea otter in the eastern Pacific Ocean. North American Fauna no. 68, xili + 352 pp. LipickEr, WILLIAM Z., JR. 1962. The nature of subspecies boundaries in a desert rodent and its implications for subspecies taxonomy. Systematic Zoology, vol. 11, pp. 160-171. LYDEKKER, RICHARD 1895. Notes on the structure and habits of the sea-otter (Latax lutris). Proceedings of the Zoological Society of London, 1895, pp. 421-423. MeErRIAM, C. Hart 1904. A new sea otter from southern California. Proceedings of the Biological Society of Washington, vol. 17, pp. 159-160. MIcter, DaNniet J. 1974. The sea otter Enhydra lutris it’s [sic] life history, taxonomic status, and some ecological relationships. State of California, Department of Fish and Game, Marine Resources Leaflet No. 7, 13 pp. PEDERSEN, RICHARD J., AND JACK STOUT 1963. Oregon sea otter sighting. Journal of Mammalogy, vol. 44, p. 415. RICKETTS, EDWARD F., AND JACK CALVIN 1948. Between Pacific tides. Revised edition. Stanford University Press (Stanford, California), xxvii + 365 pp. Roest, ArYANn I. 1971. A systematic study of the sea otter (Enhydra lutris). Proceedings of the Eighth Annual Conference on Biological Sonar and Diving Mammals, Menlo Park, California, pp. 133-135. 1973. Subspecies of the sea otter, Enhydra lutris. Contributions in Science, Natural History Museum, Los Angeles County, no. 252, 17 pp. SCHEFFER, VICTOR B., AND Forp WILKE 1950. Validity of the subspecies Enhydra lutris nereis, the southern sea otter. Journal of the Washington Academy of Sciences, vol. 40, pp. 269-272. SCHNEIDER, Kart B., AND JAMES B. Faro 1975. Effects of sea ice on sea otters (Enhydra lutris). Journal of Mammalogy, vol. 56, pp. 91-101. VANDEVERE, J UDSON B. 1970. Reproduction in the southern sea otter. Proceedings of the Seventh Annual Conference on Biological Sonar and Diving Mammals, Menlo Park, California, pp. 221-227. WILD, PAuL W., AND JAcK A. AMES 1974. A report on the sea otter, Enhydra lutris L., in California. California Department of Fish and Game, Marine Resources Technical Report no. 20, 93 pp.
PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES
Vol. XL, No. 15, pp. 439-447; 2 figs. October 3, 1975
HERALDIA NOCTURNA, A NEW GENUS AND SPECIES OF PIPEFISH (FAMILY SYNGNATH- IDAE) FROM EASTERN AUSTRALIA, WITH COMMENTS ON MAROUBRA PERSERRATA WHITLEY
by) John R. Paxton Department of Ichthyology, The Australian Museum, Sydney, Australia
Apstract: Heraldia nocturna is described as a new genus and species in the fish family Syngnathidae. The type locality is Sydney Harbour in eastern Australia. The species occurs on rocky reefs and is nocturnally active. The male has a brood area on the belly that lacks pouch folds or plates; the new genus is related to the Doryrhamphus complex of spiny, belly-pouch pipefishes. Variation in another re- lated Sydney area pipefish, Maroubra perserrata, previously known from a single specimen is described.
INTRODUCTION
In the course of a survey of the fishes of Sydney Harbour, a number of small pipefishes were collected that resembled Doryrhamphus, but differed in the trunk and tail ridges, brood pouch, and head ornamentation. The capture of specimens with quinaldine and hand net allowed aquarium observations on the habits of the new pipefishes.
Australia is richly endowed with pipefishes and seahorses; Munro (1958) lists 71 Australian species in the family Syngnathidae, including such distinc- tive forms as the endemic sea dragons Phyllopteryx and Phycodurus. Scuba collecting in both tropical and temperate waters is adding to the known fauna.
METHODS Methods for making counts and measurements follow Herald (1940). The first trunk ring is twice as long as the other rings and although actually
[439]
440 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr.
double, is counted as a single ring (Herald and Randall, 1972). Four speci- mens were maintained in aquaria for varying periods up to two months. Observations on swimming behavior were made during light and dark periods. Abbreviations of measurements are as follows: SL—standard length; TL— total length; HL—head length.
Institutional abbreviations are as follows: AMS—Australian Museum, Sydney; BMNH—British Museum Natural History, London; BPBM—Bernice P. Bishop Museum, Honolulu; CAS—California Academy of Sciences, San Francisco; QM—Queensland Museum, Brisbane; WAM—Western Australian Museum, Perth.
ACKNOWLEDGMENTS
Rudi Kuiter, diver aquarist from Sydney, collected the majority of speci- mens of the new genus and species. His sure hand with the dipnet and knowl- edgeable observations are much appreciated. Clarrie Lawler, Barry Gold- man, and Paul Zorn each collected a specimen. Helen Larson drew the figures and Bronwyn Wright typed the manuscript. C. E. Dawson and R. A. Fritzsche improved the paper with their critical comments.
Vol. XL] PAXTON: NEW PIPEFISH 441
Heraldia Paxton, new genus
Dracnosis. Very stout, Doryrhamphus-like pipefish with abdominal brood area lacking lateral membranous folds or protecting plates. Lateral trunk ridge continuous with lateral tail ridge, which in turn becomes superior tail ridge; inferior trunk and tail ridges continuous and superior trunk ridge ending free on Ist, 2nd, or 3rd tail ring. Snout without spines in either sex; head intricately sculptured; body ridges pronounced—those of first 2—4 trunk rings with rounded edges but thereafter with points at posterior edge of each ring. Base of dorsal fin very dark; caudal fin large and deeply emarginate with individual rays extending beyond web. Trunk rings 16-17; tail rings 13-15; dorsal rays 23-26 located on 512-7 rings, of which 3%-4% are trunk rings and 2-3 are tail rings. Pectoral rays 19-22; anal 4; caudal 10-11. Maximum length 74.8 mm SL, 84.2 mm TL.
Heraldia is related to the other pipefish genera with an abdominal brood area and small spines on the posterior ends of the body ring ridges. Heraldia differs from Doryrhamphus and Dentirostrum in lacking brood pouch flaps, from Dunckerocampus in its shorter snout and fewer tail rings, and from Oostethus and Maroubra in having the dorsal fin mostly on the trunk.
The type species of Heraldia, H. nocturna, is new and the only known member of the genus. The first two specimens were sent to Dr. Earl Herald, who recognized them as a new genus and provided a brief description and diagnosis before his untimely death. His help and encouragement are grate- fully acknowledged. Heraldia is named in honor of Earl Stannard Herald, whose contributions to syngnathid systematics and help to other ichthyologists were considerable.
Heraldia nocturna Paxton, new species. (Figures 1-2).
MATERIAL EXAMINED. All 14 type specimens were collected with scuba from rocky reefs at depths from 2-15 meters in Sydney Harbour, New South Wales, Australia (33° 50’S., 151° 15’E.), or Seal Rocks, N.S.W. about 190 kilometers northeast of Sydney (32° 25’S., 152° 35 E.).
Hototype. AMS I.17328-001, male, 74.8 mm. SL (84.2 mm. TL), Village Point, Watson’s Bay, Sydney Harbour, Australia, 3 m., quinaldine, R. Kuiter, 8 September 1973.
Paratypes. AMS 1.17328-002, female?, 69.5 mm. SL, collected with the holotype; AMS 1.16516-001, 64.0 mm. SL, Green Point, Sydney Harbour, 15 m., rotenone, B. Goldman, 23 Sept. 1972; AMS 1.17033—008, 67.6 mm. SL, off Store Beach, Sydney Harbour, 1-7 m., rotenone, G. Allen, D. Hoese, J. Paxton, D. Pollard, 6 April 1972; AMS 1.17112-003, 60.6 mm. SL, Seal Rocks, 10 m., quinaldine, R. Kuiter, 29 April 1973; AMS 1.17641—001, 65.5
442 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr.
mm. SL, Seal Rocks, 10 m., quinaldine, R. Kuiter, 5 May 1973; AMS I. 17642-001, 40.6 mm. SL, Green Point, Sydney Harbour, 2 m., quinaldine, R. Kuiter, 31 Dec. 1973; AMS 1.18075-001, 33.4 mm. SL, Watson’s Bay, Sydney Harbour, 5 m., quinaldine, P. Zorn, 3 Nov. 1974; BMNH 1975. 2.19.1, 60.9 mm. SL, Parsley Bay, Sydney Harbour, quinaldine, R. Kuiter, 26 Dec. 1973; BPBM 18034, 66.5 mm. SL, Seal Rocks, 12 m., quinaldine, R. Kuiter, 27 Oct. 1974; CAS 32366, 66.4 mm. SL, Camp Cove, Sydney Harbour, 5 m., hand net, C. Lawler, 13 April 1969; CAS 32367, male, 62.7 mm. SL, Green Point, Sydney Harbour, quinaldine, R. Kuiter, 10 Aug. 1974; QM 1.12920, 47.8 mm. SL, Camp Cove, Sydney Harbour, 15 m., quinaldine, R. Kuiter, 6 Jan. 1974; WAM P.25167-001, 61.0 mm. SL, Watson’s Bay, Sydney Harbour, quinaldine, R. Kuiter, 3 Aug. 1974.
Diacnosts. Counts and characteristics as given under Heraldia above; total rings 30-32; total rings plus dorsal fin rays 54-57; head in standard length 4.95.5: snout in head 2.3—2.7; dorsal fin base in head 1.4—1.6.
DerscrIPTION. The measurements of the holotype (74.8 mm. SL) in mm. are followed in parentheses by the range as percent SL for the 16 types. Head length 13.7 mm. (18.2—20.4%), snout length 5.8 mm. (5.9-8.7%), eye diameter 2.9 mm. (3.7-4.0%), dorsal fin base length 8.8 mm. (11.8-13.8%), pectoral fin length 2.8 mm. (3.4—4.4%), anal fin length 2.8 mm. (3.0-5.7%), caudal fin length 10.6 mm. (12.2—17.6%).
The head has a remarkable scrollwork of tiny ridges, some of which are not a part of the typical pipefish head ridge pattern; the ridges vary in number and development with individual specimens. The median snout ridge is with- out spines and is elevated slightly at the nostril area in front of the eyes; between the eyes it divides into two ridges, and is paralleled by at least four additional smaller ridges. The sides of the snout have a series of parallel ridges, two of which are more pronounced, those on the lateral median and on the lateral inferior surface. The supraorbital crest extends backward to the end of the eye, then divides into a series of radiating ridges. The principal opercular ridge extends diagonally completely across the upper portion of the opercle; radiating outward from it are 2-12 small ridges on the upper side
Vol. XL] PAXTON: NEW PIPEFISH 443
and 13-18 on the lower side. The supraopercular ridge has two sections, each with smaller radiating ridges present or absent. The prenuchal, nuchai, and occipital ridges are present but only slightly elevated. The pectoral cover plate has two ridges in upper and lower positions; the upper has several smaller ridges running parallel above it and the lower ridge has conspicuous ridges branching off the axis.
The brood area of the only male carrying eggs extends from the second to the penultimate (16th) trunk ring. The eggs are embedded in a gelatinous matrix; no lateral skin folds or protective plates are present and the matrix either dissolves or is absorbed shortly after the larvae are hatched. No other external sexual dimorphism is evident.
The anal and caudal fins are quite large, the latter measuring 10.6 mm. in length on the holotype; this is equivalent to a width of more than 5 trunk rings. The caudal rays number 10 in the holotype and 11 in all paratypes. There is no indication of regenerated rays.
The predominant color of the freshly killed male holotype was yellow- brown. The head is yellow-green with reddish brown blotches present on the snout, under and behind the eyes, and on the opercle. The pectoral fins are clear with a dark brown base. All of the body ridges are dark brown, an- teriorly appearing as narrow dashed lines and posteriorly as wider solid lines. The base of the dorsal fin is dark reddish brown, as are the first 2 rays; the dark pigment decreases in extent in a descending line on the sixth ray. The basal third of the middle 14 rays are light yellow-green, the outer two-thirds are clear; the posterior 2 rays are dark brown. The basal two-thirds of the anal is brown tinged with red; the distal one-third of the anal is yellow-green. The caudal is reddish brown with yellow-green basal blotches; the tips of the caudal rays are white. Three yellow blotches are present on the ventral surface of the tail rings between the anal and caudal, each blotch covering one to two rings. Three less pronounced light blotches are present on the ventral surface of the trunk rings. In preserved specimens the basic body color varies from light yellow-brown to darker brown and the ventral light areas are white.
Brotocy. Two males carrying eggs were collected in September and Oc- tober. A few hours after the holotype was placed in an aquarium, many of the eggs hatched. After formalin preservation, the remaining developing larvae fell from the egg sockets. The gelatinous matrix forming the sockets was firm and covered slightly less than half of each of the developing eggs. Sockets were present on the ventral surface of trunk rings 2-16 and extended onto the inferior lateral surfaces of rings 4-13; 173 sockets were present. The second egg-bearing male was collected on 10 September 1974 and placed in an aquarium; by 23 September all eggs had hatched or been lost and the male was preserved on 30 September. By this time almost all remnants of the
444 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.
matrix had dissolved or been absorbed; only a few dark areas representing the edges of the sockets remained. No other sexually dimorphic character was evident and none of the remaining specimens were sexed. However, the holo- type was collected under a rocky ledge with one other individual, possibly a female. Two of the Seal Rocks specimens were taken one week apart at the back of the same rocky hole and may represent a male and female. All other specimens were taken singly.
Newly hatched larvae measured 5-6 mm. TL and had fin folds but no fin rays present. The head was bent at an angle to the body, in this regard resembling a seahorse. The color was bright yellow-green with a few scattered melanophores. A bright silver and gold reflecting layer prominently surrounded the lens of the eye. All of the unhatched eggs appeared in much the same state of development, with the colors visible through the transparent egg case.
All of the Sydney Harbour specimens were taken in the outer harbor within 1.5 kilometers of the entrance, where the environment is similar enough to the open coast to support large brown kelp. Seal Rocks is on the open coast. The absence of this species from earlier collections is probably due to lack of scuba collecting from rocky regions in other areas. All specimens were collected in rocky areas—under rocky ledges, in crevices, holes, or small caves. Unlike many pipefishes, H. nocturna does not necessarily live in as- sociation with vegetation, as there was none at a number of collection sites (R. Kuiter, pers. comm.).
Four individuals were maintained in aquaria for periods of from one week to two months. During the day the pipefish remained in a small rock cave, swimming near the under surface of the overhanging rock in an upside down position. Only after some weeks in the aquarium would an individual leave the cave during periods of light and swim around the aquarium. During these brief forays, the fish righted itself and swam in a horizontal position close to the bottom, but did not come to rest. During most of this swimming, the tail was slightly bent in the region of tail rings 10-12. The large caudal was usually kept open and only rarely closed halfway like a fan, giving a pro- pulsive burst forward. The fish swam equally well backward and forward and most of the propulsion appeared to be from the pectoral fins. At night the pipefish swam freely to all corners of the aquarium in a normal horizontal posture. When the lights were turned on the fish swam to the cave and re- sumed the diurnal upside down swimming. After five minutes of darkness, swimming about the bottom of the tank was resumed. Aquarium behavior indicates Heraldia nocturna is nocturnally active, residing in protected rocky areas during the day. Doryrhamphus melanopleura from the Gulf of California exhibits the same type of upside down swimming behavior (R. Fritzsche, pers. comm.).
Vol. XL] PAXTON: NEW PIPEFISH 445
Name. The specific name of Heraldia nocturna is in reference to the noc- turnal activity of the species. The diurnal swimming posture suggests the common name ‘Upside down pipefish.’
RELATIONSHIPS. The genera Heraldia, Doryrhamphus, Dunckerocampus, Dentirostrum, Oostethus, and Maroubra appear to constitute a related group of pipefishes. Ail have abdominal brood areas, body ridges with spines, and the first body ring much longer than those remaining. The possible relation- ships of Maroubra are discussed in the next section. Heraldia, with a stout body and trunk rings more numerous than tail rings, appears most closely related to Doryrhamphus. The relationship between Doryrhamphus with brood pouch folds and Heraldia without folds is similar to that of another pair of related belly-pouch genera, Dentirostrum which has folds, and Dunck- erocampus which lacks them.
Heraldia nocturna has several characteristics that are rarely or never found in other pipefishes. For example, the basal color pattern in the dorsal fin is unique among syngnathids. The anal fin is relatively larger than that of other species. Highly emarginate caudal fin rays are known in few pipefishes, Dorichthys retzi of the tropical Indo-Pacific and juveniles of Oostethus being examples. The magnificently sculptured head of Heraldia is without equal in the pipefish world.
Maroubra perserrata Whitley, 1948.
Whitley (1948) described Maroubra perserrata as a new genus and species of pipefish from a single specimen collected from Maroubra Beach near Sydney by McCulloch in 1912. Eight specimens of this species were taken in August and November, 1974 from outer Sydney Harbour at depths from 3 to 20 m. in rocky areas. As the species previously was known only from the holotype, it is redescribed below; counts and measurements of the holotype (AMS 1.12659) are followed in parentheses by ranges of the 8 new specimens, 4 males and 4 females, 46.1—-72.9 mm. SL.
Slender pipefish with abdominal brood area lacking lateral membranous folds or protective plates; small cutaneous fold extending from midventral ridge of trunk rings in males only. Lateral trunk ridge continuous with lateral tail ridge, which in turn becomes superior tail ridge; inferior trunk and tail ridges continuous and superior trunk ridge ending free on 4th or Sth tail ring. Snout ridge without spines; small spine at anterior border of orbit. Body ridges pronounced, with points at posterior edge of each ring. Small caudal fin slightly emarginate, darkly pigmented on ventral half; other fins clear. Trunk rings 17(16); tail rings 26(24-27); dorsal rays 24(21-25), located on 5%(5—5%) total rings, of which %(0—%) are trunk rings and 5(4%4—-5'2) are tail rings. Pectoral rays 18(15-18); anal 4(4); caudal 10(9- 10).
446 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Total rings 43(40-43); total rings plus dorsal fin rays 67(61—-68); head in standard length 6.9(6.7—7.6); snout in head 2.0(1.9-2.1); dorsal fin base in head 1.6(1.4-1.7). The median snout ridge is elevated in front of the eyes; posteriorly it divides into two short, low ridges. Secondary head ridges are present, but not as well developed as those in Heraldia. The main opercular ridge extends completely across the opercle and curves dorsally posteriad. Smaller ridges radiate ventrally from the main ridge, but none are present dorsally.
In four males a small cutaneous fold is present on the midventral ridge of all trunk rings. Egg-carrying males were collected in August and November, but no eggs remained after preservation. No gelatinous matrix was present in either specimen and the way in which the low midventral skin fold protects the eggs is not clear; no lateral folds or protective plates are present. The well developed midventral ridge must separate the eggs into two groups.
In preserved specimens the reddish brown color markings of the males are slightly more pronounced than those of the females, particularly on the head. A dark stripe is present on the side of the snout from the tip to the eye and stripes and bands are present on the anterior trunk rings dorsally and laterally, while the remaining rings are evenly pigmented. One male that had been carrying eggs is much more darkly pigmented laterally with white spots in the dark pigment.
The genus Maroubra appears to be related to the doryrhamphine pipe- fishes, with an abdominal brood area, well defined body rings with posterior points, and an elongate first trunk ring. The peculiar brood area with a mid- ventral fold is a distinguishing feature, while the small caudal fin and pos- teriorly placed dorsal are not found in most of the related genera. The slender shape and large number of tail rings distinguish Maroubra from Doryrhamphus and Heraldia.
While comparing Heraldia and Maroubra with other pipefishes in the AMS collections, the holotypes of Choeroichthys suillus Whitley (1951) from Port Denison, Queensland, and Choeroichthys suillus malus Whitley (1954) from Masthead Island, Queensland were examined. The latter has all the characteristics of Doryrhamphus, and the presence of two dark stripes behind each eye indicates that C. s. malus is a synonym of D. negrosensis Herre. The holotype of C. suillus is a male with protective plates extending ventrally from the trunk rings; this feature, plus the lack of a lateral tail ridge, is diagnostic for the genus Choeroichthys.
LITERATURE CITED HERALD, E. S. 1940. A key to the pipefishes of the Pacific American coasts with descriptions of new genera and species. Allan Hancock Pacific Expeditions, vol. 9, no. 3, pp. 51-64.
Vol. XL] PAXTON: NEW PIPEFISH 447
HERALD, E. S., anp J. E. RANDALL
1972. Five new Indo-Pacific pipefishes. Proceedings of the California Academy of
Sciences, ser. 4, vol. 39, no. 11, pp. 121-140. Munro, I. S. R.
1958. Family Syngnathidae. Zn Handbook of Australian fishes, nos. 20-23, pp. 82-93. Supplement to Australian Fisheries Newsletter, vol. 17, nos. 2-5, pp. 17-20, 1720720" 17=20:
WHIttey, G. P. 1948. Studies in ichthyology, no. 13. Records of the Australian Museum, vol. 22, no. 1,
pp. 70-94.
1951. Studies in ichthyology, no. 15. Records of the Australian Museum, vol. 22, no. 4, pp. 389-408.
1954. New locality records for some Australian fishes. Proceedings of the Royal Zoological Society of New South Wales, 1952-53, pp. 23-30.
: -
INDEX TO VOLUME XL
FOURTH SERIES
(Compiled by Lillian J. Dempster)
New names and principal references in boldface type. References to figures in ztalics.
Abutilon depauperatum 183 Acacia 191
macracantha 191
rorudiana 191 Acontiinae 150, 181, 208i Acronictinae 150, 170 Acrostichum reptans 230 Acutomentum macdonaldi 130 Aeluroscalabotes 91 Aemasia 421
lichenosa 421 Agave palmeri 240, 241 Agonidae 75, 86 Agrius cingulatus 149, 152, 153 Agrotis
bosqi 163
consternans 149, 163, 164, 204, 205
fasicola 163 ipsilon 149, 163, 164 lutescens 163
subterranea williamsi 150, 163, 164
Agrotisia 179
subhyalina 179
williamsi 150, 177, 179 Allamanda 155 Alternanthera
echinocephala 182
filifolia 182 Amaranthus viridis 174 Amyna insularum 150, 180, 181, 182 Anarbylus 87, 88
switaki 87, 88, 88, 91, 91, 92 Anarchias galapagensis 417, 418, 419 Anicla
infecta 167, 208i
oceanica 150, 166, 167 Annona
montana 196
species 196 Anomis 194
editrix 151, 192, 193, 194.
erosa 151, 192, 193, 194
illita 151, 192, 193, 194.
luridula professorum 151, 192, 193, 194
professorum 194 Anthoecia cystiphora 167 inflata 167 onca 167 Anthophoridae 231 Anticarsia gemmatalis 151, 188, 189, 196 prona 151, 188, 189, 197 Anuroctonus 15 A pistes carinatus 75 species 74, 75, 85 Arctiidae 145, 149, 159 Argyrogramma verruca 151, 188, 189 Ascalapha odorata 151, 188, 189, 191 Aspidium chiapasense 229 lindenii 229 Asplenium breedlovei 209, 210, 211 cristatum 211 heterochroum 212 munehii 211, 212, 228, 229 myriophyllum 212 olivaceum 210, 211, 212 radicans 211 sessilifolium 211 soleirolioides 210, 211, 212 trichomanes 212 Atethmia subusta 181
Autoplusia egena galapagensis 151, 187, 188,
189
Bagisara repanda 150, 180, 181 subusta 181 Bendis formularis 151, 192, 193, 196 Bioculus 14 Bipes biporus 100 Bolina indomita 190 Bombyx repanda 181
[449]
450 CALIFORNIA ACADEMY OF SCIENCES
Bomolocha vetustalis 201 Brachirus 275 chloreus 275 Brachypterois serrulifer 74, 85 Brachyrus 275 Broteas 13, 14 alleni 3, 13 alleniit 2, 14 formosus 3 Broteochactas 13, 14 allenii 3 Bursera graveolens 185
Cacabus miersii 152 Caesalpinia praecox 27 section [Sappania] praecox 27 Caesalpinioideae 17, 18 Calendula 188 Callopistria floridensis 150, 172, 173 Carica papaya 155 Cassia 191 Catabena seorsa 150, 172, 173 species? 173 Catocalinae 150, 185 Celerio lineata florilega 158 Celiptera remigioides 150, 186, 187 Cercidiopsis 18 microphylla 24 Cercidium 17, 18, 19, 20, 21, 22, 23, 27, 29, 30, 32, 335 37, 375 38) 39) 40, Al, 43, 44, 45, 47, 48, 49, 50, 51, 52, 53, 54 australe 49 floridum 23, 26, 27, 29, 33, 34, 35, 38, 40, 41, 42, 43, 45, 48, 49, 51 floridum subsp. floridum 20, 23, 26, 27, 333537. 38) 40. eAON4l. 42,42, 43, 44, 45, 46, 47, 47, 48, 49, 50 floridum subsp. peninsulare 20, 22, 23, 35, 37, 38, 40, 40, 41, 42, 42, 43, 44, 45, 45, 46, 47, 47, 48, 49, 50, 53 Goldmanii 27 microphyllum 18, 19, 22, 22, 23, 24, 26, D730. 32345 GSN 7381039 139, 40; 40, 41, 42, 43, 44, 45, 406, 46, 47, 48, 49, 49, 50, 51, 52, 53, 54 molle 30, 31, 32, 33 peninsulare 35 plurifoliolatum 27 praecox 17, 19, 22, 23, 27, 28, 29, 30, 32,
[Proc. 4TH SER.
33, 37, 38, 39, 40, 40, 42, 43, 44, 45, 46, 46, 47, 48, 49, 49, 50 sonorae 33 X sonorae 19, 22, 23, 30, 31, 32, 33, 38, 40, 40, 41, 42, 43, 44, 45, 46, 46, 47, 48, 49, 49, 50, 53 spinosum 22, 27 torreyanum 29, 33, 34, 51 unijuga 27 viride 27 Cevallia sinuata 238, 240 Chactidae 13, 14 Characoma nilotica 150, 184, 185 Choeroichthys 446 suillus 446 suillus malus 446 Cirphis cooperi 170 Cissampelos pareira 174 Cissus sicyoides 157 Clerodendrum molle 154, 190 Clethra 221 Clinocottus analis 75, 86 Cobubatha numa 208i Coleonyx 91, 92 brevis 91, 92 elegans 92 reticulatus 92 variegatus 91, 92 Commicarpus tuberosus 159 Compositae 179 Convolvulus 182 Conyza species 238, 240 Cordia leucophlyctis 154, 190 lutea 154 species 190 Cottidae 75, 86 Cottoidei 63, 73, 75, 78, 80 Cottuniculidae 75, 86 Cottunculus thompsoni 75, 86 Cottus asper 74, 75, 86 Cropia infusa 150, 171, 172, 173 Crotalaria 159 Crotalus atrox 101 catalinensis 101 Cryptocarpus pyriformis 174, 176, 185, 197 Ctenitis 215, 216, 223 baulensis 213, 214, 215 bullata 213, 214 chiapasensis 229
Vor. XL]
lanceolata 229
lindenii 229
melanosticta 216
nigrovenia 215
strigilosa 213
thelypteroides 2174, 215
ursina 216, 217 Ctenosaura hemilopha 100 Cyclophora impudens 149 Cyclopteridae 75, 86
Datura meteloides 240, 241 Decelea infusa 171 Dendrochirus 266, 271, 275 barberi 265, 266, 269, 273, 274, 275, 276, 277 brachypterus 275 chloreus 266, 275, 276, 277 hudsoni 266, 275, 276, 277 zebra 74, 85, 275 Dendroscorpaena 296, 297 species 85 Dentirostrum 441, 445 Desmanthus cooleyi 238, 240 Didymocentrus 14 Diplocentridae 14 Disclisioprocta stellata 149 Dorichthys retzi 445 Doryrhamphus 439, 441, 445, 446 melano pleura 444 negrosensis 446 Drimys 221 Dryopteris futura 216, 217 munchii 216, 2/7, 218 nubigena 218 patula 220 rossit 220 tonduzii 215 Dunckerocampus 441, 445
Echidna chionostigma 417, 424 nocturna 417, 418, 419, 424 scabra 417, 424 zebra 417, 419, 420 Ectreposebastes 266, 291 imus 73, 85, 265, 266, 270, 274, 276, 290, 291, 292, 292 niger 267, 268, 291
INDEX 4
1
On
Elaphoglossum albomarginatum 218, 21/9, 220 alfredii 220 chiapense 218, 219, 220 latifolium 220 mathewsii 220 Elaphria chalcedonia 178 dubiosa 178 eneantada 150, 178, 180, 201, 202 Encelia hispida 168 Enchelycore 421 lichenosa 417, 418, 419, 421, 422 octaviana 417, 418, 419 Enhydra lutris 335, 338, 340, 342-352, 355-370, 373, 375-379, 381-388, 390, 391, 393, 395, 397, 399, 401, 403, 405, 407-409, 411-413, 429, 430 lutris gracilis 430, 433, 436 lutris lutris 429, 430, 431, 435 lutris nereis 429, 430, 431, 436 Enyo lugubris delanoi 149, 155, 156, 157 Epidromia sephyritis 151, 188, 189, 198 setophora 151, 188, 189, 197 Epiplema amygdalipennis 151 becki 149, 151, 160, 161, 201, 203 Epiplemidae 145, 149, 151, 2081 Epistor lugubris delanoi 155 Eridiphas slevini 100, 101 Erinnyis alope dispersa 149, 152, 153, 154 ello encantada 149, 155, 156, 157 obscura conformis 149, 155, 156, 157 obscura obscura 155 Eriopus floridensis 173 Eublemma obliqualis 182 recta 150, 180, 181, 182 Eublepharinae 87 Eublepharis 91 macularius 92 Eucerini 231, 239, 240, 241 Euchelia galapagensis 161 Eumorpha fasciata tupaci 149, 152, 153, 157 labruscae yupanquii 149, 152, 153, 157 Eupithecia
452 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
leleupi 149 dactylopterus 73, 85
perryvriesi 149 rufescens 312, 313 Euteliinae 150, 183 Heliocontia margana 150, 182, 184 Euxoa williamsi 163 Heliothinae 150, 167
Heliothis Feltia annexa 163 cystiphora 150, 167, 169 virescens 150, 167, 168, 169
Gambusia 312 Heliotropium angiospermum 190 Gekkonidae 87 Hemilepidotus jordani 75, 86 Geometridae 146, 149 Hemitheconyx 91
Geometroidea 145 caudicinctus 92 Glympis Heraldia 441, 442, 445, 446 ” Saereaibs Gp nocturna 439, 440, 441, 442, 444, 445 toddi 151, 192, 198, 199 Herse cingulata 152 Gonitis editrix 194 Hexagrammidae 74, 86 Conodonta nos Hexagrammos decagrammus 74, 86
Hibiscus tiliaceus 194 biarmata evadens 151, 192, 193, 195 Te ee a é : Hippomane mancinella 155 biarmata galapagensis 195
ware Hispaniscus sinensis 118 fulvangula 151, 192, 193, 195 ls mabe es O° Gramineae 185 Homoptera obsita 187 Grammitis Hoplichthyidae 74, 86 blepharodes 229 Hoplichthys langsdorfi 74, 86 harristi 230 Hydria affirmata 149
leptostoma 230 ; z Tineearitatan220N 228206 Hyles lineata florilega 149, 158, 160, 161 Hymenoptera 231
revoluta 230
Gymnapistes marmoratus 73, 85 Ie aati 181 10S Woo OO microfuliginea ‘ , 200, ; x” 417, 422, 423, 425 ‘ Gymnothorax 417, 422, f aah
aquae-dulcis 422 arae 417, 424 buroensis 417, 418, 420
vetustalis 151, 198, 199, 201, 204 Hypeninae 151, 198
castaneus 417, 418, 420 Hypolepis 223
chilos pilus 418 melanochlaena 221, 228, 229
dovii 417, 418, 420, 424 SEES GR)
funebris 418 Hypsiglena 100
mordax 417, 422
panamensis 417, 418, 420 Ingura arcigera 183
petelli 423 Inimicus cuvieri 73, 85
pictus 417, 418, 420, 424 Ipomoea 152
reticularis 423 hirsutula 238, 240, 241
rueppelliae 417, 423 pes-caprae 174
ruppeli 423 triloba 182
thomsoni 417, 424 Tracundus 266, 287
undulatus 418 signifer 73, 85, 265, 266, 267, 269, 270,
274, 276, 287, 288, 288, 289
Hadeninae 150, 168 signifer rarotongae 287, 288, 289 Hadrurus 15 signifer signifer 288
Harrisonia williamsi 179 Helicolenus Jatropha 155
Vor. XL]
Laguncularia racemosa 185, 198 Lantana 174 peduncularis 173, 190 Laphygma roseae 176 Latax lutris nereis 429 Leguminosae 17, 188 Leptocottus armatus 74, 86 Leptodeira 100 Leptotrigla alata 75, 86 Leptotyphlopidae 93 Leptotyphlops 93, 103 humilis 93, 94, 96, 98, 101, 103, 104 humilis cahuilae 96, 98, 103, 104, 105 humilis dugesi 96, 98, 104, 105 humilis humilis 96, 98, 103, 104, 105 humilis humilis X cahuilae 105 humilis humilis X utahensis 103, 106 Humilis levitoni 94, 95, 96, 98, 101, 104, 105 humilis lindsayi 96, 97, 98, 101, 104, 105 humilis segregus 96, 98, 104 humilis slevini 93, 96, 98, 101, 103, 104, 105 humilis teniculus 98 humilis tenuiculus 96, 104 humilis utahensis 96, 98, 103, 104 Letis mycerina 151, 188, 189, 191 Leucania latiuscula 168 solita 168 Liparis florae 75, 86 Lycodontis afer 418 umbra 418 xanthos pilus 417, 424 Lycophotia oceanica 167
Macroscorpius pallidus 74, 85 Magusa erema 150, 170, 171, 172, 173, 200 orbifera 170, 171 Malvaceae 195 Manduca rustica calapagensis frontispiece, 149, 152, 153, 154 sexta leucoptera 149, 152, 153 Maroubra 441, 445, 446 perserrata 439, 445 Martinapis 239, 240 bipunctata 240 luteicornis 231, 239
INDEX 453
(Martinapis) luteicornis 239 (Svastropsis) bipunctata 240 Maytenus octogona 197 Melipotis acontioides producta 151, 186, 190 harrisoni 151, 186, 191 indomita 151, 186, 190, 191 Menodora scabra 231, 238, 240, 241 Mentzelia aspera 190 pumila 238, 241 Merinthe macrocephala 294 Metalectra praecisalis 208i Metallata absumens contiguata 151, 196, 198, 199 Micra recta 182 Microgramma 230 ciliata 230 nitida 230 reptans 230 Microlepidoptera 146 Minous monodactylus 73, 85 pictus 73, 85 Mirounga angustirostris 434 Mocis incurvalis 150, 185, 186 latipes 150, 186, 187 Muraena 425 acutis 417, 424 albigutta 417, 421, 426 aquae-dulcis 417, 422, 425 argus 417, 418, 419, 421, 421, 426 clepsydra 417, 418, 419, 420 insularum 417, 425, 426 lentiginosa 417, 418, 419, 424, 425 Muraenidae 417 Murenophis marmoreus 417, 425 Myoxcephalus quadricornis 75, 86 M ythimna latiuscula 150, 168 solita 150, 168, 169, 170
Natrix valida celaeno 100 Nemapterois 275 biocellatus 275 Neogalea esula longfieldae 150, 173, 175, 7) Neomerinthe 266, 312, 313 beanorum 73, 85 hemingwayi 312
454 CALIFORNIA ACADEMY OF SCIENCES
rufescens 265, 266, 271, 274, 276, 296, Baas 3.6314 Nephrodium lanceolatum 229 Noctua dolichos 176 saucia 165 verruca 188 virescens 168 Noctuidae 145, 148, 149, 163, 2081 Noctuinae 149, 163, 208i Noctuoidea 145 Nonagria indubitans 183 Nullibrotheas 1, 2, 13, 14, 15 alee i, 74, Bs 25 By Op Wy Gy HO, WA, 13, Vs
Occella verrucosa 75, 86 Oleaceae 231 Onograceae 157 Oostethus 441, 445 Ophiderinae 151, 190, 208i Ophiodes remigioides 187 Ophiodon elongatus 86 Ophiuche lividalis 151, 198, 199, 204 minualis constans 151, 198, 199, 206 Opuntia 152 Oxydia lignata 149 Ozarba 181 consternans 150, 180, 181, 200
Pachygonia drucei 149, 157 Paectes arcigera 150, 183, 184, 185 indefatigabilis 183 isabel 183 Parascorpaena armatus 85 species 73 Parascorpaenodes hirsutus 277 Parkinsonia 18, 49, 50, 51, 52, 54 aculeata 17, 18, 19, 30, 49, 50, 51, 52, 53, 54, 190 africana 49, 51, 52, 53, 53, 54 anacantha 52, 53, 54 florida 33 microphylla 24, 26 scioana 52, 53, 53, 54 torreyana 33 Paruroctonus 15 Passiflora foetida 168 Peliala fuliginea 151, 198, 199, 201, 203, 204
[Proc. 4TH SER.
Peloropsis 310
xenops 310, 311 Peltophoropsis scioana 52, 53, 53 Perca cirrhosa 296, 297 Peridroma 165
conwayi 165
margaritosa 165
saucia 150, 165, 166, 167 Perigea
apameoides 178, 179, 180
ebba 179
mobilis 178
ruthae 179
sutor 178 Perizoma (?) perryi 149 Phalaena
(Attacus) mycerina 191
(Bombyx) odorata 191
frugiperda 176
ipsilon 163
(Noctua) eridania 174
(Noctua) ornatrix 159
00 188 Phaseolus 188 Phenacoscorpius 266, 293
megalops 266, 267, 270, 274, 276, 293,
294
nebris 265, 294 Phlebodium nitidum 230 Pholus
fasciatus tupacit 157
labruscae yupanqui 157 Phycodurus 439 Phyllopteryx 439 Physalis pubescens 154 Pisces 59, 243, 265, 417 Piscidia carthagenensis 197 Platycephalidae 74, 86 Platycephalus species 74, 86 Platysenta
mobilis 150, 178, 179, 180
ruthae 150, 177, 179
sutor 150, 178, 179, 180 Plectrogeniinae 284 Plectrogenium 266, 284.
nanum 73, 85, 265, 266, 270, 274, 276,
284, 285, 285
Pleopeltis 230
munchii 230
revoluta 230 Plusia includens 188
Voit. XL]
Plusiinae 151, 187 Plusiodonta clavifera 151, 195, 198, 199 Pocillopora meandrina 317 Polygala racemosa 238, 240, 241 Polypodium 230 astrolepis 230 blepharodes 229 ciliatum 230 harrisii 230 leptostomum 230 longepinnulatum 223 meridensis 221 munchii 230 palmeri 230 sureurrens 222, 223 Polystichum bicolor 223, 224, 225 drepanoides 225 erythrosorum 225, 226 furfuraceum 224, 225, 226 ordinatum 225 Ponometia indubitans 150, 183, 184, 185 Pontinus 266, 294, 296, 312, 314 castor 294 longis pinis 74, 85 macrocephalus 265, 266, 270, 274, 276, 294, 295, 295, 296, 314 niger 291 spilistius 266, 294, 295 Portulaca oleracea 159, 174 Powondrella cingillaria 208i Prodenia latifascia 174 Prosopis jubiflora 190 Protoparce calapagensis 154 leucoptera 152 postscripta 154 rustica calapagensis 154 rustica calapagensis ab. nigrita 154 sexta leucoptera 152 Psaphara conwayi 150, 165, 166, 167 interclusa 150, 165, 166, 167, 195, 201, 202 Pseudaletia cooperi 150, 169, 170 sequax 150, 169, 170 Pseudemys scripta nebulosa 100 Pseudoplusia includens 151, 188, 189 Pseudupeneus maculatus 124 Psorya hadesia 151, 188, 189, 197
INDEX 455
wal
Pteris chiapensis 224, 225, 226 mexicana 227 Pterois 266, 271, 272, 274, 296 barberi 275 mombasae 272 vadiata 74, 85 russelli 272 sphex 265, 266, 269, 272, 273, 274, 276 zebra 266, 275 Pteropodus sinensis 118 Pteropterus 271 Pyralidae 146 Pyralis lividalis 204 margama 182
Quercus 221
Rabula 417, 422, 425 acuta 423 davisi 417, 422, 423 fuscomaculata 422, 423 marshallensis 423 rotchit 417, 424 Remigia lutipes 187 Rhetinophloem viride 27 Rhinopias 266, 310 xenops 265, 266, 271, 274, 276, 311, 311, 312 Rhynchosia minima 197 Rivula 192 asteria 151, 192, 198, 199, 201, 202 ?dubiosa 192 Rubiaceae 158
Sagittaria 188 Salmo aguabonita 243, 244, 246, 247, 249, 251, 253, 254, 255, 256, 257 aguabonita aguabonita 243, 244, 246, 252, 254, 258, 259, 260 aguabonita gilberti 244, 259
8 oR = 8 ao Ss = x. pak i=] & = ied o d ns. bo aS WwW bo a 5 bho rs mn bo > Ro) bo On bo
’ gairdneri gilberti 243, 244, 259, (Parasalmo) 251, 25 trutta 252 whitet 259
Salmonidae 243
456
CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
Salvia reflexa 238, 241 Sarcostemma angustissima 155 Sarrothripa nilotica 185 Sarrothripinae 150, 185
Sator 101, 103
angustus 101 grandaevus 101
Sauria 87 Scalesia 146
affinis 161, 168
Scorpaena 266, 271, 296, 297, 298, 299, 310,
314, 316, 317, 320
agassizi 74, 85
albobrunnea 74, 85, 317
asperella 266, 319
ballieut 265, 266, 269, 271, 274, 276, 297, 314 ST. 319) 319
brasiliensis 74, 85
colorata 265, 266, 271, 274, 276, 314, 322, 3735 326, 327
coloratus 326
coniorta 265, 266, 271, 274, 276, 315, 316, 317
cookii 308
elongata 74, 85
frondosa 310
galactacma 265, 266, 271, 274, 314, 315
gibbosa 296, 305
guttata 74, 85
mystes 74, 85
nesogallica 296
orgila 297
parvipinnis 280
pele 265, 266, 271, 274, 276, 320, 321, B224303,9324. 325, 32680307
peruana 266, 318, 319, 320
polylepis 277
porcus 74, 85, 314
remigera 266, 289, 290
russula 74, 85
(Sebastapistes) 317
tinkhami 317
volitans 271
Scorpaenenopsis diabolus 305 Scorpaenichthys 296
marmoratus 75, 86
Scorpaenidae 59, 60, 73, 74, 75, 77, 80, 83,
109, 265, 266
Scorpaeniformes 60, 63, 74, 77, 80, 83 Scorpaeninae 312 Scorpaenodes 266, 277, 282, 283, 298
corallinus 265, 266, 269, 270, 274, 276, 280, 281, 283, 284 cortezi 110, 121 englerti 283 fowleri 297, 298, 318 guamensis 279, 282 hirsutus 265, 266, 269, 269, 274, 276, Darlrh LP DIR 6X0) kelloggi 265, 266, 269, 269, 274, 276, 278, 279, 279, 280 littoralis 265, 266, 267, 269, 270, 274, 276, 280, 281, 282, 283 parvipinnis 74, 85, 265, 266, 269, 274, 276, 279, 280, 281, 282 Scorpaenopsis 266, 268, 270, 296, 297, 298, 299, 301, 304, 305, 307, 309, 310, 314 altirostris 265, 266, 271, 274, 276, 297, 308, 309, 310, 310 brevifrons 265, 266, 271, 274, 276, 299, 300, 301, 302, 303, 304, 307, 308 cacopsis 265, 266, 271, 274, 276, 299, 304, 307, 308, 309 catacola 266 catocala 305, 306 cirrhosa 297 cotticeps 310 diabolus 265, 266, 271, 274, 276, 297, 304, 305, 306, 306, 307, 307, 308 fowleri 265, 266, 270, 274, 276, 297, 298, 298, 310 gibbosa 297, 305, 306, 307 gibbosus 305, 306 macrochir 267, 305, 307 neglecta 297, 305, 307 nesogallica 305 species 74, 85 Scorpaenopsus gibbosa 305 Scorpaenopus cacopsis 308 Scorpionida 1 Scorpius 13 OG? MN, Bs do 133 Scotia galapagosensis 163 Scutia pauciflora 171, 198 Sebastapistes 314, 317 albobrunneus 314, 316, 318 albo-brunneus 316 asperella 297, 299, 301, 318 ballieut 318 bynoensis 326 coloratus 326 coniorta 316, 317
Vor. XL]
corallicola 266, 318 galactacma 314, 315, 318 galactacme 314
INDEX
nuchalus 318 strongia 316
Sebastella littoralis 282
Sebastes 59, 60, 61, 64, 66, 67, 68, 69, 70, 71, 72, 73, 74, 76, 77, 79, 80, 86, 109, 110, fla MS 1S 129% 12595126 4272 130, 132-1136,0137, 137, 138, 139
aleutianus 64, (Allosebastes ) (Allosebastes) (Allosebastes) (Allosebastes) (Allosebastes) (Allosebastes) (Allosebastes) (Allosebastes ) (Allosebastes ) (Allosebastes) (Allosebastes ) (Allosebastes) (Allosebastes) (Allosebastes ) (Allosebastes) (Allosebastes) alutus 64, 69, atrovirens 64,
67, 69, 83, 118 118, 138, 139 cortezi 139
dallii 138 diploproa 138, 139 emphaeus 138 peduncularis 139 proriger 138 rufinanus 138 saxicola 138, 139 scythropus 138, 139 semicinctus 138 sinensis 139 variegatus 138 varispinis 139 wilsoni 138 zacentrus 138
83
(By Tis 15 383
auriculatus 64, 69, 83
aurora 64, 69, 83
babcocki 64, 69, 83
baramenuke 64, 69, 83
borealis 64, 69, 83, 109
brevispinis 64, 69, 76, 83
capensis 64, 69, 83, 138
carnatus 64, 73, 76, 79, 83
caurinus 64, 70, 72, 73, 76, 79, 83, 110
chlorostictus 64, 69, 83, 138
chrysomelas 64, 73, 76, 79, 83
ciliatus 64, 69, 83
constellatus 64, 69, 83, 138
cortezi 109, 111, 112, 113, 114, 115, 121, 230124, 124)" 125. 129. 163) AGawI35; 139
crameri 64, 69, 83
dallii 64, 69, 83, 138
diploproa 64, 69, 83, 109, 121, 124, 125, 138, 139
elongatus 64, 69, 83
emphaeus 64, 69, 83, 138
457
ensifer 64, 69, 83, 138
entomelas 64, 69, 83
eos 64, 69, 83, 138
exsul 64, 69, 83, 110, 111, 112, 113, 115, 116.118. 119.129.7106, 132. 1835.194. 135, 138
fasciatus 86
flameus 64, 69, 83
flavidus 64, 69, 79, 83
gilli 64, 69, 83
glaucus 64, 69
goodei 64, 69, 83
helvomaculatus 65, 69, 83, 138
hopkinsi 65, 69, 83
hubbsi 65, 69, 83
ijimae 65, 71, 83
inermis 65, 71, 83
iracundus 65, 69
itinus 65, 69
jordani 65, 69, 83
joyneri 65, 69, 84
kawaradae 65, 69
lentiginosus 65, 69, 84, 138
levis 65, 69, 84
longispinis 65, 69, 84
macdonaldi 65, 69, 84, 109, 110, 112, 113, Wie, TBO), ashi, si), Use, asl TBS, The, 138
macrocephalus 294, 295
maliger 65, 73, 76, 79, 84
marinus 65, 68, 69, 71, 75, 80, 84, 86
matsubarae 65, 69, 84
melanops 65, 69, 79, 84
melanostictus 65, 69
melanostomus 65, 69, 84
mentella 86
miniatus 65, 69, 84
mystinus 65, 66, 67, 69, 79, 84
nebulosus 65, 73, 76, 79, 84
nigrocinctus 65, 69, 79, 84
nivosus 65, 71, 84
notius 138
oblongus 65, 71, 84
ovalis 65, 69, 84
owstoni 65, 69, 84
pachycephalus 65, 71, 84
paucispinis 65, 70, 71, 75, 76, 79, 80, 84
peduncularis 109, 111, 174, 115, 124, 127. 128) 129) 16350194. ssyns9
phillipsi 65, 69, 84
pinniger 65, 69, 84
458
CALIFORNIA ACADEMY OF SCIENCES
proriger 65, 69, 84, 138 (Pteropodus) 76, 79, 80 rastrelliger 65, 68, 69, 76, 84 reedi 65, 69, 84
wilsoni 66, 69, 85, 138
zacentrus 66, 67, 69, 85, 138 Sebastichthys sinensis 110, 118 Sebastinae 312
rosaceus 65, 69, 84, 138 Sebastiscus
rosenblatti 65, 69, 84, 113, 138 marmoratus 74, 74, 79, 118 ruberrimus 66, 69, 84 species 85
rubrivinctus 66, 69, 84 Sebastodes
rufinanus 109, 138
rufus 66, 69, 84
saxicola 66, 69, 84, 138, 139 schlegeli 71, 79
schlegelii 66, 84
scythropus 66, 69, 84, 138, 139 (Sebastodes) 76
(Allosebastes) 118
brevis pinis 76 (Hispaniscus) 118 (Hispaniscus) sinensis 118 macdonaldi 130
proriger 130
sinensis 118
(Sebastomus) 111, 113, 115, 118, 138, 139 Sebastolobus 124 (Sebastomus) chlorostictus 138 Sebasto psis (Sebastomus) constellatus 138 galactma 314 (Sebastomus) ensifer 138 guamensis 279, 280 (Sebastomus) eos 138 kelloggi 277, 279 (Sebastomus) exsul 111, 138 parvipennis 280 (Sebastomus) helvomaculatus 138 parvipinnis 280 (Sebastomus) lentiginosus 138 scaber 279 (Sebastomus) notius 138 Sebastosemus species 74, 85 (Sebastomus) rosaceus 138 Semiothisa
(Sebastomus) rosenblatti 138 cerussata 149 (Sebastomus) simulator 138 cruciata cruciata 149 (Sebastomus) spinorbis 138 cruciata isabelae 149 (Sebastomus) umbrosus 138 Serpentes 93
semicinctus 138
Setarches 266, 289
[Proc. 4TH SER.
serranoides 66, 69, 84 guentheri 74, 85, 265, 266, 270, 274, 276,
serriceps 66, 68, 69, 75, 80, 84
simulator 66, 69, 85, 138
S7WENSIS 160. 09585, LOOM ere S 714, 105. W185 5120) 1211129) 133, 134; 135, 139
spinorbis 109, 110, 712, 113, 116, 117, IS. Me, eve IGG. AGS
steindachneri 66, 69, 85
taczanowskii 66, 70, 73, 74, 75, 76, 80, 85
thompsoni 66, 71, 85
trivittatus 66, 71, 85
umbrosus 66, 69, 85, 113, 138
variegatus 66, 69, 85, 109, 138
varispinis 109, 111, 774, 115, 125, 127, ID9IGS 134 135. 139
vexillaris 66, 73, 76, 85, 110
viviparus 86
vulpes 66, 70, 71, 85
wakijai 69
wakiyai 66
289, 290, 290, 291, 292
giintheri 289
longimanus 74, 85
remiger 289 Sida
glomerata 181
species 183 Sideria chlevastes 417, 423 Simanthedon 232, 236, 239, 240
linsleyi 231, 233, 235, 236, 239, 240,
241, 242 Solanum elaeagnifolium 238, 241 Soleirolia 212 Sorygaza didymata 201
variata 151, 198, 199, 201, 203
Spermacoce 158 Sphacelodes vulneraria 149 Sphingidae 145, 149, 152 Sphingoidea 145
VoL. XL]
Sphinx cingulata 152 tersa 158 Spodoptera 176 dolichos 150, 176, 177 eridania 150, 174, 175 frugiperda 150, 174, 175, 176 latifascia 150, 174, 175 roseae 150, 176, 177 Sporobolus virginicus 168 Spragueia creton 150, 183, 184 plumbeata 183 Svastrides melanura 240 Symplocos 221 Synanceia verrucosa 85 verrucosus 74 Synancelinae 267 Syngnathidae 429 Syngrapha egena galapagensis 187 Synhalonia 239 Syntropis 15
Taenianotus 266, 285, 296, 312 citrinellus 266, 285, 286 garretti 266, 285, 286
triacanthus 265, 266, 270, 274, 276, 285,
286, 286, 287 Tafalla clavifera 195 Tagetes 179 Tetralonia 239 Thalpochares asteria 192 Thamno phis couchi atratus 96 elegans atratus 96 Thelypteris 229 blepharis 227, 228, 229
section (Cyclosorus) blepharis 227
nubigena 228, 229
ovata 227
puberula 227
thomsoniit 229
tuerckheimii 227 Thermesia prona 197
INDEX
Thyrinteina infans 149 umbrosa 149 Tournefortia 159 psilostachya 161, 190 pubescens 161 Trachaea roseae 176 Trachea 171
cavagnaroi 150, 171, 172, 173, 200
roseae 176 Triglidae 75, 86
Uroctonus 1, 14, 15 mordax 3, 14 privus 2, 3, 14 Uropterygius necturus 417, 418, 419, 421 polystictus 417, 418, 419 species 417, 418 Urticaceae 212 Utetheisa
459
devriesi 149, 159, 161, 162, 204, 205
galapagensis 149, 159, 161, 162
ornatrix 149, 159, 160, 161
perryi 149, 159, 161, 162, 204, 205
Vaejovidae 1, 14, 15 Vaejovis 1, 14, 15 Verbena 174
Waltheria ovata 183 Wedelia 179
Xenoglossodes 239, 240
X ylophanes norfolki 149, 156, 157, 158 tersa 149, 156, 157, 158
Zale obsita 150, 186, 187 species, viridans group, 187 viridans 187 Zaniolepididae 75, 86 Zaniole pis latipinnis 75, 86 Zanthoxylum 159
460
Page
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CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
ERRATA
Line 21 from top: for Cercidium X sonorae Rose and Johnston read Cercidium X sonorae Rose and Johnston (pro sp.).
Line 20 from bottom: for wakijai read wakiyai.
Line 6 from top: for L. h. teniculus read L. h. tenuiculus.
Line 3 of Abstract: for Phenacoscorpius nebris read Phenacoscorpius megalops. Line 1 of figure caption: for Phenacoscorpius nebris read Phenacoscor pius megalops.
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