EASTERN PACIFIC FISHES OF THE
GENUS HAEMULON (POMADASYIDAE)
by
Susan L. Hong
A Thesis Submitted to the Faculty of th~
College of Science in Partial Fulfillment of the Requireme nts for the Degree
of Master of Science
Flori~a Atlantic University
Boca Raton, Florida
August, 1974 II
EASTERN PACIFIC FISHES OF THE
GENUS HAEMULON (POMApASYIDAE)
by
Susan L. Hong
This thesis was prepared under the direction of the candidate's thesis advisor, Dr. Walter R. Courtenay, Departme nt of Bi ological Sciences and has been approved by the ~embers of h e r sup e rvisory committee. It was submit ~ ed to the faculty of the College of Science and was accepted i n partia l fulfillment of the requirements for the degree of Master of Science.
SUPERVISORY COMMITTEE: hl o(t RC diAka, .\ . Thes1s Advisor 1 ~
Chairman, Department of Biological Sciences
c e d S tudies
ii Acknowledgments
I wish to thank Dr. Walter ~. Cour t enay, Jr. of
Florida Atlantic University for providing specimens, patient
suggestio n s , and helpful comments in the preparation of the
manuscript. Thanks are due the other members of my graduate
committee, Dr. Daniel Austin for loan of reference material,
Dr. Sheldon Dobkin for loan of equipment, and Dr. C. Richarc
R~bins for making available specimens from the University of
Miami Marine Laboratorj.
Appreciation is extended to the following persons at
the United States National Museum: Mr. Edgar N. Gramblin
and Dr. Ernest A. Lachner for providing specimens, work space,
and helpful suggestions; Dr. William R. Taylor for guidance
in matters of catalog number changes and type specimen
designations; and Ms. Susan Karnella and Janet Ryle for the i r
generous assistance.
Many thanks are due Ms. Doris Scu~ly for an excellent job in typing the manuscript. I am extremely grateful to my family for their patience and faithful support.
iii Abstract
Author: Susan L. Horig
Title: Eastern Pacific Fishes of the Genus
· H~emulon (Pomadasyidae)
Institution: Florida Atlantic University
Degree: Master of Science
Year: 1974
Five eastern Pacific species of the genus Haemulon
Cuvier are described, their color pattern and dentition illustrated, and a key presented for their identification.
Lythrulon Jordan and Swain and Orth6sto~chus Gill are synonymized with Haemulon. Haemulon scudderi Gill, Haemulon sexfasciatum Gill, Haemulon flavigattatum Gill, and Haemulon maculicauda (Gill) are found exclusively in the Pacific while
Haemulon steindachneri (Jordan and Gilbert) is found both in the Atlantic and the Pacific.
iv Table of Contents
Acknowledgments iii
Abstract iv
List of Tables vi
List of Figures ·viii
Introduction 1
Materials and Methods 3
Systematic account 4
Discussion 26
Literature Cited 60
v List of Tables
Table 1. Frequency distribution of dorsal fin
counts ...... •.... 28
Table 2. Frequency distribution of anal and pectoral
ray counts ...... , .. 28
Table 3. Fre quency distribution of lateral line scale
counts ...... 29
Table 4. Frequency distribution of scale counts above
and below lateral line ...... 29
Table 5. Frequency distribution of caudal peduncle
scale counts ...... 30/31
Table 6. Frequency distribution of gill raker counts .. 30/31
Table 7. Frequency distribution of spines on preopercle ...... 32
Table 8. Frequency distribution of snout length in
per cent standard length ...... •.•.... 32 Table 9. Frequency distribution of body depth in
per cent standard length .....•...... •.... 33/34
Table 10. Frequency distribution of upper jaw length
in per cent standard length ...... • 33/34
Table 11. Freque ncy distribution of interocular
distance in per cent standard length 35
Table 12. Fre que ncy distribution of eye length in
per cent st a nd a rd len g th ...... •...... •• 35
Table 13. F~ e qu e ncy d istribution of c a ud a l peduncle
depth in p e r cent s t an d a rd len g th ...... 36
vi Table 14. Frequency distribution . of caudal peduncle
length in per cent standard length ...•••..•• 36
Table 15. Frequency distribution of preanal distance
in per cent standard length ..•..•.•...••..•• 37/38
Table 16. Frequency distribution of predorsal distance
in per cent standard length ..•.••••..•••.•.• 37/38
Table 17. Frequency distribution of .head length in
per cent standard length .•••.•..•.•...•.•••• 39/40
vii List of Figures
Figure 1. Haemulori ·studderi Gill. a-~juveniles; b--adults ... 41 Figure 2. Haemulon sexfasciatum Gill. a--juveniles;
b--adults ...... 42
Figure 3. Haemulon steindachneri ...... 43
Figure 4. H~emulon flavigutt~tum Gill. a--juveniles;
b--adults ...... ~ ...... 44
Figure 5. H a ernul on_ maculicauda (Gill). a--juveniles;
b--adults ...... 45 Figute 6 . Haemulon scudderi; USNM 50365 ...... 46 Figure 7. Haemulon scudderi; UCLA C50 l-151-58 ...... 47 Figure 8. Haemulon sexfasciatum; USNM 167943 ...... 48 Figure 9. Haemulon sexfasciatum; USNM 167943 ...... 49 Figure 10. Haemulon sexfasciatum, lectotype ...... 50 Figure 11. Haemulon flaviguttatum; USNN 81148 ...... 51 Figure 12. Haemulon flaviguttatum; USNM 181298 ...... 52 Figure 13. Haemulon flaviguttatum; USNH 81144 ...... 53 Figure 14. Haemulon maculicauda, syntype ...... 54 Figure 15. Haemulon maculicauda; USNM 81143 ...... 55
Figure 16. Hae mulon scudderi ...... ••.•..•...... •....•••... ~. 56
Figure 17. Haemulon sexfasciatum ...... 56 Figure 18. Ha emulon steindachneri ...... 57 Figure 19. Haemulon flaviguttatum ...... 57 Figure 20. Hae mulon ma culicaud..:L ...... 58 Figure 21. Scatter diagram ...... 59
viii Introduction
The pomadasyid genus Haernulon Cuvier consists of seventeen species found along the tropical and subtropical
American coast. Twelve are from the western Atlantic; four, including the synonymized genera Lythrulon Jordan and Swain and Orthostoechus Gill, are from th~ eastern Pacific; and one is found along both coasts of Central America.
Thirteen species of western Atlantic grunts were treated by Courtenay (1961). This present study involves the eastern
Pacific Haemulon: H. scudderi Gill, H. sexfasciatum Gill,
H . f 1 a vi gut t _at u m Gi 11 , .!:!_. mac u 1 i cauda ( G i 11) , and H . steindachneri (Jordan and Gilbert). The first four species have been recorded from Cape San Lucas and Guaymas, Mexico to Panama. H. steindachneri has been found from Guaymas,
Mexico to Panama in the Pacific and St. Lucia, Lesser Antilles to Rio de Janeiro, Brazil in the Atlantic (Meek and Hildebrand,
1925).
The taxonomy of the Pacific grunts is in a confused state. These five species were first named~· scudderii,
H. sexfascia~~~· D:Labasi~:. stein_dachn e r~, .!!_ . flaviguttatus_, and _9rthostoe_~l:!_~s ;:naculicauda in their original descriptions.
They were later all recorded as members of the genus Diabasis_ by Jordan and Gilbert (1882). From 1884 to 1893, the trend was to place all five species into the genus Haemulon. From
1895 to 1944, _!!. flav..:!_gutt:atum wa s placed in Lythrulon and
in Orthostoech~s. More recently, workers such as Courtenay (1961) feel that Lythrulon and 2
Orthostoechus should be considered synonyms of Raemulon.
The present study utilized meristic counts, morphometric measurements, notations on adult and juv~nile pigmentation, dentition pattern, and radiographs to determine the relation ship between these five species of Pacific grunts.
4
Genus Haemulon Cuvier
Diabasis Desmarest, 1823: 23 (type species Diabasis p~rra by
subsequent designation; not Diabasis Hoffmannsegg, 1819,
preoccupied in Coleoptera).
Ha·emul on Cuvie r, 1829: 175 (type species Haemulon sciurus
(Shaw) [=Haemulon elegans Cuvier] by subsequent
designation).
Orthostoechus Gill, 1863: 255 (type species Orthostoechus . ~aculic~uda Gill by monotypy).
Haemylum Scudder, 1863: 12 (emended spelling).
Anarmostus Scudder, 1863: 12 (typ e species Diabasis
flavolineatus Desmarest by subsequent designation;
not Anarmostus Loew, 1860, preoccupied in Diptera).
Bathystoma Scudder, 1863: 12 (type species Haem~lo~
auro1ineatum Cuvier [=Haemuloti jeniguano Poey, 1860]).
Haemulum Cope, 1871 : 471 (emended spelling).
Brachygenys Scudder, 1875: 121-122 (type species Haemul c n
taeniatum Poey, 1868 [=Haemulon chrysargyreum GUnther]
by monotypy).
~t h r u 1 on J o r dan an d S w a 1 n , 1 8 8 4 : 2 8 7 ( t y p e s p e c i e s H a e:_m u 1 on
flavi.E..!:!..tt a_t us Gill).
Key to the Eastern Pacific Species of Haemulon
1. Specimens 70 rom or larger in standard length ...... 2
1. Specimens smaller than 70 mru in standard length
(juveniles) ...... 6
2. Scale rows oblique, not parallel to long axis of body. 3 5
2. Scale rows parallel to long axis of body above and
below lateral line, dark blotch on sides of scales
forming lines along scale rows, usually 13 (12 + 1)
dorsal spines ...... · ma·cul:i.cauda
3. Gill rakers 26 or less, snout length usually 10%
standard length or more 4
3. Gill rakers usually 27 or 28, snout short (8% standard
length), light spot in center of scales forming lines
along scale rows, dark blotch under angle of
preopercle ...... fl. ~viguttatum
4. No bars on body, black blotch present below free
margin of preopercle ...... 5
4. Six or seven bars present on dorsal and lateral body
surfaces, no black blotch below free margin of preopercle,
black spots on side of head ...... , ... sexfasciatum
5. Scales around caudal peduncle 26 (11-2-13), caudal spoL
present , scales with dark blot~h on sides forming lines
along scale rows steindachneri
5. Scales around caudal peduncle 22 (9-2-11), caudal spot
faded or ab sent, scales with dark blotch on sides
forming line s along scale rows ...... scudde rl
6. One lateral stripe present, running from tip of snout
to caudal peduncle, continuous with caudal spot as
in Figure 14 ...... , ...... •...... maculic auda
6. More than one lateral stripe present 7
7. Two lateral stripes and a caud al spot present 8 6
7. Three lateral stripes; first from top of head to base
of first dorsal spine; second from above eye to dorsal
surface of caudal peduncle; third from tip of snout to
caudal peduncle, continuous with caudal spot as in
Figures 8 - 9 ...... ·.. ·. ·.... ·. ·. ·... ·. ·. .· ·s e·x·f ·a s·c·i at urn
8. Anal rays 7 or 8, gill rakers usually 17 to 19,
melanophore pattern as in Figure 6 ....•....•.. s~udderi
8. ·' Anal rays 10 or 11, gill rakers usually 27 or 28,
melanophore pattern as in Figures 11 - 12 .flaviguttatum
Haemulon scudderi Gill
Mojarra Prieta, Roncador Prieto
Figures la-b, 6-7, 16a-b, 21, Tables 1-17
Haemulon s cudd e_r ii: Gill, 1863: 253-254 (original description,
Cape San Lucas).
Haemulon brevirostrum: GUnther, 1869: 418-419 (description,
Panama, Puerto Cabello, figure of head only).
Haemulon undecimale: Steindachner, 1876: 11 (Acapulco,
Panama).
Diabasis sctdderi: Jordan and Gilbert, 1882: 324 (characters
in key); 1883a: 361 (compiled); 1883d: 107 (USNM catalog
numbers); 1883e: 110 (USNM catalog numbers); 1883£:
626 (name only).
Haemulon scudderi: Jordan and Swain, 1884: 285, 299-300, 317
(emendation, synonymy, characters).--Jordan, 1885: 90
(name only); 1886: 380 (distribution); 1887: 537 (name
only).--Jordan and Bollman, 1889 : 131 (Panama).--Jordan 7
and Fesler, 1893~ 465, 472 (compiled) .--Jordan, 1895:.
458 (compiled).--Jordan and Evermann, 1896: 330 (range,
synonymy); 1898: 1293, 1299-1300 (compiled).-- Boulenger,
1899: 3 (Baie de Ste. Helene).-- Gilbert and Starks,
1904: 10'•-105, 213 (characters; comparison with
H. parrai) .-- Jordan and Everman.n, 1905: 421, 425
(compiled).-- Kendall and Radcliffe, 1912: 115 (Acapulco;
Perico Island).-- Meek and Hildebrand, 1925: 531, 542-
543 (synonymy , cha;acters) .-- Jordan, Evermann, and
Clark, 1930: 330 (compiled).-- Kumada , 1937: 46, PJ. 73,
Fig. B (characters).-- Fowler, 1944: 506 (distribution).
Materia~ examined.-- Baja California, Cape San Lucas:
Lectotype, USNM 3683 (1, 82.00); Paralectotype, USNM 212148
(1, 75. 85, formerly USNM 3683).
Western Mexico: UCLA C50 W51-58 (6, 75.90-94.85); USNM
176140 (1, 166.50); USNM 65522 (1, 184.35).
Panama: UCLA C50 W58-306 (3, 66.35-86.60); USNM 144340
(1, 26.90); USNM 81087 (1, 50.55); USNM 50365 (4, 35.15-59.65);
USNM 153616 (1, 175.45).
Equador: UCLA C50 W64-38 (21, 64.65-94.20).
Unknown: USNM 181288 (2, 152.95-170.30; USNM 176148
(3, 139.10-158.25); USNM 181299 (16, 70.60-102.85).
Diagnosis.-- Lateral line scales 46 to 50, usually 49; caudal peduncle scales 20 to 24, usually 22 (9-2-11); dorsal f in
10+1 or 11+1, usually 10+1, 14 to 17, usually 16; anal fin
3, 7 to 8, usually 9; pectoral fin 15 to 20, usually 17; gill rakers 15 to 21, usually 19 . Preserved adults with dark spot 8
at scale base on dorsal and lateral surfa c e s.
D~scription.-- Adult length has 6een recorded up to two feet (609.6 mm). Body form and pattern ar e shown in Figures la and b. The posterior margin of the maxillary in adults scarely reaches the pupil. The pre op ercle is serrated. The dorsal fin co~monly contai ns 11 o r 12 spines of whi ch the last is associated with 16 soft rays. The anal fin count is usually 3 spines and 7 or 8 rays. Fin counts are given in
Tables 1 anrl 2 . Later~! line scales are usually 49 in number.
There are usually 5 or 6 scales above and 10 or 11 scales below the latera l line. Scale rows above the lateral line are oblique and the rows below are mo r e horizontal. Frequent gill raker counts are 17, 18, and 19. Scale, gill raker, and preopercular spine counts are prese nt e d in Tables 3 to 7.
Morphometric d a ta are sho wn in Tab l es 8 to 17. Figure 21 shows the relation ship between standard length, body depth, head length, and jnterocular distance. The anterior pre- maxillary tooth pattern is indented; vome rine and palatine teeth are a b sen t; and the anteri o r lower jaw is toothed continuously .
Col.or a tion.-- I n p r es e rve d juveniles, two latera l strip e s and a caudal spot are present on each side of the body. One stripe begins above the nostrils and e x tends poste rio r ly to the dorsal surface of the caudal peduncl0 . The s e cond stripe runs from the tip of the snout to the caudal peduncl e and is continuous with the caudal s p o t . As the fish matu r es~ the more ventra l pai r of stripes fa d es first, f o llowed b y 9
the more dorsal pair of stripes a nd the caudal spot.
Dark spots develop at the base of the scales on the dorsal and lateral body surface. In pre s erved adults, these spots do not form distinct lines following the scale rows. A dark blotch is present under the free margin of the preopercle.
Melanophore pattern in preserved specimens is shown in Figure s
6 and 7.
Dentition.-- The upper jaw contains only premaxillary teeth arranged in two sets. One set lines the outer edge of the mouth and is either - discontinuous or indented at the anteriormost tip. The second set forms a smooth curve just inside the first set a£ the anterior part of the mouth. The lower jaw consists of two sets of teeth which line the edge of the mouth and are continuous at the anteriormost tip.
Figures 16a and b show the arrangement of the jaw teeth.
Remark s.-- Ex c e pt for the types of H. steindachneri, the types of the other four species of Pacific Haemulon were part of a l a rge c ollection of fishes taken by John Xantus at Cape San Lucas, Mexico under the direction of the
Smithsonian I ns titution. The collection was studie d by
Theodore Gill who descr i bed new species and published th e descriptions in the Proceedin g s of the Academy of Natural
Sciences of Philadelphia in 1863. The types were stored according to the practi c e of tha t tlrr.e which involv ed th e placement of more than one fish in t he s a me container if the s p ecimens were suffici entl y small. La ter, due to confusion at the United States Na tion a l Museum where the 10
collection was kept, many specimens were scattered or lost and the study of the undescribed portion was abandoned by
Gill. In 1883, Jordan and Gilbert examined and catalogued what remained of the Xantus collection.
In the present study, USNM 3683, labelled as the types of H. ~cudd e ri, was examined. Three specimens were present in this collect ion, two of which we r e specimens of H. scudderi and one of which was a specimen of H. sexfasciatum. No reliable museum records were found on the riumber of specimens originally submitted for placement in USNM 3683, but when
Jordan and Gilbert (1883a) looked at this collection, they reported examining three fish. This uncertainty about the types of H. scudde r i and the failure of Gill (1863) to designate a holotype in his original des~ription would necessitate the choice of a lectotype from the two syntypes o f H. scudderi in USNM 3683.
I designate USNM 3683 as the lectotype of Haemulon ~udd e ri. since this specimen more closely corresponds to Gill's original descr i ption. The other specimen of H. scudderi is designated as a paralectotype and given a new catalog number,
USNM 212148. The specimen of H. s~xfasciatum is given a new catalog number, USNM 212149. 11
Haemulon. sexfasciatum. Gill Roncador Almejero
Figures 2a-b, 8-10, 17a-b, 21 , Tables 1-17
H~e~ulbn sexfasciatus: Gill, 1863: 254 (original description,
Cape San Lucas).
Ha~mulon ma~ti l osum: Peters , 1869: 705 (description, Mazatlan).
Diabasis sexfasciatus: Jordan and Gilbert, 1882: 324
(characters in key); 1883a: 361 (compil e d); 1883c: 372
(USNM catalog numbers); 1883d: 107 (USNM catalog numbers);
1883e: 110 (USNM catalog numbers); 1883f: 626 (name only).
-- Jorda n, 1884: 287 (allied with H. m~~ulosum Peters).
Ha~mulon ~e x fasciatum: Jordan and Swain, 1884: 284, 288-289,
317 (emendation, compiled).-- Jo~dan, 1885: 90 (name only);
1886: 380 (distribution); 1887: 53 7 (name only); 1889:
330 (Tres Marias Islands).-- Evermann and Jenkins, 1891:
155 (synonymy).-- Jordan and Fesler, 1893: 464, 469
(compiled).-- Jordan, 1895: 45~ (common name, char acters).--
Jordan and Evermann, 1896: 329 (range, synonymy); 1898:
1292, 1294-1295 (characters); 1905: 421-422 (compiled).--
Kendall and Radcliffe, 1912: 114-115 (Acapulco, characters ).
-- Osburn a nd Ni chols, 1916: 163 (distribution) . -- Meek
and Hi ldebran d, 1925: 531, 543-544 (range, synonymy,
chara cters).-- Jordan, Evermann, a nd Clark , 1930: 329
(compiled).-- Breder, 1936: 27 (distribution).-- Fowler,
1944: 505 (di stribut i on). 12
Material examined.-- Baja California, Cape San Lucas:
USNM 6467 (10, 66.40-88.30); USNM 212149 (1, 47.95, formerly
USNM 3683). Lectotype, USNM 3000 (1, 78.00); Paralectotype,
USNM 212150 (damaged, formerly USNM 3000).
Western Mexico: UCLA C50 W52-112 (6, 69.90-188.40);
UCLA C50 WS8-5 (18, 34.00-88.50); USNM 167158 (1, 57.35);
USNM 37151 (1, 146.55); USNM 65523 (1, 158.90); USNM 30997
(1, 167.85); USNM 167943 =UCLA C50 W55-243 (9,44.85-55.35).
Unknown: USNM 16~942 (2, 150.65-153.70); USNM 181247
(2, 72.25-72.65.
Diagnosis.-- Lateral line scales 49 to 52, usually 50; caudal peduncle scales 22 (9-2-11); dorsal fin 10+1 or 11+1, usually
11+1, 16 to 18, rarely 18; anal fin 3, 9 to 11, usually 10; pectoral fin 15 to 18, usually 17; gill rakers 18 to 22, usually
19. Preserved adults with 6 or 7 bars on dorsal and lateral surfaces; dark spots on preopercle and opercle.
Description.-- Adults grow up to 350 ~m in length. Body form and pattern are shown in Figures 2a and b. The posterior margin of the maxillary in adults reaches the anterior edge of the pup::.l. The preopercle is serrated. The dorsal fin has mostly 12 spines the last of which is associated with 16 or 17 soft rays. The anal fin consists of 3 spines and 9
10 rays. Fin counts are given in Tables 1 and 2.
Lateral line scale count is usually 50. There are 8 or 9 scales above and 12 or 13 scales below the lateral line.
Scale rows above the lateral line are oblique and the rows below are more horizontal. Frequent gill r aker counts are 13
18, 19, and 20. Scale, gill raker, and preopercular spine counts are shown in Tables 3 to 7. Morphometric data are given in Tables 8 to 17. Figure 21 presents the relationships between standard length, body depth, head length, and inter- ocular distance. The anterior premaxillary tooth pattern is indented; the vomerine and palatine teeth are absent; and the anterior lower jaw is toothed continuously.
Coloration.-- In preserved juveniles> three stripes are present on each side of the hotly. The first runs along the median line from the top of the head to the base of the first dors al spine.
The second starts above the eye, extends across the dorsal half of the body, a nd runs along the median line on the caudal peduncle.
The third stri pe originates at the tip of the snout and extends to the caudal peduncle where it is continuous with the caudal spot. As the fisL matures, the caudal spot and the ventral~ o st stripe fade first, followed by the other two stripes. Seven bars develop on the dorsal and lateral surfaces. In preserved adults, one bar is located anterior to the dorsal spine s, two are beneath the dorsal spines, one is between the dorsal spines and rays, two are beneath the dorDal rays, and one is partly below the dorsal rays and partly on the caudal peduncle. Black spots are present on the side of the head. Melanophore pattern in preserved specimens is shown in Figures 8 to 10. Dentition.-- The upper jaw contains only ?remaxillary teeth arranged in two sets. One set l i nes the outer edge of the mouth and is either disc ontinuous or indented a t the 14
anteriormost tip. The second set forms a curve with a small indentation at the anterio~ part of the mouth. The lower jaw consists of two sets of teeth which line the edge of thi mouth and are continuous at the anteriormost tip.
The arrangement of the teeth are diagrammed in Figures 17a and b.
Remarks.-- USNM 3000, labelled as the types of ~· sexfasciatum, contained two specimens when examined in the present study.
One specimen was in go~d condition while the other was damaged and very small, making the determination of characteristics difficult. The number of specimens in the original catalog entry was three. Jordan and Gilbert (1883a) reported examining one specimen in this collection. Gill
(1863) did not designate a holotype in his original description of H. s e xfasciatum. To eliminate confusion, a lectotype must be chosen from the two specimens in USNM 3000.
I designate USNM 3000 as the leciotype of Haemulon sexfasciatum since this specimen more closely corresponds to
Gill's original description. The small, damaged specimen is designated a paralectotype and given a new catalog number,
212150.
Haemulon steind~chneri (Jordan and Gilbert)
Roncador Raiado
Figures 3, 18a-b, 21, Tables 1-17
(Pacific references only; Atlantic references in Courtenay, 196 1 )
Haemulon caudimacula: Steindachner, 1876: 15 (not of Cuvier,
1829: 176 ; description ; Acapnlco, Ri o de Janeiro, 15
Rio Grande do Sul, Maranhao).
Diabasis ste~rid~chn ~ ri: Jordan and Gi lbert, 1882: 306,
322-325 (original description, Panama, Mazatlan);
1883a: 361 (compiled); 1883c: 372 (USNM catalog
numbeTs, Colima); ] 883d: 1 07 (name only); 1883e:
110 (name only).
Haemulon steindachneri: Jordan and Swain, 1884: 285, 299-300,
317 (compiled).-- Jordan, 1885: 90 (name only); 1886:
380 (distribution); 1887: 53 7 (name only); 1895: 458
(common name , characters).-- Jordan and Evermann, 1896:
330 (range, synonymy); 1898: 1293, 1301-1302 (characters).
Gilbert and Starks, 1904: 105 (Panama, allied to
H. caudimacul a of Stei ndachner).-- Jordan and Evermann,
1905: 422, 425 (compiled).-- Kendall a n d Radcliffe,
1912: 115 (Acapul c o, Perico I s land, ch a racters).--
Osburn and Ni chols, 1916: 163 (Pichilinque Bay) . --
Meek and Hildebrand, 1925: 530, 537-538 (range, synonymy,
characters).-- Jordan, Eve rmann, and Clark, 1930: 330
(compiled).-- Kumada, 1937: 46, Pl. 31 (characte r s).-
Fowler, 1944: 247-248, 506 (distribution); 1945: 22[j
(Peru).-- Courtenay, 1961: 8 6-89, Fig. 2c, Table s 1-
10 (characters, synonymy, range).-- Dahl, 1971: 236
(characters in key).
Ha~mylum flavi g utt a tus: Bean, 1881: 96 (not of Gil l , 1863:
254; name only; Colima ). 16
Material examined.-- Pacific: Western Mexico: MCZ 882 (2,
118.15-133.70; MCZ 29565 (6, 61.65-106.65); UCLA C50 W51-41
(3, 132.65-142.65); USNM 65525 (1, 156.50); USNM 46499 (2,
99.35-106.65); USNM 46628 (1, 90.70). Syntypes, USNM 28172
(1, 133.00)"; USNM 29266 (1, 133.70); USNM 29759 (1, 121.50);
USNM 29778 ( 1 , 127.90); USNM 29795 (1, 132.70.
Panama: UCLA C50 W53-282 (15, 67.76-78.55).
Equador: USNM 41199 (1, 108.50); Argosy 64 (1, 162.50).
Peru: USNM 88731· (1, 159.45).
Atlantic: Brazil: USNM 39828 (1, 152.85); MCZ 10657
(8, 81.80··1 4 3.65); MCZ 10543 (4, 154.80-170.30).
Venezuela: UMML 5008 (1, 87.75).
Unknown: USNM 47430 (2, 111.15-121.05); UMML 11590
(2, 9175-98.40); WAL 213 (4, 11.5.10-150.70); UMML 1.1583
(1, 123.10).
Diagnosis.-- Latera l line scale s 49 to 53, mostly 52 in
Pacific specime ns, 50 in Atlantic sp e cime ns; caudal peduncle scales 22 to 26, usually 26 (11-2-13); dorsal fin 10+1 to
12+1, usually 11+1, 15 to 17, usually 16; anal fin 3, 8 to
11, usually 9 ; pectoral fin 16 to 19, usually 18 in Pacific specimens, 17 i n Atlantic specimens; gill rakers 19 to 25, usually 23 in Pacific specimens, 22 to 27, usually 24 in
Atlantic specimens. Preserved adults with stripes forme d by lines through scales following scale rows; black blotch beneath free margin of preop e rcle; caudal spot present.
D~s~ r iption.-- Adults reach a length o f ab out 350 rom. Body form and pattern are shown in F igure 3 . The posterior margin 17
of the maxillary reaches the center of the eye in adults.
The preo~ercle is serrated. The dorsal fin usually con s ists of 12 spines the last of which is associated with 16 r2ys .
The anal fin contains 3 spines and mostly 9 rays. Fin counts are given in Tables 1 and 2. The most frequent lateral line scale count is 51 or 52 in Pacific specimens and 50 or 51 in Atlantic specimens. There are usually seven or eight scales above and 13 to 15 scales below the lateral line. Scale rows above the lateral line are oblique and those below are more horizontal. Frequent gill raker counts are 22 and 23 in Pacific specimens and 23 and 24 in Atlantic specimens. Scale, gill raker, and preopercular spine counts are presented in Tables 3 to 7. Morphometric data are shown in Tables 8 to 17. Figure 21 shows the relationship between standard length, body depth, head length, and inter- ocular distance. The anterior premaxillary is toothed continuously; the vomerine and pal~tine teeth are absent; and the anterior lower jaw tooth pattern is indented.
Coloration.-- In preserved specimens, there are dark blotches on the sides of the scales which form distinct lines following th~ scale rows. A black blotch is present beneath the free margin of the preopercle. The caudal spot is present and conspicuous in adults.
Dentition.-- The upper jaw contains only premaxillary teeth arranged in two sets which line the edge of the mouth and are continuous at the anteriormost tip. The lower jaw consists of two sets of teeth which line the edge of the mouth and 18
are indented at the anteriormost tip. Figures 18a and b show the jaw teeth arrangement.
R~~~rks.-- There were slight differences between Atlantic and Pacific specimens in lateral line scale, ~ e ctoral ray, and gill raker counts. Dorsal fin, anal fin, and caudal peduncle scale counts were similar. Atlantic and Pacific specimens had similar melanophore patterns.
Ha~mulon flaviguttatum Gill
Roncador, Peixe-Fonda
Figures 4a-b, 11-13, 19a-b, 21, Tables 1-1.7
Haemulon fl a viguttatus: Gill, 1863: 254 (original
description, Cape San Lucas).
Haemulon mar gari tiferum: Gunther, 1864: 147 (description,
Panama); 1869: 419, Pl. LXV, Fig. 2 (characters,
Panama).
Haemulon flavi g uttatum: Steindachner, 1876: 42 (emendation,
characters, Acapulco, Mazatlan, Altata, Panama).--
Streets, 1877: 49 (characters, Lower California).--
Jordan and Swain, 1884: 314-315 (compiled).-- Jordan ,
1885: 89 (n a me only).-- Evermann and Jenkins, 1891:
152-153 (synonymy, common name).-- Jordan and Fesler,
189 3: 468, 479-480 (compiled).
Diabasis flaviguttatus: Jordan and Gilbe r t, 1882: 324
(compiled); 1883a: 361 (compiled); 1883b: 381 (name
only); 1883d: 107 (USNM catalog numbe r s); 1883e: 110 (USNM
catalog numbe r s); 1883f: 626 (na m ~ only). 19
Haemulon flaV6guttatum: Jordan and Swain, 1884: 287, 317
(emendation, compiled).-- Jordan, 1886: 380 (compiled);
1887: 537 (name only).
Lythrulon flaviguttaturn: Jordan, 1895: 459 (characters).--
Jordan and Evermann, 1896: 331 (range, synonymy);
1898: 1312 (characters).-- Gilbert and Starks, 1904:
105-106, 213 (allied t6 L. opalescens Jordan and Starks).-
Jordan and Evermann, 1905: 420, 429-430 (figure fair).--
Kendall and Radcliffe, 1912: 115 (Acapulco, Perico
Island, chsracters).-- Osburn and Nichols, 1916: 164
(Tiburon I., Esteban I.).-- Meek and Hildebrand, 1925:
528-529 (synonymy, characters).-- Jordan, Evermann,
and Clark, 1930: 331 (compiled).-- Breder, 1936: 2 7
(characters ).-- Kumada, 1937: 46, Pl. 32 (characters).
--Fowler, 1944: 506 (distribution).
Lythrulon opalescens: Jordan and Starks, in Jordan, 1895:
459, Pl. XL (description, Maz~tlnn).-- Jordan and
Evermann, 1896: 331 (range, synonymy); 1898: 1312-
1313 (characters).-- Jordan, Evermann, and Clark, 1930:
331 (compiled).-- Fowler, 1944: 506. (distribution).
Material examined.-- Baja California, Cape San Lucas: USNM
3682 (1, 79.45). Holotype, USNM 3681 (1, 118.30).
Western Mexico: USNM 65566 (1, 75.35); USNM 181298
(lOs 53.05-65.05).
Panama: USNM 80548 (2, 207.05-239.00; USNM 81144
(4, 49.60- 101.30); USNM 81148 (1. 39.30); UCLA C50 W53-282 20
(16, 67.95-92.70); Argosy 1 (2, 197.15-198.65); USNM 80547
(1, 186.65).
Diagnosis.-- Lateral line scales 50 to 53, usually 51; c a ud a l peduncle scales 22 (9-2-11); dorsal fin 9+1 or 11+1, rarely
9+1, 15 to 18, usually 17; anal fin 3, 10 to 11, usually 10; pectoral fin 16 to 19, usually 18; gill rakers 26 to 31, usually 28. Preserved adults with l i ght spot in center of scale forming lines along scale rows.
Description.-- The maximum lengt h of adults is about 415 mm.
Body form and pattern are shown in Figures 4a and b. The posterior margin of the maxillary reaches the center of the eye in adults. The preopercle is serrated. The dorsal fin usually contains 12 spines the last of which is associated with 17 rays. The anal fin h a s 3 spines and 10 or 11 rays. Fin counts are given in Tables 1 and 2. Lateral line scale count is usually 51. There are 7 or 8 scales above a nd 12 or 13 scales below the lateral line.
Scale rows above the lateral line are oblique and thos e below are more horizontal. Frequent gill raker counts are
27 and 28. Scale , gill raker, and preopercular spin e counts are shown in Tabl e s 3 to 7. Morphometric data are pres e n t ed in Tables 8 to 17. Figure 21 shows the relationship betwe e n standard length, body depth, head length, and interocular d~stance. The anterior premillary tooth pattern is discon t inuous ; the vomerine and palatine teeth a re ab sent; a nd the anterio r lower jaw tooth pattern is discontinuous.
Coloration.-- In preserve d juveniles, a c a ud al spot and two stri p e s are pres e nt on each side of the body. One stripe 21
begins at the tip of the snout and extends posteriorly along
the median line of the caudal ~eduncle . The second stripe
originates at the tip of the scout and runs to the caudal
peduncle where it is continuous with the c~udal spo t. As the
fish matures, the stripes and caudal spot fade while the
scales on the dorsal and lateral surfaces develop a light
colored spot in the center. The spots form lines along the
scale rows in adults. A dark blotch is present under the
angle of the preopercle. Melanophore pattern in preserved
specimens is shown in Figures 11 to 13.
De~tition.-- The upper jaw cont ains only premaxillary teeth
arranged in two sets. One set lines the outer edge of the
mouth and is indented at the antcriormost tip. The second
s et lies inside the outer set and is discontinuous at the
anteriormost tip. The lower jaw consists of two sets of
teeth which line the edge of the mouth and are discontinuous
at the anteriormost tip. Figures 19a and b show the teeth
arrangement.
Remarks.-- In the present study, USNM 3681, labelled as
the type of !i_ . flavi~tat u m, was examined. This collection
contained only one specimen and thus, it is assumed to be
the holotype according to Article 73a of the International
Code of Zoological Nomenclature. 22
Haemulon maculicauda (Gill)
Roncador Rayado, Roncador Raiado
Figures 5a-b, 14-15, 20a-b, 21, Tables 1-17
Orthostoechus maculic~uda: Gill, 1863: 255 (original
description, Cape San Lucas).-- Jordan, 189.5: 460
(compiled).-- Jordan and Evermann, 1896~ 331 (range,
synonymy); 1898: 1313-1314 (characters).-- Gilbert and
Starks, 1904: 106, 213 (characters).-- Kendall and
Radcliffe, 1912: 115-116 (Acapulco, characters).--
Meek and Hildebrand, 1925 : 526 (range, synonymy,
characters ) .-- Breder ~ 1936: 27 (distribution).-
Fowler, 194 4 : .506 (distribution).
Haemulon mazatlauum: Steindachne r, 1870: 131-134, taf. VI
(description, figure good, Ma zatlan).
Haemulon ma c ulicauda: Steiudach ner, 1376: 42-43 (Acapulco,
Mazatlan).-- Jordan and Swain, 1884: 288, 315-317
(compiled).-- Jordan, 1885: 89 (name only); 1886: 3 80
(distribution); 1887: 537 (name only).-- Evermann and
Jenkins~ 1 891: 152 (synonymy, common name).-- Jordan
and Fesler, 1893: 468, 480 (compiled).-- Jordan,
Everma nn, and Clark, 1930: 331 (compiled).
Diabas i s maculicaud 3 : Jordan and Gilbert, 1882: 325 (characters
in key); 1883a: 362 (compil e d); 1883c: 372 (USNM
catalog numbers); 1883e: 110 (USN M : atalog numbers);
1883f: 626 (name only) . 23
Material e ~a mined.-- Baja California, Cape San Lucas:
Syntypes, USNM 6557 (13, 66.25-126.30); USNM 212151 (4, 77.40-
144.05, formerly USNM 3683).
Western Mexico: MCZ 29406 (1, 83.15); USNM 65602
(12, 91.20-188.70);
Panama: USNM 78112 (1, 172.60; USNM 80551 (1, 207.75);
USNM 80550 (1, 164.05); USNM 81143 (3, 73.40-90.35); Argosy
1 (1, 148.85); UMML 27333 (1, 200.55).
Columbia: Argosy 35 (1, 108.90).
Diagnosis.-~ Lateral line scales 50 t o 52, usually 51; caudal peduncle scales 22 (9-2-11); dorsal fin ll+l to 13+1, usually
12+1, 14 to 17, usually 16; anal fin 3, 8 to 11, usually 10; pectoral fin 17 to 19, usually 18; gill rakers 22 to 29, usually 24 or 25. Preserved adults with dark blotch on scGle sides forming lines along scale rows.
Description.-- Adults grow to a len g t h o f about 230 mm. Body
form and pattern are shown in Figures Sa and b. The pos t eri or margin of the maxillary in adults does not reach the pupil.
The preoper c le is serrated. The dorsal £in commonly has 13 spines the J.a s t of wh i ch is associated wi t h 16 rays. The a n al fin contain s 3 a p ines and 9 or 10 rays. F in count s are given in Tables 1 and 2. Lateral line scale count is usually 51. There are 6 or 7 scales above and 12 or 13 scales below the lateral line. Scale : ows run parallel to the long axis of the body. Frequent g ill raker counts are 24 and 25. Scale, gill r a ker, and preop e rcul ar spine counts are 24
presented in Tables 3 to 7. Morphometric data are given in Tables 8 to 17. Figure 21 presents t he relationship between standard length, body depth, head length, and interdcular distance. The anterior premaxillary tooth pattern is discontinuous; the vomerine and palatine teeth are absent; and the anterior lower jaw tooth pattern is discontinuous.
Coldration.-- In preserved juveniles, one stripe and a caudal peduncle spot ~e present on each side of the body.
The stripe runs from the tip of the snout to the caudal spot. As the fish matures, the stripe faded quickly. The caudal spot fades, but is still visible in adults. Dark blotches develop on the sides of the scales and form lines along the scale rows. Melanophore pattern in preserved specimens is shown in Figures 14 and 15.
Dentition.-- The upper jaw contains only premaxillary teeth
~rranged in two sets. One set lines the outer edge of the mouth and is indented at the anteriormost tip. The second set lies inside the outer set and is discontinuous at the anteriormost tip. The lower jaw consists of two sets of teeth which line th e edge of the mouth and are discontinuous at the anteriormost tip. Jaw teeth arrangement is diagrammed in Figures 20a and b.
Remarks.-- In the present study, it was discovered that one of the collections containing the syntypes of H. maclicauda had the same catalog number, USNM 3683, as the type of 25
H. scudderi. Since USNM 3683 has appeared in publications as the catalog number of the type of H. ~cudderi~ it is retained as the number for the lectotype of ~· s~udderi.
The coilection containing the syntypes of H. maculicauda is given a new catalog number, USNM 212151, since USNM 3683 had not appeared in any publications as the catalog number of the types of H. maculicauda. Discussion
After e~amining the data, it seems that many meristic and morphometric characters overlapped in the five species of Pacific grunts studied. Distinct differences were not commonly found. All five species possessed scaly dorsal and anal fins, which is a characteristic of Haemulon that separates the genus from other genera in Pomadasyidae.
Studies of dentition and general body structure revealed no major differences. The scatter diagram revealed that although there were no large differences between the five species, H. ~c~dderi and H. ~teindachneri were more similar
to each other than to the other three species. H. flaviguttatu~ and H. m~culicauda showed similarities to each other and differed from the other three species in having larger standard length to body depth ratios. H. ~e~fasciatum had larger head length to interocular distance ratios than the other
Pacific grunts. H. scudderi, H. sexfasciatum, H. flavigutt~tum~ and H. ~ac~licauda had at least one lateral stripe and a
caudal spot as juveniles, thus suggesting a closer relationship
to each other and to the western Atlantic Haemulon than indicated by their adult pigmentation. As adults, H. scudEeri and H. ~~eindachneri possessed melanophore patterns that were more similar to each other than to the other Pacific grunts.
!!.· se~fasciatum, being the only species of the five which had bars on the body, exhibited the most distinctive color pattern.
26 27
Certain characteristics of H. flaviguttatum and H. maculicauda varied from those of thP other three species of
Pacific grunts and this has been used as justification by some workers for the elevation of H. ~lav~guttatum to the genus Lythrulon and~ · maculicau~a to the genus Orthostoechus.
These characteristics were the relatively short snout and high gill raker count of H. flaviguattatum and the arrangement of the scale rows parallel to the long axis of the body in
H. maculicauda. However, these differences in characteristics of H. flaviguttatum and H. maculicauda from those of the other
Pacific grunts were no greater than differences found between other species within Ha6mulo~. The short snout and high gill raker count in H. flaviguttatum compared to other Pacific
Haemulon did not deserve generic status any more ti1an those same characteristics in H. striatum or H. aurolineatum compared to H. sciurus. H. maculicauda did not differ any more from the scale pattern of Haemulon than ~id H. chrysargyreum. In
H. maculicauda, the scale rows ran parallel to the long axis of th e body both above and below the lateral line while in
H. chrysargyreum, the scale rows ran parallel only below the lateral line. Thus, there would seem to be no basis for maintaining Lythrulon and Orthostoechus as genera separate from Haemulon. 28
TABLE 1
FREQUENCY DISTRIBUTION OF DORSAL FIN COUNTS
Spines Rays 10 11 12 13 14 14 15 16 17 18
H. scudderi 45 15 2 21 32 5
H. sexfasciatum 3 49 23 27 1
H. steindachn eri 1 38 1 2 26 12 (Pacific)
H. steindachneri 21 17 4 (Atlantic)
H. flaviguttatum 1 38 1 6 28 4
H. m:aculicauda 4 34 1 1 .1 3 22 3
TABLE 2
FREQUENCY DISTRIBUTION OF ANAL AND PECTORAL RAY COUN TS
··-- -·--- -· Anal Rays Pectoral Rays 7 8 9 10 11 15 16 17 18 19 _____, __... ____ .. H. ----scudderi 28 32 1 5 34 20 H. sexfasciatum 17 33 2 2 17 24 9
H. steindachneri 7 30 3 1 14 23 2 (Pacific) ---
H. steindachn0.ri 2 16 2 1 3 13 5 (Atl;mtic)
H. f 1 a vi g_u t t at u m 20 19 1 17 18 3
H. maculicauda 1 10 27 1 2 31 6 29
TABLE 3
FREQUENCY DISTRIBUTION OF LATERAL LIN E SCALE COUNTS
46 47 48 49 50 51 52 53
H. scudderi 2 6 10 24 18
H. sexfasciatum 6 24 19 2
H. steindachneri 7 12 14 7 (Pacific)
H. steindachneri 1 10 7 2 1 (Atlantic)
H. flaviguttatum 4 16 11 8
H. maculicauda 14 17 8 1
TABLE 4
FREQU ENCY DISTRIBUTION OF SCALE COUNTS ABOVE AND BELOW LAT ERAL LINE
Above Below 5 6 7 8 9 10 11 12 13 14 15
H. scudderi 23 36 20 30 8 1
H. sexfasciatum 10 41 37 12
H. steindachneri 2 21 16 1 13 16 10 (Pacific)
H. steindachneri 18 3 8 8 4 (Atlantic)
H. flaviguttatum 1 15 22 11 11 13 4
H. maculicauda 8 31 2 22 15 30
TABLE 5
FREQUENCY DISTRIBUTION OF CAUDAL PEDUNCLE SCALE COUNTS
20 20 21 22 23 (8-2-10) (9-2-9) ( 9- 2-·10) (9-2-11) (9-2-13) -~------....· --- H. scudderi 5 2 6 39
H. sexfasciatum 48
H. s t e in d a c hr.. e r ,i 5 (Pacific)
H. steindachneri .- 1 1 (Atlantic)
H. f1aviguttatun: 36
H. macu1icauda 39
TABLE 6
FREQUENCY DISTRIBUTION OF GILL RAKER COUNTS
15 16 17 18 19 20 21
H. scudderi 1 6 15 13 23 1 1
11. sexfasciatum 11 16 12 9
H. steindachneri 1 5 (Pacific)
H. steindachneri (Atlanti c)
H. f1aviguttatum
H. maculicauda 31
TABLE 5--Continued
24 24 24 25 25 26 3 0 (9-2-13) (10-2-12)------(11-2-11) (10-2-13) (11-2-12) (11-2-13) (13-2-1 '" 1
2 1 4 2 25
1 4 1 11
TABLE 6--Continued
22 23 24 25 26 27 28 29 30 31
4
10 22 2
1 6 10 2 2
4 13 7 1
3 7 8 8 7 4 1 1 32
TABLE 7
FREQUENCY DISTRIBUTION OF SP INES ON PREOPERCLE
11-15 16-20 21-25 26-30 31-35 36-· 40 41-4 5
H. scudderi 1 5 32 19 2 1
H. sexfasciatum 14 29 7 2
H. steindachneri 12 9 13 5 1 (Pacifi c )
H. steindachneri 8 7 5 1 (Atlantic)
H. f1·avi gut ta turn 3 15 17 2 2
H. macu1icauda 5 24 7 3
TABLE 8
FREQUENCY DISTRIBUTION OF SNOUT LENGTH IN PER CENT STANDA RD LENG TH
7 8 9 10 11 12 13 14 15 _____...... _...... H. scudderi 1 1 2 5 26 20 5 2
H. sexfasciatum 3 11 20 6 6 6 1
H. steindachneri 15 9 12 4 (Pacific)
H. steindachneri 4 7 4 5 (At1an tic)
H. f1aviguttatum 2 34 2 1
H. macu1icauda 2 12 18 5 2 33
TABLE 9
FREQUENCY DISTRIBUTION 01" BODY DEPTH IN PER CENT STANDARD LENGTH ----· 26 27 28 29 30 31 32
H. scudderi 1
H. sexfasci.atum 3 5 7
H. steindachn e ri 1 3 (Pacifi c·)--
H. steindachneri (A tlan t Cc)
H. flaviguttatum 2 1 5 2 2 3
H. mac u·l i cauda 1 1 5 5 4 4 2
TABLE 10
FREQUENCY DISTRIBUTION OF UPPER JAW LENGTH IN PER CENT STANDARD LENGTH
9 10 11 12 13 11+ 15 ------· H. scudderi 10 40 11
H. sexfasciatum 9 14 27 2
H. steindachneri 1 (Pacific)
H. steindachneri (Atlantic)
H. flaviguttatum 1 4 9 23 1
H. maculicauda 10 22 6 34
TABLE 9--Continued
33 34 35 36 37 38 39 40 44
2 8 2 20 16 8 4 1 .., 10 8 4 J 5 3 1 3 1
10 9 9 3 1 2 2
3 3 5 8 1
15 3 5 . 1
7 4 5 1
TABLE 10--Continued
~ -... ·----·--- 16 17 18 19 23 26 35
1
1
27 8 4
1 1 11 6 1
1
1 35
TABLE 11
FREQUENCY DISTRIBUTION OF INTEROCULAR DISTANCE IN PER CENT STANDARD LENGTH ··---- 5 6 7 8 9 10 11 12
H. scudderi 1 8 27 23 2 1
H. sexfasciatum 2 34 13 3 1
H. steindachn .?. ri 5 22 13 (Pacific) ---
H. steindachneri 1 9 10 1 (Atlantic)
H. flaviguttatum 4 9 23 3
H. IIiaculicauda 10 21 8
TABLE 12
FREQUENCY DISTRIBUTION OF EYE LENGTH IN PER CENT STANDARD LENGTH -- - 8 9 10 11 12 13 14 15 16 17
H. scudderi 2 5 5 14 24 8 3 1
H. sexfasciatum 3 1 5 14 17 7 5 1
H. steindachneri - 6 6 4 7 13 4 (Pacific)
H. steindachneri 1 1 8 4 6 1 (Atlantic)
H. flavigut: tatum - 4 2 3 3 13 11 3
H. maculicauda 3 8 16 10 2 36
TABLE 13
;FREQUENCY DISTRIBUTION OF CAUDAL PEDUNCLE DEPTH IN PER CEN T STANDARD LENGTH
8.6- 9.1- 9.6- 10.1- 10.6- 11.1- 11.6- 9.0 9.5 10.0 10.5 11.0 11.5 12.0
H. scudderi 5 8 29 13 6 1
H. sexfasciatum 2 1 12 19 8 7 4
H. ·steindachneri 2 6 16 13 3 (Pacific)
H. steindachneri 2 5 10 1 2 (Atlantic)
H. flaviguttatum 3 4 9 11 9 3
M. ma·culicauda 4 13 15 6 1
TABLE 14
FREQUENCY DISTRIBUTION OF CAUDAL PEDUNCLE LENGTH IN PER CENT STANDARD LENGTH
10 15 16 17 18 19 20 21 22
H. scudderi 1 4 8 16 21 7 5
H. sexfasciatum 4 12 18 13 4 2
H. steindachneri 5 12 12 8 3 (Pacific)
H. steindachneri 1 4 7 7 1 (At1antfc)
H. f1aviguttatum 2 4 11 9 7 5 1
H. macu1icauda 3 9 11 6 7 2 1 37 38
TABL E 1 5--Continued
·- --·-- 68 69 70 71 72 73 74 75 76 77 78 79 80
9 9 11 9 12 2 2 2
4 9 9 10 6 1 1 1
4 5 8 5 9 1 2 1
1 1 2 5 3 5 3 1
11 6 3 1
4 11 13 5 3 1 1
TABL E 16--Cont i nue d
37 38 39 40 If 1 42 4 3 44 45 46 - -· __... --- .... -- ..... ·------1 1 5 4 16 25 9 1
6 10 12 9 7 4 5
3 7 11 8 7 2 2
1 7 8 3 2
8 16 7 2
1 4 10 13 2 5 2 1 39
TABLE 1.7
FREQUENCY DISTRIBUTION OF HEAD LENGTH IN PER CENT STANDARD LENGTH
26 27 28 29 30 31
H. scudderi
H. sex·fas·ci a tum
H. steindachneri 1 (Pacific)
H. s te indachne1·i (Atlantic)
H. flavigu~:_!:Eum 1 1 1 3
H. ma:culicauda 1 40
TABLE 17--Continued
32 33 34 35 36 37 38 39 40
3 5 18 22 12 2
6 7 13 18 7 2
2 3 12 12 6 3 1
2 7 6 6
5 14 10 1 1
1 7 16 5 8 1 41
A
B
Figure 1 . Haemulon scudderi Gill. a--juveniles , from left to right, top to bottom: USNM 144340, Pearl Islands, Panama, 26.90 mm standard length; USNM 50365, P a nama , 35. 1 5 mm standard length; USNM 81087, Balboa, Canal Zone, 53.10 mm standar d length; USN M 50365, Panama , 59.65 mm stsndard :2ngth . b--adult, USNM 65522, Acapulco , Mexico, 184 . 35 mm s tandard length. 42
A
B
Figure 2. Hae~ulon s~xfas~iatum Gill. a--juveniles, from left to right, top to bottom; USNM 167943, west Las Tetas de Cabra, Sonora, Mexico, 46.60 mm standard length; USNM 167943, west Las Tetas de Cabra, Sonora, Mexico, 54.15 mm standard length; USNM 6467, Cape San Lucas, Baja California, 77.80 mm standard length; USNM 3000, lectotype, Cape San Lucas, llaja California, 78.00 rom standard length. b--adult, USNM 65523, Acapulco, Mexico, 158.90 mm standard length. 43
Figure 3. Haemulon steindachneri (Jordan and Gilbert). top: USNM 65525, Acapulco, Mexico, 156.50 mm standard length; bottom: USNM 29759, syntype, Mazatlan, Mexico, 121.50 mm standard length. 44
A
B
Figure 4. Haemulon flaviguttatum Gill. a--juveniles, from left to right, top to bottom: USNM 81148, Naos Island, Canal Zone, 39.92 mm standard length; USNM 181298, Baja California, 55.40 mm standard length; USNM 181298, Baja C~lifornia, 55.50 mm standard length; USNM 181298, Baja Californi a, 65.05 mm standard length; USNM 81144, Panama Bay, Balboa, Canal Zone, 101.30 mm standard length. b--adult, USNM 80547, Panama, 186.65 rom standard length. 45
A
B
Figure 5. Haemulon maculicaud~ (Gill) . a--juveniles, from top to bottom: USNM 6557, syntype, Cape San Luca, Baja California, 126.30 mm standard len g tla; US f M 81143 , Toboga Island, Panama, 90.35 mm standard length; USNM 6557, syntype, Cap e San Lucas, Baja Californi a , 66.60 mm standard length . b--adult, USNM 65602, Acapulco, Mexico, 131.60 mm sLandar d length. •
~ 0'\
Figure 6. Haemulon scudderi; USNM 50365; Panama; 35.15 mm standard length. . •.• # •• \ ...... ,_ .. ·'. ·'· ...... t: •• ~ ,f • .. :: ...... ,
/
Figure 7. Haemulon scudderi; UCLA C50 W51-58; east side of Pala pt., Maz&tlan, Sinaloa, Mexico; 84 . 55 mm standard length. ~
·:.:, 'l>:l\~~'""L . ' ··...... ·'·· ~;,j,~,~~~~-~ ..,. ::n~~"~' ' ··'.-J&~-""'"'~I Figure 8. Haemulon s~xfasciatum; USNM 167943; west Las Tetas de Cabra, Sonora, Mexico; 46.60 mm standard length. .p- 1..0 Figure 9. Haemulon sexfasciatum; USNM 167943; west Las Tetas de Cabra, Sonora, Mexico; 54 . 15 mm standard length . .,,'.f;' --';.:.~' ' \":. ~~p · \.11 0 ~ <------ Figure 10 . Haemulon sexfasciatum; lectotype; USNM 3000; Cape San Lucas, Baja California; 78.00 mm standard length . V'l f-' Figure 11. Ha emulon fl~viguttatum; USNM 811'•8; Naos Island, Canal Zone; 39 . 30 mm standard length. . ~-- . . . -- . .. . -__ ·:, -: ::,.:-~'~£('· •>1::-:r.f.•·-v;.:;"-•'-"•: • "'" ·" --- Figure 12. ~~emuloh fl~vigttttatum; USNM 181298; Mexico; 65.05 mm standard length. V'1 w ~· •~ . · : ; · ...: . Figure 13. Haemulon flaviguttatum; USNM 81144; Panama Bay, Balboa, Canal Zone; 101.28 mm standard length. ,,..--- V1 .:::-- Figure 14. Haemulon maculicauda, syntype; USNM 6557; Cape San Lucas, Baja California; 66.25 mm standard length. \.11 \.11 Figure 15. Haemu1on macti1icauda; USNM 81143; Toboga Island, Panama; 90.35 mm standard length. 56 I ! A ! B ~~ ')(. -----· ;oc ~ ~ )(. )( /< Figure 16. Haemulon scudderi; USNM 181299; 83.45 rom standard length. x=larger teeth, O=smaller teeth. Othel' markings of various shapes--protrusions. a--upper jaw teeth. b--lower jaw teeth. A Figure 17. Haemulon sexfasciatum; UCLA C50 ' 1) 8-5 ; south edge of Chacala, Nayarit , Mexico; 55.70 mm standard length. x=larger teeth, O=smaller teeth. Other markings of various shapes--protrusions. a--upper jaw teeth. b--lower jaw teeth. 57 A B 0 0 ( ) Figure 18. Haeroulon steindachneri; UCLA C50 W53-282; Pan ama ; 67.75 rom standard length. x=larger teeth, O=smaller teeth. Other markings of various shapes --protrusions . a--upper jaw teeth. b--lower jaw teeth. A B Figu're 19. Haemulo_l2 flaviguttatum; UCLA C50 W53-282; Ensenada Playa Grance, Islas de San Jose , Islas Perlas, Panama; 92.70 rom standard length. x=larger teeth, O=smaller teeth. Other markings of various shapes--protrusions. a--upper jaw teeth. b--lower jaw teeth. 58 A B Figure 20. Haemulon maculicauda; MCZ 29406; Acapulco, Mexico; 83.15 mm stand a rd length. x=larger teeth, O=smaller te e th . Other markings of various shapes--protrusions . a--upper jaw teeth . b--lower jaw teeth. liVlOO c PM.I~II:. HII~MU\.0., r c S CATTEll Dl~GI\AH [ SET 1 0 !!. .1i.!!ll.LSJ c SET 2 2 W. ! £ ~f''~(I01Y tn 9,60 -c SE T ) • ~. '"' H'"' crs£8.' r :SET - X ~. llfrl 'o ! l~'ATUQJ [ ~ET ~ 5 . g.. ,.,,. c. 14 qtAY,~4 c c c a,fto -c t c ( [ ( a.oo -c t 'c c c 7.20 -c c c c c c VI &.-o -c \0 c r c c c !1.60 -c t [ t [ .[ -..eo -c c [ c [ [ ,,oo -c c ~SET 1- t [ ),20 -c------c c ·------·-·-·------·------c c c c c c cA------·------·- c 2.20 z.•o SVB:l 2.6o a.ao ), oo ),zo 3,oo 3. &o 3.aa -.oo Figure 21. Scatter diagram showing the relationship between standard length/body depth to head length/interocular distance in fiv~ species of Pacific Haemulon . 60 Literature Cited Bean, Tarleton H. 1881. Check-list of duplicates of North American fishes distributed by the Smithsonian Iristitution in behalf of the United States National Museum, 1877-1880. Proc. U.S. Natl Mus., l (1880): 75-116. Boulenger, G. A. 1899. Viaggio del Dr. Enrico Festa nell' Equador e regioni vicine. Poissons de l'Equateur. Parts 1 and 2. Boll. Mus. Zool. Anat. Torino, 14 (335): 1-8. Breder, C. M., Jr. 19~6. Heterosomata to Pediculati from Panama to Lower California. Bull. Bingham Oceanogr. Coll., 2_ (3): 1-56. Cope, E. D. 1871. Contribution to the ichthyology of the Lesser Antilles. Trans. Amer. Phil. Soc., 14: 445-483, figs. 1-3. Courtenay, Walter R., Jr. 1961. Western Atlantic fishes of the genus ~aemulon (Pomadasyidae) : systematic status and juvenile pigmentation. Bull: Mar. Sci} Gulf and Carib., .U:. (1): 66-149. Cuvier, Georges. 1829, Le regne animal distribue d'apres son organisation, pour servir de base a l'histoire nature lle des animaux et d'introduction ~ l'anat omie comparee, 2 . Paris, Deterville, 406 pp. Dahl, George. 19 71. Los peces del norte de Colombia. Institute de Desarrollo do los Recursos Naturales Renovables. 391 pp. 61 Desmarest, A. G. 1823. Premiere decade ichthyologjque, ou description complete des especes de poissons nouvelles, ou imparfaitement .connues, habitant la me r qui baigne les cotes de l'ile de Cuba. Paris, J. Tastu, 391 pp., pls. 1-6. Evermann, Barton W. and Oliver P. Jenkins. 1891. Report upon a collection of fishes, made at Guaymas, Sonora, Mexico, with descriptions of new species. Proc. U.S. Natl. Mus., 14: 121-165. Fowler, Henry W. 1944. The fishes. Lancaster, Wickersham Printing Co., 57-506 . -----. 1945. Los peces del Peru. Lima. Museo de Historia Natural "Javier Prado" Universidad Nacional Mayor de San Ma rcos. 246 pp. Gilbert, Charles H. and Edwin C. Starks. 1904. The fishes of Panama Bay. Contributions to Biology from the Hopkins Seaside Laboratory of the Leland Stanford Jr. University [reprinted from Memoirs of the California Academy of Sciences, 4], 1-304. Gill, Theo oo r e N. 1863. Catalogue of the fishes of Lower California in the Smithsonian In s titution, collected by Mr. J. Xantus, Proc. Acad. Nat. Sci. Philadelphia, (1862): 249-262. Gunther, Albert. 1864. Report of a collection of fishes made by Messrs. Dow, Godman, and Salvin in Guatemala. Proc. Zool. Soc. London, 1864 (1): 144-154. 62 Gunther, Albert. 1869. An account of the fish~s of the states of Central America, based on collections made by Capt. J. M. Dow, F~ Godman, Esq., and 0. Salvin, Esq. Trans. Zool. Soc. London, i: 377-494. Hubbs, Carl. L. and Karl F. Lagler. 1958. Fishes of the Great Lakes region. Bull. Cranbrook Ins. Sci., 26, 213 pp. > 251 figs., 44 pls. Jordan, Davi.d S. 1884. Notes on American fishes preserved in the museums of London, Paris, Berlin, and Copenhagen. Proc. Acad. Nat. Sci. Philadelphia, (1883): 281-293 • ---. 1885. A catalogue of the fishes known to inhabit the waters of North America, north of the Tropic of Cancer, with notes on the species discovered in 1883 and 1884. Rep. U.S. Comm. Fish., (1884): 1-608 . ---. 1886. A list of the fishes known from the Pacific coast of tropical America, from the Tropic of Cancer to Panama. Proc. U.S. Natl. Mus., 8 (1885): 361-39/+. ---. 1889. List of fishes collected by Alphonse Forrer about Mazatlan, with descriptions of two new species--Heros beani ctnd Po ecilia butleri. Proc. U.S. Na tl. Mus., 11 (1888): 329-334. ---. 1895. The fishes of Sinaloa. Proc. Calif. Acad. Sci., 2nd ser., 1: 377-514. Jordan, David S. and Charles H. Bollman. 1889. Scientific results of explorations by the U.S. Fish Commission steamer Albatross. Proc. U.S. Natl. Mus., 12: 149-;83. 63 Jordan, David S. and Barton W. Evermann. 1896. A check-list of the fishes and fish-like vertebrates of North and Middle America. Rep. U.S. Corum. Fish., 21 (1894-1895): 207-590. ---. J898. The fishes of North and Middle America. Bull . U.S. Natl. Mus., il (2): 1241-2183. Jordan, David S. and Barton W. Evermann. 1905. American food and game fishes; a popular account of all the species found in America north of the Equator, with keys for ready identification, life histories, and methods of capture. New York, Doubleday, Page & Co., 574 pp. Jordan, DavidS.> Barton W. Evermann, and Howard W. Clark. 1930. Check list of the fishes and fish-like vertebrates of North and Middle America north of the northern boundary of Venczue]a and Columbia. Rep. U.S. Comrn. Fish., l (1928): 1-670. Jordan, David S. and Bert Fesler. 1893. A review of the sparoid fishes of America and Europe. Rep. U. S. Corum. Fish., 27 (1889-1891): 421-544. Jordan, David S. and Charles H. Gilbert. 1882. Description of nine teen n ew spe c ies of fi s hes from the Bay of Pan ama . Bull. U.S. Fish Comrn., l (1881): 306-335 . ---. 1883a. Catalogue of the fishes col l ected by Mr. John Xantus at Cape San Lucas, which are n )w in the United States National Museum, with descriptions of eight new 64 species . Pr·oc. U.S. Natl. ,Mus., 1_ (1882): 353-371. ---. 1883b. List of fishes collected at Panama, by Rev. Mr. Rowell, now preserved in the United States National Museum . Proc. U.S. Natl. Mus., 1_ (1882): 381-382. • 1883c. List of fishes collected by Mr. John Xantus at Colima, Mexico. Proc. U.S. Natl. Mus., 5 (1882): 371-372 . • 1883d. List of fishes collected at Mazatlan, Mexico, --- . by Charles Henry Gilbert. Bull. U.S. Fish Comm., 2 (1882): 105-108 • . 1883e. List of fishes collected at Panama by Charl es Henry Gilbert . Bull. U.S. Fish Comm., ~ (1882): 109-:i.ll. . 1883f. List of fishes now in the museum of Yale College, ccllected by Prof. Frank H. Bradley at Panama, with descriptions of three new species. Proc. U.S. Natl. Mus., 1 (1882): 620-632. Jordan, David S. and Joseph Swain. 1884. A review of the species of the genus Haemulon. Proc. U.S. Natl. Mus., 7: 281-317. Kendall, William C. and Lewis Radcliffe. 1912 . The shore fishes (of the Albatross expedition, 1904-1905]. Mem. Mus. Camp. Zool. Harvard College, 12_: 75-172. 8 pls. Kumada, Tosio, ed. 1937. Marine fishes of the Pacific coast of Mexico. Odawara, Nissan Fisherie .• Institute and Co., Ltd. 102 pp., 34 pls. 65 Meek, Seth E. and Samuel F . Hildebrand. 1925. The marine fishes of Panama. Publ. l"ield Mus. Zool. 15 (2): 331-707. Osburn, Raymond C. and John T. Nichols. 1916. Shore fishes collected by the Albatross expedition in Lower California with descriptions of new species. Bull. Amer. Mus. Nat. Hist., _35: 139-181. Peters, Wilhelm. 1869. Ueber neue oder weniger bekannte Fische des Berliner Zoologischen Museums. Monatsber. Akad. Wiss. Berlin, 1869: 703-711. Scudder, Charles W. 186 3. in: List of the fishes sent by the Museum to different institutions, in exchange for other specimens, with annotations, by Frederick Ward Putnam. Bull. Mus. Comp. Zool. Harvard College, 1 (1): 2-16. Steindachner, Franz. 1870. Ichthyologische notizen VIII. Sitzber. Akad. Wiss. Wien., ~ (1869): 120-139 . . 1876. Ichthyologische Beitrage III. Sitzber. Akad. Wiss. Wien., 1l:_ (1): 1-96. 8 pls. Streets, Thomas H. 1877. Contributions to the natural history of the Hawaiian and Fanning Islands and Lower California. Bull. U.S. Natl. Mus ., 7: 43-141.