Poss & Boschung 1996 Lancelets Valid Species

ISRAEL JOURNAL OF ZOOLOGY, Vol. 42, 1996, pp.
S-13-S-66
LANCELETS (CEPHALOCHORDATA: BRANCmOSTOMATIDAE):

HOW MANY SPECIES ARE VALID?
STUART G. Poss• AND HERBERT T. BoscHUNOb

•Gulf Coast Research Laboratory, Ocean Springs, Mississippi 39566 USA
bDepartment of Biology, University ofAlabama, Tuscaloosa, Alabama 35487 USA
ABSTRACT
Fifty specific and ten generic names have been applied to lancelets living in
tropical and subtropical oceans. About 29 of the names represent valid taxa,
distinguishable on the basis of meristic variation and assignable to the genera
Branchiostoma and Epigonichthys. Branchiostoma has gonads on both sides
of the body and its bilateral metapleural folds terminate immediately posterior
to the atriopore. Epigonichthys normally has gonads present on the right side
only and the metapleural fold of the left side is confluent with the anterior
terminus of the caudal fin. Traditionally, myomere counts, counts of fin
chambers, position of atriopore and anus, and qualitative differences in noto-
chord and caudal fin shape have been used to distinguish species. Multivariate
analysis of meristic variation, using primarily American species, reveals
considerable intraspecific variability in key taxonomic features. Some species
exhibit wide variation in countable segments, whereas others are character-
ized by a narrow range. This report includes primary synonymies, diagnoses,
distributions, and nomenclatural remarks for all taxa recognized.
NAMES AND CHARACTERS

Lancelets are widely distributed in tropical and temperate seas (Figs. 1-3). Adults
typically occur in sandy and shell-sand habitats, whereas larvae may be nektonic in both
nearshore and pelagic environments. The first specimen was described as a mollusk
from the shore of southern England (Pallas, 1774). Although their similarities to
vertebrates has made them familiar to students of many branches of zoology (e.g.,
Newman, 1920; Alexander, 1975; Walker, 1975), their taxonomy and systematics have
received much less attention.
Fifty specific names have been based on extant lancelets (Table 1) and about 29 are
regarded as valid by recent workers. Various fossils have been claimed to represent early
members of the Cephalochordata (Oelofsen and Loock, 1981; Butterfield, 1990; Dzik,
1993; Briggs et al., 1994; Chen et al., 1995; Inson et al., 1995). The overall group has not
S-14 S.G. POSS AND H.T. BOSCHUNG Isr. J. Zoo!.
+ B. bermudae
,. B. bennetti
A B. californiense
CJ B. caribaeum
• B. elongatum
• B. florldae
X B. longirostrum
• B. platae
0 B. virginiae
E. lucayanus
*
Fig. I. Distribution of lancelets in the western Atlantic and eastern Pacific. Position of some
symbols displaced for clarity. Some symbols represent more than one collection, while closely
adjacent localities are not always indicated.
0 B. africae
+ B. arabiae
T B. bazarutense
v B. belcheri
..•
B.capense
B. gambiense
A B. lanceolatum
T B.leonense
• B. nigeriense
B. senegalense
*
E. lucayanus
*
m E. maldivensis
-~
Fig. 2. Distribution of lance lets in the eastern Atlantic and western Indian oceans. Position of some
Vol. 42, 1996 LANCELET TAXONOMY S-15
v B. belcher!
+ B.lndicum
11 B. lanceolatum
• B. malayanum
y B. mortonense
X B. tattersalll
ID E. australis
B E. bassanus
• E. clngalensls
0 E. cultellus
H E. hectori
'It E. lucayanus
IB E. maldlvensls
Fig. 3. Distribution of lancelets in the eastern Indian and western Pacific oceans. Position of some
received worldwide revision recently (Kirkaldy, 1895; Hubbs, 1922; Franz, 1930),
although there are several important regional taxonomic studies (Whitley, 1932; Webb,
1955, 1956a, 1956c, 1957b; Azariah, 1965a; Paulin, 1977; Nishikawa, 1981;
Richardson and McKenzie, 1994).
Three families of lancelets have been proposed and distinguished by the arrangement
of the gonads. Many earlier workers placed the larval stages in their own family, the
Amphioxididae. Most recent authors have assigned extant lancelets into the
Branchiostomatidae characterized by bilateral series of gonads, and the
Epigonichthyidae, with a single asymmetric series. Here, we recognize the distinctive
genera Branchiostoma and Epigonichthys and place them into a single family. There are
two sexes and hermaphroditic individuals are not unknown (Langerhans, 1876;
Goodrich, 1912; Orton, 1914; Chen, 1931; Chin, 1941 ). Some specimens of each genus
show quite irregular arrangement of the gonads (Franz, 1922a; Chin, 1941; Nishikawa,
l980a).
Genera have typically been established on the basis of fin organization and the
arrangement of the metapleural folds, although these too vary (Nishikawa, 1980a). Also
used have been the relative size and position of the notochord, the anatomy of feeding
structures, and overall size.
Because body proportions are often strongly affected by preservation, myotome
counts have been emphasized in taxonomy. Sundevall (1852) noted interspecific
S-16 S.G. POSS AND H.T. BOSCHUNG Isr. J. Zoot.
Table I
List of nominal extant species of lance lets in alphabetical order by species with current status
Nominal species current status
Branchiostoma africae Hubbs in Monod, 1927 valid
Heteropleuron agassizii Parker, 1904 synonym of Epigonichthys maldivensis
Branchiostoma arabiae Webb, 1957 valid
Asymmetron australis Raff, 1912 valid as Epigonichthys australis
Branchiostoma bassanum Gunther, 1884 valid as Epigonichthys bassanus
Branchiostoma bazarutense Gilchrist, 1923 probable synonym of Branchiostoma
belcheri
Amphioxus belcheri Gray, 1847 valid as Branchiostoma belcheri
Amphioxus belcherijaponica Willey, 1897 synonym of Branchiostoma belcheri
belcheri
Branchiostoma belcheri tsingtauense valid subspecies as Branchiostoma
Tchang-Si and Kwang-Chung, 1936 belcheri tsingtauense
Branchiostoma bennetti Boschung and Gunter, 1966 valid
Branchiostoma bermudae Hubbs, 1922 valid
Branchiostoma califomiense Andrews, 1893 valid
Branchiostoma capense Gilchrist, 1902 valid
Branchiostoma caribaeum Sundevall, 1853 valid
Heteropleuron cingalense Kirkaldy, 1894 uncertain as Epigonichthys cingalensis
Epigonichthys cultellus Peters, 1877 valid
Branchiostoma elongatum Sundevall, 1853 valid
Branchiostoma jloridae Hubbs, 1922 valid
Bathyamphioxus franzi Whitley, 1932 synonym of Epigonichthys cultellus
Branchiostoma gambiense Webb, 1958 valid
Branchiostoma gravelyi Prashad, 1934 synonym of Branchiostoma tattersalli
Branchiostoma haeckelii Franz, 1922 synonym of Branchiostoma lanceolatum
Heteropleuron hedleyi Haswell, 1908 probable synonym of Epigonichthys
cultellus
Heteropleuron hectori Benham, 1901 valid as Epigonichth,vs hectori
Dolichorhynchus indicum Willey, 1901 valid as Branchiostoma indicum
Limax lanceolatus Pallas, 1774 valid as Branchiostoma lanceolatum
Branchiostoma leonense Webb, 1956 valid
Branchiostoma longirostrum Boschung, 1983 valid
Branchiostoma lubricum Costa, 1834 synonym of Branchiostoma lanceolatum
Asymmetron lucayanum Andrews, 1893 valid as Epigonichthys lucayanus (see text
for nomenclatural remarks)
Asymmetron macricaudatum Parker, 1904 synonym of Epigonichthys lucayanus
Branchiostoma malayana Webb, 1956 valid as Branchiostoma malayanum
Heteropleuron maldivense Forster Cooper, 1903 valid as Epigonichthys maldivensis
Branchiostoma marambaiensis synonym of Branchiostoma caribaeum
Gon~alves da Silva, 1980
Branchiostoma minucauda Whitley, 1932 possible synonym of Branchiostoma
belcheri
Branchiostoma moretonensis Kelly, 1966 valid as Branchiostoma mortonense, but
Table I continued
Nominal species current status
possible synonym of Branchiostoma
/anceo/atum
Branchiostoma Miilleri Moreau, 1875 nomen nudum; based on unpublished
manuscript name of Kr!<lyer: probable
synonym of B. caribaeum
Branchiostoma nakagawae Jordan and Snyder, 1901 synonym of Branchiostoma belcheri
Branchiostoma nigeriense Webb, 1955 valid
Asymmetron orientale Parker, 1904 synonym of Epigonichthys lucayanus
Heteropleuron parvum Parker, 1904 synonym of Epigonichthys maldivensis
Branchiostoma pelagicum GUnther, 1889 probable larval senior synonym of
Epigonichthys lucayanus (see text for
nomenclatural remarks)
Branchiostoma platae Hubbs, 1922 valid
Epigonichthys pulchellus GUnther, 1880 nomen nudum; misspelling of E. cu/te/lus
Branchiostoma senega/ense Webb, 1955 valid
Amphioxides stenurus Goldschmidt, 1905 probable larval junior synonym of
Epigonichthys lucayanus
Branchiostoma tatter.mlli Hubbs, 1922 valid
Branchiostoma tokoradii Webb, 1956 synonym of Branchiostoma nigeriense
Amphioxides valdiviae Goldschmidt, 1905 probable larval junior synonym of
Epigonichthys maldivensis
Branchiostoma virginiae Hubbs, 1922 valid
regional differences in myotome number and proposed that myotomes be counted as

those anterior to the atriopore, those between the atriopore and the anus, and those
posterior to the anus. Nearly all subsequent workers have accepted this suggestion.
Hubbs ( 1922) first differentiated species on the basis of the number of dorsal and
preanal fin-ray chambers, a terminology almost universally employed subsequently.
These structures are fin-box coeloms that are not involved in fin support, but house
nutritional reserves in the form of neutral mucopolysaccharides that may be used in
gametogenesis (Azariah, 1965b; Welsch and Schumacher, 1984; Holland and Holland,
1990, 1991b). They are hence termed fin-chambers. Other important characteristics are
body depth, the relative position of the atriopore and anus, and the extent and shape of
the notochord.
Some species, such as Branchiostoma indicum and Branchiostoma longirostrum,
have a notochord that extends well forward of the preoral hood and forms an extremely
well developed rostral process. In most other species the anteriormost myomere attaches
closer to its anterior tip of the notochord. In species of Epigonichthys, most obviously
E. lucayanus, the notochord often extends posteriorly beyond the posterior limit of the
myotomes.
Fin shape and site of fin origin are other diagnostic characters. Fins of some species
are higher and more extensively developed than in others. Whereas the shape of the
caudal fin has been widely used in diagnoses (e.g., Whitley. 1932; Bigelow and
Farfante, 1948), it has been shown to be markedly variable in some species (Boschung
and Gunter, 1962; Azariah, 1965a); we lack quantitative studies of variablity defining
shape differences.
The number of pharyngeal slits and oral cirri have also been referred to by early
workers. Although useful, both exhibit ontogenetic variation in number and anatomical
arrangement (Webb. 1958b; Wickstead, 1964a; Nishikawa, 1980a).
There appear to be two different paths of development available to larvae, each with
different ecology (Wickstead and Bone, 1959; Wickstead, 1964a; Boschung and Shaw,
1988). An "amphioxides-type" characterizes larvae that live for extended periods in the
plankton, some at least migrating vertically each day. Such larvae may become rela-
tively large. "Amphioxus- type" larvae are characterized as smaller and many live on
the bottom, typically on sandy or sand shell bottoms along with the adults. Although
these types are not entirely distinct (Wickstead, 1964b), the number of postanal myo-
tomes increases with time of planktonic existence (Wickstead, 1964b: Nishikawa, 1981 a).
Morphological features associated with the two developmental paths seems to vary.
Variants may represent extremes of a potential continuum of environmentally induced
developmental alternatives. Not only is larval taxonomy complicated by gross morpho-
logical differences between adults and larvae, but there is the problem of matching them
(e.g .. Wickstead, 1964a; Nelson, 1968; Gosselck and Spittler, 1979). For many species
this issue, which has come to be known as the "Amphioxides problem," has never been
adequately resolved.
This report focuses on the evaluation of meristic data from numerous species,
particularly from those of the Americas, as a step toward resolving the question of just
how many species of lance lets there are. Additional understanding of morphological and
taxonomic differentiation among lancelet species worldwide will require better and
more cosmopolitan sampling. Observations based upon morphometric variation also
need to be correlated with molecular data.
METHODS
Data for six characteristics were tabulated for I, 724 specimens. using a database.
Countable features included: I) the number of myotomes anterior to the atriopore, 2) the
number of myotomes between the atriopore and anus, 3) the number of myotomes
posterior to the anus, 4) the number of dorsal fin-chambers, 5) the number of preanal fin-
chambers, and 6) the total number of myotomes. Counting methods followed Boschung
and Gunter ( 1966). being recorded separately for each specimen. Lengths are total from
anterior tip of notochord to posterior tip of caudal fin. or notochordal tip. Other
measurements followed methods of Webb ( 1955) and Boschung and Gunter ( 1962) or
were extracted from the literature. They are summarized in the diagnosis of each
species.
Vol. 42. 1996 LANCELET TAXONOMY S-19
The number of myotomes and fin-chambers did not show sexual dimorphism and the
counts of all specimens were lumped for multivariate analysis. Counts are tabulated for
a number of variables in Tables 2 and 3. which also include comparisons of values
obtained by others.
Repeatable comparisons require known materials. Lancelet taxonomy suffers from
the notable lack of reference to specific museum lots. Because species of lance lets have
sympatric and often restricted distributions. it is often difficult to ascertain whether more
recently collected material is the same as that available to previous authors. Works of
primary relevance to nomenclatural, taxonomic. and biogeographic status are listed in
the synonymies. A database. developed from collections provided by colleagues and
references to museum materials in the literature. was used to develop summary distribu-
tion maps for each species. Unfortunately. much material is of uncertain provenance.
The museums of the world contain numerous unidentified lots that remain to be
examined.
Symbolic codes for archival institutions follow Leviton et at. ( 1985 ). as emended by
Leviton and Gibbs ( 1988). The code for Naturhistorisk Riksmuseet in Stockholm has
been changed to NRM. Two additional collections are SMBL (Seto Marine Station
Laboratory) and NZOI (New Zealand Oceanographic Institute). Specimens from which
we gathered data are indicated by an asterisk preceding the catalog number. We also
repeat counts from Nishikawa (1981a). Specimens with uncertain identification are
indicated by a question mark. Most material examined by us derived from the New
World. Although some Asian. European. African. and Indo-Pacific materials were
available. much of distributional information reported for species from these areas
derives from the literature.
Countable features were analyzed using principal components computed from cor-
relations among standardized variables. Data for total myotome number were omitted
as they provided no information not already carried in the three-part myotome formula;
also preliminary analysis showed their inclusion leads to nearly identical clustering.
Principal component analysis was used to identify those counts contributing the most to
total variation and to establish the correlations among counts most important in
discriminating among taxa. Each count is treated as an axis of variation in a five-
dimensional algebraic space. The method transforms these axes into new axes. The first
principal component axis is constrained to represent the axis of maximum sample
variance. The second principal component axis characterizes maximal residual vari-
ance independent or orthogonal to the first. Computation of subsequent axes or compo-
nents is similarly constrained; most of the original total variation is represented in
fewer summary axes that are easier to visualize. The new axes are linear combinations
of the original variables. Correlations between the original variables and the new
summary axes are represented as factor loadings. These are typically presented in
tabular fashion emphasizing those original variables that contribute most or are most
coincident with a given principal component. This well-established ordination proce-
dure and its application in taxonomic analysis is introduced by Blackith and Reyment
(1971 ).
VJ
Table 2 '0->
Summary of meristic variation among species of Branchiostoma 1
Species Preatriopore Atriopore-anal Postanal Total Dorsal Preanal Source(s)
myotomes myotomes myotomes myotomes fin-chambers fin-chambers
Branchiostoma africae -43 -14.5 -12.5 -70 -300 -52 Hubbs in Monod (1927);
(42-44) (14--15) (11-14) (67-73) (-300) (50-54) Webb (1955)
Branchiostoma arabiae 38.0 16.3 10.7 65 310 77.7 This work
(n = 3) (37-39) (16-17) (10-11) (64--66) (308-312) (74--83)
39.0 16.4 9.4 64.8 333 74 Webb (1957a);
(n = 25;3) (36-41) (16-18) (8-11) (61-67) (297-366) (65-87) Dawson (1964) VJ
Branchiostoma bazarutense - - -340? 62 Gilchrist ( 1923 ); p

.,
(34--42) (15-17) (9-10) (62-69) (250-340) (35-94) Nishikawi & Mukai (1986) 0
VJ
VJ
Branchiostoma belcheri 38.0 16.8 10.4 65 290 58 Present study ;J>
Japan (B. b. tsingtauense) (35--40) (14--21) (9-12) (61-69) (253-322) (47-69) +Nishikawa & Mukai( 1986) z
0
(n =52) & China (Amoy) X
China (Amoy) (n = 148) 36.8 17.0 10.2 64 Boring and Li (1932) ~
t:J:I
(35-38) (16-18) (8-11) (62-65) - - 0
VJ
China (Amoy) (n =50) 38 17 10 65 240 72 Chin (1941) n
X
(36-39) (16-18) (9-11) (62-66) (222-298) (35-90) c:::
z
Webb (1957a) Cl
China (Amoy) (n = 25) 36.5 17.0 10.0 64 305 78
- - - -
China (Beibu Gulf= N. Gulf 37 17 II 65 316 80 Yingya eta!. (1986)
ofTongking) (n = 2,016) (36-39) (16-18) (10-13) (64--67) (284--353) (67-95)
China (B. h. tsingtauense) 39 18 10 67 316 60 Tchang-Si & Koo (1936)
(37--40) (16-18) (10-12) (65-69) (291-360) (41-80)
Sri Lanka (n =58) 37.9 17.4 9 64 - - Tattersall (1903b)
(37-39) (17-18) (8-10) (63-65) - - ~
Africa (Linga-linga) 37.1 17.7 9.9 65 287 81 Webb (1957b) ~
(n=9) (36-39) (17-19) (9-11) (63-66) (258-306) (70-88) ~

f2.
1
Mean values followed by range in parentheses.
Table 2 continued <
[2..
..,.
Species Preatriopore Atriopore-anal Postanal Total Dorsal Preanal Source(s) .""
myotomes myotomes myotomes myotomes fin-chambers fm-chambers ::0
\0
0\
Mozambique (n =I) 34 17 11 62 276 68 Webb (1957b)
(34) (17) (II) (62) (276) (68)
Philippines (n = 191) 38.2 16.5 10.3 65 301 78 Carandang (1978)
(30-44) (11-21)? (7-13) (50-75) (266-398) (44-111)
Thailand (n = 2) 36.5 18.5 10 65 301 78 Piyakamchana ( 1962)
(36-37) (18-19) (10-10) (64--66) (289-314) (72-84)
Australia (n = 26-27) 37.4 16.7 9.9 64 297 71 Richardson & McKenzie
(27-34) (15-20) (8-12) (60-69) (266-323) (63-91) (1994) t""
Branchiostoma bennetti 35.6 16.1 8.6 60 293 47 This work >

z
n
(n = 133) (34-38) (13-19) (7-10) (56-65) (189-359) (24-77) tTl
Branchiostoma bermudae 34.9 14.0 6.6 55 225 23 This work ~

(n = 38) (34-36) (11-16) (6-10) (53-59) (189-254) (16-33)
...,
Branchiostoma califomiense 44.3 16.7 9.1 70.1 355 44 This work ~
0
(n =57) (42-47) (13-19) (7-11) (64-78) (317-419) (35-59) z
0
Monterey to Gulf of Calif. - - - - - - Hubbs (1922) =:::
-<
(n = 5;22) (42-45) (16-19) (8-10) (65-71) (312-347) (-50)
California (n = 27) 45.1 17.4 8.3 72 340 32 This work
(42-53) (16-20) (8-9) (67-81) (318-381) (20-45)
Mexico (n = 23) 44.0 16.3 9.5 70 359 52 This work
(42-46) (13-20) (9-10) (64-74) (281-399) (31--68)
Guatemala to Panama 45.3 16.6 9.3 71 379 44 This work
(n = 7) (43-49) (15-18) (10-11) (66-78) (317-419) (30-56)
Branchiostoma capense 46.0 19.2 10.1 75 421 71 Webb (1957b)
(n = 3) (44-48) (18-20) (10-11) (73-77) (400-440) (62-80)
Branchiostoma caribaeum 36.9 14.9 8.9 61 295 32 This work en
(n = 164) (34-39) (11-18) {7-10) (55--65) (228-341) (22-50) ,:.,
Table 2 continued en
.:.,
w
Brazil 37.2 15.0 8.9 61 298 32 This work
(n = 94) (36-38) (14-17) (8-10) (58-65) (232-335) (22-50)
Brazil (B. marambaiense) 35.3 13.9 8.5 58 286 33 Gon~ales da Silva ( 1980)
(n =50) (32-41) (11-18) (6-10) (52-63) (248-320) (22-42)
Puerto Rico (n = 10) 35.6 15.5 8.6 60 246 36 This work
(34-37) (14-17) (7-10) (55-62) (228-292) (33-48)
Venezuela (n = 40) 36.5 14.0 8.8 60 304 32 This work en
p
(36-37) (14-16) (8-10) (59-60) (277-341) (24-43) "0
0
Branchiostoma elongatum 48.6 17.8 12.1 78 404 61 Franz (1930); en
en
(n =41) (45-54) (14-21) (10-14) (75-86) (367-443) (37-75) Webb (1957b) >
16.8 10.6 78 420 40 This work
z
Galapagos (n = 5) 50.2 0
(49-52) (15-18) (11-12) (76-79) (384-443) (37-43) ::c
;..,
Galapagos (n = 10) 48.9 18.3 10.1 77 384 53 Webb (1957b) 0:1
0
(47-50) (17-19) (9-11) (75-79) (356-420) (40-75) en
n
Peru and Chile (n = 36) 48.7 18.1 12.3 79 402 64 This work ::c
c:::
(45-54) (15-21) (12-14) (75-86) (367-431) (43-75) z
Cl
Peru (n = 16) 49.0 18.7 12.3 80 381 64 Webb (1957b)
(47-51) (18-20) (11-13) (77-84) (350-410) (37-80)
Branchiostoma floridae 36.0 15.8 8.4 60 286 41 This work
(n = 378) (33-40) (13-19) (6-11) (56-64) (206-349) (21-67)
Mississippi Sound 36.1 15.8 6.7 59 307 47 Boschung & Gunter
(n = 100) (35-37) (14-17) (6-8) (57-60) (252-359) (35-61) ( 1962)
Tampa Bay (n = 100) 35.2 15.9 7.9 59 302 46 Howell (1964)
(34-37) (15-17) (7-9) (57-61) (271-337) (35-56)
~
""'
Branchiostoma gambiense 45.3 13.9 11.8 70 362 47 Webb (1958d) N
0
(n = 17) (43-46) (12-15) (11-12) (68-72) (318-394) (39-56) ~
Table 2 continued <
~
Species Preatriopore Atriopore-anal Postanal Total Dorsal Preanal Source(s) ,!"
""""
myotomes myotomes myotomes myotomes fin-chambers fin-chambers ::0
Branchiostoma indicum 41 14 14 69 326 49 Azariah (1965a) "'a-
(n = 40) (39-43) (13-15) (13-15) (67-72) (304-395) (40-63)
Branchiostoma lanceolatum 36.1 13.8 10.7 61 212 34 This work
(n = 11) (34-38) (11-17) (10-13) (59-65) (183-237) (25-47)
Plymouth (n = 27) 36.0 14.3 12.4 63 238 38.6 Webb (1956b)
(35-37) (13-16) (10-14) (61-65) (210-270) (30-48)
Jersey (n = 16) 36.0 14.4 12.5 63 238 37.3 Webb (1956b)
(34-37) (13-16) (11-14) (61-65) (220-250) (29-42) r
Kattegat (n = II) 35.2 14.3 11.2 227 34.2 Webb (l956b)
>
60 z
(')
(34-37) (13-16) (10-13) (58-63) (210-237) (31-38) tTl
r
North Sea (n = 5) 36.2 14.4 12.4 63 227 38.4 Webb (1956b) !::l
(35-38) (13-15) (12-13) (62-64) (215-224) (35-42) ~
)<
Naples (n = 13) 34.7 14.3 11.5 61 218 35.5 Webb (1957b) 0
(33-36) (13-16) (10-13) (59-62) (202-238) (32-42) z
0
Mediterranean & Black Sea - - - - - Nogueira ( 1969) 3::
-<
(n = ?) (35-37) (14-15) (11-12) (61-62) (227-247) (32-40)
Sri Lanka (n = 10) 36.2 12.5 11.7 60 - - Tattersall (1903b)
(35-37) (11-14) (11-13) (59-61)
W. Indian Ocean 34.5 13 10 58 275 33 Webb (1957b)
(Mozambique) (n = 2) (34-35) (13) (10) (57-58) (270-280) (32-34)
India (n = 100) 35 13 13 61 252 48 Azariah ( 1965a)
(35-37) (ll-14) (11-14) (59-65) (221-288) (30-62)
Australia (B. minucauda) 37 14 12 63 238 55 Whitley (1932);
(n =I) (37) (14) (12) (63) (238) (55) Kelly (1966)
Australia (B. minucauda) 33.7 14.0 12.3 60 224 62 Richardson & Cll
(n = 27-29) (32-37) (12-15) (11-14) (57-64) (190-265) (48-76) McKenzie (1994) t..J
....,
In
Table 2 continued
~
Branchiostoma /eonense 43.2 15.5 10.4 69 382 58 VVebb(l956c; 1958a)

(n = 25) (40-45) (14-16) (10-12) (66-73) (355-tl8) (51~)
Branchiostoma /ongirostrum 36.1 16.2 9.1 61 242 40 This work

(n = 133) (34-38) (15-19) (8-10) (60-63) (209-269) (26-53)
Branchiostoma malayanum 28.3 13.8 9.8 52 201 53 VVebb (1956a);
(n = 2; n = 4) (28-29) (13-16) (8-11) (51-53) (189-221) (42-58) Gibbs & VVickstead ( 1969)
In
Branchiostoma moretonense 32.2 17.9 8.1 58 235 79 Kelly (1966) p
(n = 30) (32-33) (17-19) (7-10) (57-60) (207-262) (52-92) cg
Branchiostoma moretonense 31.3 17.4 9.6 58 211 78 Richardson & McKenzie In
In
(n=9-ll) (28-33) (15-20) (8-10) (56-60) (197-232) (67-87) (1994) >
z
Branchiostoma nigeriense 41.8 15.2 10.9 68 346 55 VVebb (1955) c
(39-44) (14-16) (10-12) (65-71) (330-376)
:r:
Lagos (n = 100) (50-58) ~
Branchiostoma nigeriense 41.7 15.6 11.0 68 343 54 VVebb (1956c) =
0
Pon Harcoun (n = 10) (40-43) (15-16) ( 10-12) (66-70) (330-360) (50-56) ~
Branchiostoma p/atae 37.0 15.0 6.8 59 278 35 Sawaya & Carvalho :r:
c
(n =64) (34-42) (9-17) (5-9) (55-65) (249-327) (19-53) (1950) z
0
Branchiostoma senega/ense 40.2 15.7 11.3 67 293 56 This work
(n = 17) (39-43) (13-17) (10-12) (65-69) (266-317) (47-68)
Branchiostoma senegalense 40.2 17.0 11.4 69 296 52 VVebb (1955; 1958d)
(n =40) (39-41) (16-18) (10-12) (67-71) (267-325) (45-59)
Branchiostoma tatter.talli 37.0 16 9.0 62 317 78 This work
(n = 2) (37) (16) (8-10) (62) (300-333) (68-87)
Branchiostoma tattersal/i 38 16.9 8.4 64 319 80? Azariah ( 1965a) ;;-
(8-9) :"'
(n = 7) (37-40) (16-18) (62-65) (310-330) (73-78?) ~
Branchiostoma virginiae 36.1 16.1 8.4 61 302 48 This work
(n = 154) (32-38) (12-18) (7-10) (56-64) (219-344) (29-66) ~
Table 3 <
Summary of meristic variation among species of Epigonichthys 1 ~
]'>
""'"
Species Preatriopore Atriopore-anal Postanal Total Dorsal Preanal Source(s) ::0
myotomes myotomes myotomes myotomes fin-chambers fin-chambers ""a-
Epigonichthys australis -33 -9 -13 55 - - Raff(l912); Scott (1962);
(n =I) (-33) (-9) (-13) (55) - - Gomon, et al. ( 1994)
Epigonichthys australis 31.6 8.1 14.3 54 189 18* Richardson & McKenzie
(n =13*,21) (29-34) (7-9) (13-17) (51-57) (158-212) (14-25) (1994)
Epigonichthys bassanus - - - - - Giinther ( 1880)
(n =?) (43-45) (13-15) (17-18) (75-76)
Epigonichthys bassanus 42.6 15.9 15.1 74 240 41 Richardson & McKenzie
r
(n =10-24) (39-46) (13-19) (13-18) (70-79) (206--275) (35-46) (1994) >
z
(j
Epigonichthys cingalensis 39 15 9 63 309 46 This work til
(39) (15) (9) (63) (309) (46) r
(n =I)? til
-l
Kirkaldy (1895 ); -l
(n =8) (38-39) (16--17) (6--8) (61--64) - - Nishikawa & Mukai( 1986) >
><
0
38.8 16.1 8 64 - - Tattersall (1903b) z
0
(n 19) = (37-39) (16--17) (8) (61--64) - - 3:
Epigonichthys cultellus 31.0 11.5 8.8 51 220 16 This work
-<
(n 19) = (30-32) (10-12) (8-10) (50-52) (201-239) (8-22)
28.6 12 9 50 201 16 Gibbs & Wickstead
(n=29) (27-30) (11-13) (9-10) (49-52) (182-228) (13-19) (1969)
29.7 11.2 8.4 49 211 24 Richardson & McKenzie
(n =54-66) (27-34) (9-13) (6--11) (46--54) (180-254) (14-31) (1994)
(as B.franzi) 26 15 7 48 229 - Franz (1922b)
(n I) = (26) ( 15) (7) (48) (229)
Epigonichthys hectori - - - - - - Paulin ( 1977)
(n = ?) (53-55) (19-20) (11-12) (84-85) (296--321) (44-49)
Vl
1 ;_,
Mean values followed by range in parentheses. VI
Cll
Table 3 collfinued N
0\
Epigonichthys lucayanus 36.5 15.9 7.7 60 307 0 This work
(n=l51) (35-52) (13-18) (6-9) (55-62) ( 167-484) 0
Epigonichthys lucayanus - - - - - 0 Nishikawa & Mukai ( 1986)
(n = ?) (38-46) (6-10) (7-14) (56-69) (142-214) 0
Australia 42.8 9.2 12.0 64* 133 0 Richardson & McKenzie
(n = 13.14*) (39-46) (8-12) (9-15) (58-71) ( 101-170) 0 (1994)
Red Sea - - - - 195 - Steinitz (1962) Cll
(n = ?) (41-44) (6-8) (7-10) (56-59) (195) - p

Persian Gulf 38.5 6.5 14 59 128 - Dawson (1964) C3
Cll
Cll
(n = 2) (38-39) (6-7) (12-16) (57-61) (114-142) - >
z
Western Atlantic - - - - - - Bigelow & Farfante 0
(n = ?) (42-46) (8-9) (11-14) (62-68) (170-180) - (1948) :c
Western Pacific 43.3 9.5 ~
12 64.5 137 0 Nishikawa ( 1980a) CD
Japan (n = 3) (42-44) (9-10) (12) (63-66) ( 101-170) (0) 0
Cll
(")
Epigonichthys maldivensis 42.3 12.8 11.8 67 285 36 This work :c
~
(n = 4) (40-44) (8-14) (11-12) (65-69) (200-331) (33-39) z
Cl
Hawaii 43 15 15 73 - - Eldredge (1967)
(n =I) (43) (15) (15) (73)
Japan 41 14 13 68 350 38 Nishikawa ( 1980a)
(n = )) (41) (14) (13) (68) (350) (38)
Madagascar 42 15 12 69 285 40 Masse ( 1964)
(n = ?) (42) (15) (12) (69) (285) (40)
Maldives - 16 - 68 - - Forster Cooper ( 1903)
(n = 4) (45-46) (16) (12-13) (68) o;
- - :"'
~
Zanzibar - - - - - - Punnett (1903)
(n = 45) (42-46) (15-17) ( 11-14) (70-76) - - ~
MULTIV ARIATE ANALYSIS

Although determination of the number of species present requires comparisons on a
specimen-by-specimen basis, computation of principal component analyses of meristics
based on character means for alllancelets provides an instructive preliminary overview.
Analysis of taxon means reveals that species are not uniformly distributed in the
morphospace defined by countable features (Fig. 4). Although Branchiostoma and
Epigonichthys differ notably with regard to the arrangement of their gonads and
metapleural folds, meristic variation does not cluster species into groups corresponding
to these genera. However, myotome formulas among species assigned to Epigonichthys
are generally more distinct than those among species of Branchiostoma. Because of lack
of adequate materials of most species of Epigonichthys, variation in this genus cannot
yet be discussed in detail.
As can be seen in the composite ordination for all specimens countable for all
variables (n = 680), those of Branchiostoma cluster in four major groups (Fig. 5; Table
4). High counts clearly distinguish a group of specimens identifiable as B. elongatum
and B. capense (Table 2), confirming the similarity of these species noted by Webb
(1957b). Although similar, B. elongatum usually has one to two more myotomes
-.
0
-- q
0
"
g.
.,.-.
Fig. 4. Principal component analysis of lancelet species based on the mean values of counts as
tabulated in Tables 2 and 3: I, B. africae; 2, B. arabiae; 3, B. bazarutense; 4, B. belcheri;
5, B. bennetti; 6, B. califomiense; 7, B. caribaeum; 8, B. floridae; 9, B. indicum; 10, B. leonense;
11, B. longirostrum; 12, B. nigeriense; 13, B. platae; 14, B. senegalense; 15, B. virginae. Other
Branchiostoma species are identified by names beside unnumbered circles; Epigonichthys spe-
cies, unnumbered squares.
S-28 S.G. POSS AND H.T. BOSCHUNG Isr. J. Zoo!.
Fig. 5. Plot of first three principal components of all Branchiostoma specimens available to this
study: B. bermudae, plus symbols; B. califomiense, open triangles (California), greyed triangles
(Mexico}, and solid triangles (Guatemala to Panama); B. capense open squares; B. elongatum,
solid squares (Chile and Peru), and greyed squares (Galapagos Is.); all other lancelets solid circles.
Some symbols may represent more than one specimen (n = 680).
Table4
First 3 meristic principal component loadings for comparison of all Branchiostoma specimens
PCI PC II PC III
Eigenvalues 2.764 0.800 0.7693
Cumulative percent of trace 55.3% 71.2% 86.6%
Preatriopore myotomes 0.8200 0.3117 0.3668
Myotomes between atriopore and anus 0.6786 -0.6014 0.0523
Postanal myotomes 0.6809 0.5025 -0.4696
Dorsal fin-chambers 0.7952 0.0299 0.4231
Preanal fin-chambers 0.73207 -0.2913 -0.4821
Vol.42,1996 LANCELET TAXONOMY S-29
anterior to the atriopore and one or two more myotomes posterior to the anus, and one
fewer between the atriopore and the anus than does B. capense. [Since Lankester
(addendum to Kirkaldy, 1895), B. elongatum has often been erroneously regarded as
conspecific with B. califomiense.] A second, although contrasting cluster with low
counts, is assignable to B. bermudae. Two larger clusters contain materials of the other
species. The first can be identified as B. califomiense. The other contains all other
nominal forms of Branchiostoma here analyzed.
Meristic variation along the second PC axis associated with the B. califomiense
cluster is greater than that for the other lancelet species here recognized. This reflects the
relatively large variation in counts for myotomes between the atriopore and anus,
postanal myotomes, and preanal fin-chambers for this species (Fig. 6; Tables 2, 5).
There are more preatriopore myotomes and myotomes between the atriopore and anus
and fewer preanal fin-chambers in some California specimens than in many specimens
from Central America. Nonetheless, the sample shows no identifiable patterns of
geographic variation or other features suggesting that it comprises more than a single
variable species.
Species from regions other than the Eastern Pacific vary less than does
B. califomiense. Collectively, these species approach the degree of variation seen in
B. califomiense. However, separate analysis (Figs. 6-8; Tables 5-7) removing variance
due to more extreme Eastern Pacific species discloses several well-defined subclusters
that suggest regional differentiation, a grouping pattern which is not simply clinal as
geographically proximate clusters are not always phenetically proximate. Fig. 6 shows
specimens of B. lanceolatum and B. senegalense falling at nearly opposite sides of the
ordination. In contrast, the geographically disparate B. arabiae, B. belcheri,
B. senegalense, and B. virginiae all cluster to one side. This suggests that at least part of
the observable difference derives from factors other than geographic variation. Such
regional differentiation seems largely to reflect species-specific differences.
Other species show complex and not easily explained regional differentiation. For
example, most specimens of B. jloridae and B. virginiae are easily distinguished (Figs.
7,8; Tables 2, 6, 7), but a small fraction cannot be discriminated by meristic variation
alone. When describing B. virginiae, Hubbs (1922) noted that it "seems not improbable
that virginiae and floridae will be found to intergrade". However, we find that most
specimens of B. virginiae can be distinguished from specimens of B. floridae using only
myotome counts.
The identity of lancelets here assigned to B. caribaeum is not fully understood. Most,
but not all, specimens from Brazil can be distinguished from both B. jloridae and
B. virginiae by having 1-4 fewer myotomes between the atriopore and the anus, by
having about 8-16 fewer preanal fin-chambers, and often one more postanal myotome.
The same is true for specimens from Venezuela and Aruba, which differ from those
from Brazil in often having one more myotome between the atriopore and the anus,
averaging about 6 more dorsal fin-chambers, and usually one fewer postanal myotome.
However, specimens from Puerto Rico average 4 more preanal fin-chambers and
average about 40 fewer dorsal fin-chambers than do Brazilian and Venezuelan ones.
Fig. 6. Plot of first three principal components of Branchiostoma specimens. minus materials of
B. califomiense, B. elongatum, and B. bermudae; B. arabiae, circle with plus symbol; B. belcheri,
inverted open triangles; B. bennetti, solid inverted triangles; B. caribaeum, plus symbols (Aruba),
open squares (Brazil), greyed squares (Puerto Rico), and solid triangles (Venezuela); B.floridae,
solid circles; B. lanceolatum, open triangles; B. longirostrum, X; B. platae, solid diamond;
B. senegalense, open diamonds; B. virginiae, open circles. Many symbols may represent more
than one specimen (n = 535).
Fig. 7. Plot of first three principal components of western Atlantic Branchiostoma, except
B. bermudae: B. bennetti, solid inverted triangles; B. caribaeum, plus symbols (Aruba), open
squares (Brazil), greyed squares (Puerto Rico), and solid triangles (Venezuela); B.floridae, solid
circles; B. longirostrum, X; B. platae, solid diamond; B. virginae, open circles. Some symbols
=
may represent more than one specimen (n 411 ).
Vol. 42,1996 LANCELET TAXONOMY S-31
Fig. 8. Plot of first three principal components for all specimens of B. caribaeum, open squares
(Brazil), plus symbols (Aruba), grayed squares (Puerto Rico), and solid triangles (Venezuela);
B. floridae, solid circles; B. virginae, open circles. Some symbols may represent more than one
specimen (n = 243).
Table 5
First 3 meristic principal component loadings for comparison of all Branchiostoma specimens,
minus materials of B. califomiense, B. elongatum, and B. bermudae
PCI PC II PC ill
Eigenvalues 1.595 1.393 1.003
Myotomes between atriopore and anus 0.6827 -0.2512 -0.4540
Dorsal fin-chambers 0.4187 -0.5528 0.6409
Preanal fin-chambers 0.8675 0.0142 -0.1763
Table 6
Meristic principal component loadings for comparison of western Atlantic Branchiostoma,
exceptB.bermudae
PCI PC II Pcm
Eigenvalues 1.828 1.207 0.962
Myotomes between atriopore and anus -0.7944 -0.1695 -0.1978
Postanal myotomes 0.4003 -0.4353 0.7170
Dorsal fin-chambers -0.2769 0.8027 0.4349
Preanal fin-chambers -0.8559 0.0043 0.3697
S-32 S.G. POSS AND H.T. BOSCHUNG Isr. 1. Zool.
Table 7
Meristic principal component loadings for all specimens of B. caribaeum, B. floridae, and
B. virginiae
Eigenvalues PC I PC II PC III
1.840 1.302 0.775
Myotomes between atriopore and anus -0.7839 -0.0079 -0.1244
Dorsal fin-chambers -0.1085 0.8242 0.3674
Preanal fin-chambers -0.6971 0.5362 -0.2269
Some Puerto Rican individuals may represent an undescribed species. However, our
samples are small and all counts have broad overlapping ranges, with some specimens
nearly identical with specimens of B. floridae, whereas others are similar to those of
B. caribaeum from Brazil. Nonetheless, our samples exhibit almost no overlap with
either B. virginiae or specimens assigned to B. caribaeum from Venezuela.
Epigonichthys is less easy to subdivide than Branchiostoma. Nonetheless, E. austra-
lis and E. lucayanus are rather readily distinguished from other species by their unique
combinations of counts (Table 3). E. cultellus and its potential synonyms have fewer
preatriopore myotomes than any other lancelets. E. maldivensis is notably more similar
to E. lucayanus in its atriopore-anal myotome counts than are E. bassanus or E. hectori;
the latter show high counts in both pre- and postatriopore segments, while sharing the
relatively high preatriopore counts seen in E. lucayanus. No such high counts are seen in
E. cultellus or its probable synonyms.
Although our material contains limited non-American material, these results gener-
ally support the groupings identified by Webb (1955, 1975) and Richardson and
McKenzie (1994). They also lend credence to the suggestions of other investigators
concerning the status of certain nominal species.
The preceding evaluation demonstrates that many species can be assigned by myo-
tome formula alone, although such discrimination often requires simultaneous examina-
tion of all 5 variables. In some cases, certain individuals require identification of more
than countable features. Nonetheless, these data show that numerous species can be
easily distinguished, and others show less well understood regional differentiation.
Whether this regional variation reflects speciation or intraspecific variation will require
corroboration from additional features and larger collections of lancelets, particularly
from Caribbean and western Pacific localities. Molecular techniques should permit
alternative means of evaluating ecological and genetic differences. They will be particu-
larly important for the evaluation of morphologically less distinctive taxa.
DISCUSSION OF TAXONOMY AND USE OF NAMES
GENERAL. Some workers (e.g., Tattersall, 1903a; Prenant, 1928; Drach, 1948) recog-
nize a few widely distributed species, whereas others (e.g., Franz, 1922a; Hubbs, 1922;
Webb, 1957a) recognize a larger number of species with relatively restricted distribu-
tions. Differences between taxonomic "Jumpers" and "splitters" are not easily resolved in
lancelets. Several of the most important characters used to differentiate species involve the
number of myotomes and their location relative to the atriopore and anus, characteristics
which are potentially related to temperature of development (Hubbs, 1922; Nishikawa.
1981 ). Nonetheless, conjunction of these and previously published studies suggest some
level of concordance even though taxonomic problems remain for some taxa.
GENERIC ASSIGNMENTS. Many authors used Epigonichthys Peters 1877 to distin-
guish E. hectori from New Zealand (Richardson, 1950; Paulin, 1977) and E. bassanus
from southeastern Australia and Tasmania (McCulloch, 1919; Waite, 1921 ; Gomon
et al., 1994), although assigning E. cultellus to Asymmetron Andrews 1893. Others (cf.
Franz, 1922b, whose revision has been widely cited) placed species described within
Heteropleuron and Epigonichthys into Asymmetron. However, Epigonichthys predates
Asymmetron.
Three features have been widely used to distinguish Asymmetron from Epigoni-
chthys. The extremely narrow caudal fin, and the posterior end of the notochord broadly
free of myotomes posteriorly (urostyloid process of Kirkaldy), give E. lucayanus a
seemingly prolonged tail. In contrast, in other species of Epigonichthys the fin is
relatively higher, more like those seen in species of Branchiostoma. However, this
distinction can not be made at small sizes and the reduced fin may represent a specializa-
tion toward a more fully pelagic existence. Nonetheless, species of Epigonichthys show
considerable variability in the shape of the caudal fin so that its evident loss in
E. lucayanus is difficult to evaluate as a diagnostic generic character. A second charac-
ter is the lack of distinct preanal fin-chambers in E. lucayanus.
E. lucayanus also has a membrane between the oral cirri (intertentacular membrane
of Kirkaldy). It is much higher ventrally than laterally, but in other species of
Epigonichthys and Branchiostoma it is only slightly higher. However, in E. maldivensis
this membrane is more similar to that of E. lucayanus than are those of other species of
Epigonichthys (Franz, 1922b; Nishikawa, 1979a; Nishikawa, 1980a). Perhaps for these
reasons, a number of authors have followed Herdman ( 1904) in placing E. maldivensis
in Asymmetron. However, no synapomorphies seem to characterize all species of
Epigonichthys exclusive of E. lucayanus. In the absence of demonstration that
E. lucayanus is the sister-taxon to all the other species of Epigonichthys, we must follow
Richardson and McKenzie ( 1994) and place Asymmetron into the synonymy of
Epigonichthys.
Heteropleuron was proposed, as a subgenus of Branchiostoma, in a summary work
by Kirkaldy (1894) and subsequently in a more extensive and influential paper
(Kirkaldy, 1895). As originally proposed, Heteropleuron included Branchiostoma
bassanum GUnther, her new species Branchiostoma (H.) cingalense (spelled as
Singalense in Kirkaldy, 1894), and Epigonichthys cultellus Peters, no type being desig-
nated. Hubbs ( 1922) subsequently designated H. cingalense as the genotype, as spelled
in the better known work by Kirkaldy (1895). Whitley (1932) "following custom", but
not Hubbs, regarded Branchiostoma bassanum the type species of Heteropleuron
Kirkaldy 1894, and a synonym of Paramphioxus Haeckel 1893.
BRANCHIOSTOMA BELCHER/. The identity of individuals identified as B. belcheri
or closely related species has not been fully resolved. There appear to be at least three
statistically definable populations of B. belcheri (Webb, 1957a, 1975; Nishikawa,
198la, 1995; Yingyaet al., 1986). One, B. belcheri belcheri, is found in western Kyushu
southward in the Western Pacific. A second, B. belcheri tsingtauensis, ranges from
Northern China and throughout the rest of Japan. Intergrades have been reported by
Nishikawa (198la) from Ariake, Shimabara, and Amasuka, Japan. A third population
occurs off the Arabian peninsula, southeastern Africa, and Madagascar (Prenant, 1928;
Webb 1957a, 1957b).
Tattersall (1903a, 1903b) proposed that B. belcheri is a subspecies of the similar
B. lanceolatum. Webb (1957a, 1957b) distinguished B. lanceolatum from B. belcheri by
its lower total myotome and dorsal fin-chamber counts. He regarded B. bazarutense as
distinct, but closely related to African populations of B. belcheri. Unfortunately, the type
specimens of B. bazarutense have been lost (Webb, 1957a). These species and related
nominal forms have not been adequately investigated based on materials from across
their entire range.
Branchiostoma moretonese could well be a synonym of B. lanceolatum, inclusive of
B. haeckelii and include B. minucauda-like specimens Nishikawa (198la). Some indi-
viduals assignable to B. belcheri are hardly distinguishable from B. minucauda,
B. bazarutense, and B. tattersalli Nishikawa (198lb).
Student's t-tests have been applied to compare Australian samples referred to
B. belcheri (Richardson and McKenzie, 1994). Although this comparison disclosed
statistically significant differences among B. minucauda, B. moretonense, B. belcheri,
and B. lanceolatum, the taxonomic significance of such pair-wise univariate compari-
sons is problematic as the observed differences are small. Conclusions were based on
limited fractions of the variation seen in either B. lanceolatum or B. belcheri. The
assignment of specimens to statistical groups is unclear and variation reported by
Nishikawa was not noted. This and other statistical studies of lancelets have the problem
that the intraspecific variation that may result from spatial autocorrelation (Sokal and
Oden, 1978a, 1978b) has yet to be distinguished from interspecific variation.
Here, B. minucauda is included in the synonymy of B. belcheri and B. moretonense is
maintained as a species. We suspect that additional materials are likely to reveal a
complex pattern analogous to that described for Caribbean species. More specimens are
needed, particularly from the equatorial western Pacific and Indian oceans. Clearly,
Tattersall's hypothesis must be more thoroughly evaluated (Nishikawa, 1981 b) and
analyses of molecular variation are desirable (Richardson and McKenzie, 1994).
BRANCHIOSTOMA LANCEOLATUM. The original description of B. lanceolatum
by Pallas has been variously dated in synonymies due to confusion of two of Pallas'
contributions to marine biology (Hubbs, 1922; Bigelow and Farfante, 1948).

Branchiostoma lanceolatum was originally described from the North Sea and the
Mediterranean. Tattersall ( 1903a, l903b) and other early workers identified specimens
from the eastern Atlantic and the Indian oceans as this species. Since Hubbs (1922),
western Atlantic materials are no longer regarded as conspecific with B. lanceolatum.
However, Hubbs' suspicion that the Indian Ocean materials likewise belong to other
species has not been shared by other authors.
Webb (1957a) noted no notable difference between a specimen taken from the Arabian
peninsula and materials examined from European waters (Webb, 1956b). Azariah (1965a)
statistically analyzed 100 specimens of B. lanceolatum from Madras. He further distin-
guished B. tattersalli from B. lanceolatum. B. tattersalli had been described by Hubbs
( 1922) on the basis of a specimen identified as B. califomiense by Tattersall ( l903b), but
without examination of the specimen. Azariah's description, based on 9 specimens, notes a
difference of only a single myomere anterior to the atriopore and one posterior to the anus.
He also mentions a notable difference in number of fin-chambers, both dorsal and preanal,
as well as in the relative height of the former. He also described a slight difference between
the position of the anus relative to the lower lobe of the caudal fin, the anus of B. tattersalli being
just inside the fin as opposed to that of B. lanceolatum which lies outside the caudal fin.
Although consistent with observations made by Webb (1956b, 1957a, 1957b) and
Azariah (1965a), our materials are inadequate to resolve this problem as they include
only limited samples from the North Sea, Mediterranean, and Red Sea populations and
two poorly preserved specimens from Sri Lanka. It is tempting to speculate that more
than one species has been sampled, but simple pairwise morphological comparisons of
populations are insufficient to resolve this issue. Because dispersal via bilge transport is
common in invertebrates (Carlton and Geller, 1993) and can not be ruled out, this form
remains a prime candidate for analysis of molecular variation. Should Indian Ocean
populations prove to represent a different species distinct from the European
B. lanceolatum, the name Branchiostoma haeckelii is available.
BRANCHIOSTOMA CALIFORNIENSE. Andrews (1893) was the first to provide a
proper binomial for B. califomiense, furnishing a description and indication of a type
specimen (see below). However, Cooper (e.g., Meek and Hildebrand, 1923) and even
Gill have often and incorrectly been cited as authors. Hubbs ( 1922) noted Cooper ( 1868)
provided an account of a species from San Diego, California, but identified it only as
"Branchiostoma -?". Eigenmann (1892a,b), misidentified it as B. elongatum. Gill
(1893) further referred to California material as Branchiostoma, but did not provide a
specific epithet. In 1893, Andrews summarized this early work and noted that one of
Gill's specimens bore the label "B. californiensis." This emendation has been referred to
subsequently (Kirkaldy, 1895; Franz, 1922; Chan et al., 1990; Oliva et al., 1995),
without noting that Andrews referred to this species as "Branchiostoma Califomiense"
and designated a type specimen, albeit listed as "Cooper's type" both on page 238listing
the species of the genus, as well as again on page 248 in his Appendix I listing materials
examined and counts.
BRANCH/OSTOMA CARIBAEUM AND BRANCH/OSTOMA PIATAE. The original
S-36 S.G. POSS AND H.T. BOSCHUNG Isr. J. Zool.
description of Branchiostoma caribaeum (Sundevall, 1853), does not fit any nominal
lancelet well. The myotome fonnula is given as 37 + 14 + 9 =60 and the caudal fin does
not surround the tip of the tail. The latter feature is characteristic of Epigonichthys
lucayanus, rather than species of Branchiostoma. However, E. lucayanus has 6-10
myotomes between the atriopore and the anus (rather than 14) and 7-14 postanal
myotomes (rather than 9). In this respect it is similar to a number of species of
Branchiostoma.
Sundevall failed to specify a type locality, although others have attempted to do so
subsequently. Tattersall (1903a: 271) stated Sundevall's "new fonn" was from Rio de
Janeiro, ignoring the references to St. Thomas and Rio de Ia Plata. Hubbs ( 1922) basing
his detenninations "with the exception of the myotome fonnula, from two small
specimens from Porto Rico", attempted to restrict B. caribaeum to the West Indies. He
gave the B. platae type locality as "off the mouth of the Rio de Ia Plata". Hubbs ( 1922)
reported a total myotome count ranging from 62--65 (n =8) for B. platae, which differed
from the 60 indicated by Sundevall, perhaps prompting Hubbs to restrict the name
B. caribaeum to specimens from the West Indies. Jordan and Evennann ( 1930) "re-
stricted" the type locality of B. caribaeum to "St. Thomas" and started that Hubbs'
B. platae was "represented on the coast of Brazil (Rio de Janeiro, etc.)", although
Hubbs' material was taken off the mouth of Rio de Ia Plata.
Despite wide use of the name, no author since Sundevall has apparently examined the
type series. Although McShine (1975) alluded to "types" and "type series" (emphasis
ours), these referred only to data drawn from Hubbs ( 1922). Her reference to dorsal fin-
chambers was provided by Hubbs and they were never mentioned by Sundevall.
McShine's counts of the "type series" were likely drawn from Andrews (1893b); they
include Sundevall's count and Andrews' count of 27 + 12 + 9 = 48 from Jamaican
materials. She used the perplexing phrase (p. 55) "as described by Sundevall" in
reference to features not mentioned by Sundevall. Actually, the description credited to
Sundevall is a verbatim quote from Hubbs ( 1922).
The types used to describe B. caribaeum were sent to the Naturhistoriska Riksmuseet
in Stockholm from St. Thomas, Virgin Islands, the harbor of Rio de Janeiro, and Rio de
Ia Plata. No catalog numbers were provided and the exact number of specimens in the
type series is unknown. Based on time and place of capture, only two lots of specimens
that might qualify as type specimens could be located in the NRM collections. One is
questionably from Rio de Janeiro (4 specimens, NRM 19368) and the other taken from
Rio de Ia Plata (23 specimens, NRM 19363). We believe these to be part of the type
series, although documentation regarding the accession of NRM 19363 is not as defini-
tive as for NRM 19368. The NRM collection now contains no specimens of
Branchiostoma from St. Thomas. We are unable to locate material collected at St.
Thomas from other collections.
Our examinations of the extant part of the "type series", indicates that it includes two
species. All in NRM 19363 are assignable to B. platae Hubbs 1922 (two are severely
damaged). All of NRM 19368 are the same as Branchiostoma marambaiensis
Gon~alves da Silva, 1980, which then becomes a junior synonym of B. caribaeum. The
evident loss of type specimens from St. Thomas, the uncertain status of Iancelets in the
Caribbean, and the difficulty in taking counts from the poorly preserved specimens in
NRM 19368, make us reluctant to select a lectotype from existing type material at this
time. However, selection of a lectotype from NRM 19368 from Rio de Janeiro would
best fit contemporary usage. It would then be left to further investigation to determine
whether those from St. Thomas are conspecific with those in Florida, currently regarded
as B.floridae.
Bigelow and Farfante (1948) reported that Amphioxus muelleri Moreau (originally
proposed as Amphioxus Miilleri) has priority over Branchiostoma platae Hubbs. As
noted by Giinther ( 1880), Moreau uses a name taken from an unpublished manuscript
provided by Kr0yer, without providing a sufficient description of the species beyond an
extensive discussion of its notochord. However, Moreau's materials were taken from the
Bay at Rio de Janeiro and not from Rio de Ia Plata. Thus, Moreau's species would likely
be a synonym of B. caribaeum and not of B. platae.
BRANCHIOSTOMA NIGER/ENS£. Wickstead ( 1964b) proposed that B. takoradii is
a synonym of B. nigeriense, which metamorphose from pelagic larvae that have grown
large through an extended pelagic existence, rather than more typical larvae. The counts
of these species are nearly identical, with B. takoradii differing only in having I or 2
more myotomes between the atriopore and anus.
EPIGONICHTHYS CINGALENSE. The status of Epigonichthys cingalensis is pres-
ently uncertain. Kirkaldy ( 1894, 1895) diagnosed this species, based on eight speci-
mens, as having the right metapleural fold continuous with the caudal fin. However, in
her 1895 work she clearly refers to material belonging to Branchiostoma. Two rather
poorly preserved specimens [CAS-SU 41417] collected and identified by A.W. Herre
have counts nearly fitting those given by Kirkaldy, but based on the descriptions given
by Azariah ( l965a), are reidentified as B. lanceolatum.
EPIGONICHTHYS CULTELLUS. The monotypic Epigonichthys cultellus (Peters,
1877) was based on 10 specimens collected at Moreton Bay, Queensland. As noted by
Kelly ( 1966 ), E. cultellus species occurs in Moreton Bay together with B. moretonense.
He further stated that Peters illustrated "the type (1876, Fig. 4 )", which has counts that
fall within the range of B. moretonense and not of E. cultellus. If Kelly is correct,
material illustrated and possibly that described by Peters may comprise two species. It
will be necessary to examine the type specimens of E. culte/lus to resolve the confusion.
Meristic variation of Heteropleuron hedleyi Haswell falls within that of
Epigonichthys cultellus. Hence, they are synonyms, as also noted by Richardson and
McKenzie ( 1994 ). The latter authors note the types of H. hedleyi (not examined) are now
so poorly preserved as to be rendered useless.
Bathyamphioxus franzi Whitley ( 1932) was named solely in reference to a 17 .5-mm
specimen collected in Shark Bay, Western Australia and based on description and figure
of Franz ( 1922b ). It was regarded as an atypical specimen of E. cultellus by Franz and by
Richardson and McKenzie (1994). The myotome formula given by Franz (26 + 15 + 7)
is within or close to values reported for E. cultellus. Franz provides no fin-chamber
counts, but his figure indicates about 229 dorsal fin-chambers and no preanal fin-
S-38 S.G. POSS AND H.T. BOSCHUNG lsr. J. Zool.
chambers, which is consistent with an immature specimen of E. culte/lus. Although only

the left side is figured, Franz does not mention gonads. The dorsal fin-chambers of this
specimen are not as high relative to their width as are those in adult E. cultellus, which
are notably higher than in most other lancelets. This may be due to the immaturity of the
specimen. However, we have seen no material in which it is quite as high anteriorly as
that figured by Peters.
Branchiostoma pelagicum Gunther, 1889, is in all likelihood based on larvae of
E. lucayanus. Since the early years oflanceletology, it has been widely regarded as distinct
from Asymmetron lucayanum Andrews, 1893. Gibson (1909}, Wickstead (l964b}, and
Nishikawa (1982) strongly suggest that these larvae belong to E. lucayanus. Our materials
further support this contention. Since Gunther's name predates that of Andrews, priority
dictates that the latter be regarded as a junior synonym. However, this severely disrupts
contemporary usage. Given the widely different morphology of the larvae and adults, direct
reference to the type material is of limited value. Determination that a single species is
involved may likely remain subject to contention. Consequently, we continue to use
Andrews' name and have applied for plenary suppression of Gunther's name.
SYSTEMATIC ACCOUNT
Lancelets belong to the Phylum Chordata, Subphylum Acrania, Class Cephalochordata,

Order Amphioxiformes, Family Branchiostomatidae. Within the family there are two
genera, Branchiostoma and Epigonichthys.
FAMILY BRANCHIOSTOMATIDAE
Synonyms include: Amphioxididae Gray, 1842: 150 (diagnosis). Amphioxini Muller,

1844: 198 (new group name). Branchiostomidae Bonaparte, 1846: 1998 (original de-
scription). Cirrhostorni Owen, 1846: 9 (new group name proposed). Amphioxoidei
Bleeker, 1859: xxxiii, 214 (family group name, includes only Amphioxus belcheri;
based on Grey, 1847a). Cirrostomi Gunther, 1870: 513 (list of species in British
Museum fish collections). Amphioxididae Goldschmidt, 1905a: 240 (includes only
Amphioxides). Branchiostomatidae Hubbs, 1922: 3, 4 (correction of Latin ending; in
key; includes Branchiostoma and Dolichorhamphus [sic]). Epigonichthyidae Hubbs,
1922: 14 (includes only Epigonichthys and Asymmetron). Asymmetrontidae Whitley,
1932: 260 (includes only Notasymmetron).
GENUS BRANCHIOSTOMA
Branchiostoma Costa, 1834

Synonyms include:
Branchiostoma Costa, 1834: 49 (type species Branchiostoma lubricum Costa 1834,
by monotypy).
Amphioxus Yarrell, 1836: 468 (type species Limax lanceolatus Pallas, 1774, by
monotypy).
Amphioxys Agassiz, 1848: 53 (unjustified emendation of Amphioxus Yarrell, 1836).
Amphipleurichthys Whitley, 1932: 256 (proposed as subgenus; type species
Branchiostoma (Amphipleurichthys) minucauda Whitley, 1932, by original des-
ignation and monotypy).
Dolichorhynchus Willey, 190 I: 269-271 (type species Branchiostoma
(Dolichorhynchus) indicum Willey, by original designation and monotypy).
Dolichorhamphus: Hubbs, 1922: 3, 13 (misspelling of genus).
GENERIC DIAGNOSIS. Gonads normally developed on right and left epipleura.
Metapleuron of each side terminates immediately posterior to atriopore.
ETYMOLOGY. From the Greek combination branchion (gill)+ stoma (mouth). Gender neuter.
SPECIES OF BRANCHIOSTOMA
Branchiostoma africae Hubbs, 1927
Branchiostoma africae Hubbs in Monod, 1927: 644 (original description; type

locality "Souelaba, Cameroons"; museum for type deposition not given, but
presumably MNHN [types probably lost]).
DIAGNOSIS Myotome formula: -43 (42-44) preatriopore + -14.5 (14--15) atriopore to
anus+ -12.5 (ll-14) postanal =-70 (67-73) total. Dorsal fin-chambers -300. Preanal
fin-chambers, -52 (65-87). Maximum number of gonads unknown. Ratio of tallest
dorsal fin-chamber height to width about 6. Ratio of postatriopore length to preatriopore
length about 0.4. Anus anterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Hubbs (1927) did not report the size of his materials.
DISTRIBUTION This species is known only off Nigeria and the Cameroons (Fig. 2).
MATERIAL ?USNM 163321 (5, NIGERIA).
Branchiostoma arabiae Webb, 1957
Branchiostoma arabiae Webb, l957a: 121-128 (original description; type locality:

"Sta. 80, 22°l3'30"N, 59°48'48"E, near Ras al Haddon the Arabian coast near the
entrance to the Gulf of Oman").
DIAGNOSIS Myotome formula: 39.0 (36-41) preatriopore + 16.4 (16-18) atriopore to
anus+ 9.4 (8-ll) postanal = 65 (61-67) total. Dorsal fin-chambers 333 (297-366).
Preanal fin-chambers, 74.0 (65-87). Maximum number of gonads unknown. Ratio of
tallest dorsal fin-chamber height to width 2.0-3.5. Ratio of postatriopore length to
preatriopore length 0.33-0.43. Anus near center or slightly posterior to center of ventral
lobe of caudal fin.
MAXIMUM SIZE61 mm.

DISTRIBUTION Possibly restricted to the Gulf of Oman and the westward end of the
Persian Gulf (Fig. 2). One poorly preserved specimen from Aldabra I. (USNM 288861)
requires confirmation.
MATERIAL BMNH 1955.12.13.1 (holotype, 52 mm, near Ras al Hadd, Gulf of Oman at
22°13'30"N., 59°48'48"E, 16-22 m, Salpa dredge, coarse sand with shell, 1654-1726 h,
30 Nov. 1933). BMNH 1955.12.13.2-4 (4, paratypes). BMNH 1955.12.13.5-15 (20,
paratypes). BMNH 1955.12.13.16-20 (listed by Webb, 1957a). BMNH 1955.12.13.21-
40 (listed by Webb, 1957a). *??USNM 288861 (I, ALDABRA ATOLL, March 1986; in
poor state of preservation). *USNM 197645 (I, PERSIAN GULF, SAUDI ARABIA,
entrance to Ras Tanura Harbor, at Buoy No. 8). *?ZMUC P.OII9 (1, BAHRAIN,
26 °'38'N, 51°10'E, 23m, 23 Jul. 1937). *?ZMUC P.OI20 (2, BAHRAIN, 26°38'N,
51°10'E, 23m, 23 Jul. 1937). ?ZMUC P.OI21 (3, BAHRAIN, 26°50'N, 51°17.0'E, 33m,
30 Jul. 1937. ?ZMUC P.OI22-31 (10, PERSIAN GULF, directly offKharg 1., 29°14'N,
50°20.0'E, 13 m, 5 March 1937).
Branchiostoma bazarutense Gilchrist, 1923

Branchiostoma bazarutense Gilchrist, 1923: 62-65, fig. (original description; type
locality: "Bazaruto Islands" [Mozambique]). Holotype probably lost (Webb,
1957a).
DIAGNOSIS Myotome formula: -36 (34-38) preatriopore + -16 (15-17) atriopore to
anus + -9 (9-10) postanal = -65 (62-69) total. Dorsal fin-chambers 340 (250-340).
Preanal fin-chambers 62 (35-94). Maximum number of gonads 28. Ratio of tallest
dorsal fin-chamber height to width about 2.0-4.0. Ratio of postatriopore length to
preatriopore length 0.39-0.54. Anus posterior to center of ventral lobe of caudal fin.
DISTRIBUTION Known only from 3 type specimens taken near the Bazaruto Islands in
southern Mozambique (Fig. 2).
MATERIAL None.
Branchiostoma belcheri Gray, 1847

Synonyms include:
Branchiostoma belcheri Gray, 1847a: 35-36 (original description; species name
originally capitalized; type locality: "Borneo" [now Sarawak]).
Branchiostoma belcheri japonicus Andrews, 1895: 57 (original description of new
subspecies; Shikajima near Fukuoka).
Branchiostoma nakagawae Jordan and Snyder, 1901: (original description; type
locality: "Koajiro Bay, Misaki").
Amphioxus japonicus: Lonnberg, 1904: 240-241 (B. nakagawae placed into syn-
onymy of B. be/cheri).
Branchiostoma lanceolatum belcheri: Tattersall, 1903b: 269-299, 272, 274-275,

279-283, 286-294, 295, 298-299 (comparisons with other Branchiostoma spe-
cies; distribution).
?Branchiostoma (Amphipleurichthys) minucauda Whitley, 1932: 257, pl. xii, fig. I
(original description; type locality: "Port Curtis, Queensland;" holotype: AMS
IA4190 [reported lost by Richardson and McKenzie, 1994]; synonymy; common
name: smalltaillancelet).
Branchiostoma belcheri tsingtauense Tchang-Shi and Koo, 1936: 76-114 (new
subspecies; original description; type locality "Kiachow Bay").
anus+ 10.4 (8-12) postanal = 65 (61-69) total. Dorsal fin-chambers 290 (222-360).
Preanal fin-chambers 58 (35-91 ). Maximum number of gonads 25. Ratio of tallest
dorsal fin-chamber height to width about 2.5-3.5. Ratio of postatriopore length to
preatriopore length 0.42-0.54. Anus in advance of center of ventral lobe of caudal fin.
MAXIMUM SIZE Webb (1956a) and Azariah (1965a) reported this species as reaching
70 mm, much larger than reported by Nishikawa (l981a) at 57 mm from extensive
records. Yingya et al. (1986) who also examined abundant material, indicate a maxi-
mum total length of 62 mm. We have not seen material larger than 55 mm.
DISTRIBUTION Southern Japan, along the Chinese coast, the Philippines, southward
and eastward to Torres Straits, and in the western Indian Ocean at Madagascar and
southeastern Africa (Figs. 2 and 3).
MATERIAL AMS IA4190 (holotype of B. minucauda, 44 mm, AUSTRALIA,
Queensland, Port Curtis, Dec. 1929). *CAS-SU 61167 (12, CHINA, Amoy, 1957-
1958). Department of Biology, De La Salle University, Manila collections (191, 15-46
mm), PHILIPPINES, Quezon Prov., Tayabas Bay, Lucena Anchorage adjacent to
Tayabas river, l3°54'N, 121 °37'E). *FAKU (3, JAPAN, Sea of Japan, Tottori Pref., off
Daisen, 16 fms). SNM 86591 (5, CHINA, Amoy). USNM 122991 (I, PHILIPPINES,
Sirun 1., S80°W, 3.80 mi., 05°35'40"N, 120°47'30"E, ALBATROSS, Sta. D5148).
USNM 130591 (313,CHINA.Amoy,Fukien). ?USNM 150796(1,PHILIPPINES,Jolo
Light, 06°06'N, 120°58'50"E, ALBATROSS Sta. D5138). ?USNM 150797 (I, PHILIP-
PINES, Anima Sola I.• l3°li'I5"N, l23°45"E, ALBATROSS Sta. D5218). *?USNM
150798 (1, PHILIPPINES, Linao Pt., 07°04'48"N, I25°39'38"E, ALBATROSS Sta.
D5253). *?USNM 168203 (I, PHILIPPINES, Gulf of Dauao, Lanao Pt., N. 22°E., 1.5
mi, 07°04'48"N, 125°39'38"E, ALBATROSS Sta. 5253). *USNM 196044 (7, PHILIP-
PINES, Manila Bay, San Minolas, dredge, sand, 2.5 fms). *USNM 210453 (II, INDO-
NESIA, MOLUCCAS IS., Abon, Poka, Beach off point on NW shore of Ambon Bay, 0-
6ft., 1130-1230 h, 22 Jan. 1973). *UAIC 7855 (10, CHINA, Fujiang).
Branchiostomtl bennetti Boschung and Gunter, 1966

Branchiostoma bennetti Boschung and Gunter, 1966: 485-489 (original description;
type locality: "Louisiana").
S-42 S.G. POSS AND H.T. BOSCHUNG lsr. J. Zoot.
DIAGNOSIS Myotome fonnula: 35.6 (34-38) preatriopore + 16.1 ( 13-19) atriopore to

anus + 8.6 (7-10) postanal = 60 (56--65) total. Dorsal fin-chambers 293 (189-359).
Preanal fin-chambers 47 (24-77). Maximum number of gonads unknown. Ratio of
tallest dorsal fin-chamber height to width 2.0-4.5. Ratio of postatriopore length to
preatriopore length 0.34-0.46. Anus near center of ventral lobe of caudal fin. Lamellae
on oral cirri elongate.
MAXIMUM SIZE 53.0 mm.
DISTRIBUTION This species is known from relatively muddy sediments ofthe northern
Gulf of Mexico, reaching depths of about 12 m (Fig. I).
MATERIAL *USNM 261424 (holotype, LOUISIANA, Grande Terre 1., West end in
mud flat, 300 yds. east of Barataria Light on the bay shore, 24 March 1982). *USNM
231730 (12, paratypes, Same data as holotype). *UAIC 1661 (10, paratypes, LOUISI-
ANA, Grand Terre, 23 Nov. 1963). *UAIC 1662 (3, paratypes, LOUISIANA, Grand
Terre, 16 Oct. 1962). *GCRL 381 (I, LOUISIANA, South of Grand Isle, Freeport Sta. I,
3-5 fms, deep plankton tow, 12.5 oc surface, 14.0 °C bottom, salinity 32.6 ppt surface,
31.9 ppt bottom, 3 March 1959). *GCRL 1053 (96, MISSISSIPPI, Mississippi Sound,
Hom Island, at Horseshoe, CEO Sta. 23, 18-20 ft, 30 Apr. 1964). *GCRL 2062 (5,
MISSISSIPPI, Mississippi Sound, Hom Island, lagoon near chimney, 3 ft, 14 May
1967). *GCRL 2319 (1, 30°02'30"N, 88°40' 15"W, plankton tow, Sta. 2, 8 Sept. 1967).
UAIC 1663 (4, LOUISIANA, 28°55'90°3l'W, 23 Jan. 1963). *UAIC 1664 (3, LOUISI-
ANA, 17 Apr. 1957). *UAIC 3620 (1. LOUISIANA, Grand Terre 1., 23 Nov. 1963).
*UAIC 3716 (3, TEXAS, Galveston 1., 22 Feb. 1966). *UAIC 3717 (36, 2 March 1966).
*UAIC 3718 (1, I Nov. 1967). *UAIC 3719 (1, 2 March 1967). *UAIC 3720 (2).
*UAIC 3721 (12). *UAIC 3723 (2, 94°54'N, 29°12'36"W, 7 Jan. 1966). *UAIC 3724 (I,
13 Dec. 1966). *UAIC 3725 (I, I Nov. 1967). *UAIC 3727 (4, 16 Jan. 1967). *UAIC
3728 (2, 13 Dec. 1966). *UAIC 3730 (21 Feb. 1966). *UAIC 3730 (4, 21 Feb. 1966).
*UAIC 3731 (4, 6 Jan. 1966). *UAIC 3734 (33, 5 Jan. 1966). *UAIC 3735 (9, 3 Feb.
1966). *UAIC 3736 (32, 4 Feb. 1966). *UAIC 3737 (2, 13 Dec. 1966). *UAIC 3738 (21,
4 Jan. 1967). *UAIC 4454 (3, LOUISIANA, Grand Isle, 12 Aug. 1972).
Branchiostoma bermudae Hubbs, 1922

Branchiostoma bermudae Hubbs, 1922: 2, 9-10 (original description; type locality:
"Bennuda;" thought most closely to resemble B. platae).
DIAGNOSIS Myotome fonnula: 34.9 (34-36) preatriopore + 14.0 (11-16) atriopore to
anus+ 6.6 (6-10) postanal =55 (53-59) total. Dorsal fin-chambers 225 (189-254).
dorsal fin-chamber height to width 3-5. Ratio of postatriopore length to preatriopore
length 0.31-0.43. Anus posterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Reported to reach 53.5 mm (Hubbs, 1922).
Vol. 42. 1996 LANCELET TAXONOMY S-43
DISTRIBUTION Restricted to Bermuda, where it is found nearshore to depths of about

13m.
MATERIAL *UMMZ 55145 (holotype, BERMUDA, donated by W. J. Crozier). *MCZ
29074 (7, BERMUDA). *MCZ 35168 (2, BERMUDA). *UAIC 3740 (13, BER-
MUDA). *UAIC 3256 (10, BERMUDA). *UAIC 3279 (15, BERMUDA). *UAIC 3281
(10, BERMUDA). *UAIC 3282 (7, BERMUDA). *UAIC 3288 (15, BERMUDA).
*UMMZ 56258 (4, BERMUDA). *UMMZ 56259 (13, BERMUDA). *USNM 21877
(1, paratype, BERMUDA, 1876-1877). *USNM 52076 (I, paratype, BERMUDA, 10
Aug. 1904). *USNM 92359 (1, paratype, BERMUDA). *USNM 92360 (1, paratype,
BERMUDA). USNM 92361 (1, paratype, BERMUDA, 1876-1877). *USNM 112510
(5, BERMUDA). *USNM 142509 (9, BERMUDA). *USNM 164691 (I, BERMUDA).
Branchiostoma californiense Andrews, 1893
Branchiostoma- Cooper, in Cronise 1868: 498 (coast of California [see taxonomic

discussion]).
Branchiostoma Californiense Andrews, l893a: 238,241 (original description; type
locality: "California, San Diego"; species name in upper case [see taxonomic
discussion]).
Branchiostoma (Amphioxus) californiensis: Kirkaldy, 1895: 312, pl. 34, fig. 4 (name
misspelled, authorship attributed to Cooper and thought close to B. elongatum).
DIAGNOSIS Myotome formula: 40 (40---45) preatriopore + 18 (14-19) atriopore to anus
+ 9 (8-9) postanal =67 (64-71) total. Dorsal fin-chambers 355 (317-419). Preanal fin-
chambers 44 (35-59). Maximum number of gonads 36. Ratio of tallest dorsal fin-
chamber height to width 4-8. Ratio of postatriopore length to preatriopore length 0.31-
0.43. Anus well posterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Reported as attaining 83.5 mm (Hubbs, 1922).
DISTRIBUTION Ranges from Chame Point, Panama in the south (Meek and
Hildebrand, 1923) to Monterey Bay (Beebe and Tee-Van, 1941) (Fig. 1).
MATERIAL USNM 8407 (holotype, received from Cooper, no other data). *CAS 84440
(I, California, Santa Cruz 1., off Kinton Pt. and Fraser Pt. l 00 ft, ORCA, 7 March 1950).
*CAS 84441 (3, San Pedro, 8 Jul. 1901). *CAS 84442 (14, MEXICO, At mouth of
Tijuana River, 12 fms, clean sand, ORCA, Oct. 1949). CAS 84443 (I, no collection
data, but probably California, collected by UC Berkeley prior to July 1947). *CAS
84444 (I, MEXICO, Baja California, San Martin 1., TEMPLETON CROCKER, 20
Aug. 1932). *CAS 3331 and 3332 (2, TEMPLETON CROCKER Sta. D 17R, 2 Aug.
1932). *CAS 20235 (1, CALIFORNIA, Monterey Bay, Haul3, 26 May 1945). *GCRL
3425 (3, PACIFIC PANAMA, Fort Kobbe Beach, 8°53'45"N, 79°34'35"W, tidepools,
sand, rock, broken shell bottom, low ebb tide, chemfish, 0-3 ft, 27 Jul. 1968). *GCRL
7452 (1, PANAMA, Panama, NE end of Naos I. Causeway, 08°56'07"N, 79°47"W,
small rocks, coarse sand and gravel, low flood tide, ichthyocide, 0---4 in., 2 Nov. 1971).
*GCRL 7590 (1, PANAMA, Panama, Farfan Point, 08°56'00"N, 79°34'00"W, sand-
sandy mud with some rock, deep tidepool, low flood tide, ichthyocide, 0-6 ft, 27 ppt, 5
Nov. 1971). *GCRL 9927 (19, PANAMA, Panama, Farfan Point, 08°56'00"N,
79°34'00"W, tidepools, mainly sand bottom, some mud and rock, ebb-low tide,
ichthyocide, 0-2 m, 29.0 oc, 34 ppt, 14 Apr. 1972). *GCRL 10502 (1, PANAMA,
Panama, Farfan Point, 08°56'00"N, 79°34'00"W, sand, sandy mud, some rock, ebb-
flood tide, ichthyocide, 0-0.75 m, 31.0 °C, 27.8 ppt, 7 Nov. 1972). *GCRL 10624 (2,
PANAMA, Panama, Venado Beach, about 3/4 way to Venado Is., tidepools and streams
on either side of connecting sandbar, small rocks, sand-silt bottomed pools, sandy
streams, low-flood tide, ichthyocide, 0-2 ft, 19 March 1972). *GCRL 10626 (1,
PANAMA, Panama, Punta Paitillo, E side, rocky tidepools,low-flood tide, ichthyocide,
0-.75 m, 28.0 °C, 35.2 ppt, 13 Apr. 1972). *GCRL 10627 {2, PANAMA, Panama,
Venado Beach along sandbar to Isla Venado, rock, sand, shell and marl bottom, ebb-
flood tide, ichthyocide, 0-0.6 m, 32 °C, 17 Apr. 1972). *GCRL 12593 (15, PANAMA,
Panama, Farfan Point, 08°56'00"N, 79°34'00"W, tidepools, mainly sand, some rock and
silt, ebb flood tide, ichthyocide, 0-2 m, 35 °C, 33.0 ppt, 6 Apr. 1973 ). *GCRL 13994 (I,
PANAMA, Panama, Naos 1., Pilot Beach, Sta. 131-2, 3 Apr. 1973). *GCRL 13995 {3,
PANAMA, Panama, Farfan Point, 08°56'00"N, 79°34'00"W, tidepools, mainly sand,
some rock and silt, ebb-flood tide, ichthyocide, 0-2 m, 35 °C, 33.0 ppt, 6 Apr. 1973).
*GCRL 13996 (1, PANAMA, Panama, Naos 1., Piolot Beach, Sta. 164, 11 Nov. 1973).
*GCRL 13997 (II, PANAMA, Panama, Farfan Point, 08°56'00"N, 79°34'00"W, 0-1.3
m, 30.0 °C, 24.0 ppt, 12 Nov. 1973). *GCRL 19120 (3, PACIFIC PANAMA, Gulf of
Panama, 08°13.2'N, 80°29.l'W, Aquapulce Tidal flats E of town, Sta. 20P, 0-2 ft, 33
ppt, 24 Apr. 1971). *GCRL 19122 (2, PACIFIC PANAMA, Gulf of Panama, Venado
Beach, rocky area, 08°53.5'N, 79°36.5'W, Sta. 23P, 25 Apr. 1971 ). *GCRL 19123 (80,
PACIFIC PANAMA, Gulf of Panama, Aguadulce, Tidal flats E of town, 08°13.2'N,
80°29.1'W, Sta. 20P, 0-2 ft, 24 Apr. 1971). *GCRL 19124 (5, PACIFIC PANAMA,
Farfan Flats, 25 °C, 4 Feb. 1977). *SIO 51-231 {1, CALIFORNIA, Santa Catalina 1).
*SIO 51-410 (14, 29.0-61.5 mm, CALIFORNIA, San Diego, Pt. Lorna). *SIO 52-8 (1,
48.0 mm, MEXICO, Baja California del Norte, 32°3I'N, II7°12'W). SIO 52-102 {2,
35.6--49.5 mm, MEXICO, Baja California del Norte, 32°24.5'N, 117° 14.2'W). *SIO 55-
107 1,42.5 mm, CALIFORNIA, San Luis Obispo County,> 35°N, 10 Dec. 1955). *SIO
58-178 {I, 72.0 mm, CALIFORNIA, La Jolla, off Beach Club, 32°5I.O'N, II7°16.0'W,
25-35 ft). *SIO 58-219 (26, 27.0-62.0 mm, MEXICO, Baja California, Isla Espiritu
Santo, 24°22.0'N, ll0°ll.O'W, II fms, 18 Apr. 1987). *SIO 58-307 (1, 72.0 mm,
MEXICO, Baja California del Norte, Punta Banda, in Papalote Bay, 31°48.0'N,
II6°42.0'W, 0-35 ft). SIO 59-199 (I, 7.5 mm, MEXICO, near Mazatlan, middle island
at Venado, 23°13.0'N, I06°25.0'W). SIO 59-234A (2, MEXICO, Baja California del
Sur; Bahia de Los Angeles, SE corner; 29°00.0'N, I I3°30.0'W, at waterline). *SIO 59-
336 (1, 5.0 mm, CALIFORNIA, Mission Bay, Ventura Bridge, 32°47.0'N, II7°15.0'W,
I m). SIO 61-227 (1, MEXICO, Cabo San Lucas, 22°52.5'N, 109°53.7'W, 0-75 ft). SIO
61-247 (17, MEXICO, off Punta Pulmo, 23°27.5'N, I09°24.4'W, 0-100 ft). SIO 61-251
{3, MEXICO, Palmas Bay, 23°38.7'N, I09°40.5'W). SIO 62-169 (I, MEXICO, Bahia de
Los Angeles, 29°00.0'N, ll3°30.0'W). *SIO 62-496 (19, 47.0-61.0 mm, MEXICO,
Baja California del Norte, off mouth of Tijuana R., 32°33.0'N, 117° IO.O'W). *SIO 64-
343 (12, 28.0-76.0 mm, CALIFORNIA, San Diego Bay, Ballast Pt., 32°46.0'N,
li7°13.0'W, 25 Sep. 1904). *SIO 65-270 (1, 20.0 mm, MEXICO, Baja California,
Canal de San Jose, 24°59.8'N, II0°46.0'W). *SIO 65-522 (4, 18.0-47.0 mm, CALI-
FORNIA, off Pt. Lorna, 32°4l.O'N, ll7°l4.0'W). SIO 67-40 (1, MEXICO, Pearl Is., Isla
Saboga, 08°40.0'N, 079°03.0'W, 0-10 ft). *SIO 73-223 (8, 65.0-41.5 mm, CALIFOR-
NIA, San Onofre Nuclear Power Plant, 33°l5'N, ll7°23.0'W, 26ft). SIO 81-169 (7,
3.0-4.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W, 10m). SIO 81-
170 (5, 4.0-5.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W, 14m).
SIO 81-171 (9, 3.0-5.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N, li3°30.0'W,
7 m). SIO 81-172 (10, 4.0-13.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N,
ll3°30.0'W, 4 m). SIO 81-173 (8, 4.0-8.0 mm, MEXICO, Bahia de Los Angeles,
29°00.0'N, ll3°30.0'W, 2m). SIO 81-175 (26, 4.0-18.0 mm, MEXICO, Bahia de Los
Angeles, 29°00.0'N, ll3°30.0'W). SIO 81-176 (59, 4.0-14.0 mm, MEXICO, Bahia de
Los Angeles, 29°00.0'N, ll3°30.0'W, 4 m). SIO 81-177 (5, 5.0-6.0 mm, MEXICO,
Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W, 27m). SIO 81-178 (4, 12.0-20.0 mm,
MEXICO, Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W, 9 m). SIO 81-179 (I, 5.0mm,
MEXICO, Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W, 31 m). SIO 81-180 (27, 3.0-
19.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W, 9 m). SIO 81-181
(2, 4.0-6.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W, 27 m). SIO
81-182 (1, 24.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W, 17m).
SIO 81-185 (31, 1.0-4.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N, ll3°30.0'W,
9 m). SIO 81-187 (130, 4.0-15.0 mm, MEXICO, Bahia de Los Angeles, 29°00.0'N,
ll3°30.0'W, 6 m). SIO 81-190 (13, 3.0-28.0 mm, MEXICO, Bahia de Los Angeles,
29°00.0'N, ll3°30.0'W, 6 m). SIO 81-191 (1,4.0 mm, MEXICO, Bahia de Los Angeles,
29°00.0'N, ll3°30.0'W, 19m). SIO 81-192 (1, 3.0-4.0 mm, MEXICO, Bahia de Los
Angeles, 29°00.0'N, ll3°30.0'W, 9-16 m). SIO 84-1233 (II, 3.0-4.0 mm, MEXICO,
near Bahia S Francisquito, 28°34.0'N, ll3°07.0'W, surface). *SIO 84-234 (1, 58.0 mm,
CALIFORNIA, 2 miles S of Pt. Lorna, 32°4l'N, ll7°l4.0'W, 24m). *SIO H49-22 (2,
53-53.5 mm, MEXICO, Baja California del Norte, Coronado I, 31 °23.6'N,
117°14.1 'W). *SIO H49-44 (7, 32.5-49 mm, CALIFORNIA, San Diego, near Pt.
Lorna). *SIO H49-152 (60, CALIFORNIA, San Diego, near Mexican Border). *SIO
H46-l23 (89, 31.0-77.0 mm, MEXICO, Baja California del Norte, S. Coronado I,
32°24.5'N, ll7°14.2'W). *SIO H46-l25 (5, 47.0-56.0 mm, MEXICO, Baja California
del Norte, S. Coronado I, 32°24'N, ll7°l4'W). *SIO H46-109 (II, 20.0-67.0 mm,
MEXICO, Baja California del Norte, near Mexico-US border, 32°3l.7'N, ll7°13.7'W).
*SIO H47-125A (73, 19.0-78.0 mm, MEXICO, Baja California del Norte, 32°24.8'N,
ll7°14.2'W). *CAS-SU 17941 (15, MEXICO, Baja California, Gulf of California,
Sefton-Stanford Exped., ORCA Sta. 9-D-l, April 1952). *CAS-SU 17942 (2,
MEXICO, Gulf of California, San Jose I. at end of Amortajada Bay, Sefton-Stanford
Exped., ORCA Sta. 2-S-1, 8 May 1952). *CAS-SU 17943 (29, mm, MEXICO, Gulf of
California, Angel de Ia Guardia 1., 29°3l'N, ll3°37'W, ll-14 fms, Sefton-Stanford
Exped., ORCA Sta. 38-D-4, 8 May 1952). *CAS-SU 18770 (I, MEXICO, Baja Califor-
nia del Sur, Bahia de Los Angeles, 28°59'N, II3°35'W, Sefton-Stanford Exped., ORCA
Sta. 39-D-3, 10 May 1952). *CAS-SU 18771 (5, MEXICO, Baja California del Sur,
passage into Bahia Los Angeles, 28°58'N, II3°29'W, ORCA Sta. 39-D-4, 10 May
1952). *CAS-SU 23029 ( 16, California, San Diego). *CAS-SU 160( 17, MEXICO, Baja
California del Sur, Louis Gonzales Bay, ALBATROSS). *CAS-SU 35949 (15, CALI-
FORNIA, San Diego Bay, 25 Jan. 1906). *CAS-SU 37362 (42, MEXICO, Baja Califor-
nia del Norte, 4 112 mi. E of Coronado 1.. VALERO Sta. 871-38, Hancock Exped., II
March 1938). *CAS-SU 48446 [formerly NYZS 27700] (2, 17-24 mm, NICARAGUA,
Corinto harbor, 27°07''N, 87° 11'37"W, ZACA Sta. 200: D-22, 1.5 fms., 7 Jan. 1938).
*CAS-SU 48447 [formerly NYZS 28469] (I, 36 mm, COSTA RICA, Golfito, Gulf of
Buice, ZACA, I ft, from sand in shallow tidal stream, 6 March 1938). *CAS-SU 48448
[formerly NYZS 28211] (2, 19-30mm, COSTA RICA, Piedra Blanca Bay, 09°51'54"N,
85°29'53"W, ZACA Sta. 218: D-2, 5 fms., dredge, 5 Feb, 1938). *CAS-SU 48449
[formerly NYZS 25482] (4, 20-48.8 mm, MEXICO. Baja California del Norte, Arena
Bank, 32°27'N, 109°24'W, ZACA Sta. 136: D-30, 35 fms, I May 1936). *USNM 39646
(I, CALIFORNIA, San Diego). *USNM 43724 (1, CALIFORNIA, San Diego).
*USNM 46683 (30, MEXICO, Baja California del, Gonzales Bay, San Louis, ALBA·
TROSS). *USNM 75530 (I, CALIFORNIA, Pacific Grove). *USNM 82702 (33,
PANAMA, Chame Point, 7 Jan. 1919). *USNM 86519 (I, CALIFORNIA, Los Ange-
les). *USNM 87105 (2, CALIFORNIA, San Pedro). *USNM 82683 (14, PACIFIC
PANAMA, Chame Point). *USNM 92121 (I, COSTA RICA, Puerto Jimenez). *USNM
94300 (3, CALIFORNIA, San Diego). *USNM I01678 (I, COST A RICA, Salinas Bay,
Hancock Exped.). *USNM 101679 (1, COSTA RICA, Salinas Bay, Hancock Exped.).
*USNM 101684 (I, COSTA RICA, PuertoCulebra, Hancock Exped.). *USNM 117648
(I, MEXICO, Baja California del Sur, Puerto San Carlos, 27°57'N, Ill 0 05'W). *USNM
11855 (I, CALIFORNIA?, Angeles Bay). *USNM 118856 (4, CALIFORNIA, San
Lucas Cove). *USNM 120127 (I, PACIFIC PANAMA, Chame Point). USNM 125097
(7, CALIFORNIA, Santa Barbara 1., ALBATROSS Sta. 2900). *USNM 128409 (1,
PANAMA, CANAL ZONE, Farfan Beach, 24 Feb. 1937). *USNM 232568 (1,
MEXICO, Sonora, Choya Bay, 31°30"N, li3°41'36"W, 20m, 21 Oct. 1967). *USNM
232569 (1, MEXICO, Sonora, 31°18'48"N, ll3°4l'I2"W, 18 m, 23 Feb. 1968).
*USNM 23570 (1, MEXICO, Sonora, 31°17'00"N, II3°41'48"W, 22m, 4 Nov. 1967).
*USNM 232571 (I, MEXICO, Sonora, Off Choya, 31 °25'48"N, II3°40'36"W,
JANTHINA V Sta. 042, 2m, 4 Sep. 1966). *USNM 232574 (1, PANAMA, CANAL
ZONE, Farfan, 23 Jul. 1978). *USNM 232575 (1, PANAMA, Whorehouse Reef,
exposed tidepool, 0.1 m, 20 July 1978). *USNM 232576 (1, PANAMA, Naos 1., Pilot
House Beach, 18 May 1982). *USNM 232577 ( 10, PANAMA, Punta Paitilla, 9 Jan.
1978). *USNM 232578 (6, PACIFIC PANAMA, Canal Zone, Farfan, Sta. 247-2, 1-1.5
m, 10 Jan. 1978). *USNM 23579 (12, PANAMA, Naos 1., 8 Jan. 1978). *USNM
256721 (2, COSTA RICA, Gulf of Nicoya, 09°52'30"N, 084°43'50"W, 10 Jul. 1980).
*USNM 232570 (I, COSTA RICA, Gulf of Nicoya, 09°54'55"N, 084°45'15"W, II Jul.
1980). *USNM 256723 (2, COSTA RICA, Gulf of Nicoya, 09°57'38"N, 084°48'20"W,
12 Jul. 1980). *USNM 256724 (6, COSTA RICA, Gulf of Nicoya, 09°55'28"N,
084°52'05"W, 13 Jul. 1980). *USNM 256725 (1, COSTA RICA, Gulf of Nicoya,
09°57'15"N, 084°50'33"W, 12 Jul. 1980). *USNM 256726 (1, COSTA RICA, Gulf of
Nicoya, 10°0 1'40"N, 084°55'57"W).
Branchiosto11Ul capense Gilchrist, 1902

Branchiostoma capense Gilchrist, 1902: 101-113 (original description; type local-
ity: "South Africa"; holotype: SAM 13727).
DIAGNOSIS Myotome formula: 47 (44-48) preatriopore + 19.2 (18-20) atriopore to
Preanal fin-chambers 71 (62-80). Maximum number of gonads about 34. Ratio of tallest
dorsal fin-chamber height to width 4.0-6.0. Ratio of postatriopore length to preatriopore
length 0.38-0.43. Anus near center of ventral lobe of caudal fin.
DISTRIBUTION Evidently restricted to southern Africa, being taken from Cape Town
to the Agulhas Banks (Fig. 2).
MATERIAL USNM 85807 (1, paratype, SOUTH AFRICA, Cape Town). ZUMC
P.Ol33-0l34 (2, SOUTH AFRICA, Agulhas Bank, 37°08'S, l8°43'E, 19 Dec. 1929).
Branchiosto11Ul caribaeum Sundevall, 1853

Synonyms include:
Branchiostoma caribaeum Sundevall, 1853: 12 (original description; type locality
[see taxonomic discussion]).
Amphioxus Miilleri Moreau, 1875: 312, 1 pl., 12 figs. (nomen nudum; based on
unpublished manuscript name of Kr~~Jyer and used in description of microscopic
anatomy of material collected from Rio de Janeiro, Brazil; external features and
counts not presented; original material in all probability lost).
DIAGNOSIS Myotome formula: 36.9 (27-37) preatriopore + 14.9 (ll-18) atriopore to
anus+ 8.9 (7-10) postanal =61 ([48] 57-65) total. Dorsal fin-chambers 295 (228-341).
dorsal fin-chamber height to width about 3. Ratio of postatriopore length to preatriopore
length 0.4. Anus near center of ventral lobe of caudal fin.
DISTRIBUTION Brazil, probably northward into Caribbean (see taxonomic discussion).
MATERIAL NRM 19368 (4, types of B. caribaeum, ?Rio de Janiero, in poor condition,
see text for discussion). ?USNM 50124 (2, PUERTO RICO, Off Vieques I., FISH
HAWK, II fms). ZUMC (6, West Indies). ?GCRL 10625 {3, PANAMA, San Bias, Isla
Pico Feo, 0-1 m, 27 °C, 34.0 ppt, 19 Apr. 1972). ?GCRL 11472 (34, PANAMA,
Panama, Farfan Point, 08°56'00"N, 79°34'00"W, 0-2 m, 35 °C, 33.0 ppt, mainly sand
tidepools, 6 Apr. 1973). ?GCRL 11562 (3, PANAMA, Panama, NE end of Naos I.
causeway, 08°56'07"N, 79°32'47"W, 0-6 em, 32 °C, 3 Apr. 1973). *?GCRL 19121 (1,
WESTERN ATLANTIC PANAMA, 09°27.45'N, 79°44.5'W, Mouth of Rio Piedras
from hwy. bridge to 200 yds. on ocean side E and W, Sta. 7A, 0-2 ft, 19 April 1971).
?GCRL 22165 (27, PANAMA, Colon, Galeta 1., reef and flats across channel from STRI
lab, 0-0.8 m, 29.5 °C, 32 ppt, 14 Aug. 1974). LACM 20459 (6, West Indies, Aruba).
?USNM 232580 (1, PANAMA, Ft. Randaloph, 1.5 m). ?USNM 232581 (1, COLOM-
BIA, Santa Marta, Beach in front of Santa Mar Hotel, 14 Oct. 1977). ?USNM 126169 (7,
Puerto Rico, Off Vieques 1., FISH HAWK Sta. 6084). USNM 232572 (1, VENEZU-
ELA, Bahia Mochima, 18 Jan. 1978). USNM 285751 (5, CARIBBEAN, Dec. 1983).
GCRL I 0697 (38, BRAZIL, Bahia, Salvador, Praia da Ribeira, 28 Oct. 1969). GCRL
16862 (8, BRAZIL, Bahia, Salvador, Praia da Ribeira, 7 Aug. 1979). UAIC 6442 (50,
BRAZIL, Sao Paulo). USNM 87817 (1, BRAZIL, Sao Sebastian). USNM 87818 (3,
BRAZIL, Paqueta 1., Paqueta). USNM 87819 (1, BRAZIL, Paqueta 1., Paqueta). USNM
87821 (11, BRAZIL, Paqueta 1., Paqueta). USNM 88043 (5, BRAZIL, San Sebastian).
USNM 132857 (I, BRAZIL, Bay of Rio). ZUMC 7 (5, BRAZIL, Bahia, 25 Aug. 1863).
Branchiostoma elongatum Sundevall, 1852

Branchiostoma e/ongatum Sundevall, 1852: 147 (original description; type locality
"Chincha Island, Peru").
dorsal fin-chamber height to width 2. 7-7 .0. Ratio of postatriopore length to preatriopore
length 0.33-0.42. Anus anterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Reported to reach 68.5 mm (Hubbs, 1922).
DISTRIBUTION Known from limited material taken from several Peruvian localities,
and from lsabela [Albemarle] I. in the Galapagos (Snodgrass and Heller, 1905) (Fig. I).
MATERIAL *GCRL 12465 (3, CHILE, Antofagasta, 0-1 m, 1973). *CAS-SU 6400 (2,
GALAPAGOS, Isabela 1., Snodgrass and Heller). *CAS-SU 37353 (23, PERU, Lobos
de Afeura Sound, VALERO Sta. 843-38, 25-30 fms, Hancock Expedition, 14 Feb.
1938). *CAS-SU 37364 (9, PERU, Bahia lndependencia, VALERO Sta. 833-38,8 fms,
Hancock Expedition, sand and shell, 1942). *CAS-SU 37365 (3, PERU, San Juan Bay,
VALERO Sta. 824-38, 15-20 fms, sand and shell, 7 Feb. 1938). *CAS-SU 37366 (53,
PERU, Sechura Bay, VALERO Sta. 846-38, 6 fms, coarse sand, Hancock Expedition,
15 Feb. 1938). *USNM 70716 (3, CHILE, Valpariso Bay). *USNM 92367 (2, CHILE,
Valpariso). *USNM 101680 (8, GALAPAGOS, Indefatigable 1., Academy Bay,
Hancock Exped.). *USNM 101681 (1, PERU, Chincha 1.). *USNM 101682 (1,
GALAPAGOS IS., Isabela 1., Tagus Cove, Hancock Exped.). *USNM 101683 (2,
GALAPAGOS IS., Isabela 1., Tagus Cove, Hancock Exped.). ?USNM 102275 (8,
GALAPAGOS IS., Charles I. [specimens probably lost, not found during 1982 inven-
tory and not on shelves). *USNM 102276 (5, GALAPAGOS IS., Chatham 1., E. of
Wreck Bay). *USNM 102277 (3, GALAPAGOS IS., James Bay, James I.). *USNM
102278 (14, GALAPAGOS IS., Hood 1., Gardner Bay). *USNM 108279 (8,
GALAPAGOS IS., Hood 1., Gardner Bay). *USNM 102280 (4, MEXICO, Baja Califor-
nia, Thurloe Bay). *USNM 102281 (1, MEXICO, Baja California, Thurloe Bay).
*USNM 108256 (4, GALAPAGOS IS., Hood 1., Gardner Bay, 01°22'00"S,
089°40'00"W, 12 fms). *USNM 109944 (25, GALAPAGOS IS., James 1., Sullivan Bay,
00°17'05"S, 090°35'10"W). *USNM 119816 (32, GALAPAGOS IS.). *USNM 127761
(5, PERU, Don Martin 1., 11 °02'00"S, 077°40'00"W).
BranchiostoiiUljloridae Hubbs, 1922

Branchiostoma jloridae Hubbs, 1922: 2, 7-8 (original description; type locality:
"Tampa Bay, Florida, by the Steamer Fish Hawk (Sta. 7121)").
anus + 8.4 (6-11) postanal = 60 (56-64) total. Dorsal fin-chambers 286 (206-349).
length about 0.4. Anus near center of ventral lobe of caudal fin.
DISTRIBUTION Gulf of Mexico from southwestern Florida westward to Texas (Fig. 1).
B. jloridae-like materials are also known from Mexico, Panama, Colombia, and Ven-
ezuela (see taxonomic discussion).
MATERIAL USNM 84466 (holotype, FLORIDA, Tampa Bay, FISH HAWK Sta. 7121,
1901). USNM 39380 (1, paratype, FLORIDA, 28°50'N, 083°00'W). USNM 43556 (1,
paratype, FLORIDA, 26°20'N, 082°39"W, GRAMPUS Sta. 5111). USNM 43557 (3,
paratypes, FLORIDA, 25°23'N, 082°32'W, GRAMPUS Sta. 5068, 1889). USNM
43877 (6, paratypes, FLORIDA, Tampa Bay, Dec. 1891). USNM 44434 (1, Paratype,
FLORIDA, Fort Tampa, 19 Feb. 1893). USNM 44660 (20, paratypes, FLORIDA,
Pensacola Harbor, Jul. 1893). USNM 49719 (101, paratypes, FLORIDA, Tampa Bay,
FISH HAWK Sta. 7121, 1901). USNM 49720 (57, paratypes, Same data as holotype).
USNM 49721 (49, paratypes, FLORIDA, Tampa Bay, FISH HAWK Sta. 7121, 1901).
USNM49722(71,paratypes,FLORIDA, TampaBay,190l,FISHHAWKSta. 7121).
USNM 49723 (165, paratypes, FLORIDA, Tampa Bay, FISH HAWK Sta. 7108,
1901). USNM 49724 (165, paratypes, FLORIDA, Tampa Bay, FISH HAWK Sta.
7108, 1901). USNM 92365 (2, paratypes, FLORIDA, Tampa Bay, Dec. 1901 [records
say reentered from 43877 but date given as Dec. 1901 rather than Dec. 1891). USNM
92366 (2, paratypes, FLORIDA, Pensacola Harbor, Jul. 1983). GCRL 3621 (13,
FLORIDA, Fort Walton Beach, under hwy. (US98) bridge between Fort Walton and
Destin, sand bottom, 2ft, 22 March 1969). GCRL 17569 (3, LOUISIANA, Ship Shoal
Lease, 28°51'34"N, 91°07'52"W, 19S-N2000, 5.0 m, 21 Sep. 1978). GCRL 17572 (1,
LOUISIANA, Ship Shoal Lease, 28°51 '34"N, 91 °07'52"W, 19S-N2000, 5.0 m, 21 Sept.
1978). GCRL 13347 (5, FLORIDA, Lee Co., Inside Boca Grande Pass, Sta. JCW 68-47,
11 March 1968). ?GCRL 23584 (93, MISSISSIPPI, West Side of Hom 1., 30°14'47"N,
088°46'56"W, 1-2m, 11 Jul. 1990). SU-IU 9396 (2, Puerto Rico, Vieques 1., U.S. Fish
Comm. Porto Rico Exped., FISH HAWK, Sta. 6084, 6 Feb. 1899). SIO 53-98 (2, 35.0-
36.0 mm, TEXAS, off Beaumont, 29°10'N, 94°03'W). SIO 92-145 (313, 8.5-49.5 mm,
FLORIDA, Old Tampa Bay, 27°45.0'N, 082°35.0'W). UAIC 468 (4, ALABAMA,
Dauphin Island, 5 Dec. 1953). UAIC 810 (450, MISSISSIPPI, Hom 1., 4 Aug. 1960).
UAIC 811 (470, MISSISSIPPI, Hom 1., 23 Aug. 1960). UAIC 1110 (1708, FLORIDA,
Tampa Bay, June 1962). UAIC 1623 (85, FLORIDA, Santa Rosa, 22 July 1963). UAIC
1627 (24, MISSISSIPPI, Hom 1., 2 Aug. 1960). UAIC 1628 (17, MISSISSIPPI, Hom 1.,
17 July 1963). UAIC 1645 (11, MISSISSIPPI, Hom 1., 1960). UAIC 1804 (100,
MISSISSIPPI, Mississippi Sound). UAIC 1805 (2, MISSISSIPPI, Petit Bois 1., 15 Aug.
1961). UAIC 1807 (8, MISSISSIPPI, Hom 1., 28 July 1960). UAIC 2195 (14, MISSIS-
SIPPI, Ship 1., 11 Feb. 1960). UAIC 2196 (2, MISSISSIPPI, Ship 1., 28 June 1957).
UAIC 2197 (3, MISSISSIPPI, Hom 1., 18 Aug. 1960?). UAIC 2198 (3, MISSISSIPPI,
Ship 1., 22 May 1960). UAIC 2199 (1, MISSISSIPPI, Ship 1., 5 May 1957). UAIC 2200
(2, MISSISSIPPI, Ship 1., 1959). UAIC 3537 (1, FLORIDA, Choctawhatchee Bay, 8
Nov. 1968). UAIC 3756 (1, FLORIDA, Santa Rosa Sound). UAIC 4160 (7, MISSIS-
SIPPI, Petit Bois 1., 27 July 1975). UAIC 4246 (53, FLORIDA, Panama City). UAIC
4249 (9, MISSISSIPPI, Petit Bois 1., 29 July 1971 ). UAIC 5868 (3, ALABAMA, Mobile
Ship Channel). UAIC 6192 (3, FLORIDA, Little Sabin Bay, Jun.-July 1961). UAIC
6193 (1, FLORIDA, Little Sabin Bay, 12 July 1961). UAIC 6194 (1, FLORIDA, Little
Sabin Bay, 9 June 1961). UAIC 6195 (1, FLORIDA, Little Sabin Bay, 9 June 1961).
UAIC 6196 (1, FLORIDA, Little Sabin Bay, 9 June 1961). UAIC 6201 (1, FLORIDA,
Little Sabin Bay, 20 Aug. 1962). UAIC 6202 (29, FLORIDA, Little Sabin Bay, 2 May
1962). UAIC 6203 (23, FLORIDA, Little Sabin Bay, 2 May 1962). UAIC 6204 (39,
FLORIDA, Santa Rosa Sound, 2 May 1963). UAIC 6205 (21, FLORIDA, Santa Rosa
Sound, 2May 1963). UIAC 6206 (5, FLORIDA, Little Sabin Bay, 21 Aug. 1963). UAIC
6207 (13, FLORIDA, Little Sabin Bay, 21 Aug. 1963). UAIC 6208 (10, FLORIDA,
Escambia Bay, 23 Aprill963). UAIC 6209 (5, FLORIDA, Santa Rosa Sound, 20 Aug.
1963). UAIC 6210 (1, FLORIDA, Little Sabin Bay, 22 Aug. 1963). UAIC 6211 (2,
FLORIDA, Little Sabin Bay, 21 Aug. 1963). UAIC 6192 (3, Florida Little Sabin Bay).
UAIC 6193 (1, FLORIDA, Little Sabin Bay). UAIC 6194 (1, FLORIDA, Little Sabin
Bay). UAIC 6195 (1, FLORIDA, Little Sabin Bay). UAIC 6196 (1, FLORIDA, Little
Sabin Bay). UAIC 6197 (1, FLORIDA, Little Sabin Bay). UAIC 6198 (3, FLORIDA,
Little Sabin Bay). UAIC 6199 (1, FLORIDA, Pensacola Bay). UAIC 6200 (4,
FLORIDA, Santa Rosa 1.). UAIC 6201 (1, FLORIDA, Little Sabin Bay). UAIC 6202
(29, FLORIDA, Little Sabin Bay). UAIC 6203 (23, FLORIDA, Little Sabin Bay). UAIC
6204 (39, FLORIDA, Santa Rosa Sound). UAIC 6205 (21, FLORIDA, Santa Rosa
Sound). UAIC 6206 (5, FLORIDA, Little Sabin Bay). UAIC 6206 (13, FLORIDA,
Little Sabin Bay). UAIC 6208 (10, FLORIDA, Escambia Bay). UAIC 6209 (5,
FLORIDA, Santa Rosa Sound). UAIC 6210 (1, FLORIDA, Little Sabin Bay). UAIC
6211 (2, FLORIDA, Little Sabin Bay). USNM 43555 (3, GRAMPUS Sta. 5066, NO
OTHER DATA). USNM 43558 (1, FLORIDA, 25°17'N, 082°49'15"W, GRAMPUS
Sta. 5064). ?USNM 39438 (1, 34°57'00"N, 075°43'30"W, ALBATROSS Sta. 2597, 15
fms). ?USNM 112509 (8, FLORIDA, Manatee Co., 3 1/2 Mi +or-, S.W. Longboat
Pass). ?USNM 112510 (5, FLORIDA, Shore of Sarasota Bay, from Whitfield Estates to
Whitaker Bayou, 10 ft). ?USNM 116415 (1, Pepperfish Key, 29°15'30"N,
083°27'30"W, FISH HAWK Sta. 7166, 5.5 fms). ?USNM 120974 (4, FLORIDA,
29°08'45"N, 083°28'00"W, FISH HAWK Sta. 7170). ?USNM 120975 (6, FLORIDA,
Off Cape Romans). USNM 68509 (14, FLORIDA, Off Port Tampa, FISH HAWK, 4
fms). USNM 89361 (25, FLORIDA, St. Petersburg). USNM 89361 (1, FLORIDA, St.
Petersburg). USNM 116838 (1, FLORIDA, Tortugas Near White Shoal). USNM
116839 (1, FLORIDA, Tortugas, S. ofChanne1 Buoy).
Branchiostoma gambiense Webb, 1958

Branchiostoma gambiense Webb, 1958d: 627-634, figs. 1-2, table. 1 (original
description; type locality: '"Cape St. Mary' Station MB6/D1, off the Gambian
coast 13°10'N., 16°59'W. Smith sampler in five consecutive hauls as close
together as possible in 18 meters").
Maximum number of gonads unknown. Preanal fin-chambers 47 (35-61). Ratio of
tallest dorsal fin-chamber height to width, 2.0-3.3. Ratio of postatriopore length to
preatriopore length 0.30-0.37. Anus posterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Reaches 41 mm.
DISTRIBUTION Known solely from the material cited by Webb (l958d) from Gambia (Fig. 2).
MATERIALBMNH 1958.3.28.15 (holotype, 29 mm, GAMBIA, l3°10'N, l6°59'W, 0.1
m2 Smith sampler, taken in 5 consecutive hauls as close together as possible, 18 m,
shelly sand, 28 Dec. 1956). BMNH 1958.3.28.11-14 (16, paratypes, 27-32 mm, same
data as holotype). BMNH 1958.3.28.8-10 (6, non-type material but same data as
holotype).
Branchiostoma indicum (Willey, 1901)

Synonyms include:
Dolichorhynchus indicum Willey, 1901: 269-271 (original description; type locality
"Ceylon").
Branchiostoma indicum Tattersall, 1903a: 275, 285-286 (comparison of lancelet
species).
DIAGNOSIS Myotome formula: 41 (39-43) preatriopore + 14 atriopore to anus+ 14
(13-15) postanal = 69 (67-72) total. Dorsal fin-chambers 326 (304-395). Preanal fin-
chambers 49 (40-63). Maximum number of gonads 29. Ratio of tallest dorsal fin-
chamber height to width 2.0-4.5. Ratio of postatriopore length to preatriopore length
0.4-0.5. Anus anterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Attains a size of at least 27 mm (Prashad, 1934).
DISTRIBUTION So far known from Sri Lanka and the southeast coast of India (Fig. 3).
MATERIAL We know of no material of this species but Azariah (l965a) cites the
existence of 165 unregistered specimens from the type locality.
Branchiostoma lanceolatum (Pallas, 1774)

Synonyms include:
Limax lanceolatus Pallas, 1774: 19, pl. I, fig. 2 a-b (original description; type
locality: Cornwall, England; generic name preoccupied by Limax Linnaeus,
1858, Mollusca). Pallas, 1778: 24-25 (redescription).
Branchiostoma lubricum Costa, 1834:49 (original description; Mediterranean near
Naples).
Amphioxus lanceolatus: Yarrell, 1836a: xxvii (name only). Yarrell, 1836b: 468-472
(description; figs.; placed in Chondropterygii, Petromyzidae).
?Branchiostoma haeckelii Franz, 1922a: 403-405, fig. 18 (original description; type
locality: "Ceylon"; in key). Franz, 1930: 573-581, table. I (comparisons of
lancelet species). Nishikawa, 1982: 119-120, table. 1 (possible synonym of B.
lanceolatum; counts).
DIAGNOSIS Myotome formula: 36.1 (34-38) preatriopore + 13.8 ( 11-17) atriopore to
anus + 11 (l 0-14) postanal = 61 (59-65) total. Dorsal fin-chambers 212 (183-288).
MAXIMUM SIZE About 61 mm (Webb, 1957a).
DISTRIBUTION B. lanceolatum is the most widely distributed species in the genus
(Figs. 2 and 3). It is known from the North Sea, westward and southward through the
English Channel and the coasts of France, Spain, and Portugal into the Mediterranean
and Black seas. It is also reported from the Suez Canal (Grovel, 1933, 1936; Tortonese,
1962) and is widely distributed in the Indian Ocean, off the East African coast (Prenant,
1928; Webb 1957a; 1957b), off Oman (Webb, 1957a), and off India and Sri Lanka
(Tattersall, 1903a; Azariah, 1965a). It appears to reach the Solomon Islands (Gibbs and
Wickstead, 1969) and materials variously identified as B. moretonense, B. haeckelii, and

B. belcheri may belong to this species or species complex (see taxonomic discussion).
MATERIAL *?CAS-SU 41417 (2, INDIA, Madras, 5 Jan. 1941). *CAS-SU 32143 (1,
GERMANY, Helgoland). *UAIC 5412.01 (9, NORTHERN RED SEA, Gulf of Suez,
1.5 m, May 1965). *USNM 22083 (2, NORWAY, Stavanger). *USNM 23020 (3,
NORWAY, Stavanger). USNM 40073 (24, ITALY, Sicily, Lago Piciriddu). *USNM
48459 (4, ITALY, Bay of Naples). *USNM 48450 (4, ITALY, Bay of Naples). *USNM
124455 (25, ITALY, Messina, Lago Piciriddu).
Branchiostoma leonense Webb, 1956

Branchiostoma leonense Webb, 1956c: 175-182, figs. 4, 6, table. 4 (original de-
scription; type locality: "mouth of the Sierra Leone River estuary, Freetown,
Sierra Leone" and "Sierra Leone estuary and in the sea off the Sierra Leone
estuary in deposits of sandy mud, muddy sand, and muddy sand with shell").
DIAGNOSIS Myotome formula: 43.2 (40--45) preatriopore + 15.5(14-16) atriopore to
anus+ 10.4 (10-12) postanal =69.2 (66-73) total. Dorsal fin-chambers 382 (355--418).
MAXIMUM LENGTH31 mm (Webb, 1956c).
DISTRIBUTION Off West Africa Guinea and Sierra Leone (Fig. 2).
MATERIAL BMNH 1955.12.13.70 (holotype, 28 mm, SIERRA LEONE, mouth of
Sierra Leone River Estuary, Freetown). BMNH 1955.12.13.71-95 (paratypes, 20 miles
up Kumrabe Creek near Freetown, 14 March 1955). ZUMC P01.18 (1, Off FRENCH
GUINEA, 09°23'N, 15°07'W, ALANTIDE Exped. Sta. 45, van Veen Grab 0.1 m2
bottom-sampler, 36 m, fine yellow sand, 2000 h, 18 Dec. 1945).
Branchiostoma longirostrum Boschung, 1983

Branchiostoma longirostrum Boschung, 1983:91-97, figs. 1-3, tabls. 1-6 (original
description; type locality: "approximately 24 km S Mobile Bay, AL, 30°03'N lat,
88°03'W long; collected in sand from 18m depth in 1963;" comparisons with
other Florida Branchiostoma; key to Western Atlantic species).
DIAGNOSIS Myotome formula: 36.1 (34-38) preatriopore; 16.2 (15-19) atriopore to
anus+ 10.4 (10-12) postanal =69 (66-73) total. Dorsal fin-chambers 242(209-269).
dorsal fin-chamber height to width 4.0-6.5. Ratio ofpostatriopore length to preatriopore
length 0.37-0.51. Anus near or posterior to center of ventral lobe of caudal fin.
DISTRIBUTION Known from the northeast coast of Florida near Jacksonville south-
ward and westward around the Florida peninsula and extending to an offshore station in
southeastern Texas and Veracruz, Mexico (Fig. I). It has been taken at depths between
II and 40 m, usually in sand strewn with shell fragments.
MATERIAL USNM 257774 (holotype, 42.7 mm, approximately 24 km due South of
Entrance to Mobile Bay, AL, 30°03'N, 88°03'W, 80ft). USNM 257772 (6, paratypes,
15.446.9 mm TL; taken with holotype), UAIC 9168.01 (8, paratypes, 21.9-40.1 mm,
30°00'N, 86°18'W, 37m, 6 Sep. 1977). ?USNM 257773 (4, MISSISSIPPI, 29°58'30"N,
088°ll'W, 15 Aug. 1968). UAIC 3740.01 (5, 24.7-40.0, approximately 30 km SSW
Mobile Bay, AL, 29°58'N, 88°ll'W, 30m, 15 Aug. 1969). UAIC 4248.01 (32, 21.8-
46.8, 29°59'N, 88°03'W, 27 m, Nov. 1976). UWF 1467 (5, 28.9-36.5, 29°50'N,
86°05'W, 37m, Oct. 1975). UAIC 6737.01 (17, 25.7-49.2, approximately 16 km E of
Jacksonville, FL, 15 m, 5 Oct. 1976). UAIC 3721.01 (12, 12.0-49.1 mm, 24°56'N,
82°19'W, approximately 30m, 21 May 1967). UAIC 3740 (13, ALABAMA, 15 mi. S.
Mobile Bay, 81ft, 1960). UAIC 4248 (45, ALABAMA, Nov. 1976). UAIC 3729.02 (1,
22.7, 29°06'N, 94°IO'W, 13 m, silty sand, 5 Jan. 1966). UWF uncat (7, 29.3-47.2,
28°15'-28°45'N, 84°00'-84°25'W, approx. 37m, 3 Nov. 1971). UWF 2888 (3, 30.2-
36.0, 25°50'N, 83°31 'W, approximately 35 m, 13 July 1976. FSBC 3873 (1, 25.6,
27°37'N, 83°28'W, 37m, 19 Jan. 1964). UAIC 3706.01, I, 28.1, 9.7 km W of Anna
Maria 1., Florida, 13m, 7 Jul 1966). UAIC 3220.01 (2, 41.3-43.5, 25°15'N, 82°23'W,
approx. 30m, 19 May 1967). UAIC 6738.02 (34, 28°00'N, 8 W, RIV Silver Bay Sta.
5252,30 m, 10 Nov. 1963).
Branchiostoma malayanum Webb, 1956
Branchiostoma ma/ayana Webb, 1956a: 121-123, fig. I (original description; type

locality: "Changi at the extreme Eastern end of the Island of Singapore"; holotype
BMNH 1955.12.13.97, paratype BMNH 1955.12.13.98).
DIAGNOSIS Myotome formula: 27.8 (27-28) preatriopore + 16 atriopore to anus+ 8
=
postanal 51.8 (51-52) total. Dorsal fin-chambers 201 (189-221 ). Preanal fin-chambers
79 (52-92). Maximum number of gonads 18. Ratio of tallest dorsal fin-chamber height
to width 2.0. Ratio of postatriopore length to preatriopore length 0.53-0.60. Anus
anterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Attains length of at least 26 mm (Piyakamchana, 1962).
DISTRIBUTION Currently known from Singapore, Gulf of Thailand, and the Solomon
Islands (Fig. 3).
MATERIAL None available.
Branchiostoma moretonense Kelly, 1966

Branchiostoma moretonensis Kelly, 1966: 259-265, tabs. l-2, figs. 1-2 (original
description; type locality: "Moreton Bay, Queensland"; holotype: AMS IB 7396).

anus+ 8.1 (7-ll) postanal = 58.2 (56-60) total. Dorsal fin-chambers 234 (197-262).
Preanal fin-chambers 79.3 (62-92). Maximum number of gonads 26. Ratio of tallest
DISTRIBUTION Reported from Queensland and the Northern Territories from Moreton
Bay to Boucaut Bay.
MATERIAL AMS 18 7396 (holotype, 28.5 mm, QUEENSLAND, Moreton Bay). AMS
IB 7397 (paratypes, QUEENSLAND). AMS IB 7398 (paratypes, QUEENSLAND): (I,
5 miles W. of Tangalooma Pt., near M.3 Red Beacon, muddy sand, 20m, I 0 Nov. 1961 );
(I, 2 miles NNW of Rous Channel Light, coarse culch of mollusc shell, coral, and
muddy sand, 20 m, I June 1962); (26, I mile SE of Hope Banks, culch, II m, I June
1962); (19, N of Naval Reserve Beacon, culch, 4 m, 2 June 1962); (3, Wend ofRous
Channel, 6 m, culch, 2 June 1962). SMBL Rare-295 (I, 33.2 mm, Arafura Sea, Boucaut
Bay, 134°33.5'E, I 1°54'S, Nov. 1940. Other materials cited by Richardson and
McKenzie ( 1994 ).
BranchiostoiiUl nigeriense Webb, 1955

Synonyms include:
Branchiostoma nigeriense Webb, 1955: 423-434 (original description; type locality
"Lagos").
Branchiostoma takoradii Webb 1956c: 173-175, 180-181, tabls. 3-4, figs. 3, 6
(original description; type locality: "' Atlantide' Expedition Station 73, 4°50'N.
I 0 40'W. near Takoradi, Gold Coast").
=
anus+ 11.0 (10-12) postanal 68 (66-72) total. Dorsal fin-chambers 346 (330-376).
MAXIMUM SIZE Grows to 35 mm in Lagos Lagoon but reaching only about a third of
this length in the open sea (Webb, 1958b).
MATERIAL ZUMC P.OI6-15, (10, NIGERIA, Port Harcourt, Niger Delta, 0.1 m2
Petersen Grab, 8 m, sand, 1230 h, 21 Feb. 1946). ZUMC P.OI.5 (l, NIGERIA, Dowes 1.,
Niger Delta, 0.1 m2 Petersen Grab, 15m, sand, 1320 h, 21 Feb. 1946. ZUMC P.Ol.l6-17
(holotype, 25.5 mm, and Paratype of B. takoradii, GOLD COAST, 04°50'N, I 0 40'W,
near Takoradi, 0.1 m2 van Veen Grab, 33 m, sand with a little mud, I 018 h, 23 Jan. 1946.
BranchiostoiiUl plalae Hubbs, 1922

Branchiostoma platae Hubbs, 1922: I 0 (original description; type locality "Steamer
Albatross at Station 2765, off the mouth of Rio de La Plata, South America (lat
36°43'S, long. 56°23'W); depth, 10.5 fathoms").
DIAGNOSIS Myotome formula: 37.0 (3~2) preatriopore + 15.0 (9 17) atriopore to
=
anus + 6.8 (5-9) postanal 59 (55-65) total. Dorsal fin-chambers 278 (249-327).
length 0.28-0.32. Anus posterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Reaches 56 mm (Hubbs, 1922).
DISTRIBUTION This species lives near the mouth of the Rio de La Plata and in
southernmost Brazil and Uruguay (Fig. I).
MATERIAL *USNM 85498 (1, holotype, ALBATROSS Sta. 2765). USNM 92362 (5,
paratypes, same data as holotype ). *NRM 19368 (23, Rio de La Plata, part of type series
of B. caribaeum, several damaged, see text for discussion).
Branchiosto11Ul senegalense Webb, 1955

Branchiostoma senega/ense Webb, 1955: 422-425 (original description; type local-
ity "Goree-Plage (Senegal)").
anus+ 11.3 (10--12) postanal = 67 (65-71) total. Dorsal fin-chambers 293 (266-325).
dorsal fin-chamber height to width, 2.7-6.5. Ratio of postatriopore length to
preatriopore length 0.33-0.54. Anus anterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE 40 mm.
DISTRIBUTION Western Sahara south to Senegal (Fig. 2).
MATERIAL BMNH 1958.3.28.1 (1, 7 mm, GAMBIA, CAPE ST. MARY Sta. MB6/
A I, l3°30'N, l6°54'30"W, 16 m, shelly sand, 0.1 m2 Smith sampler in 5 hauls, 23-26
Dec. 1956 ). BMNH 1958.3.28.5 (I, 8 mm, GAMBIA, CAPE ST. MARY Sta. MB6/A2,
13°2l'N, l7°05'W, 28m, shelly sand, 0.1 m2 Smith sampler in 5 hauls, 23-26 Dec.
1956). *GCRL 1458 (14, SPANISH SAHARA, 23°20.0'N, 16°20.6'W, grab sample, 30
m, 30 Apr. 1965). *CAS-SU 19190 (40, SPANISH SAHARA, 20 Aug. 1952). *UAIC
(6, Cape D' Arguin, 30 April 1965).
Branchiosto11Ul tattersalli Hubbs, 1922

Synonyms include:
Branchiostoma ca/iforniense (not of Andrews): Tattersall, 1903b: 224 (in part;
specimen from Ceylon).
Branchiostoma tattersalli Hubbs, 1922: 2, 12-13 (original description; type locality
"Ceylon"; based solely on Tattersall description; whereabouts of type unknown).
Vo1.42, 1996 LANCELET TAXONOMY S-57
Branchiostoma gravelyi Prashad, 1934: 333 (original description; type locality

"dredged from offCoilapatam, SWx W ofTuticorin" [India]).
DIAGNOSIS Myotome formula: 37.0 (37-40) preatriopore + 16.0 (16--18) atriopore to
anus+ 9.0 (8-10) postanal = 72 (70--74) total. Dorsal fin-chambers 317 (300--333).
MAXIMUM SIZE 42 mm (Azariah, 1965a).
DISTRIBUTION Known only from off Sri Lanka to Madras on the Indian mainland (Fig.
3).
MATERIAL ?USNM 34003 (1, SRI LANKA, F. Day). ZSI Fll675 (holotype of B.
gravelyi, 44 mm, INDIA, off Coilapatam, SW x W of Tuticorin, dredged 30 March
1928. ?USNM 149736 (3, SRI LANKA, Muttuvaratu Paar). ?USNM 149737 (3, SRI
LANKA, Muttuvaratu). ?USNM 149738 (2, SRI LANKA, Negombo). ?USNM 149739
(6, SRI LANKA, Negombo, 8 fms).
BrrmchiostoiiUI virginille Hubbs, 1922
Branchiostoma virginiae Hubbs, 1922: 2, 8-9 (original description; type locality:

"Sewell's Point, Virginia, on Chesapeake Bay").
anus+ 8.4 (7-10) postanal = 61 (56--64) total. Dorsal fin-chambers 302 (219-344).
DISTRIBUTION Garman reported this species (identified as B. caribaeum) to range
from New York to the Gulf of Mexico. However, as pointed out by Bigelow and
Farfante (1948), no records are known from farther north than Chesapeake Bay. It
appears to range northward from Vera Beach, Florida (Fig. I).
MATERIAL *GCRL 17252 ( 173, GEORGIA, 31 °45'26"N, 080°29'03"W, BLM Sta. 4c-
0190-6, 20 m, 24 Feb. 1977). *GCRL 17253 (32, GEORGIA, 31°08'00"N,
080°49'57"W, BLM Sta. 5c-0220-2, 34.2 m, 25 Feb. 1977). *GCRL 17254 (31, GEOR-
GIA, 32°54'00"N, 079° 12'00"W, BLM 2b-O 131-2, 16m, 12 Feb. 1977). *GCRL 17255
(56, GEORGIA, 31 °08'00"N, 080°49'50"W, BLM Sta. 5c-0828-3, 13m, 31 Aug. 1977).
*GCRL 17258 (63, GEORGIA, 31°45'26"N, 080°29'03"W, BLM Sta. 4c-0190-l, 20.0
m, 24 Feb. 1977). *GCRL 17259 ( 16, GEORGIA, 30°23'00"N, 080°26'00"W, BLM Sta.
6e-0265-5, 39.0 m, I March 1977). *GCRL 17260 (65, 32°54'00"N, 79° 12'00"W, BLM
Sta. 2b-013101, 16m, 12 Feb. 1977). *GCRL 17263 (32, GEORGIA, 32°54'04"N,
79°II'59"W, BLM 2b-0474-2, 14.0 m, 14 May 1977). *USNM 84465 (holotype,
VIRGINIA, Just below low water mark 4 mi E of Sewall's Point, specimen disintegrat-
ing). *USNM 2763 (I, paratype, specimen disintegrating). *USNM 28644 (I, paratype,
MARYLAND, East Shore, 1880). *USNM 92368 (2, paratypes, VIRGINIA, Sewalls
Point). *UAIC 3278 (1, MIV GILL, Cr. l, Sta. 55). *UAIC 3283 (24, MIV GILL, Cr. 2,
Sta. 44,6 May 1953). *UAIC 3284 (25, MIV GILL, Cr. 2, Sta. 44,6 May 1953). *UAIC
3285 (1, MIV GILL, Cr. 4, Sta. 36,21 Oct. 1953). *UAIC 3286 (10, M1V GILL, Cr. 4,
St. 12, 14 Oct. 1953). *UAIC 3287 (I, MIV GILL, Cr. 4, Sta. 13, 14 Oct. 1951). *UAIC
3288 (94, MIV GILL, Cr. 5, Sta. 44, 14 Feb. 1954). *UAIC 3289 (35, MIV GILL, Cr.
5, Sta. 36, 10 Feb. 1954). *UAIC 3290 (1, MIV GILL, Cr. 5, Sta. 46, 15 Feb. 1954).
*UAIC 3291 ( l, MIV SILVER BAY Sta. 3343 ). *UAIC 3292 ( l, DELAWARE, Indian
River Inlet, 29 Oct. 1960). *UAIC 3293 (1, NORTH CAROLINA, Duke Marine Lab,
Beaufort Shelf Transect, 19 Apr. 1965). *UAIC 3294 (2, NORTH CAROLINA, Duke
Marine Lab, Beaufort Shelf Transect, 19 Apr. 1965). *UAIC 3295 (I, NORTH CARO-
LINA, Duke Marine Lab, Beaufort Shelf Transect, 25 Jun. 1965). *UAIC 3296 (1,
NORTH CAROLINA, Duke Marine Lab, Beaufort Shelf Transect, 24 Jun. 1965).
*UAIC 3296 (1, NORTH CAROLINA, Duke Marine Lab, Beaufort Shelf Transect, 24
Jun. 1965). *UAIC 3297 (1, NORTH CAROLINA, Duke Marine Lab, Beaufort Shelf
Transect, 24 Jun. 1965). *UAIC 3298 (1, NORTH CAROLINA, Duke Marine Lab,
Beaufort Shelf Transect, 24 Jun. 1965). *UAIC 3299 (1, NORTH CAROLINA, Duke
Marine Lab, Beaufort Shelf Transect, 30 Sep. 1965). *UAIC 3300 (1, NORTH CARO-
LINA, Duke Marine Lab, Beaufort Shelf Transect, 30 Sep. 1965). *UAIC 3401 (44,
NORTH CAROLINA, Duke Marine Lab, Beaufort Shelf Transect, 24 Jun. 1965).
*UAIC 3402 (I, NORTH CAROLINA, Duke Marine Lab, Beaufort Shelf Transect, 24
Nov. 1965). *UAIC 3403 (1, NORTH CAROLINA, Duke Marine Lab, Beaufort Shelf
Transect, 19 Jan. 1965). *UAIC 3404 (4, NORTH CAROLINA, Duke Marine Lab,
Beaufort Shelf Transect, 19 Jan. 1965). *UAIC 3405 (4, NORTH CAROLINA, Duke
Marine Lab, Beaufort Shelf Transect, 19 Jan. 1965). *UAIC 3406 (6, NORTH CARO-
LINA, Duke Marine Lab, Beaufort Shelf Transect, 19 Jan. 1965). *UAIC 3407 (1,
NORTH CAROLINA, Duke Marine Lab, Beaufort Shelf Transect, 8 Jan. 1965). *UAIC
3408 (1, RIV GOSNOLD Sta. 1456, 19 May 1964). *UAIC 3409 (5, RIV GOSNOLD
Sta. 1465,20 May 1964). *UAIC 3410 (1, RIV GOSNOLD Sta. 1468, 20 May 1964).
*UAIC 3411 (97, RIV GOSNOLD Sta. 1478, 20 May 1964). *UAIC 3412 (1, R1V
GOSNOLD Sta. 1479,21 May 1964). *UAIC 3413 (1, RIV GOSNOLD Sta. 1484,21
May 1964). *UAIC 3414 (11, RIV GOSNOLD Sta. 1485,21 May 1964). *UAIC 3415
(78, RIV GOSNOLD Sta. 1487,21 May 1964). *UAIC 3416 (I, RIV GOSNOLD Sta.
1489, 21 May 1964). *UAIC 3417 (6, RIV GOSNOLD Sta. 1490, 21 May 1964).
*UAIC 3418 (17, RIV GOSNOLD Sta. 1491, 21 May 1964). *UAIC 3419 (90, R1V
May 1964). *UAIC 3421 (2, RIV GOSNOLD Sta. 1494,21 May 1964). *UAIC 3422
(30, RIV GOSNOLD Sta. 1495,22 May 1964). *UAIC 3423 (4, RIV GOSNOLD Sta.
1500, 22 May 1964). *UAIC 3424 (20, RIV GOSNOLD Sta. 1505, 22 May 1964).
*UAIC 3425 (21, RIV GOSNOLD Sta. 1506, 22 May 1964). *UAIC 3426 (2, RIV
May I964). *UAIC 3428 (1, R/V GOSNOLD Sta. I5II, 24 May I964). *UAIC 3429
(50, R/V GOSNOLD Sta. I525, 24 May I964). *UAIC 3430 (2, R/V GOSNOLD Sta.
I527, 24 May I964). *UAIC 343I (2, R/V GOSNOLD Sta. 30,22 May I964). *UAIC
3432 (1, R/V GOSNOLD Sta. I539, 25 May I964). *UAIC 3433 (4, R/V GOSNOLD
Sta. I540, 25 May I964). *UAIC 3434 (1, R/V GOSNOLD Sta. I543, 25 May I964).
*UAIC 3435 (25, R/V GOSNOLD Sta. I544, 25 May I964). *UAIC 3436 (1, R/V
GOSNOLD Sta. I545, 25 May I964). *UAIC 3437 (2, R/V GOSNOLD Sta. I549, 25
May I964). *UAIC 3438 (2, R/V GOSNOLD Sta. I472, 20 May I964). *UAIC 3439
(1, R/V GOSNOLD Sta. I555, 25 May I964). *UAIC 3440 (1, R/V GOSNOLD Sta.
I659, 7 Jun. I964). *UAIC 344I (I, R/V GOSNOLD Sta. I665, 7 Jun. 1964). *UAIC
3442 (48, R/V GOSNOLD Sta. 1665,7 Jun. I964). *UAIC 3443 (2 R/V GOSNOLD
Sta. I667, 7 Jun. 1964). *UAIC 3444 (66 R/V GOSNOLD Sta. I670, 7 Jun. I964).
*UAIC 3445 (7 R/V GOSNOLD Sta. I67I, 8 Jun. I964). *UAIC 3446 (1, R/V
GOSNOLD Sta. I672, 8 Jun. I964). *UAIC 3447 (1, R/V GOSNOLD Sta. I677, 10
Jun. I964). *UAIC 3448 (60, R/V GOSNOLD Sta. I678, II Jun. I964). *UAIC 3449
(25, R/V GOSNOLD Sta. I679, II Jun. I964). *UAIC 3450 (7, R/V GOSNOLD Sta.
I682,11 Jun.I964). *UAIC345I (7,R/VGOSNOLDSta.I684,II Jun.I964). *UAIC
3452 (1, R/V GOSNOLD Sta. I685, II Jun. I964). *UAIC 3453 (14, R/V GOSNOLD
Sta. I687, II Jun. I964). *UAIC 3454 (25, R/V GOSNOLD Sta. 1688, II Jun. 1964).
*UAIC 3455 (5, R/V GOSNOLD Sta. I690, 12 Jun. 1964). *UAIC 3456 (2, R/V
GOSNOLD Sta. 1695, I2 Jun. I964). *UAIC 3457 (1, R/V GOSNOLD Sta. I698, 12
Jun. I964). *UAIC 3458 (1, R/V GOSNOLDSta. I700, I2 Jun. I964). *UAIC 3459(3,
R/V GOSNOLD Sta. I702, I2 Jun. I964 ). *UAIC 3460 (2, R/V GOSNOLD Sta. 1703,
I2 Jun. I964). *UAIC 346I (2, R/V GOSNOLD Sta. 1704, I2Jun. 1964). *UAIC 3462
(7, R/V GOSNOLD Sta. I706, I3 Jun. I964). *UAIC 3463 (1, R/V GOSNOLD Sta.
I710,13Jun.I964). *UAIC3464(2,R/VGOSNOLDSta.I7I3,13Jun.I964). *UAIC
3465 (2, R/V GOSNOLD Sta. I751, I6 Jun. I964). *UAIC 3466 (2, R/V GOSNOLD
Sta. I754, 16 Jun. I964). *UAIC 3467 (2, R/V GOSNOLD Sta. 1755, I6 Jun. I964).
*UAIC 3468 (2, R/V GOSNOLD Sta. I756, I6 Jun. I964). *UAIC 3469 (3, R/V
GOSNOLDSta.I759,I6Jun.I964). *UAIC3470(10,R/VGOSNOLDSta.I759,10
Jun. I964). *UAIC 347I (2, R/V GOSNOLD Sta. I76I, I7 Jun. I964). *UAIC 3472 (2,
R/V GOSNOLD Sta. I775, I8 Jun. I964). *UAIC 3473 (II, R/V GOSNOLD Sta.
I776,18Jun.I964). *UAIC3474(1,RIVGOSNOLDSta.I778,20Jun.I964). *UAIC
3475 (1, R/V GOSNOLD Sta. I78I, 20 Jun. 1964). *UAIC 3476 (1, R/V GOSNOLD
Sta. I786, 2I Jun. I964). *UAIC 3477 (1, R/V GOSNOLD Sta. I794, 21 Jun. I964).
*UAIC 3478 (2, R/V GOSNOLD Sta. I805, 22 Jun. I964). *UAIC 3479 (1, R/V
GOSNOLDSta. 1806, 22Jun. 1964). *UAIC 3480(1I, R/VGOSNOLDSta. 1807,22
Jun. I964). *UAIC 348I (1, R/V GOSNOLD Sta. I813, 23 Jun. I964). *UAIC 3482 (2,
R/V GOSNOLD Sta. I816, 23 Jun. I964 ). *UAIC 3483 (1, R/V GOSNOLD Sta. I8I8,
23 Jun. I964). *UAIC 3484 (7, R/V GOSNOLD Sta. I821, 23 Jun. I964). *UAIC 3485
(1, R/V GOSNOLD Sta. 1822, 23 Jun. 1964). *UAIC 3486 (6, R/V GOSNOLD Sta.
1845,25 Jun. I964). *UAIC 3487 (2, R/V GOSNOLDSta. I848, 25 Jun. I964). *UAIC
3488 (1, R/V GOSNOLD Sta. 186I, 26 Jun. I964). *UAIC 3489 (1, R/V ASTERIAS
Sta. 2250, 18 May 1964). *UAIC 3490 (1, RIV ASTERIAS Sta. 2308, I Jun. 1965).
*UAIC 3491 (1, RIV ASTERIAS Sta. 2317, 6 Jun. 1964). *UAIC 3492 (2, RIV
ASTERIAS Sta. 2318, 6 Jun. 1965). *UAIC 3493 ( 156, GEORGIA, Off Altamaha R.,
26Apr. 1963). *UAIC 4394 (1, 26 34'N, 82 16'W, 65ft, II Jan. 1961). *?UAIC 4239 (2,
RIV OREGON Sta. 17776, 24 Jun. 1975). *?UAIC 42409 (I, RIV OREGON Sta.
17780, 24 Jun. 1975). *?UAIC 4241 (12, RIV OREGON Sta. 17785, 25 Jun. 1975).
*?UAIC 4242 (6, RIV OREGON Sta. 19685,20 March 1976). *?UAIC 4243 (6, RIV
OREGON Sta. 19725, 23 March 1975). *?UAIC 4244 (I, RIV BOWERS Cr. I0 I, Sta.
2, 28 50'N, 8029'W, 20Apr. 1971). *?UAIC4245 (I, RIVBOWERSCr. 109, Sta. 6, II
May 1972). *UAIC 4471 (8, GEORGIA, E. Sapelo 1., Jan. 1970). *UAIC 4472 (1,
GEORGIA, E. Sapelo 1., Feb. 1970). *UAIC 4473 (16, GEORGIA, E. Sapelo 1., Feb.
1970). *UAIC 4474 (29, GEORGIA, E. Sapelo 1., Apr. 1970). *UAIC 4475 (2, GEOR-
GIA, E. Sapelo 1., Jun. 1970). *UAIC 4476 (16, GEORGIA, E. Sapelo 1., Jan. 1970).
*UAIC 4477 (I, GEORGIA, E. Sapelo 1., Aug. 1970). *UAIC 4478 (89, GEORGIA, E.
Sapelo 1., Aug. 1970). *UAIC 4479 (33, GEORGIA, E. Sapelo 1., Sep. 1970). *UAIC
4480 (256, GEORGIA, E. Sapelo 1., Nov. 1970). *USNM 119343 (5, NORTH CARO-
LINA, Sheepshead Shoal). *USNM 119345 (I, NORTH CAROLINA, Shackelford, Off
Shore I mi SW of Beaufort). *USNM 2, NORTH CAROLINA, Shackelford, Inside,
Beaufort). *USNM 154855 (37, VIRGINIA, Hampton). ?USNM 123005 (24, NORTH
CAROLINA?).
GENUS EPIGONICHTHYS
Epigonichthys Peters, 1877
Synonyms include:
Epigonichthys Peters, 1877: 391 (type species Epigonichthys cultellum by original
designation and monotypy).
Paramphioxus Haeckel, 1893: 461 (type species Branchiostoma bassanum Gunther,
1884, by original designation and monotypy). Gill, 1895:458 (diagnosis).
Asymmetron Andrews, 1893a: 237 (type species Asymmetron lucayanum Andrews,
1893, by monotypy).
Heteropleuron Kirkaldy, 1894: 686 (no type species designated [see taxonomic
discussion]). Kirkaldy, 1895: 314 (proposed as new subgenus of Branchiostoma;
diagnosis; included B. {H.) bassanum, B. {H.) cingalense, B. (H.) cultellum).
Amphioxides Gill, 1895: 458-459 (type species Branchiostoma pelagicum Gunther
by monotypy).
Bathyamphioxus Whitley, 1932: 257-258 (type species Asymmetron australis
Raffe, 1912, by original designation).
Merscalpel/us Whitley, 1932: 259 (type species Heteropleuron hedleyi Haswell by
original designation and monotypy).
Notasymmetron Whitley, 1932: 260-261 (type species Asymmetron caudatum
Willey by original designation and monotypy).
Zeamphioxus Whitley, 1932: 264 (type species Heteropleuron hectori Benham,

1902 by original designation and monotypy).
GENERIC DIAGNOSIS Gonads present on right side only. Notochord extending poste-
rior to myotomes for a variable length (but typically free of posterior myotomes for a
greater length than in species of Branchiostoma). Right metapleural fold confluent with
ventral fin. Ventral fin with or without fin-chambers in adults. Typically found in more
open water or pelagic environments than Branchiostoma, with gonads more often devel-
oped while still living in plankton.
ETYMOLOGY From the Greek combination epi (on)+ gonos (seed)+ ichthys (fish).
Gender masculine.
SPECIES OF EPIGONICHTHYS
Epigonichthys australis (Raffe, 1912)

Asymmetron australis Raffe, 1912: 303, pl. xxxvii (original description; type locality
"South of St. Francis Island, Great Australian Bight", 35 fms).
DIAGNOSIS Myotome formula: about 31.6 (29-34) preatriopore + 8.1 (7-9) atriopore
to anus + 14.3 (13-17) postanal = 54.1 (51-57) total. Dorsal fin-chambers 189 (158-
212). Preanal fin-chambers 18.2 (14-25). Counts from Richardson and McKenzie
(1994). Maximum number of gonads 22. Ratio of tallest dorsal fin-chamber height to
width unknown. Ratio of postatriopore length to preatriopore length unknown. Anus
MAXIMUM SIZE Reported to reach 25 mm.
DISTRIBUTION Known from Great Australian Bight at St. Francis Island eastward to
Tasmania (Richardson and McKenzie, 1994) on sandy or sand shell bottoms at depths of
45-185 m.
MATERIAL AMS IA4946 (I, paratype, south of St. Francis 1., Great Australian Bight).
Other material cited by Richardson and McKenzie ( 1994).
Epigonichthys bassanus (Gunther, 1884)

Branchiostoma bassanum GUnther, 1884: 31 (original description; type locality
"Bass Strait, Australia, I 0-12 fathoms").
Preanal fin-chambers 41 (35-46). Counts based on Richardson and McKenzie (1994).
Maximum number of gonads 30. Ratio of tallest dorsal fin-chamber height to width
about 2-3. Ratio of postatriopore length to preatriopore length about 0.21. Anus well
S-62 S.G. POSS AND H.T. BOSCHUNG lsr. J. Zoot.
MAXIMUM SIZE Reported to reach 75 mm (Last, et al., 1983).

DISTRIBUTION New South Wales (north to Sydney) southward to Victoria and Tasma-
nia (Fig. 3). Gomon et al. (1994) report this species from South Australia in depths of7-
84 m, whereas Richardson and McKenzie (1994) do not show the species taken this far
westward.
MATERIAL No specimens examined. Available materials listed by Richardson and
McKenzie (1994).
Epigonichthys cingaknsis (Kirkaldy, 1894)
Synonyms include:
Branchiostoma (Heteropleuron) Singalense Kirkaldy, 1894: 686 (inadvertent origi-
nal spelling in note presenting preliminary findings; lapsus calami?; species
name in upper case).
Branchiostoma (Heteropleuron) cingalense Kirkaldy, 1895: 315, pl. 34, fig. 7
(original description; type locality "Ceylon"; spelling corrected; Heteropleuron
proposed in subheadings as subgenus of Branchiostoma).
DIAGNOSIS Myotome formula: 39 (37-39) preatriopore + 16 (15-17) atriopore to anus
+ 8 (6-8) postanal = 63 (61-69) total. Dorsal fin-chambers -309. Preanal fin-chambers
-46. Maximum number of gonads 25. Ratio of tallest dorsal fin-chamber height to width
- 2-3. Ratio of postatriopore length to preatriopore length - 0.37. Anus anterior to
center of ventral lobe of caudal fin.
DISTRIBUTION Known only from the type specimens said to have been taken from
Ceylon (Sri Lanka) and specimens reported by Tattersall (1903b).
MATERIAL None available.
Epigonichthys culteUw Peters, 1877
Synonyms include:
Epigonichthys cultellus Peters, 1877: 322-327 (original description; type locality
[Moreton Bay]; [see taxonomic remarks]).
Epigonichthys pulchellus Giinther, 1880: 696 (nomen nudum, species name mis-
spelled).
Heteropleuron hedleyi Haswell, 1908: 33 (original description; type-locality
"Murray Is., Torres Strait").
Bathyamphioxusfranzi Whitley, 1932: 260 (new name proposed for specimen from
Shark Bay, Western Australia and questionably referred to Asymmetron cultellus
by Franz, 1927b: 426, fig. 28; no description based on Franz only).
DIAGNOSIS Myotome formula: 31 (26?, 29-36) preatriopore + 15 ( 13-18) atriopore to
anus + 9 (9-13) postanal =51 (48-52) total. Dorsal fin-chambers 220 (180-254).
Preanal fin-chambers 16 (8-22). Maximum number of gonads 20. Ratio of tallest dorsal
fin-chamber height to width 2.5-8.0. Ratio of postatriopore length to preatriopore length

0.38--0.45. Anus anterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE Reported to reach 40 mm (Franz, 1922b).
DISTRIBUTION From the Solomon Islands and Queensland northward and westward
through Indonesia, the Philippines, Western Australia, Thailand, and Sri Lanka, to
Bagamoyo, Tanzania.
MATERIAL AMS 1.9254 (2, paratypes, 28.0 mm, Murray 1., Torres Strait, in 5-8 fms).
AMS IA.2810 (15, AUSTRALIA, Queensland, Michaelmas Cay, near beach, 2 fms.,
dredged, sand and mud, 1-2 June 1926). *ZMA P.0140 (1, Off New Caledonia, DANA
Sta. 3614, 22°28'S, 166°27 .5'E, at surface, 28 Nov. 1928). Other available materials
listed by Richardson and McKenzie (1994).
Epigonichthys hectori (Benham, 1901)

Heteropleuron hectori Benham, 1901: 273-280, pl. 17, figs. 1-7 (original descrip-
tion; type locality: "East Coast of the North Island of New Zealand" [footnote
indicates that "these two specimens were collected at Awanui, just south of the
East Cape; whilst Hutton in the 'Catalog' gives the locality 'Poverty Bay,' which
is a little further [sic] south;" see Paulin, 1977 for discussion of type locality]).
DIAGNOSIS Myotome formula: 53 (53-55) preatriopore + 19 ( 19-20) atriopore to anus
+ 12 (11-12) postanal =84 (84-85) total. Dorsal fin-chambers- 308 (296-321 ). Preanal
fin-chambers -46 (44-49). Maximum number of gonads 33. Ratio of tallest dorsal fin-
chamber height to width 2-4. Ratio of postatriopore length to preatriopore length about
0.38. Anus posterior to center of ventral lobe of caudal fin.
MAXIMUM SIZE 43 mm.
DISTRIBUTION Inshore waters of the north island, New Zealand and on the northern
shores of the south island.
MATERIAL All from NEW ZEALAND. OM A76.2 (holotype, Awanui, south of East
Cape, near Waiapu River mouth, 37°50'S). NMNZ 3265 (paratype, Mahia Peninsula,
39° I O'S). Other materials listed by Paulin ( 1977).
Epigonichthys lucayanus (Andrews, 1893)

Synonyms include:
?Amphioxus lanceolatus (not of Pallas): Schmeltz, 1869: 29 (brief description).
Branchiostoma pelagicum Gunther, 1889: 43, pl. 6, fig. B (original description; type
locality: "lat. 23°3'N., 156°6' W., a few degrees north of Honolulu;" compared to
B. belcheri [see taxonomic discussion]).
Asymmetron lucayanum Andrews, 1893: 213-247, fig. (original description; type
locality "Alice Town, North Bimini, Bahamas"; common name: Bahama lance-
let; larval development; comparisons; counts).
Asymmetron caudatum Willey, 1896: 219, pl. 13, fig. 1-4 (original description; type
locality "lagoon of the Deboyne groups of islands, of which Panaieti is the
largest" [Louisaide Archipelago, Papua New Guinea]).
Asymmetron macricaudatum Parker, 1904: 47-49 (original description type locality:
"Salt Key Anchorage, Fla."; type series MCZ 26282 [should be 5 specimens not
7 and length from 10.5-14.0 mm]).
Asymmetron orientale Parker, 1904: 46, pl. I, fig. 4 (original description, type
locality "Hanimadu, Tiladummati Atoll" [Maldive Is.]).
?Amphioxides valdiviae Goldschmidt, 1905: 236-240, pl. I, fig. 1 (original descrip-
tion; exact type locality not given but in open ocean "aus den Sachtzen der
deutschen Tiefseexpedition").
Amphioxides stenurus Goldschmidt, 1905: 236-240 (original description; no type
locality given except indication that specimens were collected in open ocean "aus
den Schlitzen der deutschen Tiefseexpedition" in either Atlantic, Pacific, or
Indian Branchiostoma Ocean with most materials collected in the latter ocean).
Preanal fin-chambers 42 (0-68). Maximum number of gonads 29. Ratio of tallest dorsal
fin-chamber height to width 2.0-3.5. Ratio of postatriopore length to preatriopore length
0.37-0.49. Anus anterior to center of ventral lobe of caudal fin.
DISTRIBUTION Tropical and subtropical waters of the Atlantic, Pacific, and Indian
oceans at the surface to depths of915 m (as larvae).
MATERIAL *CAS 57279 [formerly NYZS 16896](1, Bermuda, 8 mile cylinder, the
center at 32°l2'N, 64°36'W, 500 fms, net 791, 9 July 1930). *CAS 66882 (1, PALAU,
Koror 1., Mud flat off SE tip of island, between Koror and Byobu Is., mud flat covered
with occasional blades of eel grass and scattered patches of soft coral, 07°l9'36"N,
134°29'06"E, 1530-1730 h, GVF 0814, 25 Jul. 1956). *GCRL 836 (1, PERSIAN
GULF, 27°10'N, 49°5l'E, 200 yds offshore from SW end of Jaraid 1., 5 Oct. 1956).
*GCRL 4187 (1, Marshall Is., Eniwetok Atoll, Grinem 1., ll 0 22'40"N, 162°l0'15"E,
25-75 ft, 23 Sept. 1969). SIO 84-235 (1, 20.5 mm, Marshall Is., Eniwetok Lagoon,
ll 0 30.0'N, 162°l5.0'E, 20 fms). SMBL (uncataloged). Western Caroline Is., Palao,
Koror 1., 6 Jul. 1940. Okinawa. SMBL Rare-293 (900 m east of Todumai on southeast-
em coast of Sesoko 1., northwest of Okinawazima 1., 15 m, 25 ¥ 25 ¥ 5 em dredge, 1
Nov. 1978). *UAIC 4247 (45, FLORIDA). *UAIC 4456 (1, FLORIDA, Palm Beach).
*USNM 164704 (3, BERMUDA). *USNM 44845 (9, BARBADOS, N. Bimini).
*USNM 92369 (2, BARBADOS, N. Bimini). *USNM 125926 (2, PUERTO RICO).
*USNM 141724 (MARSHALL IS., Bikini Atoll, in lagoon, dredge). *USNM 164693
(12, BERMUDA). *USNM 164695 (1, BERMUDA, Castle Harbor). *USNM 164699
(3, BERMUDA). *USNM 197646 (l, PERSIAN GULF, Approx. 3/4 mi of SW end of
Janna 1., 27°20'N, 049°49'E, 4.5 fms). *USNM 232741 (1, FLORIDA, l/4 mile east of
Fowey Light, 60ft, 20 Aug. 1966). *USNM 232742 (1, FLORIDA, 1 1/2 miles West-
Northwest ofFowey Light, 20-25 ft, 20 Aug. 1966). *?USNM 292808 (2, MARSHALL
IS., Eniwetok Atoll, Aug. 1975). *ZUMC P.Ol41 (2, DANA Sta. 3582 V, l5°36'S,
168°57'W, 100m, 27 Oct. !928). *ZUMC P.Ol42 (1, DANA Sta. 3604 II, 23°32'S,
167°36'E, 100m, 24 Nov. 1928). *ZUMC P.Ol43 (3, DANA Sta. 3604 III, 23°32'S,
167°36'E, 24 Nov. 1928). *ZUMC P.Ol44 (1, DANA Sta. 3611 II, 20°53.2'S,
164°03.3'E, 26 Nov. 1928). ZUMC P.Ol45 (11, DANA Sta. 3613 II, 22°43'S,
166°05.8'E, 28 Nov. 1928). ZUMC P.Ol46 (15, 22°43'S, 166°05.8'E, DANA 3613 III,
28 Nov. 1928). ZUMC P.0417 (27, 22°43'S, 166°05.8'E, DANA Sta. 3613 IV, 28 Nov.
1928). ZUMC P.0418 (13, 22°43'S, 166°05.8'E, DANA Sta. 3613 VII, 28 Nov. 1928).
ZUMC P.l049 (8, 22°43'S, 166°05.8'E, DANA Sta. 3613 IX, 28 Nov. 1928). ZUMC
P.l050 (4, 22°43'S, 166°05.8'E, DANA Sta. 3613 X, 28 Nov. 1928). *ZUMC P.0151 (3,
27°21'S, 175°ll'E, DANA Sta. 3623 II, 9 Dec. 1928). ZUMC P.Ol52 (15, 27°21'S,
175°ll'E, DANA Sta. 3623 V, 9 Dec. 1928). ZUMC P.0153 (1, 28°17.6'S, l77°0l'E,
DANA Sta. 3624 V, 10 Dec. 1928). ZUMC P.Ol54 (9, 22°28'S, 166°27.5'E, DANA Sta.
3614, surface, 28 Nov. 1928). ZUMC P.Ol55 (1, 28°19.5'S, !76°56'E, DANA Sta. 3624
VIII, 10 Dec. 1928). ZUMC P.Ol56 (5, 22°43'S, !66°05.8'E, 28 Nov. 1928).
Epigonichthys maldivensis (Forster Cooper, 1903)
Synonyms include:
Heteropleuron maldivense Forster Cooper, 1903: 349-352, pl. XVIII, fig. 2, and
figs. 77-81 (original description; type locality: "Maldive and Laccadive Is.";
common in Zanzibar; thought Asymmetron agassizi a synonym; [type material
lost, Wickstead, 1980]).
Heteropleuron agassizii Parker, 1904: 43-44, pl. 2, fig. 5 (original description; type
locality, Maldive Is., "Malandu, Miladummadulu Atoll;" thought related to
H. bassanum and especially to H. maldivense).
Heteropleuron parvum Parker, 1904: ~5. pl. 2, fig. 6, table. 2 (original descrip-
tion; type locality, Maldive Is., "sixteen fathoms of water at Hanimadu,
Tiladummati Atoll;" compared to other Heteropleuron species).
anus+ 11.8 (11-14) postanal =70.5 (67-73) total. Dorsal fin-chambers 285 (200-340).
MAXIMUM SIZE 31.2 mm (Nishikawa & Mukai, 1986).
DISTRIBUTION Ranges widely in the Pacific and Indian oceans, from Hawaii, New
Caledonia, Japan, the Maldive and Laccadive islands, Madagascar, and Tanzania. It has
been collected from near the surface to depths of about I ,097 m (Figs. 2 and 3).
MATERIAL NSMT-Ce 1001 (1, 27.0 mm, Japan, 5 km off east coast ofTanegasima 1.,
Sta. 21, 50 m, 18 June 1975). UAIC *USNM 120417 (MALDIVE IS., Mahlosmadulu
Atoll).
ACKNOWLEDGMENTS
The authors wish to thank a number of colleagues who provided materials and kindly
assisted this study in other ways. These are: David Catania, William Eschmeyer, and
Tomio Iwamoto (CAS); Elizabeth Heal and Gordon Gunter (formerly GCRL); Meredith
Blackwell and Richard Shaw (LSU); Karsten Hartel (MCZ); Guy Duhamel (MNHN);
Chris Paulin (NMNZ); Sven Kuhlander (NRM); Keiichi Matsuura (NSMT); Cindy
Klepaldo, Richard Rosenblatt, and H. J. Walker, (SIO); Richard Mayden and Bernard
Kuhajda, UAIC; William Fink and Douglas Nelson (UMMZ); Susan Jewett and Jeffrey
Williams (USNM), Teruaki Nishikawa, Nagoya University; H. Wilkens (ZMH); J!Zirgen
Nielsen (ZMUC). W. Mike Howell, Samford University, and Hector Harima, Florida
Junior College, Jacksonville, and Matthew Thomas, Morehead State University, pro-
vided assistance in data collection. The authors also wish to thank Carl Gans and
Norman Kemp for their efforts on our behalf.
NON-LANCELET REFERENCES CITED
Lancelet references are in the bibliography of the lancelets compiled for this volume.
Blackith, R.E. and Reyment, R. 1971. Multivariate Morphometries. Academic Press, New York
and London, 412 pp.
Carlton, J.T. and Geller, J.B. 1993. Ecological roulette: the global transport of nonindigenous
marine organisms. Science 261: 78-82.
Leviton, A.E., Gibbs, R.H. Jr., Heal, E., and Dawson, C.E. 1985. Standards in herpetology and
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Poss & Boschung 1996 Lancelets Valid Species

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ISRAEL JOURNAL OF ZOOLOGY, Vol. 42, 1996, pp.

LANCELETS (CEPHALOCHORDATA: BRANCmOSTOMATIDAE):

STUART G. Poss• AND HERBERT T. BoscHUNOb

NAMES AND CHARACTERS

regional differences in myotome number and proposed that myotomes be counted as

Branchiostoma bazarutense - - -340? 62 Gilchrist ( 1923 ); p

(n=9) (36-39) (17-19) (9-11) (63-66) (258-306) (70-88) ~

Branchiostoma bennetti 35.6 16.1 8.6 60 293 47 This work >

Branchiostoma bermudae 34.9 14.0 6.6 55 225 23 This work ~

Branchiostoma /eonense 43.2 15.5 10.4 69 382 58 VVebb(l956c; 1958a)

Branchiostoma /ongirostrum 36.1 16.2 9.1 61 242 40 This work

(n = ?) (41-44) (6-8) (7-10) (56-59) (195) - p

MULTIV ARIATE ANALYSIS

DISCUSSION OF TAXONOMY AND USE OF NAMES

contributions to marine biology (Hubbs, 1922; Bigelow and Farfante, 1948).

chambers, which is consistent with an immature specimen of E. culte/lus. Although only

Lancelets belong to the Phylum Chordata, Subphylum Acrania, Class Cephalochordata,

Synonyms include: Amphioxididae Gray, 1842: 150 (diagnosis). Amphioxini Muller,

Branchiostoma Costa, 1834

Branchiostoma africae Hubbs, 1927

Branchiostoma africae Hubbs in Monod, 1927: 644 (original description; type

Branchiostoma arabiae Webb, 1957

Branchiostoma arabiae Webb, l957a: 121-128 (original description; type locality:

MAXIMUM SIZE61 mm.

Branchiostoma bazarutense Gilchrist, 1923

Branchiostoma belcheri Gray, 1847

Branchiostoma lanceolatum belcheri: Tattersall, 1903b: 269-299, 272, 274-275,

Branchiostomtl bennetti Boschung and Gunter, 1966

DIAGNOSIS Myotome fonnula: 35.6 (34-38) preatriopore + 16.1 ( 13-19) atriopore to

Branchiostoma bermudae Hubbs, 1922

DISTRIBUTION Restricted to Bermuda, where it is found nearshore to depths of about

Branchiostoma californiense Andrews, 1893

Branchiostoma- Cooper, in Cronise 1868: 498 (coast of California [see taxonomic

Branchiosto11Ul capense Gilchrist, 1902

Branchiosto11Ul caribaeum Sundevall, 1853

Branchiostoma elongatum Sundevall, 1852

BranchiostoiiUljloridae Hubbs, 1922

Branchiostoma gambiense Webb, 1958

Branchiostoma indicum (Willey, 1901)

Branchiostoma lanceolatum (Pallas, 1774)

Wickstead, 1969) and materials variously identified as B. moretonense, B. haeckelii, and

Branchiostoma leonense Webb, 1956

Branchiostoma longirostrum Boschung, 1983

Branchiostoma malayanum Webb, 1956

Branchiostoma ma/ayana Webb, 1956a: 121-123, fig. I (original description; type

Branchiostoma moretonense Kelly, 1966

DIAGNOSIS Myotome formula: 32.2 (28-33) preatriopore + 17.9 (15-20) atriopore to

BranchiostoiiUl nigeriense Webb, 1955

BranchiostoiiUl plalae Hubbs, 1922

Branchiosto11Ul senegalense Webb, 1955

Branchiosto11Ul tattersalli Hubbs, 1922

Branchiostoma gravelyi Prashad, 1934: 333 (original description; type locality

BrrmchiostoiiUI virginille Hubbs, 1922

Branchiostoma virginiae Hubbs, 1922: 2, 8-9 (original description; type locality:

Epigonichthys Peters, 1877

Zeamphioxus Whitley, 1932: 264 (type species Heteropleuron hectori Benham,

Epigonichthys australis (Raffe, 1912)

Epigonichthys bassanus (Gunther, 1884)

MAXIMUM SIZE Reported to reach 75 mm (Last, et al., 1983).

Epigonichthys cingaknsis (Kirkaldy, 1894)