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Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 21
Seven new species within western Atlantic Starksia
atlantica, S. lepicoelia, and S. sluiteri
(Teleostei, Labrisomidae), with comments on
congruence of DNA barcodes and species
Carole C. Baldwin1,†, Cristina I. Castillo1,‡, Lee A. Weigt1,§, Benjamin C. Victor2,|
1 National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 2 Ocean Science
Foundation, 4051 Glenwood, Irvine, CA 92604 and Guy Harvey Research Institute, Nova Southeastern Uni-
versity, 8000 North Ocean Drive, Dania Beach, FL 33004
† urn:lsid:zoobank.org:author:B21379DC-6FF2-4C99-89FC-F9FD5B30911D
‡ urn:lsid:zoobank.org:author:AC20D703-1D8F-460C-A9CA-5792D337A4B4
§ urn:lsid:zoobank.org:author:64AF09A5-7954-431E-A414-BFFCEB47BE59
| urn:lsid:zoobank.org:author:E06A8680-6459-4978-9C59-A8467DB761EC
Corresponding author : Carole C. Baldwin ( baldwinc@si.edu )
Academic editor : Nina Bogutskaya|Received 2 October 2010|Accepted 28 January 2010|Published 3 February2011
urn:lsid:zoobank.org:pub:03FE1035-387F-496C-8929-C529876E1082
Citation: Baldwin CC, Castillo CI, Weigt LA, Victor BC (2011) Seven new species within western Atlantic Starksia
atlantica, S. lepicoelia, and S. sluiteri (Teleostei, Labrisomidae), with comments on congruence of DNA barcodes and
species. ZooKeys 79 : 21 – 72 . doi: 10.3897/zookeys.79.1045
Now this is not the end. It is not even the beginning of the end. But it is, perhaps, the end
of the beginning. W. Churchill
Abstract
Specimens of Starksia were collected throughout the western Atlantic, and a 650-bp portion of the mi-
tochondrial gene cytochrome oxidase-c subunit I (COl) was sequenced as part of a re-analysis of species
diversity of western Central Atlantic shore shes. A neighbor-joining tree constructed from the sequence
data suggests the existence of several cryptic species. Voucher specimens from each genetically distinct
lineage and color photographs of vouchers taken prior to dissection and preservation were examined for
diagnostic morphological characters. e results suggest that S. atlantica, S. lepicoelia, and S. sluiteri are
species complexes, and each comprises three or more species. Seven new species are described. DNA data
ZooKeys 79: 21–72 (2011)
doi: 10.3897/zookeys.79.1045
www.zookeys.org
Copyright C.C . Baldwin et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
RESEARCH ARTICLE
Launched to accelerate biodiversity research
A peer-reviewed open-access journal
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
22
usually support morphological features, but some incongruence between genetic and morphological data
exists. Genetic lineages are only recognized as species if supported by morphology. Genetic lineages within
western Atlantic Starksia generally correspond to geography, such that members of each species complex
have a very restricted geographical distribution. Increasing geographical coverage of sampling locations
will almost certainly increase the number of Starksia species and species complexes recognized in the west-
ern Atlantic. Combining molecular and morphological investigations is bringing clarity to the taxonomy
of many genera of morphologically similar shes and increasing the number of currently recognized spe-
cies. Future phylogenetic studies should help resolve species relationships and shed light on patterns of
speciation in western Atlantic Starksia.
Keywords
Starksia, DNA Barcoding, new species, species complex, biogeography
Introduction
e description of six new species of Caribbean Starksia by Williams and Mounts
(2003) capped more than 100 years of systematic research on this New World labri-
somid genus. It would have been reasonable to assume after such e ort that there is
little about the group left to discover. But the utilization of modern DNA barcoding
techniques in taxonomic studies is revealing the need to reanalyze existing species clas-
si cations of many groups of animals and, in combination with traditional morpho-
logical analyses, resulting in the recognition of numerous new species (e.g., Crawford
et al. 2010, Hebert et al. 2004, Pauls et al. 2010, Pöppe et al. 2010, Ward et al. 2008,
Zemlak et al. 2009). Western Atlantic shore shes are no exception (e.g., Tornabene et
al. 2010; Victor 2007, 2010). Particularly for small cryptic reef shes such as Starksia
blennies, we do not know where we stand in terms of understanding species diversity,
and our current concepts may be surprisingly incomplete.
Starksia shes inhabit shallow to moderately deep (to ca. 30 m) rock and coral reefs
in the western Central Atlantic and eastern Paci c oceans. ey are small (Atlantic spe-
cies are generally < 40 mm SL) and cryptic, but they often exhibit bright orange or red
coloration in life. Twenty-one species are currently recognized in the western Atlantic
(Williams and Mounts 2003), six of which are considered members of the S. ocellata
species complex (Green eld 1979).
e purpose of this paper is to describe the systematic results of our recent genetic
and morphological investigations of western Atlantic Starksia, work that was prompted
by our discovery of incongruences between preliminary genetic data and the current
species classi cation. We describe seven new species within S. atlantica, S. lepicoelia,
and S. sluiteri and provide keys to the species of each of those species complexes. We
provide photographs of living and preserved pigment patterns to help in future iden-
ti cations of the included species and in distinguishing them from western Atlantic
Starksia species likely to be discovered in the future. Finally, we discuss geographical
distributions of Starksia species and comment on congruence between DNA barcod-
ing data and morphologically recognizable species.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 23
Materials and methods
Specimens used in this study were collected from Barbados, Belize, Bahamas, Curacao
(Netherland Antilles), Florida, Honduras, Panama (Atlantic), Saba Bank (Netherland
Antilles), St. omas (U.S. Virgin Islands), Tobago (Trinidad and Tobago), and Turks
and Caicos. at material and additional museum specimens examined are listed in
the appropriate species and comparisons sections. Starksia specimens included in the
genetic analysis but not in the species accounts are tabulated in Appendix 1. Institu-
tional abbreviations for collections follow Sabaj Pérez (2010).
Specimens were collected with quinaldine sulfate, rotenone, or clove oil using
snorkel gear or scuba depending on depth. Field protocol involved taking digital
color photographs of fresh color patterns and subsequently a tissue sample (mus-
cle, eye, or n clip) for genetic analysis. For many, particularly small specimens,
it was necessary to remove the posterior 1/3 to 1/2 of the body to obtain enough
tissue for genetic analysis. Voucher specimens were preserved and later used to in-
vestigate diagnostic morphological features of each recovered genetic lineage. Field
measurements of standard length (SL), to the nearest 0.5 mm, were made by viewing
specimens against a plastic ruler under a dissecting microscope. Lengths of voucher
specimens generally were not re-measured in the lab because many vouchers are
now incomplete specimens. ose that were measured in the lab were measured to
the nearest 0.1 mm with digital calipers or with the aid of an ocular micrometer in
a dissecting microscope. Lengths of head (HL) and genital papilla were measured
to the nearest 0.1 mm with the same ocular micrometer and microscope. To ensure
that we were not introducing bias due to shrinkage of specimens after preservation,
head length as a percentage of SL was calculated only for specimens in which both
measurements were made from preserved specimens. Counts of dorsal-, anal-, and
caudal- n rays were made from digital radiographs of specimens, from preserved
specimens, or from photographs of voucher specimens taken prior to dissection. We
followed Böhlke and Springer (1961) in counting the last two segmented rays of
the dorsal and anal ns separately. Lateral-line scales were not counted because too
many scales are missing on most specimens. is is likely due to the long time the
specimens were held for processing prior to preservation and the physical manipula-
tion of the specimens during processing. Pores from the circumorbital ossi cations
are either uniserial or paired; the positions of any paired pores are described based
on their position relative to the orbit as though it were a clock; on the left side, for
example, a pair of pores at 3 o’clock is on the posterior margin of the orbit, a pair at
6 o’clock is on the ventral margin.
Molecular techniques employed at the Smithsonian are as described below. Meth-
ods utilized to sequence DNA from specimens from Barbados, Honduras, Panama, and
St. omas are as outlined in Victor (2010). Tissue samples for molecular work were
stored in saturated salt bu er (Seutin et al. 1990) or in 95% ethanol. Genomic DNA
was extracted from up to approximately 20 mg minced preserved tissue via an automat-
ed phenol:chloroform extraction on the Autogenprep965 (Autogen, Holliston, Massa-
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
24
chusetts) using the mouse tail tissue protocol with a nal elution volume of 50 μL. For
polymerase chain reaction (PCR), 1 μL of this genomic DNA was used in a 10 μL re-
action with 0.5 U Bioline (BioLine USA, Boston, Massachusetts) Taq polymerase, 0.4
μL 50 mM MgCl2, 1 μL 10× bu er, 0.5 μL 10 mM deoxyribonucleotide triphosphate
(dNTP), and 0.3 μL 10 μM each primer FISH-BCL (5’-TCAACYAATCAYAAAGA-
TATYGGCAC) and FISH-BCH (5’-TAAACTTCAGGGTGACCAAAAAATCA).
e thermal cycler program for PCR was 1 cycle of 5 min at 95°C; 35 cycles of 30 s
at 95°C, 30 s at 52°C, and 45 s at 72°C; 1 cycle of 5 min at 72°C; and a hold at 10°C.
e PCR products were puri ed with Exosap-IT (USB, Cleveland, OH) using 2 μL
0.2× enzyme and incubated for 30 min at 37°C. e reaction was then inactivated for
20 min at 80°C. Sequencing reactions were performed using 1 μL of this puri ed PCR
product in a 10 μL reaction containing 0.5 μL primer, 1.75 μL BigDye bu er, and 0.5
μL BigDye (ABI, Foster City, California) and run in the thermal cycler for 30 cycles of
30 s at 95°C, 30 s at 50°C, 4 min at 60°C, and then held at 10°C. ese sequencing
reactions were puri ed using Millipore Sephadex plates (MAHVN-4550; Millipore,
Billerica, Massachusetts) per manufacturer’s instructions and stored dry until analyzed.
Sequencing reactions were analyzed on an ABI 3730XL automated DNA sequencer,
and sequence trace les were exported into Sequencher 4.7 (GeneCodes, Ann Arbor,
MI). Using the Sequencher program, ends were trimmed from the raw sequences until
the rst and last 10 bases contained fewer than 5 base calls with a con dence score
(phred score) lower than 30. After trimming, forward and reverse sequences for each
specimen were assembled. Each assembled pair was examined and edited by hand, and
each sequence was checked for stop codons. Finally the consensus sequence (655 bp)
from each contig was aligned and exported in a nexus format (sensu Swo ord 2002).
A neighbor-joining tree (Saitou and Nei 1987) and distance matrix were generated
using Paup*4.1 (Swo ord 2002) on an analysis of Kimura two-parameter distances
(Kimura 1980). e neighbor-joining tree is not intended to re ect phylogenetic re-
lationships. e labels for each entry on the tree is our DNA number, and we include
that number in the material examined sections and gure captions. Abbreviations used
in DNA numbers re ect geographical location: BAH – Bahamas, BAR – Barbados,
BLZ – Belize, BRZ – Brazil, CUR – Curacao, FLA – Florida, HON – Honduras, PAN
– Panama, SAB – Saba Bank (Netherland Antilles), STVI – St. omas Virgin Islands,
TCI – Turks and Caicos, TOB – Tobago. COI sequences are deposited in Genbank
(accession numbers HQ543038-HQ543055, HQ571151-HQ571164, HQ600864-
HQ600963).
Results
A neighbor-joining tree derived from western Atlantic Starksia COl sequences is
shown in Fig. 1. irteen of the 21 currently recognized western Atlantic Starksia
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 25
species are represented in the tree: S. atlantica, S. culebrae, S. elongata, S. fasciata, S.
guttata, S. hassi, S. lepicoelia, S. multilepis, S. nanodes, S. occidentalis, S. ocellata, S.
sluiteri, and S. starcki. Four species, S. culebrae from the U.S. Virgin Islands, S. gut-
tata from Tobago, S. occidentalis from Belize, and S. ocellata from Florida, cluster on
the tree but represent genetically distinct lineages. ose results support Green eld’s
(1979) recognition of a S. ocellata species complex with several allopatric compo-
nent species. Similarly, S. atlantica, S. lepicoelia, S. nanodes, and S. sluiteri comprise
multiple, geographically distinct, genetic lineages, suggesting that they also represent
species complexes comprising multiple allopatric species. We do not deal further with
the S. nanodes complex in this paper because no genetic data is available from the
type locality, Bahamas, and we are thus uncertain if any of the four genetic lineages
on the tree (Barbados, Belize, Panama, and Saba Bank) represents S. nanodes Böhlke
and Springer 1961. We also do not deal further with ve species, S. elongata, S. fas-
ciata, S. hassi, S. multilepis, and S. starcki (but see discussion of S. fasciata under the
S. sluiteri complex section). Each of those species is represented in our material from
only one geographical location, and material from additional geographic locations is
needed to determine if they represent species complexes. We note that our material
of S. elongata, S. fasciata, S. hassi, and S. multilepis is from the type localities of those
species or relatively close by; the type locality of S. starcki, however, is Florida, and our
specimen is from Belize.
e multiple genetic lineages within S. atlantica, S. lepicoelia, and S. sluiteri are
the focus of the species treatments below. For each complex, we discuss congruence
of the component genetic lineages with results of our morphological investigation.
When diagnostic morphological features (primarily pigment) support the genetic
data, we recognize genetic lineages as species. Green eld (1979) noted that the abil-
ity to identify individuals of the S. ocellata complex to species based on morphol-
ogy without prior knowledge of locality supports the recognition of the component
populations as species vs. subspecies. We concur, and believe that the addition of
the COl data strengthens this argument. ere are no available names for new spe-
cies within any of S. atlantica, S. lepicoelia, and S. sluiteri complexes, and the seven
unnamed species discovered are described herein as new. Keys to the species of the
S. atlantica, S. lepicoelia, and S. sluiteri complexes are provided. We suggest that
readers use the taxonomic key to western Atlantic Starksia provided by Williams
and Mounts (2003) to identify S. atlantica, S. lepicoelia, and S. sluiteri and the keys
in this paper to distinguish the members within each complex. Note that the sixth
couplet of the Williams and Mounts (2003) key contains an error: 6b should lead
the user to couplet 10, not 9 as indicated. e geographical locations listed for each
species in our keys are the type locality plus any additional localities for which we
have genetic data. Additional collecting and study are needed to determine the dis-
tributions of all western Atlantic Starksia species. Distance matrices for intra- and
interspeci c variation in COl sequences for the S. atlantica, S. lepicoelia, and S. slui-
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
26
teri species complexes are provided in tables within the text. A distance matrix for all
lineages is in Appendix 2.
Starksia atlantica Species Complex
Longley (1934) described Starksia atlantica from a single specimen from Andros
Island, Bahamas. e neighbor-joining tree derived from COI sequences (Fig. 1)
includes ve distinct genetic lineages in the S. atlantica complex. e lineages from
Barbados (BAR) and Panama (PAN) are known only from larvae or juveniles and
are not discussed further. e Panama lineage is highly divergent in COl, and it
likely represents a cryptic species within S. atlantica or one of the eight western
Atlantic Starksia species not identi ed in our material. e other three lineages—
Curacao (CUR), Saba Bank (SAB), and Bahamas/Turks and Caicos/Belize (BAH/
TCI/BLZ) comprise specimens originally identi ed as S. atlantica on the basis of
absence of an orbital cirrus. (Note: Williams and Mounts (2003) correctly noted
the absence of an orbital cirrus as diagnostic for S. atlantica in their key to western
Atlantic Starksia [p. 147], but they erroneously stated “orbital cirri present” in their
treatment of the species [p. 160].) Within the BAH/TCI/BLZ lineage, there are
three sublineages, two from Belize and one from Bahamas/Turks and Caicos Is-
lands (or four if the latter is viewed as two). We have identi ed the specimens from
Bahamas and Turks and Caicos as S. atlantica (Longley) based on the type locality
(Bahamas) and pigment pattern, speci cally the presence of two or three rows of
block-like blotches on the trunk that are irregular in size and shape (Böhlke and
Springer 1961). We found no consistent di erences between specimens from the
Bahamas and Turks and Caicos.
e two Belize sublineages di er from other members of the S. atlantica complex
by the presence of regular, vertical, brown bars on the trunk separated by narrow white
interspaces and a well-de ned horseshoe-shaped blotch on the cheek. Although those
two sublineages are genetically similar to S. atlantica, we recognize the two lineages
from Belize as a distinct species based on their strikingly di erent pigment pattern
and geographic separation. We found no consistent morphological variation between
the two Belize sublineages and treat them as a single new species. Two specimens of
this new species were illustrated as S. atlantica by Green eld and Johnson (1981:
Fig.3A,B), who noted consistent di erences in pigmentation on the body between
their material from Belize and Honduras and the description of pigmentation for S.
atlantica by Böhlke and Springer (1961). e other two genetic lineages of S. atlantica
(Fig. 1) are from Curacao (CUR) and Saba Bank (SAB). e Curacao specimens have
a distinctive pattern of pigment on the cheek and pectoral- n base, and we recognize
them as a distinct species. e single sequence from Saba Bank likely represents a new
species (Fig. 1), but additional material is needed to con dently assess its status (see
“Remarks” under “Starksia sp.” below). We describe two new species within the S. at-
lantica complex, S. sangreyae from Belize and S. springeri from Curacao.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 27
Starksia sangreyae Castillo & Baldwin, sp. n.
urn:lsid:zoobank.org:act:F61A042F-F042-48EA-B4E1-C7AD79866916
Figs 1–2, 4; Table 1
Starksia atlantica, Green eld and Johnson (1981), Fieldiana Zoology 8: Fig. 3A–B
(black and white drawings of male and female specimens from Belize)
Type Locality: Belize, Central America
Holotype. USNM 398932, BLZ 5111, male, 16.0 mm SL, sta. CB05-9, south
side of island, Carrie Bow Cay, Belize, 1–2 m, 25 April 2005, C. Baldwin, D. Smith,
L. Weigt, J. Mounts (small llet removed from right side for DNA tissue sampling).
Paratypes (all Belize). Note – posterior portion of body destroyed for DNA tis-
sue sampling of all paratypes except USNM 276147 and 321073, which are not DNA
vouchers. USNM 398939, BLZ 8031, female, 18.0 mm SL, sta. CB08-2, sand bottom
and coral heads, Curlew Cay, 16˚47'24.1"N, 88˚04'41.0"W, 5–8 m, 15 May 2008.
USNM 398933, BLZ 5033, female, 16.5 mm SL, sta. CB05-3, spur and grove, Car-
rie Bow Cay, 9–22 m, 22 April 2005. USNM 398936, BLZ 8028, male, 17 mm SL,
sta. CB08-2 (see CB08-2 above). USNM 398934, BLZ 5161, female, 17.0 mm SL,
sta. CB05-12, Curlew Cay, 15–21 m, 27 April 2005. USNM 398935, BLZ 5206,
female, 12.0 mm SL, sta. CB05-13, Belize (no other collection data available), 29
April 2005. USNM 398937, BLZ 8029, male, 17.0 mm SL, sta. CB08-2 (see CB08-2
above). USNM 398938, BLZ 8030, female, 19.0 mm SL, sta. CB08-2 (see CB08-2
above). USNM 398940, BLZ 8353, female, 16.0 mm SL, sta. CB08-32, Tobacco Cay,
16°53'23.8"N, 88°03'53.8"W, 0–5 m, 25 May 2008. USNM 276147, male, 15.0
mm SL, sta. GDJ 84-14, o northwest end of Carrie Bow Cay, 2–3 m, 7 Nov 1984.
USNM 321073, female, 18.0 mm SL, sta. GDJ 90-2, reef at and crest, coral rubble
and sand substrate, Carrie Bow Cay, 3–6 ft., 18 Sep 1990.
Additional Material (not DNA vouchers). Belize: USNM 398943, 4 specimens;
USNM 398944, 2; USNM 398945, 4; USNM 321066, 1; USNM 276068, 1; USNM
398941, 1; USNM 398942, 1.
Diagnosis. A species of Starksia distinguished by the following combination of
characters: no orbital cirrus, regular vertical brown bars on trunk separated by narrow
white interspaces, and a well de ned horseshoe-shaped blotch of dark pigment on
cheek.
Description. See Table 1. Dorsal spines XIX–XX, usually XIX (XIX in holotype);
segmented dorsal rays 7–8 (8); total dorsal elements 26–27, usually 27 (27); anal
spines II; segmented anal rays 14–16, usually 15 (15); dorsal segmented caudal- n
rays 7; ventral segmented caudal- n rays 6; dorsal procurrent caudal- n rays 5–6, usu-
ally 6 (6); ventral procurrent caudal- n rays 4–6, usually 5 (5); segmented pelvic- n
rays 2; pectoral- n rays 14–15, rarely 15 (14); vertebrae 10+21–22= 31–32, rarely
31(10+22=32); 1–4 pairs of infraorbital pores, usually 4 pairs between 3 and 6 o’clock
(4 pairs); orbital cirri absent; nape cirri present; anterior nostril cirri present; belly and
pectoral- n base naked or with only a few rows of scales anterior to the anus.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
28
Figure 1. Neighbor-joining tree derived from cytochrome c oxidase I sequences showing genetically
distinct lineages of western Atlantic Starksia.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 29
Specimens examined ranging from 12.0 to 19.0 mm SL; HL 29–34% SL (31%
in holotype); male genital papilla adhered to rst anal spine proximally; papilla length
between two-thirds and three-quarters length of rst anal spine, 0.6–1.0 mm; some
females with very small genital papilla.
Pigment. Vertical brown bars present on trunk separated by narrow white inter-
spaces; anteriormost 6 bars relatively uniform in all specimens; posterior bars often
irregular or incompletely formed. A thick horseshoe-shaped blotch of pigment pre-
sent on cheek. Bright orange pigment present on distal portions of pectoral- n rays,
and pale orange pigment usually present on distal portions of posterior anal-, caudal-,
and soft dorsal- n rays. Color pattern sexually dimorphic: males with pale red heads
(vs. females without red coloration); relatively poorly de ned horseshoe-shaped blotch
of pigment on cheek that fades posteriorly (well-de ned horseshoe-shaped blotch on
cheek that is sometimes mirrored on operculum and pectoral n base); body bars tan
and usually with some gold or green color in life (darker and without green/gold color
but some posterior bars often with some orange pigment); body bars usually terminat-
ing ventrally dorsal to ventral midline (body bars usually extending to ventral midline);
blotches of tan/gold pigment on base of dorsal n associated with body bars, and no
tan/gold color present on anal n (bright orange markings on base of dorsal n associ-
ated with body bars and several bright orange spots on base of anal n); and large dark
spot, roughly diameter of pupil or larger, on trunk at posterior end of dorsal n (two
large dark spots on trunk, one at posterior end of dorsal n similar in size to that of
males, and smaller spot at posterior end of anal n).
Color in preservative. Vertical bars on trunk, horseshoe-shaped blotch of pigment
on cheek, and spot at posterior end of dorsal n (and anal n in females) retained
in preservative; margins of at least some body bars in females with small dark spots;
Figure 2. Male and female color patterns of Starksia sangreyae: A USNM 398932, holotype, BLZ 5111,
16.0 mm SL, male B USNM 398933, BLZ 5033, 16.5 mm SL, female. C–D Diagnostic patterns of cheek
pigment of preserved female and male – C USNM 276147, 15.0 mm SL, male D USNM 321073, 18.0
mm SL, female. Photographs by Carole Baldwin, Cristina Castillo, Donald Griswold, and Julie Mounts.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
30
prominent patches of melanophores on jaws and gular region, and scattered pigment
(heavier in females) on rest of head; dorsal n ranging from overall dusky to having
concentrations of pigment on base of n associated with body bars; caudal n with
light pigment on outer rays, and pectoral n with scattered melanophores over entire
n; pelvic n clear.
Etymology. e species name is in honor of Mary Sangrey for her many years
of work coordinating the intern program at the Smithsonian’s National Museum of
Natural History. Mary brought the intern application of the second author to the
rst author’s attention and took the rst steps toward procuring funding for Castillo’s
internship.
Distribution. All material that we examined is from Belize. e range of the spe-
cies also apparently includes Honduras, as Green eld and Johnson (1981) noted that
a specimen of S. atlantica from Honduras has regular vertical bars of pigment on the
body.
Starksia springeri Castillo & Baldwin, sp. n.
urn:lsid:zoobank.org:act:495CE72B-82CD-4A2B-B192-ACAA389F40FC
Figs 1, 3–4; Table 1
Type Locality: Curacao, Netherland Antilles
Holotype. USNM 398945, female, 19.0 mm SL, sta. CUR08-10, Blue Bay, Cu-
racao, 12˚07'59.22"N, 68˚59'05.34"W, 1–25 m, 17 March 2008, C. Baldwin, D.
Smith, L. Weigt (not a DNA voucher).
Paratypes (all Curacao). USNM 399658, CUR 8148, male(?), 15.0mm SL, sta.
CUR08-03, Cas Abou, 12˚13'34.04"N, 69˚05'29.95"W, 0–4 m, 12 March 2008,
Table 1. Frequency distributions of counts among species of the S. atlantica complex.
Dorsal Spines Dorsal Soft
Rays Total Dorsal Anal Soft Rays
XVIII XIX XX 7 8 26 27 14 15 16
S. springeri 1*---1*1*--1*-
S. sangreyae - 11* 2 7 6* 5 8* 1 9* 2
S. atlantica117*- 26*35*- 44*
Pectoral Rays Dorsal
Procurrent
Caudal Rays
Ventral Procurrent
Caudal Rays Vertebrae
13 14 15 5 6 4 5 6 31 32 33
S. springeri - 3*- 1*- - 1*- - 1*-
S.sangreyae - 20* 1 3 7* 1 8* 1 1 12* -
S. atlantica118144- 44251
* Indicates count of holotype
1 Longley (1934) did not provide counts of pectoral- n rays or vertebrae for the holotype of S. atlantica
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 31
(posterior portion of body destroyed for DNA tissue sampling). USNM 399659,
CUR 8318, (sex unknown), 12.0 mm SL, sta. CUR08-05, Blue Bay, 12˚07'57.14"N,
68˚59'06.03"W, 0–25 m, 14 March 2008, (posterior portion of body destroyed for
DNA tissue sampling).
Diagnosis. A species of Starksia distinguished by the following combination of
characters: no orbital cirrus; trunk with irregular dark blotches on pale background;
pectoral- n base with relatively straight margins de ning pale gap that separates two
dark blotches; cheek with distinctive dark and pale markings: anterior portion of cheek
with prominent dark blotch, anteroventral and posterior margins of blotch well de-
ned by pale regions; posterior pale area on cheek bordered posteriorly by thin, dark,
anteroventral-to-posterodorsal streak of pigment along distal edge of preopercle.
Description. See Table 1. e female holotype is the only complete specimen
available. Counts in parentheses are those for the holotype. Few counts could be made
on partial specimens; when available, counts of partial specimens precede those of
holotype. Dorsal spines (XVIII); segmented dorsal rays (8); total dorsal elements (26);
anal spines (II); segmented anal rays (15); dorsal segmented caudal- n rays (7); ventral
segmented caudal- n rays (6); dorsal procurrent caudal- n rays (5); ventral procurrent
caudal- n rays (5); segmented pelvic- n rays 2 (2); pectoral- n rays 14 (14); vertebrae
(10+22= 32); infraorbital pores paired or unpaired, usually 1–3 pairs (3 pairs); if only
one pair of pores, pair situated at 3 o’clock; 3 pairs in holotype located at 3, 5, and 6
o’clock; orbital cirri absent; nape cirri present; anterior nostril cirri present; belly and
pectoral- n base completely naked.
Specimens examined ranging from 12.0 to 19.0mm SL; HL 25–32% SL (32% in
holotype); genital-papilla length in 15.0-mm SL paratype 0.3 mm, one-fourth length
of rst anal spine (broken); papilla adhered to spine proximally. Note: the presence of a
small but measurable genital papilla on 15.0-mm SL paratype suggests that it is a male:
although female Starksia sometimes have a small genital papilla, the 19 mm female
holotype does not. As noted below, the 15 mm paratype has a pupil-size dark spot at
posterior base of anal n, which usually characterizes females. We tentatively recognize
this paratype as a male.
Pigment. (Note: a eld photograph of the 12.0-mm SL paratype is a dorsal view
of poor quality, and only the head remains as a preserved voucher. e following
Figure 3. A Color pattern of Starksia springeri, USNM 399658, CUR 8148, paratype, 15.0 mm SL,
male(?) B diagnostic pigment pattern on cheek and pectoral- n base in preserved S. springeri, USNM
398945, holotype, 19.0 mm SL, female. Photographs by Carole Baldwin, Cristina Castillo, and Donald
Griswold.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
32
Figure 4. Comparisons among species of the Starksia atlantica complex. Left to right for each row -- S. atlantica: AMNH 241247; USNM 399621, BAH 8176,
15.0 mm SL; USNM 386971, 19.0 mm SL; USNM 386242, 17.0 mm SL. S. sangreyae: (Note: top and bottom images in rst two columns represent S. sangreyae
A and S. sangreyae B genetic sublineages, respectively.) Males – USNM 398936 (top), paratype, BLZ 8028, 17.0 mm SL and USNM 398937 (bottom), paratype,
BLZ 8029, 17 mm SL; Females – USNM 398934 (top), paratype, BLZ 5161, 17.0 mm SL and USNM 398940 (bottom), paratype, BLZ 8353, 16.0 mm SL;
preserved – USNM 276147, paratype, 15.5 mm SL; USNM 321073, paratype, 18.0 mm SL. S. springeri: USNM 399658, paratype, CUR 8148, 15.0 mm SL;
USNM 398945, holotype, CUR 08-10, 19.0 mm SL; Starksia sp. (Saba): SABA-06-01, 15.0 mm SL (no voucher). Photographs by Carole Baldwin, Cristina Castillo,
Donald Griswold, Julie Mounts, Ross Robertson, James Van Tassell, and Je rey Williams.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 33
description is based on the 15.0-mm SL paratype and the 19.0-mm SL holotype.)
Trunk with irregular dark blotches on pale background, most blotches consisting
of orange chromatophores and melanophores in paratype; two dark spots present
on trunk, large one at posterior end of dorsal n (larger than pupil diameter) and
smaller spot at posterior end of anal n. Paratype with pale orange and brown pig-
ment on head; tips of jaws with dark pigment in both paratype and holotype, but
rest of jaws and gular region distinctly barred in holotype, mottled with tiny spots in
paratype; cheek with distinctive dark and pale markings: anterior portion of cheek
with prominent dark blotch, its anteroventral and posterior margins well de ned by
pale regions; posterior pale area on cheek bordered posteriorly by thin, dark, anter-
oventral-to-posterodorsal streak of pigment along distal edge of preopercle. Bright
orange markings present on bases of dorsal n and anal ns, sometimes occurring in
pairs; bright orange pigment also present on distal portions of pectoral- n rays; pale
orange pigment present distally on at least some rays of soft dorsal, caudal, and anal
ns; pectoral- n base with relatively pale gap separating two dark blotches, margins
of gap relatively straight; dark blotches on pectoral- n base comprising orange chro-
matophores and melanophores.
Color in preservative. (Note: pigmentation on trunk in preservative based on the
only entire specimen, female holotype.) Trunk with irregular dark blotches on pale
background; spots at posterior ends of dorsal and anal ns retained in preservative.
Dark markings on head described above retained in preservative, mottled jaws and
gular region of male(?) paratype strikingly di erent from barred markings on female
holotype; top of head in both specimens covered with scattered melanophores; dark
and pale regions on cheek and pectoral- n base retained in preservative. Anal and pec-
toral ns with lightly scattered melanophores; caudal n with light pigment on outer
rays; pelvic n clear.
Etymology. Named in honor of Victor G. Springer, Senior Scientist Emeritus,
Smithsonian National Museum of Natural History, for his contributions to the sys-
tematics of blennioid shes, including Starksia, and for advice and friendship he has
bestowed upon the rst author.
Distribution. Known only from Curacao, Netherland Antilles.
Starksia sp.
Figs 1, 4
Locality: Saba Bank, Netherland Antilles
Material Examined. Specimen and photograph: USNM 388032, sta. SABA-06–
25, 9.0 mm female (not a DNA voucher), near Coral Garden at SE edge of Saba Bank,
Netherland Antilles, 17°21.10'N, 63°15.08'W, 15–18 m, 4 Jan 2006; photograph:
15.1 mm SL female (not a DNA voucher), sta. SABA-06-01, Saba Bank just south
of Poison Bank, Netherland Antilles, 17°28.47'N, 63°13.40'W, 24–27 m, 4 Jan 2006
(photographs by Je rey T. Williams).
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
34
Remarks. A DNA sequence from a single specimen collected at Saba Bank (Neth-
erland Antilles) is genetically distinct from the other members of the S. atlantica species
complex (SAB 0601019, Fig. 1). Our material includes color photographs of 9.0- and
15.1-mm SL females and the preserved 9.0 mm specimen (USNM 388032). Presum-
ably the 9.0 and 15.1 mm specimens are the same species as the specimen represented
on the tree, but we do not have tissue samples of either for genetic analysis or a pre-
served voucher of SAB 0601019 for morphological analysis.
Trunk pigment in the images and preserved specimen is similar to that of S. at-
lantica from the Bahamas and S. springeri from Curacao (i.e., mottled vs. barred as in
S. sangreyae), but the Saba specimens lack the horseshoe-shaped blotch of pigment on
the cheek characteristic of S. atlantica and the distinctive dark and pale markings on
the cheek of S. springeri. e blotches of trunk pigment in the Saba Bank specimens
are neither conspicuously block-like nor clearly organized in horizontal tiers as they are
in S. atlantica. Specimens from Saba Bank presumably represent another new species
within S. atlantica, but additional specimens are needed for comparative purposes and
description.
Comparisons among Species of the Starksia atlantica Complex (Figs 4–5, Table 1)
Comparative material. Starksia atlantica. Bahamas: USNM 386971, 1 specimen (not
a DNA voucher); USNM 386580, 1 (not a DNA voucher); USNM 386242, 6 (not
DNA vouchers); USNM 399619, 3 (not DNA vouchers); USNM 399620, BAH
8175; USNM 399621, BAH 8176; USNM 399622, BAH 8177. Turks and Caicos
Islands: USNM 399643, TCI 9044; USNM 399644, TCI 9106; USNM 399645,
TCI 9107; USNM 399647, TCI 9205. Navassa Island: USNM 360422, 3; USNM
360194, 2; USNM 359543, 2; USNM 360210, 3.
Members of the S. atlantica complex are diagnosed by the absence of an orbital cir-
rus. Starksia sangreyae is distinct in having regular vertical body bars separated by nar-
row pale interspaces and a well-de ned horseshoe-shaped blotch on the cheek. Starksia
springeri, S. atlantica, and the specimens from Saba Bank have irregular dark blotches
on a pale background on the trunk, the blotches better de ned in our S. atlantica mate-
rial than in the other species and often more clearly arranged in two or three horizontal
tiers. Starksia springeri, S. atlantica, and the Saba Bank specimens can be distinguished
on the basis of pigment patterns on the cheek: specimens from Saba Bank lack cheek
blotches; S. atlantica has a horseshoe-shaped blotch on the cheek; and S. springeri has
a prominent dark blotch on the cheek bordered anteroventrally and posteriorly by pale
areas and a thin, dark, anteroventral-to-posterodorsal streak of pigment along the distal
edge of the preopercle. Although S. sangreyae and S. atlantica are easily separated based
on trunk pigment, we note that both have a horseshoe-shaped blotch of pigment on
the cheek; the blotch is most prominent and best de ned in S. sangreyae females, often
completely faded in preserved S. sangreyae males. Starksia atlantica and S. springeri can
be separated based on pigment on the pectoral- n base: in S. atlantica, the pale gap
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 35
between two blotches of darker pigment has wavy margins, whereas in S. springeri, the
margins of the pale gap are relatively straight. Starksia springeri has XVIII dorsal spines
vs. usually XIX in the other species (Table 1), but we have only one entire specimen
of S. springeri on which to base counts. No other signi cant di erences were found in
numbers of n rays or vertebrae among species of the S. atlantica complex.
A photograph of a specimen identi ed as S. atlantica from St. Croix, U. S. Vir-
gin Islands (taken by William Smith-Vaniz) shows irregular block-like blotches on
the body arranged in roughly 3 horizontal tiers, wavy margins on the pale gap that
Figure 5. Diagnostic features in preserved A Starksia atlantica, USNM 386242, 17.0 mm SL, male—
note irregular horseshoe-shaped blotch of pigment on cheek and wavy margins of pale gap on pectoral- n
base; and B Starksia springeri, USNM 398945, holotype, 19.0 mm SL, female—note pale regions at
anteroventral and posterior margins of dark cheek blotch, thin dark anteroventral-to-posterodorsal streak
of pigment along distal edge of preopercle, and relatively straight margins of pale gap on pectoral- n base.
Photographs by Cristina Castillo and Donald Griswold.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
36
separates two darker areas on the pectoral- n base, and an irregular horseshoe-shaped
blotch of pigment on the cheek. e U.S. Virgin Islands are thus likely part of the
geographical distribution of S. atlantica Longley. Several USNM specimens identi ed
as S. atlantica from Navassa Island exhibit pigmentation that is somewhat intermediate
between that of S. atlantica and S. sangreyae: bars of pigment are present on the trunk
anteriorly as in S. sangreyae, but trunk pigment is more block-like posteriorly as in S.
atlantica; Navassa specimens also have an irregular horseshoe-shaped blotch on the
cheek as in S. atlantica. Further genetic and morphological investigation should help
clarify species issues of S. atlantica from Navassa Island.
Key to Species of the Starksia atlantica Complex
1a Body with vertical brown bars separated by narrow white interspaces ...........
.......................................................................... Starksia sangreyae (Belize)
1b Body with irregular dark blotches on pale background ................................2
2a Dark blotches on trunk often arranged in two or three horizontal tiers; pale
gap between two blotches of darker pigment on pectoral- n base with wavy
margins; cheek with irregular horseshoe-shaped blotch of pigment; no streak
of dark pigment along distal edge of preopercle ............................................
.......................................... Starksia atlantica (Bahamas, Turks and Caicos)
2b Dark blotches on trunk not conspicuously arranged in horizontal tiers;
pale gap between two blotches of darker pigment on pectoral- n base with
straight margins; cheek with prominent dark blotch bordered anteroventrally
and posteriorly by pale areas and a thin, dark, anteroventral-to-posterodorsal
streak of pigment along distal edge of preopercle ..........................................
........................................................................Starksia springeri (Curacao)
Starksia lepicoelia Species Complex
Böhlke and Springer (1961) described S. lepicoelia on the basis of numerous specimens
from the Bahamas and one from St. John, U.S. Virgin Islands. e presence of a simple
cirrus above the eye, two externally obvious pelvic- n rays, a completely scaled belly
or at least posterior half scaled, and usually 17 anal- n soft rays are diagnostic of the
species. Six genetic lineages in our data set cluster in the S. lepicoelia complex (Fig.1).
ere are no photographs or vouchers of the Barbados specimens (BAR on tree), and
that lineage is not discussed further. Clearly it represents either a cryptic species within
S. lepicoelia or one of the eight species of western Atlantic Starksia not identi ed in
our material. Two of the S. lepicoelia lineages are from the Bahamas/Turks and Caicos
(BAH/TCI), and although sequence divergence for the two is 4–6% -- much higher
than typical intraspeci c variation in western Atlantic Starksia – we were unable to nd
consistent morphological di erences between them and tentatively recognize them
together as S. lepicoelia (Fig. 6). A fourth genetic lineage comprises specimens from
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 37
Belize (BLZ), and a fth, specimens from Panama (PAN). Although those lineages
di er by only about 1% sequence divergence in COl, they are easily distinguished by
color pattern. We describe the specimens from Belize and Panama as two new species.
A sixth genetic lineage is represented in our tree by a single specimen from Saba Bank,
Netherland Antilles. Based on that specimen and several lots of non-voucher mate-
rial, we recognize the Saba Bank population as a fourth species within the S. lepicoelia
complex.
Starksia weigti Baldwin & Castillo, sp. n.
urn:lsid:zoobank.org:act:91F47395-F5D4-4160-A645-5266D10E6DBB
Figs 1, 7, 10–11; Table 2
Type Locality: Belize, Central America
Holotype. USNM 399648, BLZ 5010, male, 20.5 mm SL, sta. CB05-01, spur
and groove, Carrie Bow Cay, Belize, 6–8 m, 21 Apr 2005, C. Baldwin, D. Smith, L.
Weigt, J. Mounts (small llet removed from right side for DNA tissue sampling).
Paratypes (all Belize). USNM 399649, BLZ 5164, female, 19.0 mm SL, sta.
CB05-12, Curlew Cay, 21–25 m, 27 Apr 2005, (posterior portion of body destroyed
for DNA tissue sampling). USNM 399653, BLZ 8026, female, 17.5 mm SL, sta.
CB08-02, sand bottom and coral heads, Curlew Cay, 16˚47'24.1"N, 88˚04'41.0"W,
5–8 m, 15 May 2008, (posterior portion of body destroyed for DNA tissue sam-
pling). USNM 399652, BLZ 8025, female, 18.0 mm SL, sta. CB08-02, same col-
Figure 6. Comparison of Starksia lepicoelia specimens from Bahamas from genetically distinct lineages
(see Fig. 1): A USNM 399615, BAH 8077, 25.0 mm SL, female B USNM 399617, BAH 8079, 19.0
mm SL, female. Photographs by Carole Baldwin.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
38
lection information as above, (posterior portion of body destroyed for DNA tissue
sampling). USNM 399651, BLZ 8024, female, 19.0 mm SL, sta. CB08-02, same
collection information as above, (posterior portion of body destroyed for DNA tissue
sampling). USNM 399654, CB08-2, 2 specimens: (1) 19.5 mm SL female, (1) 19.0
mm SL female (not DNA vouchers), same collection information as above. USNM
399656, BLZ 8123, juvenile, 9.5 mm SL, sta. CB08-10, east wall drop o , Glovers
Cay, 16˚42'36.1"N, 87˚51'05.3"W, 11–23 m, 18 May 2008, (posterior portion of
body destroyed for DNA tissue sampling). USNM 399655, BLZ 8122, female, 18.0
mm SL, sta. CB08-10, same collection information as above, (posterior portion of
body destroyed for DNA tissue sampling). USNM 274922, Sta. K-103, 2 females,
20.0 and 24.0 mm SL, spur and groove, Carrie Bow Cay, 6–8 m, 10 June 1981.
USNM 276063, Sta. GDJ 84-8, 2 males, 20.5 and 23.0 mm SL, Carrie Bow Cay,
24–30 m, 5 Nov 1984.
Additional Material (not DNA vouchers, all Belize). USNM 399650, BLZ
5193, 1 specimen; USNM 365517, 4; USNM 274941, 1; USNM 328251, 2; USNM
276048, 2; USNM 327608, 1.
Diagnosis. A species of Starksia distinguished by the following combination of
characters: orbital cirrus present; belly scaled; trunk pale (pale red in life), without dis-
tinct bars or other markings; lips peppered with white spots in life; lacrimal region with
single row of small white spots in life; jaws usually with lightly scattered melanophores
in preserved specimens, without distinct banding or dark bars; entire gular region
usually covered with scattered melanophores; total dorsal elements usually 27; total
vertebrae usually 32; dorsal spines + anal soft rays + vertebrae modally 75.
Description. See Table 2. Dorsal spines XIX–XX, usually XX (XX in holotype);
segmented dorsal rays 7–8, usually 8 (7); total dorsal elements 27–28, usually 27 (27);
Figure 7. Male and female color patterns of Starksia weigti: A USNM 399648, holotype, BLZ 5010,
25.0 mm SL, male B BLZ 6121 (no voucher), 18.0 mm SL, female C–D close-up views of diagnostic
spotting on lips in life – C BLZ 6120, 24.0 mm SL (no voucher), male D USNM 399650, BLZ 5193,
24.0 mm SL, female. Photographs by Carole Baldwin and Julie Mounts.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 39
Table 2. Frequency distributions of counts among species of the Starksia lepicoelia complex1.
Dorsal Spines Dorsal Soft Rays Total Dorsal Anal Soft Rays
XIXXX7 8 9 2627282915161718
S. weigti 5 11* 5* 11 - - 10* 6 - - 6* 10 -
S. lepicoelia 2 18* 1 18* 1 - 3 16* 1 - 1 18 1*
S. williamsi 10* 3 3 10* - - 13* - - - 11* 2 -
S. robertsoni 1 7* 8* - - 1 7* - - 1 6* 1 -
Pectoral Rays Dorsal Procurrent
Caudal Rays Ventral Procurrent Caudal Rays Vertebrae
11 12 13 14 5 6 3 4 5 6 31 32 33 34
S. weigti - 3 18* - 12 2* - 1 13* - 1 12* 3 -
S. lepicoelia - - 18* 1 9 8 - - 14 2 - 2 15 1
S. williamsi 2 1 10* - 7 7* - - 13 1* 1 12* - -
S. robertsoni - - 6* 1 4* 3 1* - 4 2 2* 6 - -
Total Dorsal Elements + Anal Soft Rays + Vertebrae
73 74 75 76 77 78 79
S. weigti - 15*233-
S. lepicoelia ---21112
S. williamsi -110*2---
S. robertsoni 12*41- - -
* Indicates count of holotype
1 Böhlke and Springer (1961) did not provide counts of procurrent caudal rays or vertebrae for the holotype of S. lepicoelia
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
40
anal spines II; segmented anal rays 16–17 (16); dorsal segmented caudal- n rays (7);
ventral segmented caudal- n rays (6); dorsal procurrent caudal- n rays 5–6, rarely 6
(6); ventral procurrent caudal- n rays 4–5, rarely 4 (5); obvious segmented pelvic- n
rays 2; pectoral- n rays 12–13, rarely12 (13); vertebrae 10+21–23= 31–33, usually 32
(10+22=32); infraorbital pores usually unpaired (one pair present at 3 o’clock); orbital
cirri present; nape cirri present; anterior nostril cirri present; belly and pectoral- n base
completely scaled.
Specimens examined ranging from 9.5 mm to 24.0 mm SL; HL 30–36% SL;
length of male genital papilla two-thirds to equal length of rst anal spine, papilla
1.0–1.8 mm and free from spine.
Pigment. Both males and females with pale red to reddish brown trunk; indistinct
vertical bars, if present, more prominent dorsally; two small (less than half pupil diam-
eter) dark spots on posterior portion of trunk, one at posterior end of dorsal n and
one at posterior base of anal n. Both sexes with pale red heads, scattered small white
spots on anterior portions of lips, and single row of white spots beneath eye on lacrimal
region; white spots representing absence of chromatophores in areas otherwise covered
with pale orange to red pigment; eye with six or seven white spots around pupil, spots
separated by darker areas (e ectively a candy-stripe pattern). Males with prominent
dark blotch on cheek and with small white spots extending from anterior portions of
lips to posterior portions of jaws; females without dark cheek blotch and usually with
larger white spots, blotches, or bands on posterior portions of jaws. Males with red
pigment on dorsal n con ned to blotches at base and little red pigment on rest of n
and other median ns (but with scattered melanophores on dorsal, caudal, and anal
ns); females with red pigment extending onto entire dorsal n and with prominent
orange/red pigment on caudal and anal ns (but without prominent melanophores);
males with yellowish brown pectoral n, females with pale orange to orange pectoral
n; pelvic n clear.
Juvenile (BLZ 8123) color pattern: trunk pale orange, with some yellow mixed in;
head with dark bar from anterior portion of eye to upper and lower lips; black triangle
of pigment beneath eye; and black cap of pigment on head that extends anteriorly
to vertical through middle of eye. Dorsal, anal, and caudal ns pale orange; bases of
several dorsal- n elements with darker blotches of orange pigment; most anal- n ele-
ments with melanophore at base (typical of blennioid larvae), bases of about half of
anal- n elements also with prominent orange spot.
Color in preservative. Males mostly pale, except with very dark blotch on cheek;
trunk, belly, jaws, gular region, branchiostegals, operculum, top of head, nape, and all
ns except pelvics with scattered melanophores, pigment on trunk fairly heavy in one
male. Some females very pale, with only a few melanophores on gular region, cheek,
branchiostegals, and on all ns except pelvics; other females with poorly formed dark
blotch on cheek, fairly heavy pigment on gular region, branchiostegals, belly, dorsal
n, and anal n; and lightly scattered melanophores on trunk, jaws, operculum, top of
head, nape, caudal n, and pectoral n; pigment on head and nape usually lighter in
females than in males.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 41
Only anterior portion of body remains in juvenile voucher specimen (BZE 8123):
body mostly pale; black cap of pigment on head, dark bar from anterior portion of eye
to upper and lower lips, and black triangle of pigment beneath eye present in preserva-
tive. Etymology. e species name is in honor of Lee A. Weigt, Head of the Smithsoni-
an’s Laboratories of Analytical Biology, in recognition of his contributions to the DNA
barcoding of shes and his contributions to sh-collecting e orts in Belize, Curacao,
Florida, Tobago, and Turks & Caicos Islands.
Distribution. Known only from Belize, Central America.
Starksia williamsi Baldwin & Castillo, sp. n.
urn:lsid:zoobank.org:act:7C75F463-D33D-4411-8222-BAA0556FDEC4
Figs 1, 8, 10–11; Table 2
Type Locality: Saba Bank, Netherland Antilles
Holotype. USNM 387675, sta. SABA-06-12, 21 mmSL, male, Saba Bank (Neth-
erland Antilles), 19 m, 17°14'23"N, 63°26'55"W, 8 Jan 2006, Saba 2006 expedition
team.
Paratypes (all Saba Bank, Netherland Antilles). All paratypes are non-DNA
vouchers except USNM 397396. USNM 397396, sta. SABA-06-01, female, just
south of Poison Bank, 17°28.47'N, 63°13.40'W, 24–27 m, 4 Jan 2006 (DNA voucher
of SAB 0601010—length unknown, posterior portion of body removed for DNA tis-
sue sample); USNM 399613, sta. SABA-06-12, 3 specimens: (1) 21.5 mm SL male,
(1) 22.5 mm SL female, (1) 20.0 mm SL female, 19 m, 17o14'23"N, 63o26'55"W, 8
Jan 2006; USNM 387869, sta. SABA-06-05, 4 specimens: (1) 21.5 mm SL male, (1)
19.5 mm SL male, (1) 19.5 mm SL female, (1) 19 mm SL female, overall bank, east
side, 26–28 m, 17o24'36"N, 63o11'45"W, 6 Jan 2006; USNM 388033, sta. SABA-
06-25, 8 specimens: (1) 22.5 mm SL male, (1) 20.5 mm SL female (1) 20.0 mm SL
female (1) 19.5 mm SL female, (1) 21.5 mm SL male, (3) juveniles 8.5 -11.5 mm SL,
near Coral Garden at southeast, 15–18 m, 17o21'10"N, 63o15'08"W, 14 Jan 2006.
USNM 388444, sta. SABA-06-21, 4 specimens: (1) 18.5 mm SL female, (3) juveniles
7.5–9.0 mm SL, northeastern shallow ats, 20 m, 17o28'03"N, 63o14'59"W, 12 Jan
2006; USNM 387767, (3) females 19.5–20.0 mm SL, (4) juveniles 8.0–11.0 mm SL,
sta. SABA-06-01, just south of Poison Bank, groove in reef with sand bottom, 24–27
m, 17o28'47"N, 63o13'40"W, 4 Jan 2006.
Additional Material (not DNA vouchers, all Saba Bank, Netherland Antil-
les). USNM 388392, 6 specimens; USNM 388589, 3; USNM 387623, 1; USNM
387733, 4; USNM 388355, 2.
Diagnosis. A species of Starksia distinguished by the following combination of
characters: orbital cirrus present; belly scaled; trunk pale to tan (dark orange/tan to
bright orange in life), without distinct bars or other markings; lips without conspicu-
ous white spotting, distinct banding, or dark bars—usually with lightly scattered mel-
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
42
anophores in preserved specimens; total dorsal elements 27; total vertebrae usually 32;
dorsal spines + anal soft rays + vertebrae modally 75.
Description. See Table 2. Dorsal spines XIX–XX, rarely XX (XIX in holotype);
segmented dorsal rays 7–8, usually 8 (8); total dorsal elements (27); anal spines II; seg-
mented anal rays 16–17, rarely 17 (16); dorsal segmented caudal- n rays (7); ventral
segmented caudal- n rays (6); dorsal procurrent caudal- n rays bimodal at 5–6 (6);
ventral procurrent caudal- n rays 5–6, rarely 6 (6); segmented pelvic- n rays 2; pecto-
ral- n rays 11–13, usually 13 (13); vertebrae 9–10+22= 31–32, rarely 31 (10+22=32);
usually one pair of infraorbital pores at 3 o’clock (one specimen with all infraorbital
pores unpaired); orbital cirri present; nape cirri present; anterior nostril cirri present;
belly and pectoral- n base completely scaled.
Specimens examined ranging from 18.5 mm to 22.5 mm SL; HL 34–38% SL;
male genital-papilla length between two-thirds and three-fourths length of rst anal
spine, papilla 1.0–1.25 mm and free from spine.
Pigment. Trunk dark orange/tan to bright orange, color nearly uniform—i.e.,
without indistinct dark bars and pale areas; two small (less than half pupil diameter)
dark spots on posterior portion of trunk, one at posterior end of dorsal n and one
at posterior base of anal n. Both sexes with orange heads, a few small pale spots on
lips and lacrimal region, and six or seven white spots around pupil, spots separated by
darker areas (e ectively a candy-stripe pattern). Males with prominent dark blotch on
cheek and uniformly orange/tan lips; females without dark blotch on cheek and with
mottling of orange and pale blotches on lips. Males with red pigment on dorsal n
largely con ned to blotches at base and little red pigment on rest of n and other me-
dian ns (but with numerous melanophores on dorsal, caudal, and anal ns); females
Figure 8. Male and female color patterns of Starksia williamsi: A USNM 387869, 19.5 mm SL, male,
paratype B USNM 387767, 20.2 mm SL, female, paratype. Photographs by Je rey Williams.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 43
with bright orange spotting on dorsal, anal, and caudal ns (but without prominent
melanophores except one dark spot sometimes present in anterior portion of spinous
dorsal); males with yellowish brown pectoral n, females with orange pectoral n;
pelvic n clear.
Color in preservative. Males tan, usually with fairly heavy pigment on head,
trunk, and dorsal-, anal-, outer caudal-, and posterior portions of pectoral- n rays;
prominent dark blotch on cheek retained in preservative; no dark spots, streaks or
bars on lips. Females mostly pale, sometimes with noticeable concentrations of mel-
anophores on cheek, jaws and gular region, but no prominent dark cheek blotch;
lightly scattered melanophores usually present on branchiostegals, opercle, belly, me-
dian and pectoral ns; no conspicuous pattern of dark and pale blotches on lips, but
light bar present across lips just posterior to symphysis and sometimes a few spots
present just anterior to end of upper and lower jaws; posterior tips of upper and lower
jaws usually pale.
Etymology. Named in honor of Je rey T. Williams, Smithsonian’s National Mu-
seum of Natural History, in recognition of his work on blennioid shes, including
Starksia. Je ’s eld-collecting e orts at Saba Bank, Tobago, and Turks and Caicos re-
sulted in numerous specimens utilized in this study.
Distribution. Known only from Saba Bank, Netherland Antilles.
Starksia robertsoni Baldwin, Victor & Castillo, sp. n.
urn:lsid:zoobank.org:act:2C91C572-A7FA-4BE3-BC50-C735089B018C
Figs 1, 9–11; Table 2
Type Locality: Panama, Central America
Holotype. AMNH 249667, 22.0 mm female, sta. JVT-07-725, Islas de Las Dos
Hermanas, Portobelo, Panama, 9o35'45"N, 79o40'05"W, 2 June 2007, J. Van Tassell,
D. R. Robertson, L. Tornabene, B. Victor, E. Pena (not a DNA voucher).
Paratypes (all from Panama). USNM 399909, 21.0 mm SL male, PAN 1419,
Islas de Las Dos Hermanas, Portobelo, 9.59577N, 79.66801 W, 2 Jun 2007 ; USNM
399910, 22.0 mm SL female (not a DNA voucher), same collection information as
above; USNM 399911, 20.0 mm SL male (PAN 1418), USNM 399912, 16.0 mm SL
immature (PAN 014), Salmedina Reef, Portobelo, 9.56289 N, 79.69557 W, 31 May
2007; USNM 399913, 18.0 mm SL male (not a DNA voucher), same collection in-
formation as above; AMNH 249640, 18.0 mm SL female, sta. JVT-07-710, Salmedi-
na Reef, Portobelo, 9o33'54"N, 79o41'54"W, 30 May 2007 (not a DNA voucher);
AMNH 249642, 21.5 mm SL female, sta. JVT-07-714, Salmedina Reef, Portobelo,
9o33'46"N, 79o41'44"W, 31 May 2007.
Diagnosis. A species of Starksia distinguished by the following combination of
characters: orbital cirrus present; belly scaled; trunk pale to dark tan (dark orange/tan
to bright orange in life), without distinct bars or other markings; lips without con-
spicuous white spotting in life; ventral surface of lower jaw of males with one to three
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
44
dark blotches or bars in preserved specimens, lips without distinct banding or dark
bars; dorsal- n elements usually XX,7 – 27 total; vertebrae usually 10+22=32; dorsal
spines + anal soft rays + vertebrae modally 75.
Description. See Table 2. Dorsal spines XIX–XX, usually XX (XX in holotype);
segmented dorsal rays 7; total dorsal elements 26–27, usually 27 (27); anal spines
II; segmented anal rays 15–17, usually 16 (16); dorsal segmented caudal- n rays 7;
ventral segmented caudal- n rays 6; dorsal procurrent caudal- n rays 5–6 (5); ven-
tral procurrent caudal- n rays 3–6 (3); segmented pelvic- n rays 2; pectoral- n rays
13–14, usually 13 (13); vertebrae 10+21–22=31 or 32, usually 10+22=32 (10+21=31);
infraorbital series with one pair of pores at 3 o’clock; orbital, nape, and anterior-nostril
cirri present; belly and pectoral- n base completely scaled.
Specimens examined ranging from 16.0–22.0 mm SL; HL 32–36% SL (32); male
genital-papilla length between one-half and three-fourths length of rst anal spine,
papilla 0.6–1.9 mm and free from spine.
Pigment. Color in life known only for two females. Trunk dark orange/tan to
bright orange, color nearly uniform or with indistinct dark bars and pale areas; two
small (less than half pupil diameter), inconspicuous dark spots on posterior portion
of trunk, one at posterior end of dorsal n and one at posterior base of anal n. Head
orange, mottled with white patches; a few small, pale spots present on lips and lacrimal
region; eye with six or seven white spots around pupil, spots separated by darker areas
(e ectively a candy-stripe pattern). Bright orange spotting on dorsal, anal, and caudal
ns, and some orange pigment on pectoral n; pelvic n clear.
Figure 9. Color and preserved pigment patterns in Starksia robertsoni: A AMNH 249667, 22.0 mm SL,
female, holotype (photograph by James Van Tassell and Ross Robertson) B USNM 399911, PAN 1418,
20.0 mm SL, male, paratype (photograph by Carole Baldwin).
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 45
Color in preservative. Trunk ranging from pale to dusky, belly with fairly heavy
pigment in males and some females even if trunk pale. Males usually with prominent
dark blotch on cheek (largest male, USNM 399909, PAN1419, with dark spots on
cheek but no conspicuous blotch), females without dark cheek blotch. Underside of
lower jaw with one to three dark spots or bars in males, middle one (situated roughly
beneath a vertical through pupil) darkest and sometimes the only one noticeable; an-
terior marking, if present, sometimes extending onto lower lip as a few dark dots; no
dark spots, streaks, or bars on lips in either sex, but portions of lips uniformly covered
with melanophores in males and with at least a few spots in females; females usually
with patch or bar of pigment (small and faint in some specimens) extending from
lacrimal region across both lips. In males, branchiostegals dusky, upper part of cheek,
opercle, and top of head pale to dusky; in females, head mostly pale, with isolated
patches of spots on cheek, opercle, top of head, and branchiostegals. Dorsal, anal,
caudal, and pectoral ns dusky in males, mostly pale in females with a few scattered
spots on some ns.
Etymology. Named in recognition of the contributions by D. Ross Robertson of
the Smithsonian Tropical Research Institute to the understanding of the diversity of
shore shes of the New World and his generous facilitation of collecting in Panama.
Distribution. Known only from Panama (Atlantic)
Comparisons among Species of the Starksia lepicoelia Complex (Figs 10–11)
Comparative material. Starksia lepicoelia. Bahamas (DNA vouchers): USNM 399615,
BAH 8077; USNM 399616, BAH 8078; USNM 399617, BAH 8079. Bahamas (not
DNA vouchers): USNM 399923, 1 specimen; USNM 399924, 1; USNM 399925,
1; USNM 399926, 1; USNM 399927, 9; USNM 399928, 1; USNM 399929, 1;
USNM 399930, 1; USNM 399931, 1; USNM 399932, 1; USNM 399933, 1;
USNM 399934, 1; USNM 399921, 1; USNM 399922, 1; USNM 386919, 3 speci-
mens; USNM 386972, 15; USNM 386383, 1; USNM 386402, 8; USNM 386651,
2; USNM 386581, 3; USNM 386500, 4; USNM 387026, 3; USNM 386244, 13;
USNM 387069, 6; USNM 399618, 1; USNM 399614, 2; Turks and Caicos Islands
(DNA vouchers): USNM 399638, TCI 9291; USNM 399639, TCI 9292; USNM
399640, TCI 9293; USNM 399641, TCI 9294; USNM 399636, TCI 9112; Turks
and Caicos Islands (not DNA vouchers): USNM 399637, 7; USNM 399642, 1.
Navassa Island (not DNA vouchers): USNM 359448, 5; USNM 359699, 19. U.S.
Virgin Islands, St. Croix (not DNA vouchers): UF 149809, 11; UF 149815, 33; UF
149814, 10.
Comparisons. Starksia lepicoelia and S. starcki are the only previously described
western Atlantic Starksia with the combination of an orbital cirrus, two externally ob-
vious pelvic- n rays, and a scaled belly (Williams and Mounts 2003). Starksia starcki
is easily distinguished from the species of the S. lepicoelia complex by the presence of
eight or nine irregular dark bars on the body and usually 19 segmented anal- n rays.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
46
Figure 10. Comparisons among species of the Starksia lepicoelia complex. Left to right: S. lepicoelia (BAH): USNM 399928, BAH 10050, 25.0 mm SL; USNM
399617, BAH 8079, 19.0 mm SL; USNM 399921, BAH 9103, 26.0 mm SL; USNM 386972, 14.0 mm SL; S. lepicoelia (TCI): USNM 399638, TCI 9291, 23.5
mm SL; USNM 399641, TCI 9294, 25.5 mm SL; USNM 399642, 23.0 mm SL; USNM 399641, TCI 9294, 25.5 mm SL; S. weigti: BLZ 6120, 24.0 mm SL (no
voucher); USNM 399650, BLZ 5193, 24.0 mm SL; USNM 399648, BLZ 5010, holotype, 20.5 mm SL; USNM 274922, paratype, 20.0 m SL; S. williamsi: USNM
387767, 19.8 mm SL; USNM 387767, 20.2 mm SL; USNM 387675, holotype, 21.0 mm SL; USNM 387869, paratype, 19.5 mm SL; S. robertsoni: AMNH
249642, paratype, 21.5 mm SL; USNM 399909, PAN 1419, paratype, 21.0 mm SL; AMNH 249667, holotype, 22.0 mm SL. Photographs by Carole Baldwin,
Cristina Castillo, Donald Griswold, Ross Robertson, James Van Tassell, and Je rey Williams.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 47
In life, S. weigti is easily distinguished from S. lepicoelia, S. williamsi, and S. robert-
soni by the conspicuous pale round spots on the lips. In preservative, S. lepicoelia males
are distinctive in having at least some very dark spots, streaks, or bars on the lips and
lower jaw, and S. robertsoni males have at least one (up to three) dark spots or bars on
the ventral portion of the lower jaw (but not on the lips). Although the di erences are
subtle, preserved males of S. williamsi typically can be separated from preserved males
of S. weigti in having the lips uniformly covered with melanophores except for the
pale anterior tips. In S. weigti males, lip pigment is variable, but there are usually one
or two thin, faint, poorly formed bars of pigment following the pale anterior portions
of the lips; posteriorly, the lips may be uniformly covered with melanophores as in S.
williamsi or be quite pale.
Preserved female S. lepicoelia also have a distinctive lip pattern—alternating pale
and dark areas. Although this banding pattern appears to be present in color images of
S. williamsi, S. weigti, and S. robertsoni, it is not present in preserved females of those
species, suggesting that in S. lepicoelia the banding comprises both chromatophores
and melanophores whereas in females of the other species it comprises only chroma-
tophores and thus is not retained in preservative. As in males, di erences in head pig-
ment between preserved female S. williamsi and S. weigti are subtle, but S. williamsi
females have a relatively well-formed bar of pigment from the anterior portion of the
lacrimal across both lips, whereas S. weigti females typically have only a light scattering
of melanophores on the upper lip beneath the anterior portion of the lacrimal. Ad-
ditionally, S. williamsi females tend to have a bit of dark pigment at the posteroventral
corner of the orbit and another bit just ventral to posteriormost point of orbit; S. weigti
females usually have more widely scattered pigment on the cheek -- sometimes in a
fairly cohesive spot. e head pigment of female S. robertsoni is very similar to that
of S. williamsi, but modal di erences in n-ray counts separate them, and they are
geographically distinct. Speci cally, S. williamsi—from the eastern Caribbean—typi-
cally has XIX,8 dorsal- n elements, whereas S. robertsoni—from Panama—typically
has XX,7.
Modal di erences in some counts also help separate other species: S. lepicoelia
modally has 28 total dorsal- n elements, 33 vertebrae, and 78 total dorsal elements +
anal soft rays + vertebrae (vs. 32, 27, and 75, respectively, in S. williamsi and S. weigti).
Starksia williamsi modally has XIX dorsal- n spines, whereas S. lepicoelia and S. weigti
modally have XX.
We examined color photographs and numerous preserved specimens from St.
Croix, U.S. Virgin Islands, but we do not have genetic data for that material. Fresh
specimens lack the diagnostic white spots on the lips of S. weigti. Preserved specimens
most closely resemble S. lepicoelia in pattern of pigment on the lips and lower jaw, with
females typically having at least some alternating pale and dark areas (nearly identical
to that of S. lepicoelia in some specimens, not distinctive at all in others). Although
most males have fairly uniform pigment on the lips and lower jaw, at least some males
have the distinctive dark bars, spots, or streaks characteristic of male S. lepicoelia. If
the St. Croix specimens represent one of the known S. lepicoelia species, it seems likely
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
48
Figure 11. A Comparisons of head pigment of preserved males and females among species of the Stark-
sia lepicoelia complex. S. lepicoelia: A USNM 399921, BAH 9103, 26.0 mm SL, male B USNM 399617,
BAH 8079, 19.0 mm SL, female; S. weigti: C USNM 399648, BLZ 5010, holotype, 20.5 mm SL, male
D USNM 399651, BLZ 8024, paratype, 19.0 mm SL, female; S. williamsi: E USNM 387675, holotype,
21.0 mm SL, male F USNM 387869, paratype, 19.5 mm SL, female; S. robertsoni: G USNM 399913,
paratype, 18.0 mm SL, male H AMNH 249667, holotype, 22.0 mm SL, female. Photographs by Carole
Baldwin, Cristina Castillo, and Donald Griswold.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 49
based on geography and pigmentation that they are S. lepicoelia. However, we note
that S. lepicoelia typically has 28 total dorsal elements and 17 anal- n soft rays, whereas
the St. Croix specimens (15 counted) typically have 27 and 16, respectively (but 28
dorsal elements and 17 anal rays are not uncommon counts). Additional investigation,
including genetic analysis, is needed.
Key to Species of the Starksia lepicoelia Complex
1a Lips with distinct dark bars or blotches in preserved males; lips and lower jaw
with alternating pale and darker areas in preserved females; total vertebrae
modally 33; total dorsal elements + anal soft rays + vertebrae modally 78 .....
....................................................S. lepicoelia (Bahamas, Turks and Caicos)
1b Lips without distinct dark markings in preserved males; lips and lower jaw
without conspicuous alternating pale and darker areas in preserved females;
total vertebrae modally 32; total dorsal elements + anal soft rays + vertebrae
modally 75 ..................................................................................................2
2a Preserved males with one to three small dark spots or bars on ventral portion
of lower jaw; dorsal- n elements modally XX,7 ............................................
................................................................... S. robertsoni Panama (Atlantic)
2b Preserved males without dark markings on ventral portion of lower jaw; dor-
sal- n elements modally XIX,8 or XX,8 ......................................................3
3a Lips with conspicuous pattern of white spotting in life; dorsal- n spines mo-
dally XX (also see “Comparisons,” above) .......................... S. weigti (Belize)
3b Lips with few or no white spots in life; dorsal- n spines modally XIX (also see
“Comparisons,” above) ..........S. williamsi (Saba Bank, Netherland Antilles)
Starksia sluiteri Species Complex
Metzelaar (1919) described Brannerella sluiteri from two specimens from Bonaire,
Netherland Antilles. Longley (1934) synonymized Brannerella with Starksia Jordan
and Evermann (type species Labrisomus cremnobates Gilbert, from the eastern Paci c).
Böhlke and Springer (1961) concurred with Longley’s synonymy, noting that Bran-
nerella is distinctive in a single character, and generic recognition of one-character
di erences would require the erection of several new genera within Caribbean Starksia.
Our material includes three genetic lineages originally identi ed as S. sluiteri based
on the taxonomic key of Williams and Mounts (2003) — one from Curacao, one
from Tobago, and one from Belize/Honduras/Panama. Specimens in all three lineages
modally have 13 pectoral- n rays, 20 or fewer dorsal- n spines, and two or three rows
of dark spots or blotches along the body—features typical of S. sluiteri. We have identi-
ed our genetic lineage from Curacao (CUR in Fig. 1) as S. sluiteri (Metzelaar) based
on geography and morphology. In particular, the second row of dark markings (middle
row when there are three) are distinctly round in S. sluiteri and in our Curacao speci-
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
50
mens, whereas those markings are usually vertically elongate in our specimens from
Belize (BLZ), Honduras (HON), and Panama (PAN). Additionally, although Metze-
laar (1919) illustrated a male specimen in his original description, he did not mention
any round, pale markings on the head—prominent diagnostic features in males of our
specimens from Tobago (TOB) that are lacking in our male S. sluiteri from Curacao.
We recognize the genetic lineage from Tobago, as well as that from Belize/Honduras/
Panama, as new species within the S. sluiteri complex and provide descriptions below.
Böhlke and Springer (1961) noted that counts of dorsal- and anal- n elements in
specimens of S. sluiteri they examined from o Colombia and Venezuela (XIX dorsal
spines and 15–16 anal rays) di er from those given by Metzelaar (XX and 17). Based
on pigment, their Colombian and Venezuelan specimens appear to be S. sluiteri. Our
specimens from Curacao, as well as Böhlke and Springer’s two Venezuelan specimens
(USNM 195750), have XIX dorsal spines and 15–16 anal rays. ere is thus a discrep-
ancy between counts in our material and those reported by Metzelaar for the holotype.
We examined a photograph of the holotype, and there appear to be XX dorsal- n
spines as noted by Mezelaar; XX is likely a non-modal count for S. sluiteri. We note
that there is more variation in dorsal- and anal- n counts in some Starksia species than
suggested by Metzelaar’s description; for example, S. green eldi has XVIII–XX dorsal
spines, 7–9 dorsal rays, and 14–16 anal rays.
Starksia green eldi Baldwin & Castillo, sp. n.
urn:lsid:zoobank.org:act:CFD1A620-8C85-4DC3-82A9-8A86BAE66C2A
Figs 1, 12, 15; Table 3
Starksia sluiteri Williams and Mounts 2003, Aqua 6(4): Fig. 9 (male and female speci-
mens from Tobago)
Type Locality: Tobago, Trinidad and Tobago
Holotype. USNM 320832, male, 19.0 mm SL (not a DNA voucher), sta. JTW
90-9, vertical wall just north of Charlotteville on east side of North Point, Tobago,
5–12 m, 8 Sep 1990, J. T. Williams, J. Howe, S. Blum, D. Johnson, S. Love, M.
Schotte.
Paratypes (all from Tobago). USNM 398919, male, 22.0 mm SL (not a DNA
voucher), same locality information as for holotype; USNM 398922, TOB 9282, fe-
male, 19.0 mm SL, sta. TOB09-8, rock/coral outcrops on sand, Pirate’s Bay, Charlot-
teville, < 3 m, 11o19.300'N, 60o32.977'W, 18 Mar 2009 (small llet removed from
right side for DNA tissue sample). USNM 398921, TOB 9275, male, 17.0 mm SL,
collected in same station, TOB09-8, as USNM 398922 above (small llet removed
from right side for DNA tissue sample); USNM 398920, TOB 9212, male, 15.0 mm
SL, sta. TOB09-6, Buccoo Reef, 9–11 m, 11o11.167'N, 60o50.761'W, 17 Mar 2009
(posterior portion of body destroyed for DNA tissue sample); USNM 398924, sta.
TOB09-11, 4 specimens: (1) 12.0 mm SL juvenile, (2) 18.0 mm SL females, (1) 19.5
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 51
mm SL female (not DNA vouchers), Store Bay, 5–9 m, 11o09.349'N, 60o50.535'W, 16
Mar 2009; USNM 398923, sta. TOB 09-1, (1) 17.0 mm SL male (not a DNA vouch-
er), coral heads/coral rubble o Mt. Irvine Beach (Hotel Beach), < 1 m, 11o11.786'N,
60o47.768'W, 15 Mar 2009; USNM 320829, sta. JTW 90–11, female, 22.0 mm SL
(not a DNA voucher), coral rubble/sand, Buccoo Reef (reef crest and lagoon side of
reef), 1–3 m, 11o11'12"N, 60o49'30"W, 10 Sep 1990.
Additional Material (all Tobago). USNM 398925, TOB 9213; USNM 398926,
TOB 9214; USNM 398918, 19 specimens; USNM 398917, 16; USNM 320823, 5.
Diagnosis. A species of Starksia distinguished by the following combination of
characters: orbital cirrus present; two to three rows of dark blotches on side of body,
blotches in middle row (or ventral row if only two rows) mostly circular, never verti-
cally elongate or oval; white (or pale), mostly round spots (absence of melanophores
against a darker background) on at least portions of cheek, opercle, and gular region,
this spotting pattern more prominent in males; males with dark blotch of pigment on
anterior portion of spinous dorsal n; rst anal- n spine one-half to three-quarters
length of male genital papilla; belly naked.
Description. See Table 3. Dorsal spines XVIII–XX (XIX); segmented dorsal rays
7–9, modally 8 (7); total dorsal elements 26–28, modally 27 (26); anal spines (II); seg-
Figure 12. Male and female color patterns of Starksia green eldi: A USNM 398920, TOB 9212, 15.0
mm SL, male B USNM 398922, TOB 9282, 19.0 mm SL, female C–E Diagnostic features of preserved
S. green eldi - C USNM USNM 398919, paratype, male, 22.0 m SL, note pale spots on head D USNM
320832, holotype, male, 19.0 mm SL, note pale spots on head and dark blotch in anterior portion of
spinous dorsal n E USNM 320829, female, 22.0 mm SL, note pale spots on head. Photographs by Car-
ole Baldwin, Cristina Castillo, and Donald Griswold.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
52
Table 3. Frequency distributions of counts among species of the Starksia sluiteri complex1.
Dorsal Spines Dorsal Soft Rays Total Dorsal Anal Soft Rays
XVIII XIX XX 789262728151617
S. green eldi 7 14* 1 5* 12 5 6* 15 1 6* 16 -
S. langi - 10*1 6*4 - 5*5 - 7*1 -
S. sluiteri - 51*2*3114*1211*
Pectoral Rays Dorsal
Procurrent
Caudal Rays
Ventral
Procurrent
Caudal Rays Vertebrae
1213145656313233
S. green eldi - 24* 1 8* 10 16* 1 1 11* 2
S. langi - 12* - 3* 4 6* - 4* 3 -
S. sluiteri 14- 1- 1- - 2-
* Indicates count of holotype
1 Metzelaar (1919) did not provide counts of pectoral- n rays or vertebrae for the holotype of S. sluiteri
mented anal rays 15–16, modally 16 (15); dorsal segmented caudal- n rays 7; ventral
segmented caudal- n rays 6; dorsal procurrent caudal- n rays 5–6 (5); ventral procur-
rent caudal- n rays 5–6, rarely 6 (5); segmented pelvic- n rays 2; pectoral- n rays
13–14, rarely 14 (13); vertebrae 10+21–23=31–33, usually 10+22=32 (10+22=32);
infraorbital pore arrangement variable—unpaired (condition in holotype), one pair at
3 o’clock, two pairs (3 and 6 o’clock), and one specimen with three pairs (3, 4, and 5
o’clock); orbital, nape, and anterior-nostril cirri present and; belly and pectoral- n base
completely naked.
Specimens examined ranging from 11.0–23.0 mm SL; HL 30–36% SL (36%);
length of male genital papilla 19–26% SL in specimens 19.0 mm SL and larger (26%),
12–14% in specimens 15.0–17.0 mm SL; papilla adhered to rst anal- n spine and
extending well beyond it, spine one-half length of papilla in most males, greater than
three-quarters in smallest males.
Pigment. Head and body pale yellow to pale orange, generally more orange in
males, more yellow in females; posterior margins of most body scales covered with
yellow or orange chromatophores mixed with melanophores, resulting in background
pattern of chain-link or diamond-shaped markings. Two or three rows of dark mark-
ings on trunk in mature specimens, markings di use in some specimens: dorsalmost
row with 7–10 roughly square blotches that extend onto bases of dorsal- n elements
(another dark blotch on nape in line with this row of markings); second row with
6–7 circular blotches situated just above lateral midline; lower row, if present, with
1–4 di use, round to oblong blotches. A few to many white, mostly round spots on
at least portions of cheek, opercle, and gular region and sometimes lower jaw; this
pattern resulting from the absence of melanophores against a darker background and
typically signi cantly more prominent in males. Males also di ering from females in
having dark blotch of pigment on anterior portion of spinous dorsal n. Distinctive,
dark-orange markings usually present on proximal portion of dorsal n where dark
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 53
blotches in dorsalmost row of markings on body extend onto dorsal n; where those
dark blotches extend onto two (vs. one) dorsal- n element, dark orange markings dis-
tinctly paired. Orange pigment also present on distal portions of pectoral- n rays and
lighter orange pigment present on at least distal portions of second dorsal-, caudal-,
and posterior anal- n rays; sometimes orange blotches present intermittently along
lengths of second dorsal-, caudal-, and anal- n rays forming wavy stripes or bars of pig-
ment on those ns. Orange pigment present on top of head, in bright ring around eye,
and on nasal cirrus. Some specimens with dark orange pigment on snout, in blotches
radiating from pupil, on operculum, and on dorsal portions of pectoral- n base. In
one specimen most chromatophores on head and body yellow to yellowish orange, but
those on nasal cirrus, around eye, and on ns distinctly orange.
Color in preservative. Diagnostic dark markings on trunk present as described
above; diagnostic white, round spots on head described above present as distinctive
pale markings in preserved specimens—head markings especially prominent in large
males; trunk largely tan and peppered with dark dots, especially along posterior mar-
gins of scales; lips with mottled or barred pigment pattern; a fairly uniform covering
of melanophores on snout, branchiostegals, pectoral- n base, and belly; eye sometimes
surrounded by dark ring of pigment; top of head and nape usually darker than rest
of head, pigment on nape usually in form of dark saddle extending over dorsal mid-
line; two concentrations of melanophores usually visible on brain; dorsal and anal ns
dusky, dark body blotches in upper row usually extending onto base of dorsal n; dor-
sal n of males with dark blotch between spines II–IV; caudal- n rays edged with dark
pigment, outer rays with more uniform scattering of melanophores; proximal portion
of pectoral n covered with scattered melanophores, distal portion with dark edging
along rays; males sometimes with pigment on membranes between some pectoral rays
distally; pelvic n clear.
Etymology. e species name is in honor of David W. Green eld, in recognition
of his work on blennioid shes, particularly his work on the Starksia ocellata complex.
Distribution. Known only from Tobago
Sarksia langi Baldwin & Castillo, sp. n.
urn:lsid:zoobank.org:act:3C78FE0F-BFD6-4F14-9E91-4DD3825A67AE
Figs 13–15, Table 3
Type Locality: Belize, Central America
Holotype. USNM 398927, female, 17.0 mm SL (not a DNA voucher), sta.
CB08-19, inside and outside of Curlew Reef, Belize, 0–3 m, 21 May 2008, C. Bald-
win and Z. Foltz.
Paratypes. USNM 398928, BLZ 8062, female, 17.0 mm SL, sta. CB08-5,
patch reef at south end of Carrie Bow Cay, Belize, 0–3 m, 16 May 2008 (posterior
portion of body removed for DNA tissue sample). USNM 398929, BLZ 8131,
female, 16.0 mm SL, sta. CB08-11, coral heads on sand bottom, Glover’s Reef, Be-
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
54
lize, 0–3 m, 16o43'08.4"N, 87o53'13.1"W, 18 May 2008 (posterior portion of body
removed for DNA tissue sample); USNM 398930, BLZ 8216, female, 11.5 mm SL,
sta. CB08-20, south end of Carrie Bow Cay, Belize, 0–3 m, 21 May 2008 (posterior
portion of body destroyed for DNA tissue sample); USNM 398931, BLZ 8266,
male, 18.0 mm SL, sta. CB08-27, south end of Carrie Bow Cay, 0- m, 23 May 2008
(posterior portion of body removed for DNA tissue sample); USNM 349080, male,
18.0 mm SL (not a DNA voucher), reef crest in front of Carrie Bow Cay, Belize,
16 July 1991; USNM 399917, HON 050, male, 16.3mm SL, Utila, Bay Islands,
Honduras, 3 Jul 2008.
Additional Material. Belize: USNM 317476, 1 specimen (not a DNA voucher).
Colombia (Cayos del Este): UF 223370, 5 (not DNA vouchers)—counts made from
1 male and 1 female, both 16.0 mm SL included in Table 3. Colombia (Isla Providen-
cia): MZUSP 107860, 1 (not a DNA voucher). Panama (San Blas Islands): USNM
399918, PAN 018.
Diagnosis. A species of Starksia distinguished by the following combination of
characters: orbital cirrus present; two rows of prominent, very dark blotches on side of
body, at least some of those in lower row vertically elongate to oval, rarely round; males
with dark, fat, crescent-shaped marking on cheek and without dark blotch on ante-
rior portion of spinous dorsal n; females with scattered dark spots on lower half of
head and on pectoral- n base; rst anal- n spine in males two-thirds to three-quarters
length of male genital papilla; belly naked.
Figure 13. Male and female color patterns of Starksia langi: A USNM 398931, paratype, BLZ 8266,
18.0 mm SL, male B USNM 398929, paratype, BLZ 8131, 16.0 mm SL, female C–E Diagnostic features
of preserved S. langi – (C and D) USNM 398931, paratype, BLZ 8266, male, 18.0 mm SL, note dark
marking on cheek and absence of dark blotch in anterior portion of spinous dorsal n E USNM 398928,
paratype, BLZ 8062, female, 17.0 mm SL, note small dark spots on head. Photographs by Carole Bald-
win, Cristina Castillo, and Donald Griswold.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 55
Description. See Table 3. Dorsal spines XIX–XX, rarely XX (XIX); segmented
dorsal rays 7–8 (7); total dorsal elements bimodal at 26–27 (26); anal spines II; seg-
mented anal rays 15–16, rarely 16 (15); dorsal segmented caudal- n rays 7; ventral
segmented caudal- n rays 6; dorsal procurrent caudal- n rays 5–6 (5); ventral pro-
current caudal- n rays 5; segmented pelvic- n rays 2; pectoral- n rays 13; vertebrae
10+21=31,10+22=32, or 11+21=32 (10+21=31); infraorbital pore arrangement vari-
able—unpaired (condition in holotype), one pair at 3 o’clock, or two pairs (3 and 4
o’clock); orbital, nape, and anterior-nostril cirri present; belly and pectoral- n base
completely naked.
Specimens examined ranging from 9.0–19.0 mm SL; HL 29–33% SL (29%);
length of male genital papilla 19–22% SL; papilla adhered to rst anal- n spine and
extending well beyond it, spine two-thirds to three-quarters length of papilla.
Pigment. Head and body pale orange; posterior margins of most body scales covered
with yellow or orange chromatophores mixed with melanophores, resulting in back-
ground pattern of chain-link or diamond-shaped markings. Two rows of dark markings
on trunk: dorsal row with 9 roughly circular blotches that extend onto bases of dorsal-
n elements (another dark blotch on nape in line with this row of markings); ventral
row with 6–7 blotches along middle of trunk, at least some vertically elongate to oval in
shape; blotches generally not round, although one or more within row may be roughly
so. Females with small dark spots on cheek, operculum, branchiostegals, lower jaw, gu-
lar, and pectoral- n base; spots smaller than pupil (several would t in pupil) but much
larger than tiny dark dots that pepper most of head and trunk; males with dark, fat,
crescent-shaped marking on cheek; orange chromatophores associated with head mark-
ings in both sexes. Both males and females lacking dark blotch of pigment on anterior
portion of spinous-dorsal n. Prominent orange markings present on bases of dorsal- n
elements above dark blotches along dorsal portion of trunk; where dark blotches extend
onto bases of two dorsal- n elements, orange markings distinctively paired; other orange
pigment including chromatophores on top of head, around eye, on nasal cirrus, and on
tips of pectoral-, dorsal-, caudal-, and anal- n rays; those on pectoral n bright orange.
Color in preservative. Diagnostic dark blotches on trunk present as described
above; diagnostic small dark spots on head in females and large blotch on cheek in
males also distinctive in preserved specimens; body overall tan to dark tan. Males with
uniform scattering of spots on lips and rest of head and pectoral- n base; dorsal, cau-
dal, anal, and pectoral rays dusky – i.e., with pigment on membranes between n rays.
Females with dark spots on lips, chin, snout, circumorbitals, and pectoral- n base; top
of head and nape densely covered with melanophores; dorsal, caudal, anal, and pecto-
ral rays edged in dark spots, but little or no pigment on membranes between n rays.
Dark blotches on dorsal portion of trunk extending onto dorsal- n rays in both sexes;
belly pale to lightly pigmented; pelvic n clear.
Etymology. Named in honor of Michael A. Lang, Director of the Smithsonian
Marine Science Network (MSN) and Smithsonian Science Diving Program, in grati-
tude for the support MSN has provided for our Caribbean sh diversity studies and in
recognition of the contributions Michael has made to science diving.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
56
Distribution. Known from Belize, “Colombia,” Honduras, and Panama (see “Re-
marks” below).
Remarks. A tissue sample from a single specimen o Honduras (HON 050 on
tree in Fig. 1) produced a COl sequence very similar to those of our Belize speci-
mens, and one from Panama (PAN 018) is approximately 1% di erent. e Honduras
specimen (Fig. 14A) has the diagnostic pigment on the cheek of male S. langi, and the
Panama specimen (Fig. 14B) has the diagnostic small dark dots of female S. langi. We
recognize the Honduras and Panama specimens as S. langi.
We lack tissue samples of Colombian specimens, but the ve specimens in UF
223370 from Cayos del Este (San Andrés) and a 16-mm SL specimen from Isla Provi-
dencia (Fig. 14C) appear to have the vertically elongate pigment blotches on the trunk
diagnostic of S. langi. Pigment is somewhat faded in the UF specimens, but the 16-
mm SL female in the lot has dark spots on the head as in female S. langi. Although
we include “Colombia” in the distribution list of this species above, we note that the
Colombian specimens are from the Archipelago of San Andrés, Providencia, and Santa
Catalina, a group of islands nearly 800 km from Colombia but only 220 km from
Nicaragua. We have no material from continental Colombia, but S. sluiteri replaces S.
langi o Venezuela.
Comparisons among Species of the Starksia sluiteri Complex (Fig. 15)
Comparative material. Starksia sluiteri. Curacao (all DNA vouchers): USNM 399623,
CUR 8162; USNM 399624, CUR 8226; USNM 399625, CUR 8227; USNM
399626, CUR 8271. Los Roques, Venezuela (not DNA vouchers): USNM 195750,
2 specimens. Dominica (not DNA vouchers): USNM 198263, 15. Puerto Rico (not
a DNA voucher): USNM 219143, 1. Antigua (not a DNA voucher): UF 11344, 1.
Mexico (not DNA vouchers): UF 209342, 2. Starksia fasciata, Turks & Caicos Islands
(all DNA vouchers): USNM 399681, TCI9204; USNM 399683, TCI 9349; USNM
399684, TCI 9350; USNM 399685, TCI 9714. Starksia sp. Navassa Island (not DNA
vouchers): USNM 361059, 2.
Starksia langi is easily distinguished from S. green eldi and S. sluiteri based on
pigmentation of the trunk, head (females), and rst dorsal n (males). e trunk pig-
ment of S. langi comprises both larger and more prominent markings than that of
S. green eldi and S. sluiteri, and only in S. langi are the markings in the second row
vertically elongate (generally round in the other species and sometimes considerably
more di use in S. green eldi). Starksia green eldi lacks dark markings on the head in
both sexes, and S. sluiteri lacks them in females; S. langi males have a prominent dark
blotch on the cheek, and females have numerous small, discrete, dark spots. Males of
S. langi lack a dark blotch on the anterior portion of the dorsal n, whereas this blotch
is present in S. green eldi and S. sluiteri.
Starksia green eldi can be distinguished from S. langi and S. sluiteri by the white
(or pale), mostly round spots (absence of melanophores against a darker background)
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 57
on at least portions of cheek, opercle, and gular region. is pattern is present in both
sexes but is often much more prominent in males. Williams and Mounts (2003) noted
that S. sella, another species of Starksia known only from Tobago, has small pale spots
on the head, but that species lacks dark blotches along the trunk, lacks a dark blotch in
the anterior dorsal n of males, and may be larger (Williams and Mounts specimens of
S. sella are 13.7–27.7 mm SL, our specimens of S. green eldi are 11.0–23.0 mm SL).
S. sluiteri (Metzelaar) is most easily distinguished from S. langi by having the sec-
ond row of trunk blotches almost perfectly round (vs. vertically elongate), in lacking
conspicuous dark spots on the head (females), and in having a dark marking on the
anterior portion of the dorsal n (males). From S. green eldi, S. sluiteri di ers in lack-
ing pale round spots on the head. Although S. sluiteri and S. langi have very similar
Figure 14. Starksia langi. A Male from Honduras, USNM 399917, HON 050, paratype, 16.3 mm SL
(right side, reversed) B Female from Panama (Atlantic), USNM 399918, PAN 018, 14.5 mm SL C Male
from Isla Providencia, Colombia, MZUSP 107860, 16 mm SL. Photographs by Carole Baldwin.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
58
Figure 15. Comparisons among species of the Starksia sluiteri complex and S. fasciata. S.green eldi, left to right: USNM 398921, paratype, TOB 9275, 17.0 mm
SL; USNM 398922, paratype, TOB 9282, 19.0 mm SL; USNM, 320832, holotype, 19.0 mm SL; USNM 320829, paratype, 22.0 mm SL. S. langi: USNM 398931,
BLZ 8266, 18.0 mm SL; USNM 398928, BLZ 8062, 17.0 mm SL; USNM 349080, paratype, 18.0 mm SL; USNM 398927, holotype, 17.0 mm SL. S. sluiteri:
USNM 399626, CUR8271, 16.5 mm SL; USNM 399624, CUR8226, 18.5 mm SL; USNM 195750, 16.9 mm SL. S. fasciata: USNM 399681, TCI 9204, 14.0
mm SL; USNM 399683, TCI 9349, 18.0 mm SL. Juveniles/small adults: S. green eldi, USNM 398925, TOB 9213; S. langi, USNM 398930, paratype, BLZ 8216;
S. sluiteri, USNM 399625, CUR 8227. Photographs by Carole Baldwin, Cristina Castillo, Donald Griswold, and Je rey Williams.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 59
chromatophore patterns, S. sluiteri appears to have more orange pigment on the sec-
ond dorsal, caudal, and anal ns.
In their descriptions of S. leucovitta, S. melasma, S. multilepis, S. rava, and S. sella,
Williams and Mounts (2003) noted that those species belong to the S. sluiteri complex.
Large genetic distances separate the species of the S. sluiteri complex, and our S. multilepis
samples from Brazil are nearly as similar genetically to S. sluiteri as S. langi is (Fig. 1). We
have no tissue samples of the other proposed members of the S. sluiteri complex for com-
parative purposes. ose species are not very similar to S. sluiteri in trunk pigment, par-
ticularly in lacking any bold markings. Starksia fasciata from the Turks and Caicos Islands
(TCI 9204, TCI 9349, TCI 9350) is embedded within our S. sluiteri complex (Fig. 1),
and S. fasciata is morphologically similar to species in that complex (Fig. 15). In Williams
and Mounts (2003) diagnostic key, S. fasciata and S. sluiteri are in the same couplet, sepa-
rated by pattern of pigment on the trunk (bars of trunk pigment in the former, rows of
dark blotches in the latter). Male and female S. fasciata from the Turks and Caicos Islands
(Fig. 15) are very similar to male and female S. langi from Belize in head pigmentation
and in having prominent orange markings along the base of the dorsal n. More mate-
rial is needed to determine if S. smithvanizi, a species that Williams and Mounts (2003)
considered part of the S. fasciata complex, also is genetically aligned with the S. sluiteri
complex. We reiterate that our neighbor-joining tree (Fig. 1) is not intended to re ect
phylogenetic relationships, and a species-level phylogeny derived from multiple genes
should help resolve species and supra-speci c relationships in the S. sluiteri complex.
Museum specimens examined from the Lesser Antilles (Dominica) and Puerto Rico
appear to be S. sluiteri based on trunk pigment (round vs. elongate blotches in the sec-
ond row of markings) and no conspicuous round pale spots on the cheek. e pigment
is somewhat faded in those specimens, however, and more material, including tissue
samples for genetic analysis, is needed. Two female specimens from Navassa (USNM
361059) are not S. sluiteri, as the markings in the second row of trunk blotches are elon-
gate, not round. However, those markings are rectangular in the Navassa specimens,
and the markings in the upper row are square—much more so than in our material of
S. langi from the western Caribbean. e larger of the two females has some dark spots
on the head as in S. langi. More material is needed. Other museum material examined
(e.g., the UF specimens from Antigua and Mexico) are too faded to identify to species.
Key to Species of the Starksia sluiteri Complex
1a Body with two rows of sharply contrasting dark blotches along sides of trunk,
at least some markings in lower row vertically elongate; males without dark
blotch in anterior portion of spinous dorsal n, females with conspicuous
round dark spots on head ..................... S. langi (Belize, Honduras, Panama)
1b Body with two or three rows of di use to sharply contrasting dark blotches
along sides of trunk, those in second row mostly round; males with dark
blotch in anterior portion of spinous dorsal n, females with tiny dots but
without conspicuous round dark spots on head ..........................................2
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
60
2a Portions of head (at least cheek, operculum, gular region) with conspicuous
pale round spots, this spotting pattern often much more prominent in males
than females ..............................................................S. green eldi (Tobago)
2b Head without conspicuous pale round spots S. sluiteri (Netherland Antilles)
Discussion and conclusions
Gilbert (1965) and Green eld (1979) noted that some species of Starksia can only be
distinguished on the basis of color patterns—i.e., they exhibit no other morphological
di erences except sometimes modal di erences in counts. Green eld (1979) surmised
that color patterns on the lips and sides of the head may be important in species recog-
nition in blennioid shes, which often live in cryptic habitats, in some cases (e.g., some
chaenopsids) with only the heads typically visible. Our morphological investigation of
the multiple genetic lineages within S. atlantica, S. lepicoelia, and S. sluiteri resulted
in similar ndings—i.e., most of the member species within the three complexes are
distinguished from one another solely on the basis of pigment patterns, sometimes
only di erences in pigment on the lips and cheeks. All di erences in counts are modal.
Morphological di erences other than pigmentation separate some of the species
complexes; for example, members of the S. atlantica complex lack an orbital cirrus,
and those of S. lepicoelia have a scaled belly. Genetic divergence among species within
each complex is generally smaller than that between complexes: 2–14% within S. at-
lantica, 1–9% within S lepicoelia, and 7–19% within S. sluiteri vs. 17–22% between S.
atlantica and S. lepicoelia, 17–24% between S. lepicoelia and S. sluiteri, and 17–23%
between S. atlantica and S. sluiteri (Tables 4–7). e genetic distances separating spe-
cies of the S. lepicoelia complex are particularly small, and those species are separated
on the basis of minor di erences in pigmentation on the head. Larger genetic distances
separate most species of the S. sluiteri complex, and more prominent di erences in
trunk pigmentation separate some of those species. ere is thus a correlation between
small di erences in COl sequences and minor di erences in pigmentation, suggesting
that pigment patterns may be among the rst morphological changes accompanying
speciation in Starksia. Green eld (1979) did not have the bene t of genetic data for
comparative purposes, but our COl data for four species in his S. ocellata complex
(Fig. 1, Appendix 2) support his decision to recognize species almost entirely on the
basis of minor di erences in pigment. Although species recognition based on such
limited morphological data may in general be a questionable practice, the congruence
between Green eld’s (1979) S. ocellata species and the COl data supports this practice
in Starksia.
ere is not, however, universal congruence between genetic divergence and rec-
ognizable morphological di erences in our data set. One S. green eldi specimen, TOB
9312, is 2% di erent from other S. green eldi, and one S. fasciata, TCI 9204, is 2%
di erent from other S. fasciata. Both of those values are high for intraspeci c variation
in shes in general (often well less than 1%), but we nd no morphological evidence
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 61
Table 4. Average (and range) Kimura two-parameter distance summary for the Starksia atlantica species
complex based on cytochrome c oxidase l (COl) sequences of individuals represented in the neighbor-
joining tree in Figure 1. Intraspeci c averages are shown in bold. n/a = no average (one specimen). BAR
– Barbados, SAB – Saba Bank, PAN – Panama.
Starksia atlantica
(n=7) sangreyae A
(n=6) sangreyae B
(n=6) BAR
(n=2) SAB
(n=1) springeri
(n=2) PAN
(n=2)
atlantica 1%
(0-2) ------
sangreyae A2%
(2-3) 1%
(0-2) -----
sangreyae B2%
(2-3) 2%
(2-3) 1%
(1-0) ----
BAR 9%
(9-10) 10%
(10-12) 9%
(9-10) 0%
(0) ---
SAB 9%
(8-10) 9%
(9-10) 9%
(8-9) 3%
(3) n/a
n/a --
springeri 9%
(8-10) 10%
(9-10) 9%
(8-10) 5%
(5-6) 5%
(5) 0%
(0) -
PAN 13%
(12-14) 13%
(12-14) 13%
(12-13) 11%
(11) 11%
(10-11) 12%
(11-12) 0%
(0)
Table 5. Average (and range) Kimura two-parameter distance summary for the Starksia lepicoelia species
complex based on cytochrome c oxidase l (COl) sequences of individuals represented in the neighbor-
joining tree in Figure 1. Intraspeci c averages are shown in bold; n/a = no average (one specimen).
Starksia lepicoelia A
(n=7) lepicoelia B
(n=2) robertsoni
(n=3) weigti
(n=12) williamsi
(n=1)
lepicoelia A1%
(0-2) ----
lepicoelia B5%
(4-6) 1%
(1) ---
robertsoni 7%
(6-7) 7%
(7-8) 0%
(0-1) --
weigti 6%
(5-8) 6%
(6-7) 2%
(1-2) 0%
(0-1) -
williamsi 8%
(8-9) 8%
(8) 7%
(7) 7%
(7) n/a
n/a
supporting the genetic divergences. Similarly S. sangreyae comprises two sublineages
that are as genetically distinct in COl (2–3%) as S. sangreyae is from S. atlantica (2–
3%), yet no consistent morphological di erences were discovered, not even minor
di erences in color pattern. Even more puzzling, the two genetic sublineages of S. lepi-
coelia are 4–6% di erent in COl, yet we found no morphological di erences between
them (Fig.6). Very little material of one of those lineages is available, and further in-
vestigation is needed. Specimens in the two lineages were taken in the Bahamas at the
same station, in 20–40 ft. of water o Great Stirrup Cay.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
62
In contrast to the examples above, very little sequence divergence in COl exists
between S. sangreyae from Belize and S. atlantica from Bahamas/Turks and Caicos (2–
3%), yet those species are easily distinguished on the basis of trunk pigment. Similar
incongruences between COl data and morphology have been documented. For exam-
ple, Baldwin et al. (2009b) found two morphological (pigment) variants of the goby
Coryphopterus venezuelae, yet those morphs are not genetically distinct. Victor (2010)
pointed out incongruences between COl data and morphologically recognizable spe-
cies in greenbanded gobies (Elacatinus spp.). Speci cally, he noted that E. multifascia-
tus from the eastern Caribbean and E. panamensis from Panama are morphologically
extremely similar, but exhibit 11.3% sequence divergence in COl; he further noted
that despite prominent di erences in color pattern between E. rubrigenus and E. pana-
mensis, those species exhibit only 3.3% sequence divergence in COl.
ose examples notwithstanding, the general congruence between COl lineages
and morphologically recognizable species in western Atlantic Starksia is remarkable,
and we have found the same to be true in our genetic and morphological investigations
of other shore sh genera (e.g., Baldwin et al. 2009a, Baldwin et al. 2009b, Tornabene
et al. 2010). A paper summarizing Smithsonian investigations of western Central At-
lantic shore sh diversity and the utility of DNA Barcoding in this work is in prepa-
ration. Cases in which incongruences exist between genetic and morphological data
ultimately will be further investigated; because DNA barcoding involves sequencing a
Table 6. Average (and range) Kimura two-parameter distance summary for the Starksia sluiteri species
complex based on cytochrome c oxidase l (COl) sequences of individuals represented in the neighbor-
joining tree in Figure 1. Intraspeci c averages are shown in bold.
Starksia green eldi
(n=6) fasciata
(n=4) sluiteri
(n=4) langi
(n=6)
green eldi 1%
(0-2) ---
fasciata 8%
(7-9) 1%
(0-2) --
sluiteri 14%
(13-14) 15%
(14-17) 1%
(0-1) -
langi 17%
(17-19) 16%
(16-19) 16%
(16-17) 1%
(0-2)
Table 7. Range Kimura two-parameter distance summary for the Starksia atlantica, S. lepicoelia, and S.
sluiteri species complexes based on cytochrome c oxidase l (COl) sequences of individuals represented in
the neighbor-joining tree in Figure 1. Within-complex ranges are shown in bold.
S. atlantica complex
(n=26) S. lepicoelia complex
(n=25) S. sluiteri complex
(n=20)
S. atlantica complex 2-14% --
S. lepicoelia complex 17-22% 1-9% -
S. sluiteri complex 17-23% 17-24% 7-19%
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 63
relatively short segment of a single mitochondrial gene, adding additional genetic data
may help resolve some con icts. On the morphological side, adding information from
early life history stages may be of value: the pelagic larval stages of many marine shes
o er a suite of characters for study not present in adults.
A striking element of our COl data for Starksia (Fig. 1) is the correlation be-
tween genetic lineages and geography within the S. atlantica, S. lepicoelia, S. sluiteri,
and S. ocellata species complexes. Specimens from Bahamas, Belize, Curacao, Saba
Bank, and Tobago never occur in more than one genetic lineage within each com-
plex, yet the species complexes themselves are broadly distributed (Fig. 16). Starksia
nanodes also appears to be a broadly distributed species complex, with geographi-
cally distinct genetic lineages in Panama, Barbados, Saba Bank, and Belize (Fig. 1).
Green eld (1979) proposed superspecies status (sensu Amadon 1966, Mayr 1963)
for the S. ocellata complex based on its six allopatric component species, and the S.
atlantica, S. lepicoelia, and S. sluiteri species complexes described herein could be
categorized likewise (we note, however, that the superspecies category has not been
widely adopted in systematic treatments of shes). It is not clear what evolutionary
mechanisms are driving speciation within Starksia, but the life history of the group is
characterized by a short pelagic phase of about two weeks (Victor, unpublished data).
Although pelagic larval duration (PLD) is not always a good indicator of genetic
structure (e.g., Bowen et al. 2006), a short PLD combined with restricted movement
of adults may support the evolution of numerous allopatric species within a group
by restricting gene ow among populations. It is premature to conduct a phylogeo-
graphic analysis of western Atlantic Starksia, but we concur with Green eld (1979)
that the division of some Starksia species into multiple allopatric component species
is not typical of western Atlantic shore shes in general. As noted by Floeter et al.
(2008), Briggs’ (1974) two major biogeographic provinces of the Caribbean (western
Caribbean plus Florida and West Indian/eastern Caribbean) are largely supported
by recent genetic and biogeographical studies. Starksia is not the only exception to
this general trend. Colin (2010) described ve eco-morphological suites of western
Atlantic Elacatinus goby species that are similar to our Starksia species complexes
in that each comprises multiple species usually with allopatric distributions, and
the suites themselves are broadly distributed. Considerably more studies of diversity
and distribution of speciose genera of small, cryptic, Caribbean reef shes and other
Caribbean marine life are needed to determine if there are subdivisions of the major
biogeographic provinces and, if so, what evolutionary mechanisms may be support-
ing them. Rocha et al. (2005) suggested that ecological speciation, in which natu-
ral selection in di erent environmental conditions in adjacent locations may drive
populations along separate evolutionary pathways, could help explain high levels
of species diversity in marine shes in the absence of su cient physical barriers to
account for that diversity. Colin (2010) suggested that faunal breaks in Elacatinus
species may correlate well with observed ocean currents, and he proposed to further
investigate known sh distributions and actual dispersal potential as estimated from
satellite-tracked current drifters.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
64
Figure 16. Distribution of species in the S. atlantica, S. lepicoelia, and S. sluiteri complexes. Only loca-
tions for genetically analyzed specimens plotted. Additional locations for some species discussed in text.
For Starksia, future investigation must include more taxonomic and geographic
coverage. Increased sampling will assuredly result in the recognition of new species
and likely of new species complexes. e faunal breaks that separate members of the
species complexes are unknown. In S. atlantica and S. lepicoelia, our specimens from
Bahamas and Turks and Caicos represent the same species, and in S. sluiteri, specimens
from Belize, Honduras, and Panama appear to be the same. Specimens in close prox-
imity geographically thus tend to cluster into recognizable species. As better coverage
is attained, it will be interesting to see if the same geographical boundaries character-
ize more than one of the species complexes or if the boundaries are di erent for each.
Likewise it will be interesting to compare geographic boundaries of Starksia species
with faunal breaks in other reef shes such as Elacatinus. Future phylogenetic studies in
which relationships among species and species complexes of Starksia and other groups
are hypothesized should help shed light on patterns of speciation in small reef shes of
the western Atlantic.
Because we do not know how much more investigation is required to obtain a
reasonably complete picture of Starksia biodiversity and biogeography, the words of
Winston Churchill included as an epigraph in this paper seem particularly appropriate.
e study of Starksia must continue.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 65
Acknowledgments
J. Van Tassell, D.R. Robertson, W. F. Smith-Vaniz, and J. T. Williams provided color
images of various Starksia species. D. Griswold assisted with photography and radi-
ography of preserved specimens. A. Driskell and A. Ormos provided laboratory and
logistical assistance. D. Smith provided eld assistance and contributed in numerous
other ways to the project. K. Murphy and D. Pitassy provided assistance with catalog-
ing and database issues. J. Bagley, C. Caldow, M. Carpenter, K. Clifton, A. Driskell,
M. Fagan, Z. Foltz, B. Holt, J. Lang, L. Lang, B. Langton, and J. Mounts provided
eld assistance. B. Brown, A. Carvalho, L. Jordan, J. Lamkin, R. Robins, A. Shiroza,
and H. Valles provided specimens. M. van Oijen and R. de Ruiter provided images and
information on the holotype of Brannerella sluiteri. D. Munn and D. Wilson provided
funding for the second author’s internship at the Smithsonian. Research in Florida was
conducted pursuant to SAL # 07SR-1024B to the rst author. A. Gazit, K. Wilson, and
M. Kunen facilitated collecting in Curacao through the CARMABI laboratory. Field-
work in the Bahamas was conducted under the auspices of the Perry Institute of Ma-
rine Science, with logistical assistance from B. Gadd, E. Lamarre, and D. O’Donnell.
A portion of the research in the Bahamas was funded by a donation from C. B. Lang
to the Smithsonian Institution in memory of D. E. Baldwin and R. A. Lang. R. Lang-
ton, J. Gobin, and K. Caesar facilitated collecting in Tobago, and B. Holt provided
support and logistical help for research on South Caicos Island. D. R. Robertson and
the Smithsonian Tropical Research Institute, the Government of Panama, the Kuna
people of the Kuna Yala of the Comarca of San Blas, R. Nemeth and the MacLean
Marine Science Center at the University of the Virgin Islands, and M. Shivji at the Guy
Harvey Research Institute at the Nova Southeastern University Oceanographic Center
provided cooperative assistance. e Smithsonian Marine Science Network provided
major funding for eldwork through a grant to the rst author, and the Smithsonian
DNA Barcoding Initiative provided funding for COl analyses. Additional DNA bar-
coding was facilitated by B. Hanner and supported through funding to the Canadian
Barcode of Life Network from Genome Canada (through the Ontario Genomics In-
stitute), NSERC, and other sponsors listed at www.BOLNET.ca. is is contribution
number 898 of the Caribbean Coral Reef Ecosystems Program (CCRE), Smithsonian
Institution, supported in part by the Hunterdon Oceanographic Research Fund, and
Smithsonian Marine Station at Fort Pierce (SMSFP) Contribution No. 841.
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68
Appendix 1
Starksia specimens included in the genetic analysis (Fig. 1) but not examined for the species
accounts. e voucher specimens from Barbados (BAR), Brazil (BRZ), Panama (PAN),
Florida (FLA 090), and St. omas (STVI) are part of B. Victor’s personal collection.
Species DNA Standard Length (mm) Specimen voucher
S. atlantica PAN 091 9.2 PAN-LIDB091
PAN 092 9.3 PAN-LIDB092
BAR 009 9.9 BAR-LIDB009
BAR 062 9.1 BAR-LIDB062
S. occidentalis BLZ 6271 17.5 USNM 399661
BLZ 6273 30 USNM 399662
BLZ 6304 27 No
BLZ 7335 13 USNM 399663
BLZ 7749 25 USNM 399664
BLZ 8248 24 USNM 399669
BLZ 8249 27 USNM 399670
BLZ 8250 23 USNM 399671
BLZ 8317 25 USNM 399672
BLZ 8375 33 USNM 399665
BLZ 8376 30 USNM 399666
BLZ 8377 9 No
BLZ 8378 15 USNM 399667
BLZ 8379 22 USNM 399668
PAN 058 19.2 PAN-BVCOR058
S. ocellata FLA 090 8.3 FLA-LIDB090
FLA 7391 35 USNM 399686
S. guttata TOB 9016 25 USNM 399630
TOB 9017 25 USNM 399631
TOB 9018 23.5 USNM 399632
TOB 9127 11.5 USNM 399633
S. culebrae STVI 1336 20 USVI-LIDM1336
STVI 019 16 USVI-LIDB019
STVI 121 30.4 USVI-LIDMA121
S. elongata TCI 9115 24 USNM 399634
S. starcki BLZ 8297 11 USNM 399660
S. hassi CUR 8096 28 USNM 399627
CUR 8097 24 USNM 399628
CUR 8319 19 USNM 399629
CUR 8320 10 No
CUR 8349 10 No
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 69
Species DNA Standard Length (mm) Specimen voucher
S. multilepis BRZ 1323 23.2 NOR-LIDM1323
BRZ 1324 19.3 NOR-LIDM1324
BRZ 1325 19.4 NOR-LIDM1325
S. nanodes BAR 011 9 BAR-LIDB011
BAR 013 12.9 BAR-LIDB013
BLZ 5076 12 No
BLZ 5104 13 USNM 399673
BLZ 5105 14 USNM 399674
BLZ 5162 15 USNM 399675
BLZ 5163 12 USNM 399676
BLZ 5277 8 No
BLZ 5424 15.5 USNM 399677
BLZ 6124 13 USNM 399678
BLZ 6125 13 USNM 399679
BLZ 7330 8.5 No
BLZ 7680 15 No
BLZ 8121 15 USNM 399680
BLZ 8289 8.5 No
BLZ 8391 12.5 USNM 399954
PAN 053 14.8 PAN-BVCOR053
PAN 054 13.3 PAN-BVCOR054
PAN 056 14 PAN-BVCOR056
PAN 1247 18.4 PAN-LIDM1247
SAB 0606056 NA * USNM 397404
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
70
Starksia species 12345678
green eldi
(n = 6) 11%
(0-2) -------
fasciata
(n = 4) 28%
(7-9) 1%
(0-2) ------
sluiteri
(n = 4) 3 14%
(13-14) 15%
(14-17) 1%
(0-1) -----
langi
(n = 6) 4 17%
(17-19) 16%
(16-19) 16%
(16-17) 1%
(0-2) ----
multilepis
(n = 3) 5 23%
(22-24) 21%
(20-21) 21%
(21) 17%
(17-18) 0%
(0) ---
hassi
(n = 5) 6 23%
(22-25) 21%
(21-22) 22%
(22-23) 21%
(21-22) 22%
(21-22) 1%
(0-1) --
occidentalis
(n = 15) 7 23%
(22-24) 23%
(22-24) 22%
(21-23) 21%
(21-22) 20%
(20-21) 21%
(20-22) 0%
(0-1) -
ocellata
(n = 2) 8 22%
(22-23) 22%
(22-23) 21%
(20-22) 21%
(21-22) 20%
(20) 22%
(21-22) 3%
(3-3) 0%
(0)
guttata
(n = 4) 9 21%
(20-22) 21%
(21-22) 19%
(18-20) 21%
(20-22) 21%
(20-21) 19%
(18-20) 4%
(3-4) 5%
(5)
culebrae
(n = 4) 10 23%
(23-24) 24%
(23-24) 21%
(20-21) 22%
(21-24) 20%
(20) 21%
(21-22) 6%
(5-6) 6%
(5-6)
elongata
(n = 1) 11 20%
(19-20) 20%
(19-20) 20%
(19-21) 19%
(18-20) 21%
(21) 23%
(22-24) 16%
(15-16) 15%
(15-16)
lepicoelia A
(n = 7) 12 20%
(19-21) 20%
(19-21) 19%
(18-20) 22%
(21-22) 22%
(22-23) 23%
(22-24) 19%
(18-20) 18%
(17-19)
lepicoelia B
(n = 2) 13 20%
(19-21) 20%
(20-21) 18%
(17-19) 21%
(20-22) 23%
(22-23) 22%
(22-23) 18%
(18) 19%
(18-19)
robertsoni
(n = 3) 14 21%
(20-22) 22%
(22-23) 20%
(19-21) 21%
(20-22) 22%
(22) 23%
(22-23) 19%
(19-20) 19%
(19-20)
weigti
(n = 12) 15 20%
(19-23) 22%
(22-24) 20%
(19-20) 21%
(20-22) 22%
(22-23) 22%
(21-23) 19%
(19-20) 19%
(18-19)
williamsi
(n = 1) 16 20%
(20-21) 21%
(21) 20%
(20-20) 21%
(21-22) 22%
(22) 20%
(19-20) 19%
(19-20) 19%
(19)
BAR
(n = 3) 17 20%
(19-21) 20%
(19-21) 22%
(21-23) 21%
(20-21) 23%
(23) 22%
(21-22) 21%
(20-22) 21%
(21-22)
starcki
(n = 1) 18 23%
(23-24) 24%
(23-25) 24%
(24) 22%
(22-24) 26%
(26) 25%
(25-26) 22%
(22-23) 22%
(22)
atlantica
(n = 7) 19 19%
(19-21) 21%
(20-22) 22%
(21-22) 20%
(20-22) 21%
(20-22) 22%
(21-23) 21%
(20-21) 20%
(20-21)
sangreyae A
(n = 6) 20 20%
(19-21) 20%
(19-21) 22%
(21-23) 21%
(20-22) 22%
(21-22) 23%
(22-24) 21%
(21-22) 20%
(20-21)
sangreyae B
(n = 6) 21 20%
(19-22) 21%
(20-22) 22%
(22-23) 20%
(19-20) 21%
(21-22) 23%
(22-24) 21%
(21-22) 21%
(20-21)
BAR
(n = 2) 22 19%
(18-20) 20%
(20) 20%
(20-21) 18%
(17-19) 23%
(23) 20%
(19-20) 20%
(20) 20%
(20)
SAB
(n = 1) 23 19%
(18-20) 20%
(20-21) 21%
(20-22) 18%
(18-19) 24%
(24) 20%
(19-20) 20%
(20-21) 20%
(20)
springeri
(n = 2) 24 18%
(17-19) 20%
(20-21) 21%
(20-21) 19%
(18-20) 24%
(24) 23%
(20-21) 19%
(19-20) 19%
(19-20)
PAN
(n = 2) 25 20%
(20-21) 21%
(20-21) 22%
(21-23) 20%
(20-21) 23%
(23-24) 23%
(22-24) 18%
(18-19) 19%
(19)
nanodes PAN
(n = 5) 26 24%
(23-25) 22%
(21-23) 22%
(21-23) 21%
(20-22) 21%
(20-22) 20%
(20-21) 23%
(22-) 23%
(22-23)
nanodes SAB
(n = 1) 27 24%
(24-25) 24%
(23-24) 23%
(22-23) 21%
(21) 21%
(21-21) 25%
(25-26) 22%
(22-24) 22%
(22-22)
nanodes BAR
(n = 1) 28 24%
(24-25) 24%
(24-25) 22%
(22-23) 20%
(20-21) 22%
(22) 25%
(25-26) 23%
(23-23) 23%
(22-23)
nanodes BLZ
(n = 14) 29 25%
(24-28) 23%
(22-24) 22%
(21-24) 21%
(20-22) 20%
(20-21) 25%
(24-26) 23%
(22-25) 23%
(22-24)
Appendix 2
Average (and range) Kimura two-parameter distance summary for Starksia based on
cytochrome c oxidase l (COl) sequences of all individuals represented in the neighbor-
joining tree in Figure 1. Intraspeci c averages are shown in bold. n/a = no average (one
specimen). BAR – Barbados, SAB – Saba Bank, PAN – Panama, BLZ – Belize.
Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri... 71
9 101112131415161718
1----------
2----------
3----------
4----------
5----------
6----------
7----------
8----------
91%
(0-1) ---------
10 6%
(5-6) 0%
(0) --------
11 16%
(16) 17%
(17) n/a
n/a -------
12 19%
(17-20) 19%
(18-19) 24%
(23-24) 1%
(0-2) ------
13 18%
(17-18) 18%
(18) 23%
(22-23) 5%
(4-6) 1%
(1) -----
14 20%
(19-21) 19%
(18-19) 22%
(22) 7%
(6-7) 7%
(7-8) 0%
(0-1) ----
15 20%
(19-20) 19%
(19-20) 23%
(22-24) 6%
(5-8) 6%
(6-7) 2%
(1-2) 0%
(0-1) ---
16 20%
(20-21) 19%
(19) 23%
(23) 8%
(8-9) 8%
(8) 7%
(7) 7%
(7) n/a
n/a --
17 20%
(19-21) 21%
(21-22) 21%
(21-22) 14%
(13-15) 15%
(15) 14%
(14-15) 13%
(13-15) 13%
(13-14) 1%
(1) -
18 23%
(23-24) 24%
(24) 23%
(23) 24%
(24-25) 25%
(25) 24%
(24) 24%
(23-25) 24%
(24) 25%
(24-26) n/a
n/a
19 20%
(20-21) 20%
(19-21) 21%
(20-21) 19%
(18-20) 18%
(18-19) 18%
(17-19) 18%
(17-20) 19%
(18-20) 22%
(21-23) 22%
(21-23)
20 21%
(20-22) 19%
(19-20) 21%
(21-22) 19%
(19-20) 19%
(18-20) 18%
(17-18) 18%
(17-21) 18%
(18) 22%
(21-22) 22%
(22-23)
21 21%
(21-22) 20%
(20-21) 21%
(20-21) 20%
(19-21) 20%
(19-20) 18%
(17-18) 18%
(17-20) 19%
(19-20) 22%
(21-23) 22%
(22-23)
22 19%
(18-19) 19%
(19) 20%
(20) 21%
(20-21) 19%
(19-20) 18%
(18) 19%
(18-21) 19%
(19) 22%
(21-23) 22%
(22)
23 19%
(19) 19%
(19) 19%
(19) 20%
(20-21) 19%
(19) 19%
(18-19) 19%
(18-20) 18%
(18) 22%
(21-22) 21%
(21)
24 18%
(18-19) 18%
(18-19) 18%
(18) 21%
(20-21) 19%
(19-20) 18%
(18-19) 19%
(19-21) 18%
(18) 21%
(21-22) 21%
(20-21)
25 19%
(18-19) 18%
(18-19) 21%
(21) 21%
(20-22) 20%
(20) 20%
(19-20) 20%
(19-20) 19%
(19) 21%
(21) 25%
(25-26)
26 22%
(21-23) 23%
(22-24) 23%
(23) 22%
(21-23) 22%
(21-22) 22%
(22-23) 23%
(22-24) 22%
(22) 24%
(23-24) 27%
(26-27)
27 23%
(23-24) 21%
(21) 24%
(24) 23%
(22-24) 22%
(22-23) 22%
(22) 22%
(22-23) 23%
(23) 24%
(24-25) 29%
(29)
28 24%
(23-24) 22%
(22) 25%
(25) 24%
(23-24) 21%
(21) 23%
(22-23) 24%
(23-24) 23%
(23) 25%
(24-25) 29%
(29)
29 23%
(22-24) 24%
(23-25) 22%
(21-22) 23%
(22-25) 22%
(21-23) 24%
(23-25) 23%
(22-25) 23%
(24-25) 22%
(22-23) 26%
(26-28)
Appendix 2, cont.
Carole C. Baldwin et al. / ZooKeys 79: 21–72 (2011)
72
19 20 21 22 23 24 25 26 27 28 29
1-----------
2-----------
3-----------
4-----------
5-----------
6-----------
7-----------
8-----------
9-----------
10-----------
11-----------
12-----------
13-----------
14-----------
15-----------
16-----------
17-----------
18-----------
19 1%
(0-2) ----------
20 2%
(2-3) 1%
(0-2) ---------
21 2%
(2-3) 2%
(2-3) 1%
(1-0) --------
22 9%
(9-10) 10%
(10-12) 9%
(9-10) 0%
(0) -------
23 9%
(8-10) 9%
(9-10) 9%
(8-9) 3%
(3) n/a
n/a ------
24 9%
(8-10) 10%
(9-10) 9%
(8-10) 5%
(5-6) 5%
(5) 0%
(0) -----
25 13%
(12-14) 13%
(12-14) 13%
(12-13) 11%
(11) 11%
(10-11) 12%
(11-12) 0%
(0) ----
26 19%
(18-20) 20%
(19-21) 20%
(19-21) 20%
(20-21) 20%
(20-21) 23%
(22-23) 22%
(22-23) 1%
0(1-) ---
27 20%
(19-20) 20%
(19-20) 21%
(20-21) 20%
(19-20) 20%
(20) 23%
(22-23) 20%
(20) 12%
(11-12) n/a
n/a --
28 19%
(19-20) 20%
(19-21) 20%
(20-20) 20%
(20) 19%
(19) 22%
(21-22) 21%
(21) 11%
(10-11) 3%
(3) n/a
n/a -
29 18%
(17-19) 19%
(18-19) 19%
(18-19) 21%
(20-22) 21%
(20-22) 20%
(20-21) 21%
(21-23) 12%
(11-14) 11%
(11-12) 11%
(11-12) 0%
(0-1)
Appendix 2, cont.
