77
A new ca e-dwelling species o T imma (Teleos ei, Gobiidae)
om he Red Sea, wi h no es on Red Sea endemism in T imma spp.
Vik o N. Peinemann1, Lucia Pombo-Ayo a1,2 , Wal e A. Rich1, Michael D. Fox1, Da en J. Coke 1,3
1 Ma ine Science P og am, Di ision o Biological and En i onmen al Science and Enginee ing, King Abdullah Uni e si y o Science and Technology, Thuwal
23955, Saudi A abia
2 Depa men o Ma ine Science, The Uni e si y o Texas a Aus in, Ma ine Science Ins i u e, 78373 Po A ansas, Texas, USA
3 KAUST Co al Res o a ion Ini ia i e (KCRI), King Abdullah Uni e si y o Science and Technology (KAUST), 23955 Thuwal, Saudi A abia
Co esponding au ho : Vik o N. Peinemann ( ik o [email p o ec ed])
Copy igh : © Vik o N. Peinemann e al.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
A new species o he gobiid genus T imma is desc ibed om he Fa asan Banks in he
sou heas e n Red Sea. The new species is cha ac e ized by ha ing a p edo sal mid-
line wi h 7–8 scales, he i h pel ic- in ay unb anched, cheek and ope cle scaleless,
all pec o al- in ays unb anched, and a do sal in VI + I,7, wi hou elonga e spines. In
li e, he species is b igh yellow h oughou , wi h a dis inc i e yellow-g een longi udinal
band in he cen al hi d o he do sal ins. The new species inhabi s ca es on exposed
o sho e ee s a dep hs be ween 15 and 30 m whe e i occu s in small g oups o up o
10 indi iduals. The new species appea s o be sis e o T. winchi om he wes e n Indian
Ocean. We also p esen a mul ilocus phylogeny (COI, 16S, P , S7I1), including all known
Red Sea T imma and 21 non-Red Sea species, and an expanded supe ma ix ee wi h
93 species o place Red Sea endemism in b oade con ex . This b ings he o al numbe
o T imma species known om he Red Sea o 10, wi h eigh appea ing o be endemic
o he egion. The high p opo ion o endemism in he genus is no ewo hy, e en o he
Red Sea, which has one o he highes p opo ions o endemic ee ishes in he Indo-Pa-
ci ic. Mo eo e , K2P dis ances in he wo widesp ead species sugges hey may also
ep esen c yp ic endemic species, bu u he analyses a e needed. The new species is
cu en ly known only om he Fa asan Banks egion despi e ex ensi e sampling along
he Saudi A abian Red Sea coas .
Key wo ds: Biodi e si y, biogeog aphy, COI, c yp oben hic, Indian Ocean, phylogeny,
axonomy
In oduc ion
The gobiid genus T imma Jo dan & Seale, 1906 is one o he mos speciose
gene a o co al ee -associa ed ishes in he Indo-Paci ic, wi h o e 110 alid
desc ibed species. Commonly known as pygmygobies due o hei small body
size ( ypically <30 mm SL), membe s o his genus can be ound on mos In-
do-Paci ic co al ee s. Along wi h he simila ly di e se and small-bodied E io-
a Jenkins, 1903, hey equen ly ep esen one o he mos abundan gobiids
on ha d subs a es o co al ee s (Win e bo om 2019). Like E io a, species o
Academic edi o : Tihomi S e ano
Recei ed:
18 May 2025
Accep ed:
22 Augus 2025
Published:
9 Oc obe 2025
ZooBank: h ps://zoobank.
o g/13A27F9A-2CD1-44B1-911C-
C08ABC79EFB0
Ci a ion: Peinemann VN, Pombo-
Ayo a L, Rich WA, Fox MD, Coke DJ
(2025) A new ca e-dwelling species
o T imma (Teleos ei, Gobiidae) om
he Red Sea, wi h no es on Red Sea
endemism in T imma spp. ZooKeys
1255: 77–93. h ps://doi.o g/10.3897/
zookeys.1255.159341
ZooKeys 1255: 77–93 (2025)
DOI: 10.3897/zookeys.1255.159341
78
ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
T imma equen ly show s ong si e ideli y and mic ohabi a specializa ion,
likely con ibu ing o hei apid di e si ica ion. New species a e being disco -
e ed equen ly, and molecula analyses sugges ha many widesp ead spe-
cies may ep esen complexes o mul iple c yp ic species wi h es ic ed geo-
g aphic dis ibu ions (Win e bo om e al. 2020).
The Red Sea ha bo s a dis inc i e assemblage o T imma species, wi h mos
being endemic o he egion (Bogo odsky and Randall 2019). Mos o hese spe-
cies show s ong associa ions wi h speci ic ee zones o mic ohabi a s. T im-
ma la icauda um (Go en, 1982) and T. a ido i (Go en, 1978) o en ep esen he
dominan c yp oben hic species on many Red Sea ee s, in some cases wi h
dozens o indi iduals pe squa e me e (Coke e al. 2018). Howe e , hei niche
o e lap is limi ed, wi h ypically only one o he species domina ing in each hab-
i a . Some species, including T. ba alli Win e bo om, 1995 and T. sheppa di
Win e bo om, 1984, p ima ily inhabi dep hs o a leas 30 m, while o he s, such
as T. mendelssohni (Go en, 1978), a e common in he uppe 10 m o he ee .
Associa ions wi h ca es, which a e common o T imma species h oughou he
Indo-Paci ic, a e also p esen in he Red Sea. Bo h T. ishelsoni Go en, 1985 and
T. nuba um Win e bo om, Bogo odsky & Alpe mann, 2023 li e in ca e-associa -
ed g oups. The o me p e e s small g oups o low densi y, while he la e can
be obse ed in dense schools o upwa ds o 30 indi iduals, ho e ing abo e he
subs a e nea he en ances o ca es (pe s. obs. o i s au ho ).
Du ing biodi e si y su eys o o sho e ee s in he Fa asan Banks egion
o he Saudi A abian Red Sea, we disco e ed a dis inc i e, yellow-colo ed spe-
cies o T imma inhabi ing ca e sys ems a dep hs be ween 15–30 m. While
o en co-occu ing wi h o he ca e-associa ed T imma species, his axon was
ypically es ic ed o a eas sligh ly deepe inside he ca e and mo e closely
associa ed wi h he ca e subs a e. He ein we desc ibe his new species based
on mo phological and molecula e idence and discuss i s ela ionship o o he
membe s o he genus, pa icula ly i s appa en sis e species T. winchi Win e -
bo om 1984 om he wes e n Indian Ocean.
Ma e ials and me hods
Mo phological measu emen s and me is ic coun s ollow s anda d p o ocols
es ablished by Richa d Win e bo om o ensu e compa abili y wi h p e ious
wo k (Win e bo om 2019; Win e bo om e al. 2023 and e e ences ci ed he e-
in). Likewise, he o ma o he diagnosis and desc ip ion ollows he sequence
o cha ac e s in he key o he genus (Win e bo om 2019) and o he ecen
species desc ip ions (Win e bo om e al. 2023). We documen ed cephalic
senso y papillae ollowing es ablished ow naming con en ions (Win e bo om
2011; Win e bo om and E dmann 2015). Cha ac e anges a e epo ed as min-
imum – holo ype – maximum, ollowed by he mean and numbe o specimens
in pa en heses. Values o he holo ype a e p esen ed in bold.
Specimens we e collec ed om ca e habi a s a dep hs o 15–30 m using
clo e oil anes he ic and hand ne s. Li e and esh colo a ion was documen -
ed h ough pho og aphs aken immedia ely a e collec ion and du ing in si u
obse a ions. Ex-si u pho og aphs we e aken in an up igh pho o ank wi h a
Nikon Z7 came a body, Nikon Z MC 105 mm /2.8 mac o lens, and a Godox
MF12 Mac o lash. To enhance isibili y o key axonomic ea u es, pa icu-
79
ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
la ly he cephalic senso y papillae, specimens we e empo a ily s ained wi h
cyanine blue 5R (acid blue 113).
Fo molecula analyses, we ex ac ed genomic DNA om issue samples us-
ing he Qiagen DNeasy Blood & Tissue Ki . A egion o he cy och ome c oxidase
subuni I (COI) gene was ampli ied using he p ime se s FishF2 and FishR2
(Wa d e al. 2005) and GOBYL6468 and GOBYH7696 (Thacke 2003). Resul -
ing sequences we e hen cleaned and aligned in Geneious P ime 2025.0.3 o
p oduce a 648-bp alignmen . Pai wise gene ic dis ances be ween species we e
calcula ed using MEGA11 using he Kimu a 2-pa ame e (K2P) model (Kimu a
1980; Tamu a e al. 2021) on he COI alignmen . We also ampli ied h ee addi-
ional loci om he same specimens: mi ochond ial 16S RNA using p ime s
16Sa L and 16Sb H (Palumbi e al. 1991), nuclea S7 ibosomal p o ein in on
1 (S7I1) using p ime s S7I1F and S7I1R (Chow and Hazama 1998), and nuclea
P o ease III (P ) using p ime s P F2 and P R2 (Yamada e al. 2009). P was
included because i can help esol e deepe spli s and has been used in he
simila ly speciose genus E io a (To nabene e al. 2013).
The inal conca ena ed alignmen consis ed o 648 bp COI, 588 bp P , 904
bp S7I1, and 608 bp 16S. The mul i-locus da ase comp ised specimens om
ou s udies and collec ions, including all desc ibed Red Sea T imma, 21 non-
Red Sea T imma, and wo likely undesc ibed axa: one Red Sea species simila
o T. anaima Win e bo om, 2000 and one Madagasca species simila o T.
quad imacula um Hoese, Bogo odsky & Mal, 2015. These specimens a e cu -
en ly being wo ked on by he au ho s, bu hey we e included he e due o hei
ele ance o Red Sea endemism in T imma spp.
A maximum-likelihood ee was in e ed using W-IQ-TREE (T i inopoulos e
al. 2016) ia he web in e ace a h p://iq ee.cibi .uni ie.ac.a /. P o ein-cod-
ing genes (COI and P ) we e pa i ioned by codon posi ion; all loci we e also
pa i ioned by gene. The bes - i subs i u ion model was au oma ically selec ed.
B anch suppo was assessed wi h 1000 ul a as boo s ap eplica es. Sue io a
ae hon Peinemann, Pombo-Ayo a & To nabene, 2024 was included as an ou -
g oup o oo he ee. The phylogene ic ee was isualized using In e ac i e T ee
O Li e (iTOL) 6 (Le unic and Bo k 2024) and inalized in Adobe Illus a o 29.2.1.
In addi ion o he mul i-locus da ase , we cons uc ed a supe ma ix ha com-
bines he ull conca ena ed da a o ou specimens (COI, P , S7I1, 16S) wi h
COI-only e minals o T imma species no ep esen ed in ou collec ion. We que-
ied he Ba code o Li e Da a Sys em (BOLD) o all such species, selec ing one
ep esen a i e COI sequence pe species, wi h p e e ence o longe sequences
and o eco ds iden i ied by ecognized expe s. These public COI sequences
we e aligned o he COI pa i ion o ou conca ena ed da ase ; he emaining
h ee loci o hose added axa we e coded as missing da a. This p ese es he
highe signal o ou mul i-locus sampling while placing i in he con ex o he
majo i y o desc ibed species o T imma. Supp. ma e ial 1 lis s, o each added
species, he species name as used in he ee, he BOLD P ocess ID and, whe e
a ailable, he co esponding GenBank accession numbe , he name o he spec-
imen iden i ie , and he collec ion locali y. Supp. ma e ial 2 lis s in o ma ion o
specimens in ou collec ion used o he mul i-locus da ase and he COI and
16S GenBank accession numbe s gene a ed o his s udy. The inal supe ma ix
ee included all 33 T imma species om ou mul i-locus da ase plus 60 addi-
ional COI-only species om BOLD, o a o al o 93 T imma species.
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ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
Type specimens ha e been ca alogued and deposi ed in he ich hyology col-
lec ion o he Royal On a io Museum (ROM). All ieldwo k was conduc ed unde
app o al 20IAUCUC05 issued by he KAUST Ins i u ional Animal Ca e and Use
Commi ee (IACUC).
Resul s
T imma be umeni sp. no .
h ps://zoobank.o g/9759ABBF-2807-4C55-9925-7598731B789B
Figs 1, 2
Mikey’s golden pygmygoby
Type locali y. Saudi A abia, Red Sea: Fa asan Banks, 18.5041°N, 40.6606°E.
Type ma e ial. Holo ype • ROM 116871, 13.6 mm SL male, Red Sea, Saudi
A abia, Fa asan Banks, 18.5041°N, 40.6606°E, inside ca e o exposed ee wall,
27 m dep h, collec ed wi h clo e oil and hand ne , V. N. Peinemann, 5 June 2024.
Pa a ypes • ROM 116872, n = 2, 13.3 mm SL male, Red Sea, Saudi A abia,
Fa asan Banks, 18.5041°N, 40.6606°E, inside ca e o exposed ee wall, 27 m
and 15 m dep h espec i ely, collec ed wi h holo ype • ROM 116873, 13.8 mm
SL male, Red Sea, Saudi A abia, Fa asan Banks, 18.8581°N, 40.3788°E, inside
ca e o exposed ee wall, 15 m dep h, V. N. Peinemann, 6 June 2024 • ROM
116874, n = 3, 10.7 mm SL emale, 10.9 mm SL male, and 10.7 mm SL male,
Red Sea, Saudi A abia, Fa asan Banks, 19.8375°N, 39.9297°E, inside ca e o
exposed ee wall, 30 m dep h, V. N. Peinemann, 19 May 2022.
Diagnosis. A species o T imma dis inguished by he ollowing combina ion
o cha ac e s: p edo sal midline wi h 7–8 scales; i h pel ic- in ay unb anched
and 40–42% leng h o ou h ay; cheek and ope cle scaleless; all pec o al- in
ays unb anched; body b igh yellow h oughou and wi hou ba s in li e; do sal
in VI + I,7; no elonga e spines in do sal in; bony in e o bi al wid h 81–84%
o pupil diame e ; i is golden-yellow wi h melanopho es excep o golden ing
a ound pupil and in e ed golden iangle ex ending om en al ma gin o pu-
pil; ins anslucen wi h yellow-g een longi udinal band in cen al hi d o do sal
ins; a hin black midla e al line om abo e pec o al- in base o caudal- in base,
isible in li e and in p ese a i e.
Figu e 1. F eshly collec ed T imma be umeni sp. no ., holo ype, ROM 116871. Pho og aph: V. N. Peinemann.
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ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
Figu e 2. Close-up o key ea u es o T imma be umeni sp. no ., p ese ed in e hanol, ROM 116871 (A, B, C, D, E, G), ROM
116874 (F). A. La e al iew o head showing cephalic senso y papillae, specimen s ained wi h cyanine blue, papillae high-
ligh ed wi h black do s, papilla ows connec ed by do ed lines and labeled acco dingly; B. Do sal iew o head, showing
cephalic senso y papillae and nasal appa a us, specimen s ained wi h cyanine blue, papillae highligh ed wi h black do s,
papilla ows connec ed by do ed lines and labeled acco dingly; C. Ven al iew o head, showing cephalic senso y papil-
lae, gill opening, and pel ic- in base; D. Caudal in wi h senso y papillae; E. Male u ogeni al papilla; F. Female u ogeni al
papilla, specimen s ained wi h cyanine blue; G. La e al iew o whole ish (p ese ed in 75% e hanol).
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ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
Desc ip ion. Based on holo ype and 6 pa a ypes 10.7–13.8 mm SL. Do sal
in VI + I,7, second spine longes bu no elonga e ex ending o base o second
do sal- in spine when adp essed; all ays o second do sal in b anched; sec-
ond do sal in no elonga e, eaching pos e io ly 23–26–30 (27%, 7) o caudal
peduncle leng h; anal in I, 8, all ays b anched; anal in no elonga e, eaching
pos e io ly 23–30 (27%, 7) o caudal peduncle leng h; pec o al- in ays 14–15,
all unb anched; pec o al- in eaching pos e io ly o e ical abo e u ogeni al
papilla; pel ic in I, 5, i h ay unb anched and 40–42 (41%, 7) leng h o ou h
ay, which eaches pos e io ly o be ween base o i s and hi d anal- in ay, pel-
ic ays 1 o 4 wi h one b anching poin each; basal memb ane app oxima ely
9–10% o ou h pel ic- in ay; no aenum; caudal in wi h 2 do sal and 2–3
en al segmen ed unb anched ays, and 6 do sal and 5 en al segmen ed
b anched ays; caudal in wi h ou e ically aligned papillae a base and h ee
ows o 9–10 papillae each ex ending om he e ically aligned papillae o
pos e io ma gin o caudal in (Fig. 2D); male u ogeni al papilla elonga ed and
na ow (Fig. 2E; emale u ogeni al papilla sho and bulbous (Fig. 2F).
La e al scales 24; an e io ans e se scales 7–8, pos e io ans e se scales
7–8; cheek and ope cle scaleless; 7–8 scales on p edo sal midline; body scales
c enoid; ci cumpeduncula scales 12; scale ows in en al midline be ween
base o las anal- in ay and i s p ocu en caudal- in ay 7–8.
Gill opening ex ending an e o en ally o be ween middle and pos e io hi d
o pupil (Fig. 2C); an e io na is ubula eaching an e io ly ac oss uppe lip o
pos e io ma gin o lowe lip; pos e io na is o al and po e-like wi h aised im,
sepa a ed om bony on o o bi by 1.6–2 imes i s diame e (Fig. 2B); bony
in e o bi al wid h 81–84% o pupil diame e (82%, 7); no de mal idge on mid-
line o nape ex ending an e io ly om o igin o i s do sal in.
Caudal peduncle dep h as pe cen age o caudal peduncle leng h 30–41
(37.2%, 7); head leng h as pe cen age o SL 27–29–31 (29.3%, 7); ho izon al
eye diame e 40–45 (43.1%, 7); snou leng h 16–20 (18.9%, 7); and uppe -jaw
leng h 32–33 (32.6%, 7) as pe cen age o head leng h
Numbe o papillae in each ow (Fig. 2A): a = 6 (4); b = 1–2 (5); c = 5 (5); d =
3–4 (5); d’ = 3–4 (5); p = 6 (5); e-an e io = 7–9 (5); e-pos e io = 7–9 (5); i-an e-
io = 6–7 (5); i-pos e io = 7 (5); cs” = 3 (5); o = 7–8 (4); oi = 3–4 (4); u = 4 (3);
cp = 1 (5); n = 1 (5); = 2–3 (5).
Colo pa e n, li e (Fig. 1). Body b igh yellow h oughou . Head p ima ily
yellow wi h o ange colo a ion a ound he jaws, snou , in e o bi al egion and
below he en al ma gin o he eye. Indi iduals can ac i a e i e anslucen
whi ish ba s along do sal ma gin o body, om abo e he pec o al in base o
he caudal peduncle, some imes isible in si u. Head and body wi h nume ous
small melanopho es sca e ed h oughou , unning along scale pocke s in he
do sal hi d o body, and sca e ed wi hou pa e ns below he midla e al line.
Specimens smalle han 11 mm SL wi h ew melanopho es on en al hal o
body. A hin black midla e al line unning along body om abo e pec o al- in
base o caudal- in base. I is golden-yellow, peppe ed wi h melanopho es, ex-
cep in a golden ing su ounding he pupil and an in e ed golden iangle ex-
ending om en al ma gin o he pupil. Do sal ins anslucen wi h dense ed,
whi e, and black ch oma opho es along basal hi d, a yellow-g een longi udinal
band unning along cen al hi d, and spa se ed, whi e, and black ch oma o-
pho es along dis al hi d. Anal in mos ly anslucen wi h ed, whi e, and black
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ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
ch oma opho es, wi hou yellow-g een pigmen a ion. Do sal and en al i hs
o caudal in anslucen wi h sca e ed ed and whi e ch oma opho es, cen al
sec ion o caudal in yellow o yellow-g een. Pec o al ins and pel ic ins hyaline.
Colo pa e n, p ese ed (Fig. 2G). Head and body p ima ily ligh yellow in
la ge and ecen ly p ese ed specimens, slowly ading o whi e in small spec-
imens olde han a yea . Melanopho es emain sca e ed h oughou . A hin
black midla e al line jus unde scales, ex ending om abo e pec o al- in base
o caudal- in base. I is densely packed wi h melanopho es, appea ing mos ly
black. Do sal and anal ins hyaline wi h melanopho es sca e ed in he pos e-
io hi d o each in, g owing dense owa ds he pos e io ma gin o he in.
Pec o al, pel ic, and caudal ins hyaline.
E ymology. Named in hono o Michael L. Be umen in ecogni ion o his sub-
s an ial con ibu ions o ou unde s anding o he ecology and biodi e si y o Red
Sea co al ee s. Mikey’s golden pygmygoby is sugges ed as he common name.
Dis ibu ion and habi a . T imma be umeni sp. no . inhabi s ca es o ex-
posed o sho e ee s, whe e i mo es along he su ace o ca e oo s and walls
in small g oups o h ee o en indi iduals (occasionally i is also soli a y). I is
ypically ound deep wi hin ca es, a ely seen wi hin he i s 80 cm o a ca e
en ance. The habi a is simila o ha o i s sis e species, T imma winchi, in
he Seychelles (Ryan Daly pe s. comm.).
Specimens we e collec ed a dep hs be ween 15 and 30 m. While only wo
specimens we e obse ed shallowe han 20 m, he species is ela i ely com-
mon in ca es a 30 m. Ou collec ions and su eys we e limi ed o 30 m, bu i
is likely ha he species ex ends o g ea e dep hs. We obse ed his species
exclusi ely on ee s wi h s eep walls ha ex end well in o he mesopho ic.
The species is p esen h oughou much o he Fa asan Banks in he sou h-
eas e n Red Sea (Fig. 3). Despi e ex ensi e collec ions in simila habi a s along
0500 1,000 km
T imma be umeni sp. no .
T imma winchi
R
R
e
e
d
d
S
S
e
e
a
a
A
A
a
a
b
b
i
i
a
a
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n
S
S
e
e
a
a
I
I
n
n
d
d
i
i
a
a
n
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O
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c
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Figu e 3. Map showing collec ion locali ies o T imma be umeni sp. no . ( ed diamonds;
p esen s udy) and i s sis e species T. winchi (o ange diamonds; Win e bo om 2019) in
he Red Sea and Indian Ocean.
84
ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
he Saudi A abian coas line no h o he Fa asan Banks, om Jeddah o Ti an,
we ha e no obse ed his species anywhe e bu he Fa asan Banks. I is possi-
ble ha he species is endemic o he sou he n hal o he Red Sea.
Compa isons
In he key o he genus (Win e bo om 2019), T. be umeni keys ou o couple
71 bu does no ma ch ei he o he wo species in he couple . T imma be u-
meni di e s om bo h in ha ing 7 do sal in ays ( s 9 in T imma imaii Suzuki &
Senou, 2009 and 8 in T imma ma sunoi Suzuki Sakaue & Senou, 2012). I addi-
ionally di e s om T. imaii in ha ing a bony in e o bi al wid h wide han 80%
o pupil wid h ( s 40% in T. imaii) and om T. ma sunoi in ha ing 7–8 p edo sal
midline scales ( s 6 in T. ma sunoi). The gene al body, in, and eye colo is also
clea ly dis inc in each species.
In o e all mo phology, he T. be umeni is mos simila o T. winchi, which is
cu en ly known om he Seychelles and he Chagos A chipelago (Fig. 4A). They
sha e a uni o m yellow body colo , ha e a simila habi a , and a geog aphically
close bu non-o e lapping dis ibu ion. The COI sequences o T. be umeni and T.
winchi di e by a K2P dis ance o 6.4% (see Suppl. ma e ial 3 o K2P dis ances
o Red Sea T imma and sis e axa in Fig. 5B). Se e al ai s can be used o sepa-
a e he wo species. These a e summa ized in Table 1. No ably, he wo Chagos
( ype locali y) male specimens o T. winchi ha e an elonga e do sal spine, which
is absen in Ryan Daly’s pho og aphs o T. winchi om he Seychelles. I has no
been e alua ed whe he his ep esen s a dis inc species o in aspeci ic a ia-
ion. In Table 1 we ollow he o iginal desc ip ion in lis ing do sal-spine elonga-
ion o T. winchi, bu he a ia ion wa an s u he in es iga ion.
Wi hin he Red Sea, T. ishelsoni bea s some simila i y o T. be umeni. Bo h
species ha e a p ima ily yellow body colo , a yellow longi udinal band along
he do sal ins, and a p ima ily yellow caudal in. Bo h species also ha e an
Table 1. Compa ison o key cha ac e s be ween T imma be umeni sp. no . and T. winchi. Da a o T. winchi based on
Win e bo om (1984), Win e bo om (2019), and examina ion o a pho og aph o holo ype.
T imma be umeni sp. no . T imma winchi
Geog aphic ange Fa asan Banks, sou he n Red Sea Seychelles and Chagos A chipelago
Do sal in VI + I,7 VI + I,8
Elonga e do sal spines None; second spine longes eaching o base o
second do sal- in spine when adp essed
Second spine elonga e, eaching pos e io ly jus
beyond base o las do sal- in ay when adp essed
(absen in Seychelles specimens)
Second do sal- in leng h Las ay no elonga e, eaching pos e io ly 23–30%
o caudal peduncle leng h
Las ay elonga e, eaching pos e io ly wo- hi ds o
caudal peduncle leng h
Anal- in leng h Las ay no elonga e, eaching pos e io ly 23–30%
o caudal peduncle leng h
Las ay elonga e, eaching pos e io ly wo- hi ds o
caudal peduncle leng h
Do sal- in colo (ali e) Black longi udinal basal s ipe composed o la ge
melanopho es wi h sca e ed i idocy es; single
yellow longi udinal band abo e i ; dis al hi d hyaline
Black longi udinal basal s ipe composed o la ge
melanopho es wi h sca e ed i idocy es; a cen al
yellow longi udinal band, hen a na ow hyaline
band; dis al qua e yellow
Anal- in colo (ali e) No yellow pigmen a ion, mos ly anslucen wi h
ed, whi e, and black ch oma opho es
Mos ly yellow, wi h basal band o melanopho es
and i idocy es
Caudal- in colo (ali e) Yellow wi h anslucen do sal and en al ma gins Yellow h oughou wi h dis al ma gin o
melanopho es
85
ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
Figu e 4. Compa ison o species simila o T imma be umeni sp. no . in he Red Sea and Indian Ocean. A. T. winchi, ho-
lo ype, ROM 41477, esh, Salomon A oll, Chagos A chipelago; B. T. ishelsoni, esh, S ai s o Ti an (no he n Red Sea /
Gul o Aqaba), Saudi A abia; C. T. ishelsoni, li e, Obs uc ion Ree (Thuwal, cen al Red Sea), Saudi A abia. Pho og aphs:
A. by R. Win e bo om; B, C. by V. N. Peinemann.
o e lapping dis ibu ion in he cen al o sou he n Red Sea and a e associa ed
wi h ca es. T imma be umeni can be dis inguished om T. ishelsoni by ha -
ing no elonga e do sal spines ( s second spine elonga e and ilamen ous in
92
ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
Yamada T, Sugiyama T, Tamaki N, Kawaki a A, Ka o M (2009) Adap i e adia ion o go-
bies in he in e s i ial habi a s o g a el beaches accompanied by body elonga ion
and excessi e e eb al segmen a ion. BMC E olu iona y Biology 9(1): 145. h ps://
doi.o g/10.1186/1471-2148-9-145
Supplemen a y ma e ial 1
COI-only axa included in he expanded 93-species phylogene ic da ase
Au ho s: Vik o N. Peinemann, Lucia Pombo-Ayo a, Wal e A. Rich, Michael D. Fox, Da en
J. Coke
Da a ype: xlsx
Explana ion no e: Fo each species, he able lis s he species name as used in he ee,
he BOLD P ocess ID, and whe e a ailable he co esponding GenBank accession
numbe , he name o he specimen iden i ie , and he collec ion locali y.
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License
(h p://openda acommons.o g/licenses/odbl/1.0/). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow use s o eely sha e, modi y, and
use his Da ase while main aining his same eedom o o he s, p o ided ha he
o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/zookeys.1255.159341.suppl1
Supplemen a y ma e ial 2
Specimens included in he mul i-locus da ase gene a ed o his s udy
and used in he phylogene ic analyses, wi h associa ed ouche numbe s,
collec ion locali ies, and GenBank accession numbe s o COI and 16S
Au ho s: Vik o N. Peinemann, Lucia Pombo-Ayo a, Wal e A. Rich, Michael D. Fox, Da en
J. Coke
Da a ype: xlsx
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License
(h p://openda acommons.o g/licenses/odbl/1.0/). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow use s o eely sha e, modi y, and
use his Da ase while main aining his same eedom o o he s, p o ided ha he
o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/zookeys.1255.159341.suppl2
93
ZooKeys 1255: 77–93 (2025), DOI: 10.3897/zookeys.1255.159341
Vik o N. Peinemann e al.: A new ca e-dwelling T imma om he Red Sea
Supplemen a y ma e ial 3
Pai wise gene ic dis ances (Kimu a 2-pa ame e ) be ween COI sequences
o T imma species om he Red Sea and Indo-Paci ic
Au ho s: Vik o N. Peinemann, Lucia Pombo-Ayo a, Wal e A. Rich, Michael D. Fox, Da en
J. Coke
Da a ype: xlsx
Explana ion no e: The new species, T. be umeni, is highligh ed in bold. Values shown as
pe cen ages. GenBank accession numbe s and collec ion locali ies a e p o ided o
each sequence. Cells a e colo -coded om yellow (high di e gence) o g een (low
di e gence) o aid isualiza ion.
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License
(h p://openda acommons.o g/licenses/odbl/1.0/). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow use s o eely sha e, modi y, and
use his Da ase while main aining his same eedom o o he s, p o ided ha he
o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/zookeys.1255.159341.suppl3
