Larva and pupa of Amyna axis (Guenee, 1852) and affirmation of its taxonomic placement in Bagisarinae (Lepidoptera, Noctuidae)

Abstract

The larva and pupa of Amyna axis (Guenee, 1852) are described and illustrated, and observations are provided on the insect’s life history and larval biology. Larval, adult, and life history characters support the transfer of Amyna Guenee from Acontiinae Guenee, 1841 to Bagisarinae Crumb, 1956. The phylogenetic placement of the Bagisarinae is enigmatic; some adult and larval features indicate that the subfamily is a basal trifid proximate to Acontiinae, whereas other larval and life history characters suggest an association with Scoliopteryginae, a basal quadrifid group. Larvae exhibit a green-to-black color polyphenism presumably linked to larval density, with darker phenotypes occurring during outbreak densities. Parallel color shifts that occur in other Lepidoptera are briefly discussed.

Full text

A new species of Odaginiceps Fiers, 1995 (Copepoda, Harpacticoida, Tetragonicipitidae)... 1
A new species of Odaginiceps Fiers, 1995
(Copepoda, Harpacticoida, Tetragonicipitidae)
from the Mediterranean coast of Turkey
Süphan Karaytuğ1,†, Serdar Sak2,‡, Alp Alper2,§
1 Mersin University, Faculty of Arts and Science, Department of Biology, 33343, Mersin, Turkey 2 Balıkesir
University, Faculty of Arts and Science, Department of Biology, 10145, Balıkesir, Turkey
† urn:lsid:zoobank.org:author:D8DB71BC-2009-42DA-9C9A-81E6996AACF5
‡ urn:lsid:zoobank.org:author:96F0B01E-E714-42F7-A19B-304D9F0C6426
§ urn:lsid:zoobank.org:author:4B6E6F46-CA51-46FB-967E-E17A280A4589
Corresponding author : Süphan Karaytuğ ( suphankaraytug@gmail.com )
Academic editor: Danielle Defaye|Received 19 January 2010|Accepted 22 July 2010|Published 27 August2010
urn:lsid:zoobank.org:pub:7FD0C11F-0732-41AD-B48A-C3D8979E167D
Citation: Karaytuğ S, Sak S, Alper A (2010) A new species of Odaginiceps Fiers, 1995 (Copepoda, Harpacticoida,
Tetragonicipitidae) from the Mediterranean coast of Turkey. 53 : 1 – 12 . doi: 10.3897/zookeys.53.389
Abstract
Male and female of Odaginiceps korykosensis sp. n. (Copepoda, Harpacticoida, Tetragonicipitidae), collect-
ed in the intertidal zone of Kızkalesi beach along the Mediterranean coast of Turkey (Mersin Province), are
described.  e new species is the  fth member of the genus and can easily be distinguished from the other
species by the presence of four setae/spines on the second endopodal segment of P4 and by the structure of
the caudal rami. Previously, representatives of the genus Odaginiceps have been reported from Gulf of Mex-
ico, o Bermuda and Kenya. O. korykosensis sp. n. is the  rst record of the genus in the Mediterranean Sea.
Keywords
Harpact icoida, Tetragonicipitidae, Odaginiceps, taxonomy, new species
Introduction
 e genus Odaginiceps Fiers, 1995 is one of the 12 genera currently recognized in the
family Tetragonicipitidae and now comprises four species (Wells 2007).  e genus was
 rst created by Fiers (1995) in one of his comprehensive and excellent papers on the
ZooKeys 53: 1–12 (2010)
doi: 10.3897/zookeys.53.389
www.pensoftonline.net/zookeys
Copyright S. Karaytug, S. Sak, A. Alper. 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
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Süphan Karaytuğ, Serdar Sak & Alp Alper / ZooKeys 53: 1–12 (2010)
2
Tetragonicipitidae from the Yucatecan continental shelf (Mexico). In this paper, Fiers
(1995) described O. clarkae Fiers, 1995 from the West Central Atlantic (Quintana Roo
State, Nichupte Lagoon, Cancun, Mexico) and designated it as the type species of the
genus. In the same paper Fiers (1995) also described O. xamaneki Fiers, 1995 from
the Western Central Atlantic (western region of Yucatecan continental shelf, Mexico).
 e third species (O. elegantissima Fiers, 1995) was also described and allocated to the
genus by Fiers (1995) after reexamining the material previously identi ed by Coull
(1970) as Diagoniceps laevis Willey, 1930 from Castle Harbour (Bermuda). Fiers and
De Troch (2000) later described the fourth species, O. immanis Fiers and De Troch,
2000, from the Indo-Paci c (Gazi Bay, Kenya). No other report of Odaginiceps has ap-
peared in the literature since then. But the intensive investigation (carried out between
2000–2010) of over 500 phytal and interstitial harpacticoid samples taken from nearly
200 di erent stations along the mediolittoral zone of rocky shores and sandy beaches
of almost all Turkish coasts (unpublished data) revealed a new species of Odaginiceps
which was found only in a single locality. Both sexes of this new species were described
in detail below.
Material and methods
Samples were collected using the Karaman-Chappuis method (Delamare Deboutte-
ville 1953) from the type locality on three di erent dates (Table 1) but only one male
and one female were obtained at the  rst sampling (April 09, 2007). Prior to dissec-
tion, the habitus was drawn from whole specimens temporarily mounted in lactophe-
nol. Specimens were dissected in lactic acid and the dissected parts were mounted in
lactophenol mounting medium. Broken glass  bres were added to prevent the animal
and appendages from being compressed by the coverslip and to facilitate rotation and
manipulation, allowing observation from all angles. Preparations were subsequently
sealed with Entellan® (Merck). All drawings were prepared using a camera lucida on
Olympus BX-51 di erential interference contrast microscope. Total body length was
measured from the anterior margin of the rostrum to the posterior margin of the
caudal rami. Measurements were made with an ocular micrometer. Scale bars in il-
lustrations are in μm.  e descriptive terminology is adopted from Huys et al. (1996).
Abbreviations used in the text are: ae, aesthetasc; P1–P6, for swimming legs 1–6; exp
Date 09 April 2007 27 July 2007 26 November 2007
pH 8,16 7,29 8,14
Temperature (°C) 18,3 29,8 19,1
Conductivity (ms) 45,7 57,1 54,0
Salinity (ppt) 34,5 37,8 35,5
Oxygen (mg/L) 2,78 1,11 1,71
Table 1. Some physical and chemical parameters of interstitial water on di erent sampling dates at the
type locality.

A new species of Odaginiceps Fiers, 1995 (Copepoda, Harpacticoida, Tetragonicipitidae)... 3
(enp)-1 (-2, -3) to denote the proximal (middle, distal) segment of a ramus. Material
was deposited in the Mersin University Zoology Museum (MUZM) at Mersin, Turkey.
Physical and chemical parameters of the interstitial water in the sampling pit are sum-
marized in Table 1. Parameters were measured with an YSI 85 Handheld Dissolved
Oxygen and Conductivity Instrument (YSI Inc.), with the exception of the pH which
was measured with an Orion 3-star ( ermo Fisher Scienti c Inc.) Portable pH Meter.
Results
Order Harpacticoida Sars, 1903
Family Tetragonicipitidae Lang, 1944
Genus Odaginiceps Fiers, 1995
Odaginiceps korykosensis sp. n.
urn:lsid:zoobank.org:act:8EF0600A-D85B-4032-9661-26A514C0ADDD
Figs 1–6
Type locality . Turkey, Mediterranean coast, Mersin Province; intertidal zone of
Kızkalesi beach (36°27.473'N, 34°08.647'E).  e type locality is  ne sand beach.
Material examined : Holotype ♀ dissected on seven slides. Allotype ♂ dissected
on six slides. Legs. S. Karaytuğ, S. Sak, A. Alper and S. Sönmez.
Description . Female (Fig. 1A, B). Total body length 770 μm, with largest width
measured at cephalothorax. Integument of cephalic shield smooth, of all other somites
ornamented with irregular pattern of hardly visible spinules. Body surface with sensilla
pattern as  gured. Posterior margin of the body somites with serrate hyaline frills.
Rostrum (Fig. 1A) large, widest at base, extending halfway along second anten-
nular segment; with two delicate sensillae and a mid-dorsal pore.
Urosome (Figs 1A,B) 5-segmented, comprising P5-bearing somite, genital dou-
ble somite and three free abdominal somites. Genital double-somite longer than
wide; with transverse surface ridge dorsally and laterally (Fig. 1B) extending ventrally
(Figs1A; 2A), indicating original segmentation. Genital  eld (Fig. 2A) with small
copulatory pore located in median triangular depression. A spermatophore attached to
the copulatory pore. First and second somite of genital double-somite and second and
third abdominal somites with continuous spinules near distal margin dorsally (spinules
of  rst somite of genital double-somite interrupted midway) (Fig. 1A); genital double
somite and second abdominal somite with spinular row near distal margin ventrally,
interrupted midway; third abdominal somite with continuous spinular row near distal
margin ventrally (Fig. 2A). Anal somite (Fig. 1A) with distal spinular row extending
dorsally to the either side of anal operculum; operculum smooth, slightly convex.
Caudal rami (Figs 1C; 3D, E) tapering posteriorly with 4–5 dorsal spinules distally
near the base of seta V; 1.7 times longer than wide; inner margin ornamented with spi-
nules (Fig. 3D); with a pore on proximal third of dorsal surface (Fig. 3D), another pore

Süphan Karaytuğ, Serdar Sak & Alp Alper / ZooKeys 53: 1–12 (2010)
4
Figure 1. Odaginiceps korykosensis sp. n. Female. A habitus, dorsal B habitus, lateral C right caudal ra-
mus with terminal complement, dorsal. Male. D P2 endopod, anterior E P3 terminal endopod segment,
anterior F P4 endopod, anterior.
present on ventral surface near the base of seta III (Fig. 2A); with seven setae (Fig. 3D,
E), seta I minute located near the base of seta II; setae II–III bare; setae IV–V strongly
developed and bipinnate; seta V with swollen sinuate base; seta VI short and as long as
seta III; seta VII tri-articulated.
Antennule (Fig. 3A) 9-segmented; segment 1 with a long plumose seta at antero-
distal margin, a spinular row on anterior surface, long spinules along inner margin,
small tube-pore on dorsal surface near inner margin. Segment 2 with long spinules
on caudal margin. Segment 4 with long aesthetasc fused basally to seta. Segment 9
ADB
E
C
F
A–C 100
50
D–F

A new species of Odaginiceps Fiers, 1995 (Copepoda, Harpacticoida, Tetragonicipitidae)... 5
longest, bears an apical acrothek consisting of a short aesthetasc and two setae. Arma-
ture formula 1-[1 plumose], 2-[6+3 plumose], 3-[6+1 plumose], 4-[3+(1+ae)], 5-[2],
6-[2+2 plumose], 7-[2], 8-[1+1 plumose], 9-[5+acrothek].
Antenna (Fig. 3B). Coxa small and smooth. Basis with 2 long spinules at outer
margin. Exopod 1-segmented with one lateral pinnate seta, apical armature consists
of one pinnate seta and one pinnate spine; a few spinules present around outer distal
corner and midway along outer margin. Endopod 2-segmented;  rst endopod segment
with one plumose seta at proximal third of outer margin. Distal endopod segment with
various spinular rows as  gured and with two abexopodal unipinnate spines laterally
(both spines with subapical tubular extension). Apical armature of enp-2 consisting of
Figure 2. Odaginiceps korykosensis sp. n. A female, abdomen, ventral B male, abdomen, ventral.
A
B
A
B50
50

Süphan Karaytuğ, Serdar Sak & Alp Alper / ZooKeys 53: 1–12 (2010)
6
two pinnate setae, and  ve geniculate setae; longest geniculate seta with large spinules
and fused at base to long pinnate seta.
Labrum (Fig. 4D). Free margin straight, with spinular row at distal corners and
 ne spinular row subdistally on ventral surface.
Mandible (Fig. 4A, B). Coxa robust, gnathobase with one pinnate seta at dorsal
corner and several blunt multicuspidate teeth along distal margin. Palp biramous; basis
strong, with three plumose setae along inner margin, ornamented with a group of long
spinules proximally. Exopod 3-segmented,  rst segment with two plumose setae, sec-
Figure 3. Odaginiceps korykosensis sp. n. Female. A antennule, dorsal B antenna C P5, anterior D left
caudal ramus, dorsal E right caudal ramus, lateral.
A
B
C
E
D
I
II
II
III
III IV
IV
VI
VI V
V
VII
VII
I
A, B
C
D, E
20
50
20

A new species of Odaginiceps Fiers, 1995 (Copepoda, Harpacticoida, Tetragonicipitidae)... 7
ond segment with two bare setae and third segment with two bare setae fused at base.
Endopod 1-segmented with two lateral and six distal bare setae (two inner distal setae
and two outer distal setae fused at base).
Maxillule (Fig. 4E). Praecoxal arthrite with seven spines around distal margin, with
spinules as  gured; anterior surface with two bare setae; posterior surface with three plumose
and one pinnate setae. Coxal endite with two smooth and four plumose setae. Basis with
seven bare and one unipinnate setae. Long endopod segment and square exopod segment
each with three plumose setae. Endopod and exopod with a row of  ne marginal setules.
Figure 4. Odaginiceps korykosensis sp. n. Female. A mandible B distal margin of gnathobase C maxilla,
posterior D labrum, ventral E maxillule, anterior F maxilliped, posterior.
A
B
C
E
F
D
A–F
20

Süphan Karaytuğ, Serdar Sak & Alp Alper / ZooKeys 53: 1–12 (2010)
8
Maxilla (Fig. 4C). Syncoxa ornamented with spinules as  gured; with three en-
dites. Proximal endite with one plumose and three unipinnate setae; middle endite
with three unipinnate setae; distal endite with two unipinnate and one plumose setae.
Allobasis drawn out into pinnate claw; accessory armature consisting of two bare setae
and one curved spine. Endopod 2-segmented; proximal segment with one bare seta,
distal segment with one geniculate and two bare setae.
Maxilliped (Fig. 4F). Subchelate and ornamented with spinular rows as  gured.
Syncoxa with three inner bare setae subdistally. Basis with two bare setae along inner
margin. Endopod with one small accessory seta, one bare and two plumose setae.
Figure 5. Odaginiceps korykosensis sp. n. Female. A P1, dorsal B P2, anterior C P3, anterior D P4,
anterior.
B
C
D
A–D 50

A new species of Odaginiceps Fiers, 1995 (Copepoda, Harpacticoida, Tetragonicipitidae)... 9
P1 (Fig. 5A). Intercoxal sclerite rectangular and smooth. Small praecoxa triangu-
lar and bare. Coxa with complex spinular ornamentation anteriorly as  gured. Basis
narrower than coxa; anterior surface with pore near the base of outer bipinnate spine;
inner side with long slender spinules. Exopod 3-segmented, segments with spinular
rows along inner and outer margins. Endopod 2-segmented; enp-1 reaching almost
middle of exp-3, with spinular row along inner and outer margins; long inner seta
plumose and located subdistally; enp-2 slightly shorter than enp-1, with spinules
along inner and outer margins and with two long articulated setae and one small
inner apical seta.
Figure 6. Odaginiceps korykosensis sp. n. Male. A habitus, dorsal B P5, anterior C rostrum and anten-
nule, dorsal D terminal segments of antennule.
A
B
C
D
A
B
C, D
100
20
20

Süphan Karaytuğ, Serdar Sak & Alp Alper / ZooKeys 53: 1–12 (2010)
10
P2–P4 (Fig. 5B–D). Intercoxal sclerite unornamented. Coxa and basis with com-
plex spinular ornamentation as  gured. Exopod 3-segmented. Endopod 2-segmented.
Endopodal and exopodal segments with spinular rows along inner and outer margins.
Exp-1 of P2-P3 without spinules along inner margin. Enp-1 of P2 without inner seta.
Terminal outer seta of P2 enp-2 bare. P4 exp-2 with one plumose inner seta. With a pore
on anterior surface of enp-2 and anterior surface of exp-2 and -3. Exp-1 and -2 (and -3
in P4) with a posterior spinule patches. Enp-2 with a posterior spinular row subdistally.
Armature formula of swimming legs:
P1 P2 P3 P4
Exp. Enp. Exp. Enp. Exp. Enp. Exp. Enp.
0.0.022 1.120 0.0.023 0.021 0.0.023 1.021 0.1.322 (♀)
0.1.222 (♂)1.121
P5 (Fig. 3C). Baseoendopod and exopod covered with  ne spinules on anterior
surface, with long slender spinules along inner and outer margins. Exopod 3.6 times
longer than wide, with 6 setae; seta 1 longest and plumose; seta 3 smallest and bare.
Baseoendopod longer than wide; with 2 unipinnate and 3 plumose setae.
P6 (Fig. 2A) represented by a small segment with one outer plumose seta and two
slender bare setae.
Description . Male (Fig. 6A). Total body length 510 μm. Body smaller and more
slender than female, largest width measured at midway of cephalothorax. Body orna-
mentation generally as in female. Sexual dimorphism observed in antennule, P2-P6
and genital segmentation.
Antennule (Fig. 6C, D) indistinctly 10-segmented, sub-chirocer. Segment 1 short,
with small tube-pore on dorsal surface and with long spinules along caudal margin.
Segment 2 longest. Segment 4 with partial suture line dorsally. Segment 5 with long
aesthetasc fused basally to seta. Segment 10 bears an apical acrothek consisting of
a short aesthetasc and two slender setae. Armature formula 1-[1 plumose], 2-[7+3
plumose], 3-[4], 4-[4+1 plumose], 5-[4+1 spine+(1+ae)], 6-[1+2 spines], 7-[2], 8-[1],
9-[2], 10-[5+acrothek].
P2 enp-2 (Fig. 1D); outer terminal spine more robust than female; middle ter-
minal seta bare, shorter than female and as long as outer spine; inner terminal seta
minute. P3 enp-2 (Fig. 1E); inner terminal seta modi ed to a short spine (arrowed
in  g. 1E). P4 exp-3 (Fig. 1F) with 6 setae, inner terminal seta of female (arrowed in
 g.5D) absent in the male.
P5 biramous (Fig. 6B), fused medially. Baseoendopod ornamented with patch of
spinules as  gured; with two pores (one near the base of outer basal seta and the other
near the base of inner terminal spine of endopodal lobe); endopodal lobe with one
lateral and two distal spines. Exopod with three outer bare setae, 1 terminal bare seta
and two inner unipinnate setae; with two pores (one tube pore near the base of outer
proximal bare seta and the other near the base of outer median bare seta). P6 vestiges
asymmetrical (Fig. 2B); each P6 with one plumose inner seta and two long bare setae.

A new species of Odaginiceps Fiers, 1995 (Copepoda, Harpacticoida, Tetragonicipitidae)... 11
Etymology .  e speci c name refers to “korykos” which is the historical name of
Kızkalesi province (Mersin, TURKEY).
Discussion
 e new species can be attributable to the genus Odaginiceps by the absence of in-
ner seta on the proximal segments of P2-P4 exopod and P2 endopod, the short sec-
ond antennulary segment, the large prominent rostrum, the presence of pinnate setae
on the second and third antennulary segments, and the two-segmented P1 endopod
with three armature elements on the second segment (Fiers 1995; Fiers and De Troch
2000).  us far, four species have been assigned to the genus Odaginiceps: O. clarkae,
O. xamaneki, O. elegantissima and O. immanis.  e new species can easily be distin-
guished from its congeners by the presence of four setae/spines on the second endopo-
dal segment of P4, by the shape of the caudal rami and by the occurrence of four setae
on the proximal endite of maxilla.  e new species is most closely related to the O. xa-
maneki. Both species di er from the 3 other congenerics by the presence of short, and
compact caudal rami (long and semi-cylindrical in the other species) and the spinular
posterodorsal ornamentation of the urosomites (spinular rows present only ventrally
in the 3 other species). O. korykosensis sp. n. (female) di ers from O. xamaneki by the
presence of spinules (interrupted midway) along the posterioventral margin of genital
double-somite, by the absence of a central spinular patch on ventral surface of second
abdominal somite, by the longer outermost seta of P5 baseoendopod, by the much
longer terminal plumose setae of P2-P3 enp-2, by the longer P3 enp-2, as well as other
minor di erences observed on the spinular ornamentations of various appendages.
Fiers (1995) assumed that the presence of an inner pectinate element on the P3
enp-1, the presence of an inner seta on P4 exp-2, and the distinctly shorter caudal
rami in both sexes could be su cient grounds to erect a new genus to accommodate
O. xamaneki.  e above mentioned potential generic characters for O. xamaneki are
also observed in O. korykosensis which supports the previously formulated assumption.
However, until the speci c and generic importance of these characteristics are better
understood and evaluated, both species are retained in the genus Odaginiceps.
 e new species lacks a seta (arrowed in the Fig. 5D) on the third exopodal seg-
ment of the P4 in the male.  e absence of this seta in the male supports the assump-
tion of Fiers (1995) that almost all tetragonicipitid males bear one seta less on the third
exopodal segment of the P4.
Note on the ecology and distribution. Examination of the extensive interstitial
samples taken from almost all sandy beaches along the Turkish coasts revealed that the
new species occurs at the type locality only. On the other hand three samples taken
from the type locality in di erent seasons revealed no true interstitial forms.  e ab-
sence of true interstitial forms but the presence of O. korykosensis sp. n. in the type
locality can be explained by the very low oxygen levels measured at the type locality

Süphan Karaytuğ, Serdar Sak & Alp Alper / ZooKeys 53: 1–12 (2010)
12
(Table 1). Most harpacticoids are sensitive to reduced oxygen supply, which restricts
their occurrence to the upper sediment layers and favours epibenthic life. It can be
discerned from the general body shape of Odaginiceps spp. (see Fig. 1A, B; Fiers 1995;
Fiers and De Troch 2000) that they are not truly interstitial, but are probably epi-
benthic forms crawling on/in the upper surface of sediment along the shallow/deeper
waters, meaning that two specimens of O. korykosensis sp. n. accidentally entered into
the interstitial sample.  e e ect of the oxygen supply on the horizontal composition
of harpacticoid species in the sediment can be supported by the presence of several in-
terstitial forms (such as Arenosetella Wilson, 1932) found in 95 % of stations sampled
along the Mediterranean coast of Turkey (unpublished data).  e oxygen levels in the
sediments in these localities were much higher than those observed at the locality of
Odaginiceps korykosensis sp. n.
Acknowledgements
 is study was funded by TÜBİTAK under the project number 106T590. We also
would like to thank Serdar Sönmez for his help in collecting the material.
References
Coull BC (1970) Shallow water meiobenthos of the Bermuda Platform. Oceologia (Berlin) 4:
325–357.
Delamare Deboutteville C (1953) Recherches sur l’écologie et la répartition du mystacocaride
Derocheilocaris remanei Delamare et Chappuis, en Méditerranée. Vie et Milieu 4: 321–380.
Fiers F (1995) New Tetragonicipitidae (Copepoda, Harpacticoida) from the Yucatecan con-
tinental shelf (Mexico), including a revision of the genus Diagoniceps Willey. Bulletin de
l’Institut Royal des Sciences naturelles de Belgique Biologie 65: 151–236.
Fiers F, De Troch M (2000) New Tetragonicipitidae (Copepoda, Harpacticoida) from the Indo-
Paci c. Hydrobiologia 434: 97–144.
Huys R, Gee JM, Moore CG, Hamond R (1996) Marine and Brackish Water Harpacticoid
Copepods. Part 1. In: Barnes RSK, Crothers JH (Eds) Synopses of the British Fauna (New
Series) Field Studies Council, Shrewsbury 51: i-viii, 1–352.
Wells JBJ (2007) An annotated checklist and keys to the species of Copepoda Harpacticoida
(Crustacea). Zootaxa 1568: 1–872.