Table Of Content2004. The Journal of Arachnology 32:22-30
BEHAVIORAL REPERTORY OF THE NEOTROPICAL
HARVESTMAN ILHAIA CUSPIDATA
(OPILIONES, GONYLEPTIDAE)
Wilton Pereira, Abner Elpino-Campos and Kleber Del-Claro: Laboratorio de
Ecologia Comportamental e de Interagoes (LECI), Instituto de Biologia, Universidade
Federal de Uberlandia, CP 593, 38400-902 Uberlandia, MG, Brazil
Glauco Machado1: Museu de Historia Natural, Instituto de Biologia, Universidade
Estadual de Campinas, CP 6109, 13083-970 Campinas, SP, Brazil
ABSTRACT. In this study, we provide an ethogram for the harvestman Ilhaia cuspidata and describe
the daily activity pattern of captive individuals. We also provide a comparison between the behavioral
repertory of this species with that of the syntopic Discocyrtus oliverioi. Five females and four males of
/. cuspidata were maintained in the same terrarium from November 1999-November 2000 for qualitative
and quantitative observations. Twenty behavioral acts were recorded, classified in seven categories and
the relative frequency of each was determined: exploration (69.8%), resting (16.7%), feeding (6.3%),
grooming (4.4%), social interactions (2.6%), reproduction (0.1%) and others (0.3%). There was a marked
difference in the frequency of the behavioral categories between sexes: females fed more frequently than
males and males were involved in social interactions more frequently than females. During most of the
daylight hours, individuals remained inside shelters and became active from 19:00-09:00 h. Although I.
cuspidata and D. oliverioi showed almost the same behavioral acts, there were quantitative differences in
their repertories: the relative frequency ofbehavioral categories “resting” and “social interactions” were
higher for /. cuspidata whereas “reproduction” and “grooming” were higher for D. oliverioi. The main
qualitative difference between these two species was related to the forms of parental care: females ofD.
oliverioi guard their eggs and first instarjuveniles, whereas females of I. cuspidata scatter their eggs in
time and space and do not actively protect their offspring. Since both species share the same habitats
(sometimes in multi-species aggregations), the behavioral differences between them may be explained by
particular morphological and physiological characteristics of the species, as well as by phylogenetic con-
straints.
Keywords: Activity pattern, defensive behavior, Discocyrtus ethogram, gregariousness, parental care
,
The Opiliones constitutes a highly diversi- maintained in captivity, where they behave in
fied order in terms of morphology and habits, a similarway to that observed in the field (e.g.
and includes approximately 5000 species Capocasale & Bruno-Trezza 1964; Elpino-
&
widespread throughout the world (Adis Campos et al. 2001; Willemart 2001), facili-
Harvey 2000). Harvestmen are particularly tating quantitative and qualitative compari-
suitable for behavioral studies because indi- sons between species.
viduals ofmany species are abundant and easy An ethogram is a set of comprehensive de-
to observe, both in the field and laboratory. scriptions ofthe behavioral repertory ofa spe-
Selected species may be good model organ- cies (Brown 1975). Allied with field obser-
isms for experimental manipulations since vations, ethograms are important starting
they allow researchers to test hypotheses points for any ethological research and for un-
about the ecological meaning of certain be- derstanding the biology and ecology ofa wide
havioral patterns (e.g. Mora 1990; Machado
& Oliveira 1998; Machado et al. 2002). Rep- rolainvgeerioofianHi.maSlosar(eLeshn1e9r45194(0)G.onDyilsecpotciydrateu,s
resentatives ofthe suborderLaniatores may be
Pachylinae) was the first harvestman to have
its behavioral repertory studied and is so far
1 Author for correspondence. the only species available for comparison (El-
22
.
PEREIRA ET AL.—BEHAVIORAL REPERTORY OF ILHAIA CUSPIDATA 23
pieo-Campos et aL 2001). This harvestman lating the natural lighting at the time of the
species showed 25 behavioral acts that were study. —
divided Into six major categories: exploration, Activity pattern. The activity scheduleof
resting, foraging, social interactions, self- L cuspidata was quantified by observing cap-
grooming, and reproduction. The individuals tive individuals at 1 h intervals during a 24 h
were inactive during the day and foraged and period. Sampling at each interval consisted of
Interacted with other Individuals at night. Fe- counting the number of individuals accom-
males reproduced in captivity and showed ma- plishing the different types of behavioral acts
ternal care toward the eggs and newly hatched during a 5 min period. These observations
offspring (Elpieo-Campos et aL 2001). were repeated on three non-consecutive days
Discocyrtus oliverioi is very abundant in in January 1999 totaling 72 h of sampling.
urban areas ofUberlaedia, state ofMinas Ger- The main aim of these samplings was to de-
ais, Brazil, where it occurs syntopically with termine the relative frequency ofthe majorbe-
two other harvestmen, Discocyrtus sp. and II- havioral categories throughout the day. At this
hoio cuspidata Roewer 1913 (Elpino-Campos point, we were not interested in determining
et ah 2001). In this paper, we provide a de- the biological rhythm of the species.
tailed ethogram for I. cuspidata (Gonylepti- Behavioral repertory and ethogram.—In
dae, Gonyleptinae) and describe the daily ac- order to identify and describe the behavioral
tivity pattern of captive individuals. Some acts of I. cuspidata, six hours of field obser-
comments on the foraging, defense, gregari- vations and another 6 h ofobservations In the
ouseess, and reproduction of this harvestman laboratory (20 sessions of 36 min ad libitum
are also presented for the first time. In addi- sampling, sensu Altmann 1974) were done at
tion, we have made qualitative and quantita- night when the animals were most active
tive comparisons between the behavioral rep- (18:00-00:00 h) in January 1999. The de-
ertories of L cuspidata and D. oliverioi scription of all behavioral acts and their clas-
sification into behavioral categories were as
METHODS described by Elpino-Campos et al. (2001). In
addition quantitative data only on captive in-
Collection of harvestmen and mainte-
— dividuals were obtained in 30 sessions of ob-
nance In the laboratory. Individuals of /. servations from November 1999-November
cuspidata were collected in the Experimental
2000. Each session consisted of scan samples
Garden of the Ueiversidade Federal de Uber-
m of all individuals performed at 50 min inter-
landia (18° 53'S, 48° 15'W; 863 ele.), an vals over 200 min. For each sampling all in-
urban area in the state ofMinas Gerais, south- dividuals were scanned twice. These obser-
eastern Brazil. The behavioral observations vations were made between 19:00 and 03:00
were based on nine individuals (5 females and h with a red lamp to avoid disturbing the an-
4 males) maintained in the same terrarium imals (cf. Machado & Oliveira 1998; Elpino-
(20.5 X 44.5 cm X 27.5 cm high) containing Campos et al. 2001). Ten minutes before each
soil, leaves and two pieces of tree fern trank behavioral sampling, 10 termite workers and
(8X10 cm each) as shelters. The shelters had three pieces of earthworm (1 cm each) were
the shape of a gabled roof, so that the indi- offered as food. The data recorded in these
viduals could slip under them, finding protec- samplings were used to construct the etho-
tion against light. Temperature and moisture gram. The absolute frequencies of the major
conditions in the laboratory were similar to behavioral categories were compared between
those in the field (nearly 25 °C and 70-80% sexes and between I. cuspidata and D. oliv-
relative humidity). In addition, nine individ- erioi using a G test (Sokal & Rohlf 1995). The
uals ofD. oliverioi and one ofDiscocyrtus sp. null hypothesis was of no difference between
were also placed in the same terrarium. The sexes and between species. Some specific be-
harvestmen were fed live termite workers (In- havioral acts (see results) were also compared
secta, Isoptera), freshly chopped pieces of using a chi square test done on the raw data
earthworms (Annelida, Oligochaeta), honey (Sokal & Rohlf 1995).
solution and an artificial diet for ants (Bhatkar Voucher specimens ofmales and females of
& Whitcomb 1970). A light schedule of 14L: I. cuspidata, D. oliverioi, and Discocyrtus sp.
10D was maintained in the laboratory, simu- were deposited in the Museu de Biodiversi-
n n
24 THE JOURNAL OF ARACHNOLOGY
D U 0
Resting E3 Exploration Feeding Grooming HU Others
0
Time (h)
—
Figure 1. Daily activity schedule of the harvestman Ilhaia cuspidata. The data are based on three
different days ofsampling and were obtainedfrom nine captive individuals. The moon and the sun indicate
dusk and dawn, respectively.
dade do Cerrado (MBC, Uberlandia, MG, outside the shelters sometimes during the day,
Brazil) and at the Museu de Zoologia da drinking water or walking—about (Fig. 1).
Universidade de Sao Paulo (MZSP, Sao Paulo, Behavioral repertory. Twenty behavior-
SP, Brazil). al acts, classified into seven major groups of
RESULTS activities, were recorded for I. cuspidata:
— feeding, social interactions, resting, explora-
Activity pattern and foraging. During tion, grooming, reproduction and others (Ta-
most of the daylight hours, individuals of Il- ble 1). Exploration was the most common ac-
haia cuspidata were found hidden inside shel- tivity, followed by resting and feeding (Table
ters (Fig. 1). Individuals became active with 1). Although field observations indicated that
the onset of darkness (about 18:00 h), when /. cuspidata is a gregarious harvestman (see
they left the shelters to forage (Fig. 1). They below), in captivity individuals rested alone
accepted both live (termites) and dead (pieces more frequently than in groups (males and fe-
of worm) animals, with the food item being males combined; x2 = 8.33; d.f. = 1: P <
grasped and taken to the shelter to be con- 0.01). Self-grooming activities, known as leg
sumed. Fights for food and aggressive inter- threading in harvestmen (cf. Edgar 1971), in-
actions were observed among conspecifics ( volved mainly legs II (males and females
= 7) and with Discocyrtus oliverioi (n = 3) combined; x2 = 12.43; d.f. = 3; P < 0.001)
in the shelters. Sometimes, two or more con- (Table 1). Grooming behavior occurred when
specifics were seen feeding simultaneously on the harvestmen were moving around the en-
the same piece offood ( = 3). Most individ- vironment and, more frequently, after feeding
uals of 1 cuspidata became inactive at about (when both legs and pedipalps were cleaned).
.
9:00 h, i.e., four hours after sunrise (Fig. 1). There was a marked difference in the fre-
Notwithstanding, individuals were recorded quency of behavioral categories between
1
PEREIRA ET AL.—BEHAVIORAL REPERTORY OF 1LHAIA CUSPIDATA 25
—
Table L Behavioral repertory and frequency ofeach behavioral act for nine captive individuals ofthe
harvestman Ilhaia cuspidata. n = number of individuals observed and, in parentheses, the number of
behavioral acts observed for each sex.
Frequency of behavioral acts (%)
Males {n = 4) Females (n = 5) Total (n - 9)
Behavioral acts (881) (1291) (2172)
Feeding 5.1 7.05 6.27
Feed on termites 1.82 1.94 1.89
Feed on pieces of worm 2.61 3.87 3.36
Feed on honey solution 0.23 0.31 0.28
Fight for food 0.45 0.93 0.74
Social interactions 4.32 1.47 2.61
Touching conspecific with legs 1.70 0.70 1.10
Touching individual of D. oliverioi with legs 0.68 0.23 0.41
Attacking conspecific 1.14 0.54 0.78
Attacking individual ofD. oliverioi 0.80 0 0.32
Resting 17.59 16.11 16.72
Alone 10.33 9.14 9.62
In multi-species aggregation 7.26 6.97 7.10
Exploration 68.10 70.73 69.65
Walkabout 18.50 19.60 19.15
Touching the substrate with first legs 16.80 17.82 17.40
Touching the substrate with second legs 32.80 33.31 33.10
Self-grooming 4.54 4.26 4.37
—
Leg threading—first pair 1.02 0.93 0.97
Leg threading—second pair 1.70 1.63 1.66
Leg threading—third pair 0.68 0.93 0.83
Leg threading fourth pair 0.80 0.54 0.64
Cleaning the pedipalps 0.34 0.23 0.27
Reproduction 0 0.15 0.11
Oviposition 0 0.15 0.11
Other 0.34 0.23 0.27
Pressing the body against the substrate 0.34 0.23 0.27
Total 100.0 100.0 100.0
males and females (G = 20.25; d.f. = 4; P < ulated, the harvestmen rarely released repug-
0.001). Doe to the low frequency of behav- nant substances or attacked the aggressor with
ioral acts the categories “reproduction” and the pedipalps.
“others” were not included in this analysis. Oviposition of I. cuspidata was observed
The frequency of feeding and exploration ac- twice (January and February) in the labora-
tivities was higher in females than in males, tory. The females repeatedly touched a given
whereas males were involved in social inter- point on the substrate with the first two pairs
actions more frequently than females (Table of legs before ovipositing. Subsequently, a
1). — single egg was laid and the female spent 5-
Notes on natural history. When dis- 10 s attaching debris and soil particles on the
turbed in the field, individuals of /. cuspidata egg surface with the first two pairs of legs.
remained motionless with the legs retracted Females abandoned the oviposition site, leav-
over the body. In this position, the darkbrown ing the egg unprotected. Another four eggs
coloration ofthe animals was extremely cryp- (similar in size and color) were found unpro-
tic against a darkbackground. Individuals also tected in the terrarium and were probably laid
showed thanatosis in which the legs werefully by females of /. cuspidata. Two eggs were
extended laterally and the animal became rig- covered by debris, while the other two had no
id for up to 5 min. Even when persistently particles attached.
mechanically disturbed with forceps ormanip- In the field, five multi-species aggregations
26 THE JOURNAL OF ARACHNOLOGY
—
| Male—s C] Females
—i—
100 —i -r-
1
90
Feeding Social Resting Exploration Grooming Reproduction
interactions
Behavioral categories
—
Figure 2. Comparison of the relative frequency of the behavioral categories observed for males and
females of the harvestmen llhaia cuspidcitci in captivity.
were found under fallen tree trunks (Fig. 3). time and space and did not actively protect
The mean number of individuals was 21.6 ± their offspring. The relative frequency of the
20.5 for /. cuspidata, 5.0 ± 2.2 for D. oliv- major behavioral categories differed statisti-
erioi and 2.8 ± 1.8 for Discocyrtus sp. Fe- cally between the two species (G = 86.29; d.f.
,
males were always most frequent within the = 6; P < 0.001). Individuals of /. cuspidata
aggregations, and accounted for55.6%-76.9% rested and interacted with other individuals
of the aggregated individuals of /. cuspidata. more frequently than D. oliverioi (Fig. 4). In
C—omparisons with Discocyrtus oliver- turn, the frequency of reproductive and clean-
ioi. There were both qualitative and quanti- ing activities were higher in D. oliverioi than
tative differences between the behavioral rep- in I. cuspidata (Fig. 4).
ertories of 7. cuspidata and D. oliverioi (Fig. DISCUSSION
4). These two harvestmen species showed the
same behavioral acts in the categories feeding, Although the subfamily Gonyleptinae is
social interactions, resting, exploration, and one of the largest among the Gonyleptidae
self-grooming (Table 1; Elpino-Campos et al. (Pinto-da-Rocha 1999), there are few studies
2001). In the category reproduction, however, describing the ecology and behavior ofits rep-
D. oliverioi showed a more diversified reper- resentatives (but see Machado & Pizo 2000;
tory, including seven acts, whereas /. cuspi- Machado & Vidal 2001). Some behavioral
tada showed only one (Table Elpino-Cam- patterns of llhaia cuspidata are very similar
1;
pos et al. 2001). In contrast to D. oliverioi to other gonyleptids, especially those of the
,
whose females laid a batch of eggs and re- subfamily Pachylinae (see Acosta et al. 1993,
mained over the eggs, protecting them for 1995; Capocasale & Bruno-Trezza 1964; El-
about 22 days (Elpino Campos et al. 2001), pino-Campos et al. 2001). Individuals of /.
females of /. cuspidata scattered their eggs in cuspidata are mainly nocturnal, generalist
PEREIRA ET AL.—BEHAVIORAL REPERTORY OF ILHAIA CUSPIDATA 27
—
Figure 3. Multi-species aggregation of harvestmen under a fallen trunk in the Experimental Garden
ofthe Universidade Federal de Uberlandia, in the state ofMinas Gerais, southeastern Brazil. MD = male
ofDiscocyrtus oliverior, FD = female of D. oliverioi; MI = male ofIlhaia cuspidata', FI = female of/.
cuspidata.
predators that carry their food to shelter be- fungi or other free-loading organisms before
fore feeding, and may be found in small ag- they have time to penetrate the cuticle, and to
gregations during the day, like many gonylep- restore sensory receptors to their full sensing
&
tids (see Capocasale Bruno-Trezza 1964; status (Edgar 1971).
Acostaet al. 1993, 1995; Gnaspini 1996; Hoe- Males and females of /. cuspidata differed
nen & Gnaspini 1999; Machado et al. 2000; in the relative frequency ofthe behavioral cat-
Elpino-Campos et al. 2001; Santos & Gnas- egories, with females feeding more frequently
pini 2002). The defensive behavior, however, than males. It is known that the respiratory
is more similar to that of cosmetids (Eisner et rate of female harvestmen increases when
al. 1978), manaosbiids (Cokendolpher 1987), they are producing and maturing eggs (Phil-
and trogulids (Hillyard & Sankey 1989), lipson 1962, 1963). This physiological change
which also feign death and rarely emit gland during egg development probably accounts for
secretions. the higher requirement for food (Phillipson
The most frequent behavioral act for the 1962, 1963; Gnaspini 1996). Indeed, in the
captive individuals of /. cuspidata was to cavernicolous harvestman Goniosoma spe-
touch the substrate with the second pair of laeum (Mello-Leitao 1923), females bearing
legs. In harvestmen these legs possess numer- eggs leave the cave to forage more frequently
ous sense organs and are used to touch the than other adult individuals in the population
substrate, food, and other organisms (Edgar (Gnaspini 1996). The month in which the
1963). During such activities, offensive organ- quantitative observations were conducted in
isms and debris come into contact with the our study (January) coincided with the period
sense organs. Thus, the main functions of leg when females of/. cuspidata were laying eggs
threading probably are to remove pathogenic in the laboratory. Thus, the higher frequency
28 THE JOURNAL OF ARACHNOLOGY
Discocyrtusoliverioi Ilhaiacuspidata
Feeding Social Resting Exploration Grooming Reproduction
interactions
Behavioral categories
—
Figure 4. Comparison of the relative frequency of the behavioral categories observed for the har-
vestmen Ilhaia cuspidata and Discocyrtus oliverioi in captivity.
of feeding activities in females compared to to rest alone more frequently than in groups
males may be related to the accumulation of in the laboratory.
energy for egg production and maturation. The main qualitative difference between
Males of/. cuspidata interacted aggressive- these two harvestmen species was related to
ly with conspecifics at a higher frequency than the forms ofparental care: females ofD. oliv-
females. Possible explanations for this behav- erioi guarded their eggs and first instar juve-
ioral pattern include intolerance among males, niles, whereas females of I. cuspidata scat-
competition for females, and territorial fights tered their eggs in the substrate and did not
(see Macias-Ordonez 1997). More details of show any additional interaction with the off-
the natural history of this species in the field spring. Since eggs of/. cuspidata and D. oliv-
are needed to assess the causes of aggressive erioi are laid in the same environment and
behavior in males. The aggressiveness among supposedly face similar selective pressures
males, however, may explain why females ac- (such as abiotic factors and predators), how
count for most of the aggregated individuals. can we explain the differences in the forms of
Under natural conditions, /. cuspidata and parental care?
D. oliverioi live in the same habitat, use the The evolution of the different forms of pa-
same microhabitats to forage, take shelter and rental care is a complex function of many in-
reproduce, and show a considerable overlap in teracting factors including morphology, defen-
their periods ofactivity (see also Elpino Cam- sive behavior, and phylogenetic constraints
pos et al. 2001). Our results, however, showed (see Tallamy & Wood, 1986). Harvestmen
that, at least in captivity, these species differ with a short ovipositor, like all Laniatores, are
in their allocation of time and energy to the unable to hide their eggs in deep cavities
behavioral categories studied. Individuals of/. where they are inaccessible to most predators.
cuspidata rest more and seem to be more ag- In these cases, the physiology of egg produc-
gressive than D. oliverioi, which may explain tion, life span, and defensive strategies may
why individuals of the former species tended offer a partial explanation on when subsocial
—
PEREIRA ET AL. BEHAVIORAL REPERTORY OF ILHAM CUSPIDATA 29
behavior may evolve (review in Machado & The defensive secretions of Pachyloidellus go-
Raimuedo 2001). Maternal care Is a viable liath (Opiliones: Laniatores: Gonyleptidae). Bon-
strategy only when females lay eggs aggre- ner Zoologischer Beitrage 44:19-31.
gated in the time and space, live long enough Adis, I. & M.S. Harvey. 2000. How many Arach-
to benefit one or more batches, and have de- nida and Myriapoda are there world-wide and in
Amazonia? Studies on Neotropical Fauna and
fensive strategies that enable them to protect
Environment 35:139-141.
the offspring against predators. Species in Altmann, I. 1974. Observational study ofbehavior:
which females do not fit these features (or are sampling methods. Behaviour, 49:227-265.
constrained by phylogenetic inertia) present Bhatkar, A. & W.H. Whitcomb. 1970. Artificial diet
alternative forms of parental care including for rearing various species of ants. Florida En-
egg hiding and/or egg covering. This is prob- tomologist 53:229-232.
ably the case of I. cuspidata that, despite liv- Brown, J.L. 1975. The Evolution ofBehavior. W.W.
ing more than one year as adult, may be phys- Norton, New York.
iologically constrained to iteroparity andrelies Canals, J. 1936. Observaciones biologicas en arac-
nidos del orden Opiliones. Revista Chilena de
mainly on evasive strategies of defense.
Historia Natural 40:61-63.
Egg hiding by females is probably the most &
Capocasale, R. L.B. Brano-Trezza. 1964. Biolo-
common form of parental investment among gia deAcanthopachylus aculeatus (Kirby, 1819),
harvestmen of the suborder Laeiatores (Ma- (Opiliones: Pachylinae). Revista de la Sociedad
chado & Raimundo 2001). Generally, non- Uruguaya de Entomologia 6:19-32.
subsocial harvestmen hide their eggs inside Cokendolpher, J.C. 1987. Observations on the de-
small natural crevices or cover them with de- fensive behaviors of a neotropical Gonyleptidae
bris and leave the offspring without additional (Arachnida: Opiliones). Revue Araehnologique
care (Canals 1936; Cokendolpher & Jones 7:59-63.
1991; Juberthie 1965, 1972; Willemart 2001). Cokendolpher, J.C. & S.R. Jones. 1991. Karyotype
Laying eggs in several batches or scattering and notes on the male reproductive system and
single or few eggs over a very wide area may natural history of the harvestman Vonones sayi
(Simon) (Opiliones: Cosmetidae). Proceedingsof
be advantageous and confer protection to the
theEntomological Society ofWashington93:86-
offspring because detection by natural ene-
91.
mies would be reduced (Edmunds 1974; Wil- Edgar, A.L. 1963. Proprioception in the legs of
lemart 2001). phalangids. Biological Bulletin, Woods Hole
ACKNOWLEDGMENTS 124:262-267.
Edgar, A.L. 1971. Studies on the biology and ecol-
The authors thank R. Pinto-da-Rocha and ogy of Michigan Phalangida (Opiliones). Mis-
E. G. Vasconcelos forIdentification ofthe har- cellaneous Publications, Museum of Zoology,
vestmen species. Drs. P. Gnaspiei, J.C. Cok- University of Michigan 144:1-64.
eedolpher, R. Macias-Ordonez, and T. Novak, Edmunds, M. 1974. Defence in Animals: A Survey
and two anonymous reviewers provided help- of Antipredator Defences. Longman, New York.
ful comments on an early draft. We are es- Eisner, T, D. Alsop & J. Meinwald. 1978. Secre-
tions of opilionids, whip scorpions and pseudo-
pecially indebted to Dr. Gail Stratton forhelp-
scorpions. Pp. 87-99. In Handbook of Experi-
ful suggestions and careful editing of the
mental Pharmacology. Yol. 48 (S. Bettini, ed).
manuscript. The authors were supported by
Springer-Veriag, Berlin.
fellowships from CNPq (to AEC and KDC), Elpino-Campos, A., W. Pereira; K. Del-Claro & G.
Fapemig (to KDC), and FAPESP (to GM; no. Machado. 2001. Behavioral repertory and notes
02/00381-0). WP and AEC are grateful forthe on natural history ofthe neotropical harvestman
logistical support provided by the Instituto de Discocyrtus oliveriot (Opiliones: Gonyleptidae).
Riocieecias da Ueiversidade Federal de Uber- Bulletin of the British Arachnological Society
land!a 12:44-150.
Gnaspini, P. 1996. Population ecology of Gonioso-
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30 THE JOURNAL OF ARACHNOLOGY
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