Tree Use by Harvestmen

Download as pdf or txt
Download as pdf or txt
You are on page 1of 6

Tree Use by Harvestmen (Arachnida: Opiliones) in the

Rainforests of Trinidad, W. I.
Author(s): Jessica A. Burns, Rebecca K. Hunter and Victor R. Townsend, Jr.
Source: Caribbean Journal of Science, 43(1):138-142.
Published By: University of Puerto Rico at Mayagüez
DOI: http://dx.doi.org/10.18475/cjos.v43i1.a13
URL: http://www.bioone.org/doi/full/10.18475/cjos.v43i1.a13

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the


biological, ecological, and environmental sciences. BioOne provides a sustainable online
platform for over 170 journals and books published by nonprofit societies, associations,
museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content
indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/
terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercial
use. Commercial inquiries or rights and permissions requests should be directed to the
individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers,
academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research.
NOTES

Caribbean Journal of Science, Vol. 43, No. 1, 138-142, 2007 development in the soil or leaf litter (Edgar
Copyright 2007 College of Arts and Sciences
University of Puerto Rico, Mayagüez 1971). After the final molt, adults of many
species may undertake vertical migrations
Tree Use by Harvestmen into shrubs or trees in search of food or
mates (Bristowe 1949; Todd 1949; Williams
(Arachnida: Opiliones) in the 1962; Edgar and Yuan 1968; Edgar 1971;
Rainforests of Trinidad, W. I. Goodnight and Goodnight 1976; Adams
1984; Morse 2001). Individuals have been
J ESSICA A. B URNS , R EBECCA K. H UNTER , observed to climb to considerable heights,
AND V ICTOR R. T OWNSEND , J R . Virginia reaching the forest canopy in several in-
Wesleyan College, 1584 Wesleyan Drive, stances (Edgar 1971; Gunnarson and Hake
Norfolk, Virginia 23502 USA 1999). Adult harvestmen may spend weeks
Email: [email protected] or months on the trunks or branches of
trees (Todd 1949; Edgar 1971). In some spe-
ABSTRACT.—Relatively little is known about the cies, individuals may undertake daily ver-
microhabitat preferences of most species of Neo- tical migrations, ascending trees under low
tropical harvestmen. We investigated the use of light conditions and returning to caves be-
trunks, buttresses, and the leaf litter in the immedi- fore dawn (Gnaspini 1996; Machado et al.
ate vicinity of trees by multiple species of harvest- 2000). Use of arboreal microhabitats may
men in a Trinidad, W. I. rainforest. A total of 238 represent selection for favorable microcli-
individuals were collected. This included adults of
mates as these realms generally provide
species from the families Cosmetidae, Manaosbi-
idae, Sclerosomatidae, and Stygnidae. Our results in- shelter from wind, heavy rainfall, and di-
dicate that the cosmetids (2 species), especially rect, extended exposure to sunlight (Todd
Cynortula sp., were the most abundant species to 1949; Cockerill 1988; Machado and Vascon-
occur on the trees. We also found significant, posi- celos 1998; Machado et al. 2000).
tive correlations for several species with regard to In contrast to temperate species, rela-
tree size and the number of individuals present. We tively little is known about the ecology or
hypothesize that harvestmen may use the surfaces of natural history of most tropical harvestmen
trees as well as the leaf litter in the immediate vicin-
species. In particular, there is a general
ity of the buttresses as either shelters from potential
predators or as areas that provide favorable micro- paucity of knowledge concerning the biol-
climates (e.g., relatively high humidity). ogy of Caribbean species, especially those
occurring on the island of Trinidad (Cok-
KEYWORDS.—Cosmetidae, ecology, Manaosbiidae, endolpher and Camilo-Rivera 1989). The
natural history, Sclerosomatidae, Stygnidae vast majority of field and laboratory stud-
ies of Neotropical harvestmen have focused
The behavior, ecology, and natural his- primarily on Brazilian species from or near
tory of temperate species of harvestmen the Mata Atlantic forest (Goodnight and
(Arachnida: Opiliones) have been investi- Goodnight 1976; Gnaspini 1996; Machado
gated in a number of studies (Bristowe and Vasconcelos 1998; Machado and Pizo
1949; Todd 1949; McAlister 1962; Williams 2000; Machado et al. 2000; Mestre and Pinto
1962; Edgar and Yuan 1968; Edgar 1971; da Rocha 2004; Pereira et al. 2004). These
Fowler and Goodnight 1974; Adams 1984). studies have revealed that tropical harvest-
Most species of harvestmen are considered men prefer humid microhabitats (e.g., leaf
to be primarily nocturnal, generalist preda- litter and caves) and adults are generally
tors/scavengers that feed primarily upon nocturnally active with regard to feeding,
worms (oligochaetes), insects, other har- mating, and movement (Goodnight and
vestmen, and occasionally vegetation, par- Goodnight 1976; Gnaspini 1996; Machado
ticularly flowers and fruits (Halaj and Cady et al. 2000). In Trinidad, anecdotal observa-
2000; Machado and Pizo 2000). In most spe- tions have been reported that indicate
cies of temperate regions, individuals un- adults of several species (most notably spe-
dergo at least some period of ontogenetic cies from the families Agoristenidae and
138
NOTES 139

Cosmetidae) are active in the leaf litter dur- vations of harvestmen behavior prior to
ing the day and at night (Kury and Pinto- collection.
da-Rocha 2002). Field studies of tropical This study was conducted from 12-15
harvestmen have also demonstrated that July 2005 in the forests adjacent to the
individuals tend to occupy spaces beneath beaches of Petite Tacarib and Gran Tacarib
rocks and within or under logs as well as along the northern coast of Trinidad (10ºN
the leaf litter and the trunks of trees (Good- 47⬘38.8⬙, 61ºW 13⬘32.7⬙; datum: WGS84).
night and Goodnight 1947, 1976; Gnaspini Between 07:30 and 12:30 hrs, the external
1996; Machado and Vasconcelos 1998; surfaces of the trunks and buttresses of 30
Machado and Pizo 2000; Machado et al. randomly selected trees were examined.
2000; Mestre and Pinto-da-Rocha 2004; For each tree, diameter at breast height
Pereira et al. 2004). (DBH) was measured using a tree diameter
In Trinidad, 24 described species of har- tape. Individual harvestmen were collected
vestmen are known to occur including taxa by hand from ground level to a height of
representing two suborders and eight approximately 2 m. Harvestmen were im-
families (Goodnight and Goodnight 1947; mediately preserved in 10% buffered for-
Cokendolpher and Camilo-Rivera 1989; malin and later examined to determine sex
Pinto-da-Rocha and Kury 2003; Kury 2003). and maturity status (juvenile or adult). In
Of these species, only Ethobunus tubercula- addition to the surfaces of the tree, the leaf
tus (Zalmoxidae: Goodnight & Goodnight litter inside the buttresses or cavities within
1947), Paecilaema inglei (Cosmetidae: Good- the trunk was also thoroughly searched by
night & Goodnight 1947), Phareicranaus hand. This sampling technique for leaf lit-
calcariferus (Cranaidae: Simon 1879), Priono- ter is effective for harvestmen with rela-
stemma insulare (Sclerosomatidae: Roewer tively large body size (i.e., cosmetids, cra-
1953), Prionostemma vittatum (Sclerosomati- naids, manaosbiids, sclersomatids, and
dae: Roewer 1910), Rhopalocranaus albilineatus stygnids). In addition, it is also considered
(Manaosbiidae: Roewer 1932), Santinezia to be an adequate technique for sampling
serratotibialis (Cranaidae: Roewer 1932) are taxa of relatively small body size (e.g., ago-
known from additional locations outside of ristenids, samoids, and zalmoxids) as well
Trinidad, primarily Venezuela (Goodnight as those which utilize immobility as a de-
and Goodnight 1947; Cokendolpher and fensive behavior. Thus, although we did
Camilo-Rivera 1989; Pinto-da-Rocha and not collect representatives of these species
Kury 2003). The purpose of this study was in our field study, our inferences regarding
to provide insight into the ecology and the use of the leaf litter adjacent to trees by
natural history of the harvestmen that oc- agoristenids, samoids, and zalmoxids are
cur in the rainforests of the Northern Range limited by our sampling technique. During
in Trinidad. Specifically, we investigated each sampling day, we recorded air tem-
the use of trunks, buttresses, and the leaf perature, relative humidity, and other per-
litter in the immediate vicinity of the trees tinent weather conditions. Data for tree size
by harvestmen in crappo-cocorite seasonal (DBH) were log transformed and correla-
rainforest (Murphy 1997). This type of rain- tions between tree size and the number of
forest features a relatively high discontinu- harvestmen were examined using Pear-
ous canopy (30-46 m), an abundance of li- son’s product moment correlation. Inter-
anas, orchids, and bromeliads, as well as a specific variation in the number of harvest-
developed shrub and understory layer men occurring on trunks, buttresses, and
(Murphy 1997). Most trees in this type of the litter in the immediate vicinity of the
forest have a relatively smooth bark and trees were assessed using a nonparametric
well-developed buttresses. In addition to Kruskal-Wallis Analysis of Variance (␣ =
determining which species of harvestmen 0.05), followed by multiple comparisons
occupy trees, we examined the relationship tests (Siegel and Castellan 1988). Voucher
between tree size and harvestmen abun- specimens were deposited into the natural
dance. When possible, we also made obser- history collections of the California Acad-
140 NOTES

emy of Sciences (CAS) and the Louisiana harvestmen in Trinidad were those by
State Arthropod Museum (LSAM). Goodnight and Goodnight (1947) and Kury
Our results indicate that five species of and Pinto-da-Rocha (2002). In addition to
harvestmen make extensive use of trees in providing insight into habitat use by sev-
the seasonal crappo-cocorite rainforests of eral taxa, our study is the first to report
Trinidad (Table 1). The harvestmen species gregarious behavior for the family Stygni-
were Cynortula sp. (Cosmetidae), Paecilaema dae. The relatively low number of juveniles
inglei (Cosmetidae), Prionostemma sp. (Scle- (2 out of 240) that we collected may indicate
rosomatidae), Rhopalocranaus albilineatus an ontogenetic shift in the use of trees simi-
(Manaosbiidae), and Stygnoplus clavotibialis lar to the pattern observed in temperate
(Stygnidae: Goodnight and Goodnight species (Edgar 1971). Alternatively, the
1947). We collected significantly more number of juveniles in our study may sim-
adults of Cynortula sp. (KW = 69.1, df = 4, ply reflect the reproductive biology of these
p < 0.001) than the other four species. How- species and the fact that there were rela-
ever, there were no other significant differ- tively few immature individuals in the
ences for the number of adults between the populations of these species at the time of
other species of harvestmen. We collected our sampling. Our examinations revealed
238 adult harvestmen (Table 1) and 2 juve- that the sex ratio for each of the five species
niles (Cynortula sp., n = 1 and Paecilaema that we collected on the trees was nearly
inglei n = 1). We found significant, posi- 1:1. In addition, multiple females of both
tive correlations between tree size (DBH) Cynortula sp. and Rhopalocranaus albilineatus
and harvestmen number for Cynortula sp. were found to be gravid (2 out of 6 and 3
(r = 0.431), P. inglei (r = 0.147), and R. albi- out of 6, respectively), indicating that these
lineatus (r = 0.315). The number of adults species were either breeding or about to be-
was not significantly correlated with DBH gin breeding. Dissections of 4-6 adult fe-
for either Prionostemma (r = 0.012) or S. males of the other three species (Paecilaema
clavotibialis (r = 0.005). During the collection inglei, Prionostemma sp. and Stynoplus clavo-
of our data, we did not directly observe tibialis), however, did not yield similar re-
feeding or mating behavior. However, on sults.
one tree (DBH = 35.3 cm), we did find two The most abundant species at our field
different aggregations (25 and 32 individu- site was the cosmetid Cynortula sp., al-
als, respectively) of adults of S. clavotibialis. though we did observe and capture species
Also, near two trees (DBH = 80.6 cm and representing other families. Owing to the
76.6 cm), three adult cranaids (two Santine- general lack of activity and movement that
zia serratotibialis and one Phareicranaus cal- we observed, we hypothesize that these
cariferus) were found in adjacent leaf litter. harvestmen use trees as shelters, at least
However, we did not observe individuals during daylight hours. During our study,
of either of these two large species of har- we observed adult Cynortula sp. and Priono-
vestmen using tree trunks or buttresses stemma sp. actively moving through the leaf
during our study. litter in the late afternoon and early
Prior to this field study, the only pub- evening hours. On four occasions, we also
lished observations of the natural history of observed adult Prionostemma sp. in the leaf

TABLE 1. Adult harvestmen colllected from 30 trees (mean DBH: 73.2 cm, SD = 177.038, range = 8.6-1000 cm)
from rainforest along the Northern coast of Trinidad.

Total number of Standard


Species specimens collected Mean deviation Range
Cynortula sp. 140 4.7 4.7 0-20
Paecilaema inglei 12 0.4 0.9 0-4
Prionostemma sp. 14 0.5 0.8 0-3
Rhopalocranaus albilineatus 10 0.3 0.6 0-2
Stygnoplus clavotibialis 62 2.1 10.4 0-57
NOTES 141

litter feeding upon adult or larval insects (Phalangida) in Great Britain and Ireland, with
(several dipterans and one ant). An isolated notes on their names, enemies and food. J. Animal
Ecol. 18:100-114.
instance of florivory was noted as well. In
Clawson, R. C. 1988. Morphology of defense glands of
addition, we found both of these species the opilionids (daddy longlegs) Leiobunum vittatum
commonly inhabiting spaces beneath logs. and L. flavum (Arachnida: Opiliones: Palpatores:
Further study of the behavior and ecology Phalangiidae). J. Morphol. 198:363-381.
of the harvestmen of Trinidad are required Cokendolpher, J. C. 1987. Observations on the defen-
before their patterns of microhabitat use sive behaviors of a Neotropical Gonyleptidae
and selection can be fully understood. (Arachnida, Opiliones). Revue Arachnologique 7:59-
Trees are generally believed to provide 63
Cokendolpher, J. C., and G. R. Camilo-Rivera. 1989.
favorable microclimates (high relative hu- Annotated bibliography to the harvestmen of the
midity and reduced exposure to direct sun- West Indies (Arachnida: Opiliones). Occasional Pa-
light) for harvestmen. In addition, arboreal pers Florida St. Coll. Arthropods 5:1-20.
perches may enable harvestmen to avoid Cockerill, J. L. 1988. Notes on aggregations of Leiobu-
potential lethal interactions with large ar- num in the southern U.S.A. J. Arachnol. 16:123-126.
thropod or vertebrate predators in the leaf Edgar, A. L., and H. A. Yuan. 1968. Daily locomotor
litter. Defenses include inactivity, crypsis, activity in Phalangium opilio and seven species of
Leiobunum (Arthropoda: Phalangida). Bios 39:167-
leg autotomy (Guffey 1998), dorsal spines,
176.
and chemicals extruded from repugnatory Edgar, A. L. 1971. Studies on the biology and ecology
glands (Roth and Eisner 1962; Cokendol- of Michigan Phalangida (Opiliones). Misc. Publ.
pher 1987; Clawson 1988; Gnaspini and Univer. Michigan Mus. Zool. 144:5-63.
Cavalheiro 1998; Machado et al. 2005). All Fowler, D. J., and C. J. Goodnight. 1974. Physiological
of these may facilitate co-existence with populations of the arachnid, Leiobunum longipes
generalist predators (e.g., lizards, on or (Opiliones: Phalangiidae). Syst. Zool. 23:219-225.
Goodnight, C. J., and M. L. Goodnight. 1947. Studies
near the surfaces of trees). of the phalangid fauna of Trinidad. Am. Mus. Nov.
1351:1-13.
Acknowledgments.—We are grateful to the Goodnight, C. J., and M. L. Goodnight. 1976. Obser-
Wildlife Section of the Forestry Division of vations on the systematics, development, and hab-
its of Erginulus clavotibialis (Opiliones: Cosmeti-
Trinidad and Tobago, W. I. for granting dae). Trans. Amer. Micros. Soc. 95:654-664
permission to conduct this research. We Gnaspini, P. 1996. Population ecology of Goniosoma
thank Paul Resslar, Mike Moore, and Ste- spelaeum, a cavernicolous harvestmen from South-
phen Broadbridge for assistance in the eastern Brazil (Arachnida: Opiliones: Gonylep-
field. We also thank Jessica Tibbetts and tidae). J. Zool. London. 239:417-435.
Daniel Proud for help with the identifica- Gnaspini, P., and A. J. Cavalheiro. 1998. Chemical and
behavioral defenses of a Neotropical cavernicolous
tion and sorting of specimens. We thank
harvestman: Goniosoma spelaeum (Opiliones, Lania-
C. Guffey and two anonymous reviewers tores, Gonyleptidae). J. Arachnol. 26:81-90.
for their comments and suggestion. This re- Guffey, C. A. 1998. Leg autotomy and its potential
search was supported by a grant from the fitness costs for two species of harvestmen (Arach-
Virginia Federation of Independent Col- nida, Opiliones). J. Arachnol. 26:296-302.
leges (RKH), a Virginia Wesleyan College Gunnarson, B., and M. Hake. 1999. Bird predation af-
Faculty Summer Development Grant fects canopy-living arthropods in city parks. Can. J.
Zool. 77:1419-1428.
(VRT), and the Natural Science Under- Halaj, J., and A. B. Cady. 2000. Diet composition and
graduate Research Fund at Virginia Wes- significance of earthworms as food of harvestmen
leyan College. Specimens were legally col- (Arachnida: Opiliones). Am. Midl. Nat. 143:487-491.
lected and returned to U.S. under export Kury, A. B. 2003. Annotated catalogue of the Laniatores of
permit number 000541. the New World (Arachnida, Opiliones). Revista
Iberica de Aracnología, Zaragoza, vol. especial
monográfico, n° 1: 1-337.
LITERATURE CITED Kury, A. B. and R. Pinto-da-Rocha. 2002. Opiliones, In
Amazonian Arachnida and Myriapoda, ed. J. Adis, pp.
Adams, J. 1984. The habitat and feeding ecology of 345-362. Sofia: Pensoft Publishers.
woodland harvestmen (Opiliones) in England. Machado, G., P. C. Carrera, A. M. Pomini, and A. J.
Oikos 42:361-370. Marsaioli. 2005. Chemical defense in harvestmen
Bristowe, W. S. 1949. The distribution of harvestmen (Arachnida, Opiliones): Do benzoquinone secre-
142 NOTES

tions deter invertebrate and vertebrate predators? reference to those of the Oxford district. J. Animal
J. Chem. Ecol. 31:2519-2539. Ecol. 18:209-229.
Machado, G., R. L. G. Raimundo, and P. S. Oliveira. Williams, G. 1962. Seasonal and diurnal activity of
2000. Daily activity schedule, gregariousness, and harvestmen (Phalangida) and spiders (Araneida)
defensive behavior in the Neotropical harvestmen in contrasted habitats. J. Animal Ecol. 31:23-42.
Goniosoma longipes (Opiliones: Gonyleptidae).
J. Nat. Hist. 34:587-596.
Machado, G., and M. A. Pizo. 2000. The use of fruits by
the Neotropical harvestmen Neosadocus variabilis
(Opiliones: Laniatores, Gonyleptidae). J. Arachnol.
28:357-360.
Machado, G., and C. H. F. Vasconcelos. 1998. Multi-
species aggregations in Neotropical harvestmen
(Opiliones, Gonyleptidae). J. Arachnol. 26:389-391.
McAlister, W. H. 1962. Local movements of the har-
vestman Leiobunum townsendi (Arachnida: Phalan-
gida). Texas J. Sci. 14:167-173.
Mestre, L. A. M., and R. Pinto-da-Rocha. 2004. Popu-
lation dynamics of an isolated population of the
harvestmen Ilhaia cuspidata (Opiliones: Gonylepti-
dae), in Araucaria Forest (Curitiba, Parana, Brazil).
J. Arachnol. 32:208-220.
Morse, D. H. 2001. Harvestmen as commensals of crab
spiders. J. Arachnol. 29:273-275.
Murphy, J. C. 1997. Amphibians and Reptiles of Trinidad
and Tobago. Malabar, Florida: Krieger Publishing,
Co.
Pereira, W., A. Elpino-Campos, K. Del-Claro, and
G. Machado. 2004. Behavioral repertory of the
Neotropical harvestmen Ilhaia cuspidata (Opiliones:
Gonyleptidae). J. Arachnol. 32:22-30.
Pinto-da-Rocha, R., and A. B. Kury. 2003. Phylogentic
analysis of Santinezia with description of five new
species (Opiliones, Lanitores, Cranaidae). J. Arach-
nol. 31:173-208.
Roewer, C. F. 1910. Revision der Opiliones Plagi-
ostethi (= Opiliones, Palpatores). 1 Teil: Familie der
Phalangiidae. (Subfamilien: Gagrellini, Liobunini,
Leptobunini). Abhandlunge aus dem Gebiet der
Naturwissenchaften hrsg. vom naturwissenschaftlichen
Verein in Hamburg 19:1-294.
Roewer, C. F. 1932. Weitere Webernechte VII. Ergän-
zung der “Webernechte der Erde”, 1923 (Cra-
nainae). Archiv für Naturgeschichte, N. F. 1:275-350.
Roewer, C. F. 1953. Neotropische Gagrellinae. (Opil-
iones, Arachnida). Mitteilungen aus dem Zoolo-
gischen Museum in Berlin 29:180-264.
Roth, L. M., and T. Eisner. 1962. Chemical defenses of
arthropods. Ann. Rev. Entomol. 7:107-136.
Siegel, S., and N. J. Castellan, Jr. 1988. Nonparametric
Statistics for the Behavioral Sciences, 2nd edition.
New York: McGraw-Hill, Inc.
Simon, E. 1879. Essai d’une classification des Opil-
iones Mecostethi, Remarques synonymiques et de-
scriptions d’espèces nouvelles. Annales de la Sociètè
entomologique de Belgique 22:183-241.
Stradling, D. J. 1994. Distribution and behavioral ecol-
ogy of an arboreal “tarantula” spider in Trinidad.
Biotropica 26:84-97.
Todd, V. 1949. The habitats and ecology of the British
harvestmen (Arachnida, Opiliones), with special

You might also like