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Invertebrate animals extracted from native Tillandsia (Bromeliales : Bromeliaceae) in Sarasota County, Florida

June 2004
Florida Entomologist 87(2):176-185

June 2004
87(2):176-185

DOI:10.1653/0015-4040(2004)087[0176:IAEFNT]2.0.CO;2

Authors:

J.Howard Frank

University of Florida

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Twenty four epiphytic bromeliads belonging to four species (Tillandsia fasciculata Swartz, T recurvata (L.), T setacea Swartz, and T. utriculata L.) were collected in Sarasota County, Florida, in October-November 1997. Macroscopic invertebrate animals were extracted from each by washing in water, filtering, and preserving in 75% ethanol. Plant sizes were measured in several ways, and their substrate was identified. Invertebrates were sorted, counted, and identified as far as possible to the species level. Two species (T. fasciculata, T. utriculata) that impound water in their leaf axils housed aquatic dipteran larvae and pupae (Psychodidae, Culicidae, Ceratopogonidae, Chironomidae, Muscidae, and Aulacigastridae) representing 7 species in 6 genera. Only T. utriculata had a clear relationship between plant size and number of invertebrates, which was steeper when only aquatic insect larvae were counted. Plants of all four species housed terrestrial invertebrates, representing minimally an additional 82 species in 75 genera and 63 families, very few of which are known to have an obligate relationship with bromeliads, but showing that these plants support a diverse invertebrate fauna. The presence of ant nests in some bromeliads complicated analysis. Such a list of terrestrial invertebrates, identified to the species level, has not before been compiled for bromeliads in Florida. Some collaborating taxonomists obtained specimens of species that they could not identify, including probably undescribed species.

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176

Florida Entomologist

87(2) June 2004

INVERTEBRATE ANIMALS EXTRACTED FROM NATIVE

TILLANDSIA

(BROMELIALES: BROMELIACEAE) IN SARASOTA COUNTY, FLORIDA

J. H. F

RANK

, S. S

REENIVASAN

, P. J. B

ENSHOFF

, M. A. D

EYRUP

, G. B. E

DWARDS

, S. E. H

ALBERT

A. B. H

AMON

, M. D. L

OWMAN

, E. L. M

OCKFORD

, R. H. S

CHEFFRAHN

, G. J. S

TECK

, M. C. T

HOMAS

T. J. W

ALKER

AND

W. C. W

ELBOURN

Entomology and Nematology Department, University of Florida, Gainesville, FL 32611-0630

Marie Selby Botanical Gardens, 811 South Palm Avenue, Sarasota, FL 34236

Myakka River State Park, 13207 SR 72, Sarasota, FL 34241

Archbold Biological Station, P.O. Box 2057, Lake Placid, FL 33862

Florida State Collection of Arthropods, 1911 SW 34th St., Gainesville, FL 32608-1268

Department of Biological Sciences, Illinois State University, Normal, IL 61790-4120

Fort Lauderdale Research and Education Center, University of Florida, 3205 College Ave.

Ft. Lauderdale, FL 33314-7799

BSTRACT

Twenty four epiphytic bromeliads belonging to four species (

Tillandsia fasciculata

Swartz,

T. recurvata

(L.),

T. setacea

Swartz, and

T. utriculata

) were collected in Sarasota County,

Florida, in October-November 1997. Macroscopic invertebrate animals were extracted from

each by washing in water, ﬁltering, and preserving in 75% ethanol. Plant sizes were mea-

sured in several ways, and their substrate was identiﬁed. Invertebrates were sorted,

counted, and identiﬁed as far as possible to the species level. Two species (

T. fasciculata

utriculata

) that impound water in their leaf axils housed aquatic dipteran larvae and pupae

(Psychodidae, Culicidae, Ceratopogonidae, Chironomidae, Muscidae, and Aulacigastridae)

representing 7 species in 6 genera. Only

T. utriculata

had a clear relationship between plant

size and number of invertebrates, which was steeper when only aquatic insect larvae were

counted. Plants of all four species housed terrestrial invertebrates, representing minimally

an additional 82 species in 75 genera and 63 families, very few of which are known to have

an obligate relationship with bromeliads, but showing that these plants support a diverse in-

vertebrate fauna. The presence of ant nests in some bromeliads complicated analysis. Such

a list of terrestrial invertebrates, identiﬁed to the species level, has not before been compiled

for bromeliads in Florida. Some collaborating taxonomists obtained specimens of species

that they could not identify, including probably undescribed species.

Key Words: Bromeliads, phytotelmata, insects,

Tillandsia utriculata

, bromeliad inhabitants

ESUMEN

Se colectaron 24 bromeliáceas epíﬁtas que pertenecen a cuatro especies (

Tillandsia fascicu-

lata

Swartz,

T. recurvata

(L.),

T. setacea

Swartz, y

T. utriculata

) en el Condado de Sara-

sota, Florida, durante octubre-noviembre de 1997. Se extrayeron los animales invertebrados

macroscópicos de cada planta lavándola en agua y ﬁltrando, seguido por preservación de los

especímenes en etanol de 75%. Se midieron los tamaños de las plantas por varios

parámetros, y se identiﬁcó su sustrato. Los invertebrados se ordenaron, contaron e identiﬁ-

caron tanto posible al nivel de especie. Las dos especies (

T. fasciculata

T. utriculata

) que em-

balsan agua entre sus axilas de hojas alojaron larvas y pupas acuáticas de moscas

(Psychodidae, Culicidae, Ceratopogonidae, Chironomidae, Muscidae y Aulacigastridae) rep-

resentando 7 especies en 6 géneros. Solo

T. utriculata

tuvo una relación clara entre tamaño

de planta y cantidad de invertebrados, la cual fue más fuerte cuando se contaron solamente

las larvas de insectos acuáticos. Plantas de las cuatro especies alojaron invertebrados ter-

restres, representando un mínimo de 82 especies adicionales en 75 géneros y 63 familias,

muy pocas de las cuales se conocen tener una relación obligada con bromeliáceas, pero de-

muestran que estas plantas sostienen una diversa fauna de invertebrados. La presencia de

nidos de hormigas en algunas bromeliáceas complicó el análisis. Tal lista de invertebrados

terrestres, identiﬁcados al nivel de especie, no ha sido recopilado anteriormente para bro-

Frank et al.: Invertebrates from Florida

Tillandsia

Bromeliads 177

meliáceas en Florida. En este proyecto, varios taxónomos obtuvaron especimenes no-identi-

ﬁcados, incluyendo especies probablemente no-descritas.

Translation provided by the authors.

Bromeliads (Bromeliaceae) are a family of at

least 2500 species of monocotyledonous plants, al-

most restricted to the Neotropical region, but in-

cluding all of Mexico and southernmost USA. The

complex architecture of some species traps water

in leaf axils (forming phytotelmata) and harbors

many species of invertebrate animals. There are

thus three types of associations of invertebrates

with these plants: (a) those that feed on the

plants, (b) organisms aquatic at least in their im-

mature stages, and (c) those terrestrial organisms

for which bromeliads provide concealment, hu-

midity, or prey (Frank 1983). Within all three

groups are specialists, associated only with bro-

meliads, as well as generalists that occupy simi-

lar (non-bromeliad) habitats.

Four approaches have been followed in at-

tempts to unravel the mysteries of bromeliad

fauna. They may be termed (i) brief reports of new

discoveries, (ii) in-depth studies (behavioral or eco-

logical or taxonomic) of selected taxa, (iii) whole-

fauna inventories, and (iv) broad-scale hypothesis

tests. Major difﬁculties with the last two ap-

proaches are the need to involve teams of special-

ist taxonomists, and of distinguishing transient

species from those that have some kind of obligate

or at least usual relationship with bromeliads.

In Florida, an inventory of the macroscopic

aquatic invertebrate fauna in bromeliad tanks

(phytotelmata) is contained in an unpublished

Ph.D. dissertation (Fish 1976). A little of the con-

tent of that work was reviewed in Frank (1983).

An introduction to, and a bibliography of, studies

of the fauna and microﬂora of bromeliad phytotel-

mata, in Florida and abroad, are WWW-published

(Frank 1996 a, b). A complete illustrated key to

all developmental stages of all aquatic inverte-

brates in bromeliad phytotelmata in Florida can-

not now be prepared because some species are yet

undescribed (unknown to science). In contrast,

there are only 16 native species of bromeliads in

Florida, identiﬁable by color photographs online

as part of Frank & Thomas (1996) or (for the more

botanically adept) by a key in Wunderlin (1998).

In Florida, there has been one inventory of the

entire invertebrate fauna in the bromeliad

Tillandsia utriculata

L. (Sidoti 2000), but most of

its identiﬁcations reached only the level of order.

There are works on some insects that feed on and

harm bromeliads. Detection of a moth, whose lar-

vae destroy pods of the bromeliad

Tillandsia fas-

ciculata

Swartz, led to a publication about larvae

of several moths that occasionally are collected

from native bromeliads in Florida (Heppner &

Frank 1998). Detection of a Mexican weevil,

Metamasius callizona

(Chevrolat), in Florida led

to several publications about bromeliad-eating

weevils, reviewed and augmented by Frank

(1999) and expanded in two webpages (Larson &

Frank 2000; Larson et al. 2001) and two websites

(Frank & Thomas 1996; Larson 2000). Notable

studies in other countries are by Picado (1913) in

Costa Rica, and Beutelspacher (1971a, b) and

Palacios-Vargas (1981, 1982) in Mexico.

The Marie Selby Botanical Gardens (MSBG),

Sarasota, Florida, have an “Intern Program.” Un-

der this program, students interested in plant

ecology and other aspects of botany are brought

from elsewhere to conduct a short-term (few

months) research project in one of these subjects.

Margaret Lowman (Research Director, MSBG)

and Sheeba Sreenivasan (an intern from Trinidad

and Tobago) in 1997 designed a project that was

to be a quantitative examination of the inverte-

brate fauna associated with native bromeliads in

Sarasota County. One of us (JHF) was asked to

visit MSBG to explain to Sheeba how to extract

invertebrates from bromeliads and preserve them

for examination, and also to receive her in his lab-

oratory and provide her with literature that

would help her to make preliminary identiﬁca-

tions of the invertebrate specimens. These were

limited to insects, arachnids, myriapods, mol-

luscs, annelids, and the larger crustaceans. Fur-

ther development depended upon specialist

taxonomists to take the preliminary identiﬁca-

tions as far as possible to the species level.

In November 1997, Sheeba visited JHF’s labo-

ratory at the University of Florida, and used a

leaf-area-area meter for about 3 days to measure

the leaf-areas of the bromeliads she had collected.

To help complete the project, he sorted Sheeba’s

specimens, some to family, but others only to the

level of order. He provided genus- or species-level

identiﬁcation of the immature mosquitoes and a

few other dipteran larvae with which he was fa-

miliar and, much later, drafted a manuscript for

review by the other contributors. All the remain-

ing specimens had to be sent to specialist taxono-

mists for reliable identiﬁcation, and the contacts

were made and specimens shipped before the end

of 1997. Fortunately, taxonomists of the Florida

State Collection of Arthropods (FSCA) were recep-

tive to providing help. Here is an account of these

invertebrates. This account recognizes the essen-

tiality of the contributions of several taxonomists,

who were offered co-authorship (some declined).

ATERIALS

AND

ETHODS

Twenty four bromeliads were collected from

sites in Sarasota County, principally from old-

178

Florida Entomologist

87(2) June 2004

growth hammocks in the Myakka River State

Park, and secondarily two sites in Sarasota. They

were removed from various substrates including

living trees, dead trees (snags), and a gate (Table

1). While it was being collected, each plant was

kept as upright as possible to prevent spillage.

The plant was then placed into a polyethylene bag

and brieﬂy sprayed with insect repellent before

fastening the bag.

Invertebrates were extracted from bromeliads

by a variant of the method of Frank et al. (1976).

Each plant was cleaned with a jet of water from a

hose, with the washings directed into a bucket.

The plant was repeatedly submerged and shaken

in the bucket before being returned to its bag. The

water in the bucket was then ﬁltered with a tea-

strainer (mesh size 500 µm) and the residue ex-

amined for invertebrates with a dissection micro-

scope. Collected invertebrates were preserved in

vials containing 75% ethanol for subsequent iden-

tiﬁcation.

The collected bromeliads included whole spec-

imens of the epiphytic species

Tillandsia utricu-

lata

T. fasciculata

T. setacea

, and

T. recurvata

These include all the most widespread of Florida’s

16 native species except

T. usneoides

(L.). Only

the ﬁrst two of these impound water in leaf axils,

and they do this only when they have reached a

certain minimal size (exceeded by the specimens).

The volumetric capacity of the water-impounding

leaf axils of

T. utriculata

has been related to

length of longest leaf and to age (in one habitat)

by Frank & Curtis (1981), so length of longest leaf

was one of the measurements made (Volumetric

capacity in ml = 0.003251

leaf length in cm

2.7799

Other measurements were made by dismantling

each plant, leaf by leaf, starting from the outer-

most and working inward. This was done on a

white background to facilitate detection of any re-

maining invertebrates. A component part was

considered to be either a leaf or an infructescence

(the fruiting phase of an inﬂorescence). Each ele-

ment was further designated as live or dead. Each

bromeliad’s live and dead leaves were counted.

All components were refrigerated until leaf and

infructescence areas were measured with an area

meter (LI-COR Portable Area Meter, model LI

3000, LI-COR Inc., Lincoln, NE, USA). They were

then oven-dried for 48 hr before weighing. Table

1, which lists the measurements, is thus a habitat

description rather than results.

ESULTS

Table 2 lists the invertebrates collected to the

level of family (with number of specimens) for

each of the 24 bromeliads sampled. Identification

of the invertebrates to the species level, where

possible, is given below. Comments are made

where deemed appropriate. Three vials contain-

ing Mollusca and four with Thysanoptera were

mislaid somewhere in the Florida State Collec-

tion of Arthropods; details of their contents would

not substantially change the conclusions.

Mollusca

None was identiﬁed. The three missing vials (6

specimens) may be located in FSCA. No mollusc

has a known, obligate relationship to bromeliads.

However, H. E. Luther (pers. comm.) has observed

snails eating bromeliad trichome caps in the ﬁeld

and greenhouse. Assume minimally one family,

one genus, and one species.

Isopoda (identiﬁed by G. B. Edwards)

Oniscidae: genus and species unidentiﬁed (17

specimens).

Rhyscotidae: genus and species unidentiﬁed (9

specimens).

Diplopoda (identiﬁed by G. B. Edwards)

Chilognatha: family and genus unidentiﬁed,

species 1 (12 specimens), species 2 (1 specimen).

Pselaphognatha: Polyxenidae:

Polyxenus fas-

ciculatus

(Say) (5 specimens).

These were collected from all three

Tillandsia

species in MRSP. They have no known relation-

ship to bromeliads. All of the Chilognatha were

immature, so could not be identiﬁed reliably.

Chilopoda (identiﬁed by G. B. Edwards)

Lithobiidae: ?

Neolithobius

sp. (1).

This immature specimen was collected from

T. utriculata

in MRSP. It has no known relation-

ship to bromeliads.

Araneae (identiﬁed by G. B. Edwards)

Segestriidae:

Ariadna bicolor

(Hentz) (2).

Theridiidae:

Phoroncidia americana

(Emer-

ton) (1 immature), ?genus (1 immature).

Mysmenidae:

Mysmenopsis cymbia

Levi (10).

Linyphiidae:

Ceraticelus

phylax

Ivie & Bar-

rows (1 female).

Tetragnathidae:

Dolichognatha pentagona

(Hentz) (1), ?

D. pentagona

(1 immature, dam-

aged), ?

Tetragnatha

sp. (2 immatures).

Lycosidae: ?genus (2 immatures).

Pisauridae: ?

Dolomedes

sp. (1 immature).

Agelenidae: ?

Agelenopsis

sp. (1 immature).

Hahniidae:

Hahnia okeﬁnokensis

Chamberlin

& Ivie (1).

Dictynidae:

Emblyna capens

Chamberlin (1),

Emblyna

sp. (2 immature),

Lathys delicatula

Gertsch & Mulaik (2).

Anyphaenidae:

Lupettiana mordax

(O.P. Cam-

bridge) (1).

Liocranidae:

Scotinella pintura

(Ivie & Bar-

rows) (3),

Scotinella

sp. (1 immature).

Frank et al.: Invertebrates from Florida

Tillandsia

Bromeliads 179

Clubionidae:

Clubiona pygmaea

Banks (3),

Clu-

biona

sp. (1 immature),

Elaver excepta

(L. Koch)

(10).

Gnaphosidae:

Litopyllus cubanus

Bryant (1),

Sergiolus

sp. (1 immature).

Sparassidae:

Pseudosparianthis cubana

Banks

(1 immature).

Thomisidae:

Bassaniana ﬂoridana

(Banks) (1),

Bassaniana

sp. (4 immatures).

Salticidae:

Anasaitis canosa

(Walckenauer) (2).

The spiders seem to represent 21 species, in 21

genera and 17 families. For only one spider (

Pele-

grina tillandsia

[Kaston]) is a bromeliad (

Tilland-

sia usneoides

) in the southern USA known to be

the preferred habitat. In the Neotropical region,

however, other spiders typically inhabit bromeli-

ads and even are semi-aquatic in bromeliad leaf

axils.

Pseudoscorpiones (identiﬁed by G. B. Edwards)

Chthoniidae: genus and species unidentiﬁed (2

immatures).

One specimen was from

T. fasciculata

and the

other from

T. setacea

. They were unidentiﬁable

because immature. Pseudoscorpions have no

known obligate relationship to bromeliads.

Acari (identiﬁed by W. C. Welbourn)

Liodidae: Liodes sp. 1 (16), Liodes sp. 2 (13), Li-

odes sp. 3 (3).

Ascidae: Lasioseius sp. (2).

Haplozetidae: genus and species unidentiﬁed

(1).

Oripodidae: genus and species unidentiﬁed (1).

Uropodidae: Uropoda sp. (2).

Oppiidae: genus and species unidentiﬁed (1).

Orbataloid: genus and species unidentiﬁed (1).

Histiostomatidae: Hormosianoetus sp. (37).

None of these 10 species in 8 genera and 8 fam-

ilies is known to have any obligate relationship

with bromeliads. There is a pressing need for more

basic taxonomic work on Floridian Acari other

than those of economic importance; only then will

specimens be identiﬁable to the species level.

TABLE 1. BROMELIAD MEASUREMENTS (NUMBER OF LIVE LEAVES, DEAD LEAVES, LONGEST LEAF, AREAS OF LIVING AND

DEAD

LEAVES, OF LIVING AND DEAD INFRUCTESCENCES, AND DRY WEIGHTS) MATCHED TO SAMPLE NUMBER

(CODE), SUBSTRATE ON WHICH IT GREW, AND IDENTIFICATION.

Code Substrate

No. live

leaves

No. dead

leaves

Longest

leaf (cm)

Live leaf

area (cm

)

Dead leaf

area (cm

)

Live infr.

area (cm

)

Dead infr.

area (cm

)

Dry wt

(g)

Tillandsia fasciculata

8 Cephalanthus 64 4 58.4 2621.99 60.18 0 0 57.5

9 Cephalanthus 65 20 62.0 2794.81 316.20 0 0 71.5

10 Cephalanthus 51 11 46.4 898.86 66.24 0 0 21.5

15 snag 55 13 44.5 2002.08 199.07 0 0 33.0

17 Ulmus 33 3 44.5 881.81 44.55 0 0 10.5

18 fallen branch 55 2 22.8 350.46 37.23 0 0 6.5

19 rooted in soil 58 10 29.2 475.15 27.79 0 0 11.5

20 Quercus 76 13 55.4 5006.87 371.90 0 0 189.5

Tillandsia utriculata

1 snag 41 6 32.0 531.62 38.44 0 0 8.0

2 snag 56 29 46.5 1793.91 237.25 0 0 32.0

3 snag 55 22 33.0 824.92 63.65 0 0 11.5

4 snag 62 11 17.9 164.69 14.34 0 0 2.5

16 Quercus 60 12 91.1 9285.81 909.66 0 0 147.0

24 Quercus 59 11 93.3 7350.48 0 0 0 152.5

Tillandsia recurvata

21 Sabal 1140 71 10.1 672.1 48.60 0 0 10.5

22 wooden gate 80 1 9.3 51.69 2.37 6.02 0 1.0

23 Quercus 537 58 13.5 305.72 82.25 34.36 23.00 5.0

Tillandsia setacea

5 snag 196 82 19.0 151.37 62.86 0 0 3.5

6 snag 349 24 18.6 236.57 17.15 0 0 3.0

7 snag 158 22 27.2 210.06 27.90 28.20 5.58 8.0

11 Cephalanthus 419 63 32.1 556.39 60.18 67.68 7.97 16.5

12 Cephalanthus 338 29 28.9 437.23 62.32 18.87 16.13 9.5

13 Cephalanthus 81 34 23.5 185.37 36.46 26.92 0 3.0

14 Quercus 379 32 29.8 298.02 18.81 4.93 5.33 8.5

180 Florida Entomologist 87(2) June 2004

TABLE 2. ORIGIN (CODE/PLACE) OF TILLANDSIA SPECIMENS AND NUMBERS OF ARTHROPODS EXTRACTED, TO FAMILY

LEVEL

. ALL WERE COLLECTED IN MID-OCTOBER TO MID-NOVEMBER 1997 IN SARASOTA COUNTY, FL.

CD/PL Fauna to level of family, and number of specimens Sum

Tillandsia fasciculata

08/ M CRUSTACEA: Isopoda: Rhyscotidae (2), ARACHNIDA: Araneae: Theridiidae (1), Mys-

menidae (10), Hahniidae (1), Liocranidae (1), INSECTA: Homoptera: Aphididae (1), Lepi-

doptera: Tineidae (larvae, 1), Diptera: ?Muscidae (larva 1), Hymenoptera: Formicidae (1)

09/ M ARACHNIDA: Araneae: Tetragnathidae (1), Dictynidae (1), Liocranidae (1), Clubionidae

(2), Salticidae (1), INSECTA: Isoptera: Kalotermitidae (4), Blattodea: Blatellidae (2), Pso-

coptera: Lepidopsocidae (nymphs 2), family indet. (nymphs 2), Diptera: ?Muscidae (larva 1),

Hymenoptera: Formicidae (1)

10/ M CRUSTACEA: Isopoda: Rhyscotidae (6), ARACHNIDA: Araneae: Linyphiidae (1), Age-

lenidae (1), Clubionidae (2), Acari: Uropodidae (1), INSECTA: Orthoptera: Gryllidae (3),

Psocoptera: Lepidopsocidae (1), Archipsocidae (1), Peripsocidae (2), family indet. (1), Co-

leoptera: Tenebrionidae (1), Hymenoptera: Formicidae (1)

15/ M DIPLOPODA: Pselaphognatha: Polyxenidae (1), ARACHNIDA: Araneae: Segestriidae (1),

Pisauridae (1), Acari: Histiostomatidae (19), INSECTA: Thysanoptera (1), Psocoptera: Lep-

idopsocidae (1), Orthoptera: Gryllidae (2), Coleoptera (larvae 3, of 3 families), Diptera: Psy-

chodidae (larvae 32), Culicidae (larva 1), Ceratopogonidae (larvae 3)

17/ M CRUSTACEA: Isopoda: Rhyscotidae (1), DIPLOPODA: Chilognatha: family indet. (1),

ARACHNIDA: Araneae: Clubionidae (2), Pseudoscorpionida: Chthoniidae (1), INSECTA:

Blattodea: Blatellidae (4, and one egg case), Lepidoptera: family indet. (larvae 5), Co-

leoptera: Carabidae (larva 1), Diptera: Ceratopogonidae (larva 1)

18/ M ARACHNIDA: Araneae: Anyphaenidae (1), Thomisidae (1), Salticidae (1), Acari (?1), IN-

SECTA: Orthoptera: Gryllidae (1), Coleoptera: Scirtidae (1), Hymenoptera: Formicidae (1)

19/ M CRUSTACEA: Isopoda: Oniscidae (3), DIPLOPODA: Pselaphognatha: Polyxenidae (3), Pse-

laphognatha: family indet. (3), ARACHNIDA: Araneae: Lycosidae (2), INSECTA: Blattodea

(egg case 1), Homoptera: Ortheziidae (1), Coleoptera: Brentidae (1), Hymenoptera: Formi-

cidae (128 plus brood), Ichneumonidae (2)

144

20/ M MOLLUSCA (3), CRUSTACEA: Isopoda: Oniscidae (10), ARACHNIDA: Araneae: Dic-

tynidae (1), Sparassidae (1), INSECTA: Blattodea: Blattidae (1), Orthoptera: Gryllidae (1),

Lepidoptera: family indet. (larva 1), Diptera: Psychodidae (larvae 2), Ceratopogonidae (lar-

vae 8, pupae 2) Chironomidae (larvae 4), Aulacigastridae: (larva 1, pupa 1)

Tillandsia utriculata

01/ M ARACHNIDA: Araneae: Dictynidae (1), Acari: Liodidae (1) 2

02/ M CRUSTACEA: Isopoda: Oniscidae (2), DIPLOPODA: Chilognatha: family indet. (5),

ARACHNIDA: Araneae: Tetragnathidae (1), Liocranidae (2), Clubionidae (1), Acari: Ascidae

(2), INSECTA: Blattodea: Blatellidae (1), Psocoptera: Pseudocaeciliidae (1), Liposcelidae

(1), family indet. (nymph 1), Diptera: Ceratopogonidae (larva 1), Psychodidae (larvae 16),

Hymenoptera: Formicidae (5)

03/ M DIPLOPODA: Pselaphognatha: Polyxenidae (1), ARACHNIDA: Araneae: Clubionidae (1),

Thomisidae (1), INSECTA: Psocoptera: Caeciliusidae (1), Archipsocidae (1), Liposcelidae (1),

Lepidopsocidae (nymph 1), family indet. (nymph 1), Blattodea: Blattidae (2), Homoptera:

Coccidae (1), Coleoptera: Curculionidae (3), Diptera, Ceratopogonidae (larvae 2)

04/ M DIPLOPODA: Chilognatha: family indet. (2), ARACHNIDA: Araneae: Clubionidae (1), Th-

omisidae (2), Acari: Liodidae (1), INSECTA: Blattodea: Blatellidae (2), Coleoptera: Curcu-

lionidae (1), Diptera: Chironomidae (1)

16/ M CHILOPODA: Lithobiidae (1), DIPLOPODA: Chilognatha: family indet. (1), ARACHNIDA:

Acari: Histiostomatidae (18), INSECTA: Blattodea: Blattidae (1), Psocoptera: family indet.

(1 nymph), Thysanoptera (2), Lepidoptera: family indet. (larva 1), Diptera: Psychodidae:

(larvae 108, pupa 1), Culicidae: (larvae 28), Ceratopogonidae (larvae 47), Chironomidae (lar-

vae 3), Aulacigastridae: (5), Cecidomyiidae (adults 2, pupa 1), Hymenoptera: Formicidae (6)

226

24/ S ARACHNIDA: Araneae: Dictynidae (1), Gnaphosidae (1), INSECTA: Collembola: Entomobry-

idae (22), Psocoptera: Trogiidae (2), family indet. (nymph 1), Coleoptera: Coccinellidae (1),

Diptera: Psychodidae (larvae 31), Hymenoptera: Formicidae (76, of 2 spp., each with brood)

135

Tillandsia recurvata

21/ S INSECTA: Psocoptera: Lepidopsocidae (1), Coleoptera: Elateridae (1), Hymenoptera: For-

micidae (2)

22/ S No animals were collected 0

Frank et al.: Invertebrates from Florida Tillandsia Bromeliads 181

Collembola (identiﬁed by R. J. Snider)

Entomobryidae: Seira steinmetzi Wray (22).

Hypogastruridae: Xenylla sp. (5).

The specimens of Xenylla represent an unde-

scribed species and were retained by R. J. Snider.

Orthoptera (identiﬁed by T. J. Walker)

Gryllidae: Cycloptilum trigonipalpum (Rehn &

Hebard).

All 8 specimens of Orthoptera were identiﬁed

as of this species or were unidentiﬁable because

immature, but probably belong to this species.

Seven of them were collected from T. fasciculata

and one from T. setacea, all within MRSP.

Blattodea (identiﬁed by M. C. Thomas)

Blattellidae: Cariblatta sp. prob. lutea (Saus-

sure & Zehnter) (12).

Blattidae: Eurycotis ﬂoridana (Walker) (4).

Neither species has any known obligate rela-

tionship with bromeliads.

Isoptera (identiﬁed by R. H. Scheffrahn)

Kalotermitidae: Crytotermes ?cavifrons Banks

(4).

No termite is known to have an obligate rela-

tionship with bromeliads. Most likely these had

fallen from a dead tree limb.

Psocoptera (identiﬁed by E. L. Mockford)

Trogiidae: Cerobasis guestfalica (Kolbe) (4).

Lepidopsocidae: Echmepteryx (Thylacopsis)

madagascariensis (Kolbe) (3), Echmepteryx sp. (1),

Nepticulomima sp. (1), unidentiﬁed nymphs (4).

Liposcelidae: Liposcelis ornata Mockford (1),

Liposcelis sp. (2).

Archipsocidae: Archipsocus sp. (2), unidenti-

ﬁed nymphs (1).

Peripsocidae: Peripsocus sp. (5).

Caeciliusidae: Valenzuela indicator (Mockford)

(= Caecilius indicator Mockford) (1).

Pseudocaeciliidae: Pseudocaecilius citricola

(Ashmead) (1).

Epipsocidae: Mesepipsocus niger (New) (3), un-

identiﬁed nymphs (4).

None of these 12 species in 9 genera and 8 fam-

ilies is known to have an obligate relationship to

bromeliads. As in so many other groups, it is dif-

ﬁcult to identify nymphs reliably to the species

level. The surprise among these specimens was

the ﬁnding of specimens of C. guestfalica from

two Tillandsia specimens (T. recurvata, T. utricu-

lata) from the city of Sarasota; it is an adventive

species.

Thysanoptera

None was identiﬁed. The four missing vials (6

specimens) may be located in FSCA. Assume min-

imally one family, one genus, and one species.

23/ S MOLLUSCA (1), ARACHNIDA: Araneae: Gnaphosidae (1), INSECTA: Hemiptera: Miridae

(nymphs 2), Psocoptera: Trogiidae (1), Hymenoptera: Formicidae (1)

Tillandsia setacea

05/ M Diplopoda: Pselaphognatha: Polyxenidae (1), ARACHNIDA: Acari: Liodidae (4), Haploz-

etidae (1), Oripodidae (1), INSECTA: Psocoptera: Peripsocidae (1), Archipsocidae (1

nymph), Lepidopsocidae (1 nymph), Orthoptera: Gryllidae (1)

06/ M ARACHNIDA: Araneae: Clubionidae (2), Thomisidae (1), Acari: Liodidae (9), INSECTA:

Psocoptera: Peripsocidae (2), Homoptera: Aphididae (4), Hymenoptera: Aphelinidae (1)

07/ M MOLLUSCA (2), ARACHNIDA: Araneae: Segestriidae (2), INSECTA: Blattodea: Blatel-

lidae (2), Lepidoptera: family indet. (larvae 2), Hymenoptera: Formicidae (52)

11/ M ARACHNIDA: Araneae: Dictynidae (1), Clubionidae (1), Acari: Liodidae (2), INSECTA:

Thysanoptera (1), Coleoptera: Scirtidae (1)

12/ M ARACHNIDA: Araneae: Theridiidae (1), Tetragnathidae (2), Thomisidae (1), Acari: Lio-

didae (2), Oppiidae (1), ‘orbataloid’ (1), Pseudoscorpionida: Chthoniidae (1), INSECTA: Col-

lembola: Hypogastruridae (5), Thysanoptera (2), Psocoptera: Peripsocidae (nymph 1),

Hymenoptera: Formicidae (1)

13/ M CRUSTACEA: Isopoda: Oniscidae (2), ARACHNIDA: Araneae: Clubionidae (1), Acari: Lio-

didae (13), INSECTA: Psocoptera: Liposcelidae (1), Diptera: Cecidomyiidae (1)

14/ M ARACHNIDA: Araneae: Clubionidae (1), Acari: Uropodidae (1), INSECTA: Lepidoptera:

Gelechiidae (larva in ﬂower) (1), Hymenoptera: Formicidae (1)

Notes: CD = code number (collection no.)/PL = place (M = Myakka River State Park, S = Sarasota). SUM = total number of in-

vertebrate animals of the groups sampled. Presence of immature stages suggests that development was taking place in the brome-

liad unless the individuals fell from the tree above.

TABLE 2. (CONTINUED) ORIGIN (CODE/PLACE) OF TILLANDSIA SPECIMENS AND NUMBERS OF ARTHROPODS EXTRACTED, TO

FAMILY

LEVEL. ALL WERE COLLECTED IN MID-OCTOBER TO MID-NOVEMBER 1997 IN SARASOTA COUNTY, FL.

182 Florida Entomologist 87(2) June 2004

Hemiptera/Homoptera (identiﬁed by S. E. Halbert)

(Ortheziidae by A. B. Hamon)

Miridae: unidentiﬁed nymphs.

Aphididae: Myzocallis sp. (4), unidentiﬁed ge-

nus (1 cast skin).

Ortheziidae: Orthezia sp. (1 nymph).

Coccidae: genus and species unidentiﬁed (1

adult male).

The Myzocallis aphids are known to feed on

oak; one of the specimens was parasitized by Aph-

elinus sp. (Hymenoptera: Aphelinidae) (det. G. A.

Evans, FSCA). The only species of Orthezia re-

ported from Tillandsia in Florida is O. tillandsiae

Morrison, but the specimen obtained (from T. fas-

ciculata in MRSP) is immature and could not be

identiﬁed with certainty. The Miridae were uni-

dentiﬁable because they were immature.

Coleoptera (identiﬁed by M. C. Thomas)

Scirtidae: Ora sp. (1), Cyphon sp. (1).

Elateridae: Conoderus amplicollis (Gyllenhal)

(1).

Tenebrionidae: Glyptotus cribratus LeConte (1).

Brentidae: Apion sp. (1).

Curculionidae: Acalles clavatus Say (3);

Conotrachelus maritimus Blatchley (2).

Larvae of Scirtidae are aquatic. It is possible

that the two adults of Scirtidae are associated

with bromeliad phytotelmata, but both specimens

were found in T. setacea, which does not form phy-

totelmata. Alternatively, their larvae may de-

velop in treeholes. Specimens mislaid include one

adult of Coccinellidae and several beetle larvae, of

which one is of Carabidae.

Lepidoptera (identiﬁed by D. H. Habeck)

Tineidae: genus and species unidentiﬁed (1

larva).

Gelechiidae: genus and species unidentiﬁed (1

larva).

Family uncertain: (8 larvae).

Lepidoptera were represented only by larvae of

“primitive” families. Their identiﬁcation was un-

certain. It is not clear whether these larvae were

feeding on bromeliads, on debris in the bromeli-

ads, or on the tree canopy (or epiphytes) above.

The gelechiid larva was found clinging to a ﬂower

(which was not a bromeliad ﬂower) in the brome-

liad, but there is no evidence it was feeding on that

ﬂower. There is a clear need to rear lepidopteran

larvae encountered in bromeliads, to allow identi-

ﬁcation of adults and associate larvae with adults.

Diptera (identiﬁed by J. H. Frank)

(Cecidomyiidae by R. J. Gagné, Ceratopogonidae by

G. J. Steck)

Cecidomyiidae: Campylomyza sp. (1 adult),

Lestodiplosis laticaulis Gagné (2 adults and 1

pupa).

Psychodidae: Alepia sp. (190 larvae and pu-

pae).

Culicidae: Wyeomyia mitchellii (Theobald) (25

larvae), W. vanduzeei Dyar & Knab (3 larvae).

Ceratopogonidae: Forcipomyia sp. or spp. (62

larvae, 2 pupae)

Chironomidae: Monopelopia tillandsia Beck &

Beck (53 larvae), (1 damaged unidentiﬁed adult).

Aulacigastridae: Stenomicra (7 larvae).

?Muscidae: genus and species unidentiﬁed (2

larvae) (perhaps this is the Neodexiopsis sp. of

Fish [1976]).

Specimens of Cecidomyiidae were shipped in

vials of alcohol to R. A. Gagné; he found both spe-

cies interesting and retained one specimen of each.

He reports that L. laticaulis is known as a preda-

tor of Diaspis echinocacti (Bouché) (Homoptera:

Diaspididae) a scale insect on Opuntia cacti—so

its presence in T. utriculata is unexpected. Larvae

of Psychodidae, Culicidae, many Ceratopogonidae,

Chironomidae, and Aulacigastridae are aquatic

and, expectedly, were found only in T. fasciculata

and T. utriculata. Identiﬁcation of these larvae

was made by J. H. Frank (who makes no claim to

be an expert on larvae of Diptera), either by prior

experience (larvae of Wyeomyia mosquitoes), by

use of keys to larvae of Chironomidae (Epler

2001), or (for the other families) according to the

brief descriptions by Fish (1976). Although 27

years have passed since Fish (1976) reported his

collections, the “Neurosystasis” (Psychodidae) and

“Stenomicra” (Aulacigastridae) occurring in Flor-

ida bromeliads have not yet been formally de-

scribed. G. J. Steck (FSCA) questioned the name

Neurosystasis (he identiﬁed them as belonging to

Telmatoscopus) and suggested contacting Larry W.

Quate (Poway, California), a specialist in the fam-

ily. Quate requested adult specimens reared from

ﬁeld-collected larvae in order to make a precise

identiﬁcation and, if necessary, prepare a formal

description. Thereupon, JHF (with help from M.

M. Cutwa and G. F. O’Meara, Florida Medical En-

tomology Laboratory, Vero Beach) obtained larvae

from bromeliads in southeastern Florida and pro-

vided them to G. J. Steck who from them reared a

few adults and shipped them to Quate in 1999-

2000. Quate reported that they represent the ﬁrst

Nearctic record for a member of the genus Alepia,

and he was pleased to see the associated larvae.

Tragically, Larry Quate died in January 2002. It

should be fairly easy to obtain more larval speci-

mens and rear more adults to replace those unre-

turned by his estate.

Hymenoptera (ants identiﬁed by M. A. Deyrup)

Formicidae: Camponotus ﬂoridanus (Buckley)

(2), Camponotus planatus Roger (26), Cremato-

gaster ashmeadi Mayr (64), Paratrechina longi-

cornis (Latreille) (50), Pheidole megacephala

(Fabricius) (5), Pheidole moerens Wheeler (129).

Frank et al.: Invertebrates from Florida Tillandsia Bromeliads 183

Ichneumonidae: genus and species unidenti-

ﬁed (2) (det. L. A. Stange, FSCA).

Aphelinidae: Aphelinus sp. (1) (see under

Hemiptera and Homoptera).

Ant nests with brood were detected in T. utric-

ulata (C. planatus and P. longicornis), T. fascicu-

lata (P. moerens), and T. setacea (C. ashmeadi).

The other ant specimens doubtless were foraging

from nests elsewhere. It has long been known

that ants will nest in the dry, outer leaf axils of

bromeliads such as T. fasciculata and T. utricu-

lata that hold water in their inner axils. One

plant of T. utriculata in Sarasota provided space

for nests of two species: C. planatus and P. longi-

cornis. Paratrechina longicornis, P. megacephala,

P. moerens, and C. planatus are adventive species.

Ants were identiﬁed from Tillandsia spp. in

various Neotropical countries and Florida by

Wheeler (1942). However, the Tillandsia were not

identiﬁed to species level, nor were the localities

in Florida nor dates of collection speciﬁed.

Table 2 arranges the collection data by sample

number, with invertebrates identiﬁed to the level

of family. This arrangement was designed to allow

extraction of numerical data for statistical analy-

sis. However, the Table suggested few patterns

that would yield useful analysis. To further com-

plicate the table by including species names

would have been unwieldy.

A simple analysis was made by contrasting the

content of the three smallest with three largest

plants within each species (Tables 1 and 2), a

valid statistical method. For T. fasciculata the

smallest plants were nos. 17, 18, and 19 (with 17,

7, and 144 invertebrates). The three largest were

8, 9, and 20 (with 19, 18, and 36 invertebrates).

The presence of an ant nest in plant 19, with 128

adult ants was the cause of the high count in a

small plant. Even if all data for ants were omitted,

the evidence for relationship of plant size to num-

ber of invertebrates would have been negligible.

For T. utriculata, the three smallest plants

were 1, 3, and 4 (with 2, 16, and 10 invertebrates),

and the three largest were 2, 16, and 24 (with 39,

226, and 135 invertebrates). In plant 24, ants ac-

counted for 76 of the invertebrates. Whether or

not we exclude data for ants, the largest plants

clearly have more invertebrates, and these were

mainly aquatic dipteran larvae (Ceratopogonidae,

Culicidae, and Psychodidae; except in plant 24). If

we exclude ants and aquatic insect larvae, the

three smallest plants had 1, 14, and 9 inverte-

brates whereas the three largest had 17, 28, and

28; again there is a relationship between plant

size and number of invertebrates, but it is shal-

lower than when including the aquatics. If we in-

clude only the aquatic invertebrates, then the 3

smallest plants had 0, 2, and 1 invertebrates,

whereas the three largest had 17, 191, and 31; the

larger plants clearly had many more, but variance

is huge. We might expect that the number of

aquatic dipteran larvae would best be associated

with volumetric capacity of bromeliad axils (calcu-

lated from length of longest leaf). But intraplant

variance in numbers of contained invertebrates

warns us that the ﬁtting of regressions will suffer

from high sums-of-squares errors. The presence of

ant nests adds greatly to variance.

For the three T. recurvata plants sampled, the

number of invertebrates was not related to plant

size. For T. setacea, the three smallest plants were

5, 6, and 13 (with 11, 19, and 18 invertebrates)

and three largest were 11, 12, and 14 (with 6, 18,

and 4 invertebrates); there was no relationship of

plant size to number of invertebrates.

ISCUSSION AND CONCLUSION

The total number of invertebrates in leaf axils

of T. utriculata was related to plant size, but the

number of aquatic insect larvae increased more

strongly with plant size. The numbers of inverte-

brates were not or not clearly related to plant size

in the other three bromeliad species, although

such a relationship is something that would be

expected given a very large number of samples

(because larger plants provide more habitat).

Data in Tables 1 and 2 could be the materials for

hundreds of regression analyses, should anyone

wish to do these.

This study scratches the surface of Florida’s

bromeliad fauna. It reafﬁrms that larvae of sev-

eral aquatic Diptera (Psychodidae, Culicidae,

Ceratopogonidae, Chironomidae, Muscidae, and

Aulacigastridae), perhaps one species of scale in-

sect (Ortheziidae), and perhaps one or more spe-

cies of Lepidoptera (Tineidae and/or Gelechiidae)

have an obligate relationship with bromeliads.

The null hypothesis for all the remaining species

is that they “just happened to be there” and may

additionally be found in tree canopies or in leaf

litter on the ground. This null hypothesis cannot

now be tested for lack of studies of the canopy

fauna or the leaf litter fauna in Myakka River

State Park.

This in no way discounts the importance of

bromeliads as habitat for large numbers of inver-

tebrate species: how many other small plants

have such a diversity of invertebrates? At least 70

families with 82 genera and 90 species are repre-

sented in the few (24) samples. Further sampling

should yield very many more species (and genera

and families) at least in Coleoptera, and perhaps

some other orders, including species that just

happen to be represented in the bromeliads at the

time of sampling.

If sampling is to be repeated, this should be (a)

with very many more samples to allow more rep-

lication and thus a more useful comparison be-

tween the faunas of the four bromeliad species,

(b) with prior agreement (probably involving

funded written subcontracts for expenditure of

184 Florida Entomologist 87(2) June 2004

time) from numerous specialist taxonomists to

devote time to the project, (c) with the collector

charged with rearing representatives of all the

immature arthropods to the adult stage. The ad-

vantage of having more samples will be the avail-

ability of a series of adult specimens of every

species represented, except perhaps a few of the

transients. The advantage of rearing the imma-

ture arthropods will be that adult specimens will

be available for identiﬁcation, and identiﬁers will

then have immature specimens reliably associ-

ated with the adults; thereafter, the specialists

may be able to provide identiﬁcation keys to the

immature stages. The collector should be proﬁ-

cient in invertebrate classiﬁcation, and should

have the time to rear immature arthropods to the

adult stage.

Raw data used to compile Table 2, on inverte-

brates associated with the 24 plants, will be of-

fered to the “Bromeliad tank dwellers database”

on the website of the Florida Council of Bromeliad

Societies (http://www.fcbs.org). It records any an-

imal species detected in or on a bromeliad, not

just tank dwellers (the aquatic species in tanks).

This could lead to detection of other animals fre-

quently associated with bromeliads, even if it

takes tens of thousands of records. It was not easy

to obtain identiﬁcations to the species level of in-

vertebrates collected from bromeliads in Florida,

and we were only partially successful, and only

for some groups. We warn investigators who

would like to conduct similar studies in the Neo-

tropics that they will encounter severe taxonomic

problems. The effort to collect the specimens is

small compared with the effort required to iden-

tify the specimens reliably to the species level.

Identiﬁcation not made to the species level is

worth rather little. Taxonomists need to be con-

vinced that the project is worth their support. In

this project, some taxonomists obtained useful

and interesting specimens, at least of Xenylla (an

undescribed species), Cerobasis guestfalica, Cam-

plomyza sp., Alepia (the ﬁrst Nearctic record),

and various mites of uncertain identity.

The sampling method did not collect micro-

scopic aquatic organisms. For these, it would be

better to use a siphon or large syringe (such as an

“oven baster”) to extract the water from leaf axils

of T. fasciculata and T. utriculata, and to decant

this water directly into Petri dishes for micro-

scopic examination. Such a method should collect

bacteria, Fungi Imperfecti, algae, rotifers, nema-

todes, platyhelminthes, annelids, ostracods and

copepods. But identiﬁcation of these would have

been beyond the skills of the taxonomists in-

volved in the present study.

Future projects of this nature in Florida with

all four of these plant species are unlikely in the

near future. This is because the weevil Metama-

sius callizona was detected in Myakka River

State Park in September 1999 and, since then,

has been relentlessly destroying the park’s popu-

lations of T. utriculata and T. fasciculata (T. M.

Cooper in Larson 2000). Similar destruction has

been detected in almost all southern Florida

counties. These two bromeliad species are right-

fully listed in the Florida Administrative Code as

endangered species. Recovery of their populations

is unlikely unless the weevil can be brought un-

der biological control.

CKNOWLEDGMENTS

We thank M. M. Cutwa and G. F. O’Meara (Florida

Medical Entomology Laboratory, Vero Beach) for collect-

ing psychodid larvae from bromeliad leaf axils, G. R.

Buckingham (USDA-ARS, Gainesville) for loan of a leaf-

area meter, G. A. Evans (FDACS-DPI, Gainesville) for

identiﬁcation of an aphelinid specimen, L. A. Stange

(FDACS-DPI, Gainesville) for family-level identiﬁca-

tion of two wasp specimens, D. H. Habeck (Entomology

and Nematology Dept., University of Florida, retired)

for identiﬁcation of lepidopteran larvae, R. J. Gagné

(Systematic Entomology Laboratory, USDA-ARS) for

identiﬁcation of cecidomyiids, R. J. Snider (Michigan

State University) for identiﬁcation of Collembola, E. E.

Baskerville and C. Gambetta (MSBG) and J. Y. Miller

(Allyn Museum of Entomology, Sarasota) for transport,

E. E. Baskerville, B. K. Holst and H. E. Luther (MSBG)

for plant identiﬁcation, H. E. Luther and J. Y. Miller for

access to literature, H. E. Luther for permission to cite

his observation on snails eating bromeliad trichome

caps, R. E. Rivero (MSBG) for lending a dissecting mi-

croscope, M. A. Blanco, E. Krötz and D. Mondragón

(MSBG) and staff of MRSP for help, B. C. Larson and

R. D. Cave (UF-IFAS) for critical reviews of the manu-

script, and R. D. Cave for preparation of a Spanish ab-

stract. This is Florida Agricultural Experiment Station

journal series R-09639.

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Bromeliad habitat regulates the richness of associated terrestrial and aquatic fauna

Article

Mar 2021

Bromeliads harbour a great diversity of fauna species. The epiphytic habit of bromeliads is subject to higher temperatures and higher incidence of light than is the epigeic habit, so we expected individuals of the Vriesea neoglutinosa species to differ in terms of the composition, richness and abundance of their fauna associated with different bromeliad habits. We carried out a study in a restinga area on Marambaia Island, Brazil. We collected 32 bromeliads, of which 16 were epigeic and 16 were epiphytic, and found a total of 1125 individual animals of 88 morphospecies. Arthropods stood out in the sample, among which 68.2% were insects. Aquatic fauna was richer in epigeic bromeliads, and terrestrial fauna was richer in epiphytic bromeliads. As expected, species composition was different for aquatic and terrestrial fauna. The heterogeneity of the restinga environment and differences in the development stages of the fauna may be factors that influence its composition. Both bromeliad habits sustain fauna with several trophic guilds that compose a complex food web and depend on the bromeliad environment. As different habits lead to different fauna types, both epigeic and epiphytic bromeliads play a role in the maintenance of local biodiversity and are important for macrofauna conservation in restingas.

Diversidad estacional de Chrysomelidae (Coleoptera) en Tillandsia recurvata L. (Bromeliaceae) en el Parque Nacional El Cimatario, Querétaro, México

Article

Full-text available

Jul 2020

RESUMEN. El género Tillandsia (Bromeliaceae) representa un grupo de plantas epífitas cuya importancia como microhábitat para los insectos ha sido documentada, pero se desconoce su relación con las variaciones estacionales en selvas secas en las comunidades de crisomélidos (Chrysomelidae: Coleoptera). En este trabajo, se documentó la fauna de Chrysomelidae presentes en Tillandsia recurvata en una selva baja caducifolia dentro del Parque Nacional El Cimatario, mediante la recolección de plantas en árboles seleccionados al azar, desde febrero de 2017 hasta junio de 2018. La variación estacional de la abundancia y diversidad de crisomélidos fue evaluada mediante pruebas no paramétricas, números de Hill, y análisis de correspondencia simple. Se encontraron 94 ejemplares de Chrysomelidae, distribuidos en cinco subfamilias, nueve géneros y 16 especies. El mayor número de ejemplares y especies se obtuvo para la subfamilia Chrysomelinae, en particular en el género Calligrapha. La abundancia y diversidad se incrementaron de forma significativa durante la temporada seca. Los resultados sugieren que algunas especies de Chrysomelidae seleccionan los microhábitats proporcionados por Tillandsia, en ausencia de lluvias y ausencia de fuentes alimenticias durante las temporadas secas.

Ecologia de campo: Serra e Litoral Catarinense

Book

Full-text available

Jan 2017

Este livro é uma publicação do Programa de Pós-Graduação em Ecologia, do Centro de Ciências Biológicas, da Universidade Federal de Santa Catarina (PPGECO-UFSC). O curso de Ecologia de Campo foi realizado de 8 a 14 de novembro de 2016 na Fa- zenda Reunidas Campos Novos, em Bom Retiro, SC, e de 15 a 21 no Parque Municipal da Lagoa do Peri, em Florianópolis, SC. A atividade de campo é uma prática fundamental ao Ecólogo. Neste curso de Eco- logia de Campo, oito Mestrandos e cinco Doutorandos realizaram trabalhos de curta duração, utilizando modelos biológicos e sistemas distintos dos quais dedicam sua formação acadêmica. O curso foi organizado a fim de proporcionar aos alunos uma imersão teórica e prática em Ecologia. Os trabalhos foram elaborados e orientados por professores do departamento de Ecologia e Zoologia da UFSC e pós-doutorandos do PPGECO-UFSC. Também participaram da elaboração e orientação dos trabalhos a Prof. Dra. Tatiana Silva Leite (Depto. De Oceanografia e Limnologia - UFRN), e os Professores Dr. Elisandro Ricardo Drechsler dos Santos e Dr. Pedro Fiaschi (Depto. de Botânica - UFSC). Neste livro você encontrará 22 publicações do 9º Curso de Ecologia de Campo do PPGECO-UFSC. Foram orientados nove trabalhos em Bom Retiro e seis trabalhos no Parque Municipal da Lagoa do Peri. Outras sete publicações são resultados de proje- tos livres, desenvolvidos em conjunto por alunos e professores. Os trabalhos foram desenvolvidos em diversos ambientes: riachos, vegetação ribeirinha e de encosta, Floresta Ombrófila Densa e Floresta Ombrófila Mista, restinga, áreas com vegetação em regeneração e sob diferentes intensidades de manejo, lagos, praias e costões ro- chosos. Foram estudadas plantas, peixes, crustáceos, macroinvertebrados aquáticos, besouros e outros insetos, artrópodes, raias, até pescadores. Esta publicação se destina a alunos de cursos de Graduação e Pós-Graduação e profissionais da área de biodiverisade e de áreas afins. Esperamos que os trabalhos aqui apresentados inspirem outros cientistas e apoiem tomadores de decisões na conserção da biodiversidade.

A record of Disparalona hamata (Birge, 1879) (Cladocera: Chydoridae) in phytotelmata of Tillandsia aguascalentensis Gardner, 1984 (Poales: Bromeliaceae)

Article

Full-text available

Mar 2019

Bromeliads (Bromeliaceae) are an extremely diverse family of the angiosperms widely distributed in the tropical and subtropical regions of the Americas and West Africa. They often serve as phytotelmata, accumulating rainwater between leaves. Such water reservoirs can be inhabited by diverse organisms. But to date not much attention is paid to inventory of these organisms, with careful identification of each taxon. We found a microcrustacean Disparalona hamata (Birge, 1879) (Crustacea: Cladocera) in the bromeliad Tillandsia aguascalentensis Gardner, 1984 in Mexico. Investigated population included parthenogenetic females, gamogenetic females and males. Hereby the population apparently can pass the full life cycle under conditions of phytotelmata. Along with ecological observations, we provide additional taxonomic notes on the genus Disparalona Fryer, 1968 itself. Recently a series of morphological revisions was conducted for this genus. It was subdivided into two subgenera: Disparalona s.str. and Mixopleuroxus Hudec, 2010. But, in fact, the second taxon is a junior synonym of Leptorhynchus Daday, 1905. In this regard here we provide an updated list of taxonomic synonyms for the subgenus Leptorhynchus.

First steps to study the demography of vascular epiphytes in cities

Article

Full-text available

Jan 2024
Braz J Biol

Urban ecosystems could jeopardize the existence of vascular epiphytes (VE) given that their occurrence is linked to phorophyte availability and particular climatic conditions. Despite reports of VE in cities, nothing is known about their demography. A first step in this direction is to describe their population structures (PS). We established the PS of VE present in urban parks in Oaxaca City (Mexico), addressing the following questions: 1) what is their demographic status? and 2) are there differences in the structure of populations growing in native versus exotic phorophytes? During 2021, we censused all the trees in six urban parks, recording their origin (native or exotic), the epiphytic species found on them and the development stages present in each VE population. Overall, five VE species were documented: Tillandsia ionantha, T. makoyana, T. sp., T. schiedeana and T. recurvata (Bromeliaceae); the first three with only one individual and the latter two with 95 and 5,694, respectively. A MANOVA test indicated significant differences in PS between T. recurvata (type I structure, suggesting a growing population) and T. schiedeana (type III structure, suggesting a senile population) (Wilkes' λ= 0.821, F-Radio= 11.96 P<0.001). PS showed no differences related to tree origin. Our results indicate that it is necessary to conduct demographic studies to have a more accurate idea of the current condition of vascular epiphytes in cities. For instance, even though we found five VS species, only one of them seems to have viable populations in Oaxaca city.

Aquatic invertebrate diversity in tank bromeliads in an enclaved wet forest in Brazil’s semiarid region

Article

Full-text available

Apr 2022

Phytotelmata bromeliads provide microhabitats for many organisms and maintain high aquatic diversity within Neotropical forests. However, they are more vulnerable to different environmental stressors that shape community structure. While most studies have been carried out in large rainforests, gaps remain about the bromeliad-dwelling community for the wettest points in the Caatinga domain (Brazilian semiarid region). Here, we investigated the influence of abiotic and temporal variables and bromeliad size on community structure between urban and forest sectors. Between 2014 and 2017, we randomly selected 20 urban and 40 forest bromeliads and compared functional groups of invertebrates for richness, abundance, and diversity. We recorded 41 morphospecies, especially Rotifera (19) and Insecta (13). Urban bromeliads exhibited higher richness of filter-feeder, while forest fragment bromeliads were dominated by predators, shredders, and gatherers. We did not observe differences in community structure regarding bromeliad size. However, the wetter period exhibited higher species richness, and higher diversity was associated with the highest rainfall. This study presents for the first time a list of aquatic microinvertebrate species and the influence of environmental parameters on the community structure associated with bromeliads of Caatinga. We have also expanded the list of species associated with phytotelmata in the Neotropical region.

A new species of Liposcelis (Insecta: Psocoptera: Liposcelididae) from Belarus

Article

Full-text available

Mar 2020

In 2019 a strikingly colored species of Liposcelis (Insecta: Psocoptera: Liposcelididae), unknown from the Palaearctic region, was collected by A. Ostrovsky in an ant's nest near Gomel city (Belarus). It is here described as Liposcelis aleksandrowiczi n. sp., a bisexual sister species to the parthenogenetic (thelytokous) New World Liposcelis ornata Mockford, 1978. The new species can be distinguished from L. ornata by slight differences in body coloration, and by some details of pronotal chaetotaxy and head pilosity.

Potential bottom‐up control of Metamasius callizona in Florida, USA

Article

Feb 2024

Metamasius callizona (Chevrolat) (Coleoptera: Curculionidae) is an invasive weevil destroying Tillandsia utriculata L. (Bromeliaceae) populations in Florida, USA. A T. utriculata population in Belize coexists with M. callizona without adverse effects. Tillandsia utriculata and M. callizona from Belize and Florida were studied in the laboratory to determine how Belize T. utriculata can coexist with Belize M. callizona, using Florida and Belize T. utriculata , as well as pineapple, Ananas comosus (L.) Merr. (Bromeliaceae), as host plants. Florida M. callizona on pineapple leaves produced more eggs per female per day and had a shorter developmental time and pupation period, as well as a smaller adult width, than Belize M. callizona . Florida and Belize weevils did not differ significantly in rates of egg hatch, pupation, and adult emergence. Florida M. callizona laid similar numbers of eggs per day on Florida and Belize T. utriculata . Egg hatch rates were similar on pineapple and Florida and Belize T. utriculata . Pupation rates were similar on pineapple and Florida T. utriculata leaves; no larvae on Belize T. utriculata attained fourth instar. Field and laboratory observations revealed that Belize and Florida M. callizona mined Belize T. utriculata without destroying the meristem, allowing plants to survive, but Belize M. callizona destroyed the core of pineapple tops. Soluble solids were greatest in pineapple and least in Belize T. utriculata . Leaf toughness was greatest in pineapple and least in Florida T. utriculata . Metamasius callizona biology and behavior and T. utriculata plant characteristics may drive the severity of weevil damage on populations of T. utriculata . The discovery of a T. utriculata population in Belize coexisting with M. callizona offers a possible way to control M. callizona in Florida.

First steps to study the demography of vascular epiphytes in cities

Preprint

Full-text available

Sep 2022

Urban ecosystems could jeopardize the existence of vascular epiphytes (VS), given that their occurrence is linked to phorophyte availability and particular climatic conditions. Despite reports of VS in cities, nothing is known about their demography. A first step in this direction is to describe their population structures (PS). We established the PS of VS present in urban parks in Oaxaca City, addressing the following questions: what is their demographic status? and are there differences in the structure of populations growing in native versus exotic phorophytes? During 2021, we censused all the trees in six urban parks, recording their origin (native or exotic), the epiphytic species found on them and the development stages present in each VS population. Overall, five VS species were documented: Tillandsia ionantha , T. makoyana , T. sp , T. recurvata and T. schiedeana ; the first three with only one individual and the latter two with 5,694 and 95, respectively. A MANOVA test indicated significant differences in PS between T. recurvata (type I structure, suggesting a growing population) and T. schiedeana (type III structure, suggesting a senile population) (Wilkes' λ = 0.821, F-Radio = 11.96 P < 0.001). PS showed no differences related to tree origin. Our results indicate that it is necessary to conduct demographic studies to have a more accurate idea of the current condition of vascular epiphytes in cities. For instance, even though we found five VS species, only one of them seems to have viable populations in Oaxaca city.

Systematics, phylogeny, evolution and biogeography of eu-Bromelioideae and genome size evolution in Bromeliaceae

Thesis

Full-text available

Jan 2018

Sascha Heller

Phylogenetic trouble unleashed The first part of my thesis deals with a comprehensive phylogeny of the Bromelioideae subfamily. The family Bromeliaceae is subdivided into eight subfamilies, one of them is the Bromelioideae. Phylogenetic relationships among the Bromelioideae are still poorly understood and many of the extant genera are suspected to be not monophyletic. Especially Aechmea, the largest and most polymorphic genus constitutes many questions and the genus was used as a depot for taxonomically problematic species. The phylogenetic study presented here is the most comprehensive one so far, covering about half of the known species (434 of 965, Table 1) of Bromelioideae. The phylogeny was generated using plastid (atpB-rbcL, matK, rps16, ycf1_1, ycf1_6) and nuclear (AGT1_exon, ETS, G3PDH, PHYC, RPB2) genetic markers. The markers were analysed individually as well as combined using maximum likelihood and Bayesian analysis. The comparison of plastid vs. nuclear data revealed significant differences which were discussed in detail and hypothesised to indicate hybridisation in certain lineages. Nevertheless, the combination of both datasets increased the overall resolution of the phylogeny and was used to discuss the results in the light of previous studies. The entire phylogeny was divided into 32 groups for discussion. These groups represent potential genera or starting points for further studies in order to reorganise the polyphyletic genera of Bromelioideae into monophyletic lineages. Many extant genera of the eu-Bromelioideae were found to be not monophyletic. Monophyly was observed for the genera Acanthostachys, Billbergia, Cryptanthus, Disteganthus, Hoplocrypanthus, Lapanthus, Orthocryptanthus, Orthophytum, Rokautskyia, Ronnbergia, Sincoraea, Wittmackia and the monotypic ones (Deinacanthon, Eduandrea, Fascicularia, Hohenbergiopsis, Pseudananas). The genus concept proposed by Smith and Downs (1979) is therefore rejected, as well as the taxonomic utility of petal appendages, which were mainly used to delimit genera. In summary, this study and recent studies highlighted other morphological characters (e.g. pollen morphology, stigma type) as much more informative. However, no single character should be used to delimit genera and combinations of relevant characters are required. Even the petal appendages can pose a taxonomical important character at certain taxonomic level. The combination of biogeography and phylogeny revealed that species of some groups which co- occur in a biome or region are also phylogenetically closely related. These groups were not recognised before because the misinterpretation of homoplastic characters led to wrong taxonomical conclusion. For example, the recent re-organisation of the Cryptanthoid group and the re-establishment of Wittmackia with the former Hohenbergia subgen. Wittmackiopsis species highlighted, among other characters, the importance of biogeography. Another case is the subgenus Neoregelia subgen. Hylaeaicum which is geographically and phylogenetically separated from the Nidularioid group and therefore has to be excluded. 5 The large phylogeny presented here gives evidence for multiple invasions of the Brazilian biomes (Amazon Forest, Atlantic Forest, Cerrado, Caatinga) as well as of Central America and the Greater Antilles. It is important to note that the phylogeny is lacking resolution in the deeper nodes. Confident assumptions are therefore hindered and the historical biogeography of Bromelioideae remains cryptic. Anyway, the Atlantic Forest is nowadays the diversity hotspot of the core Bromelioideae and critically endangered. Extensive conservation efforts are required to protect the diverse flora, including the bromeliads. The genetic markers used so far in bromeliad phylogenies provided only limited variation resulting in often unresolved complexes. The search for additional suitable genetic markers in bromelioid phylogenies yielded the nuclear marker AGT1. The amplified fragment consists of one well conserved exon region as well as a highly variable intron. The intron was too variable for aligning it across the entire bromelioid set. On the other hand, the intron provides relevant information for inferring phylogenies of closely related species groups (e.g. in Ananas, Cryptanthoid group). Furthermore, AGT1 is proposed as a genetic barcode in Bromelioideae because it poses much more information then the commonly used ones (e.g. matK). Does size matter? The second part of this thesis deals with the genome size evolution within the family Bromeliaceae. Samples from seven subfamilies were screened with the emphasis on the subfamily Bromelioideae. The data were combined with data from literature and the observed patterns were discussed in relation to known phenomena (e.g. correlations to environment and life form). In the second sub-chapter I have chosen the species Tillandsia usneoides to study the intraspecific genome size variation in combination with morphology and biogeography. Genome size and base composition were measured using the flow cytometry technique. Bromeliaceae comprises mostly diploid species with predominantly 50 small chromosomes (2n), small genome sizes (0.59-4.11 pg) and normal GC content (36.46-42.21 %) compared to other families. Polyploidy was observed so far in the subfamilies Bromelioideae, Tillandsioideae and Pitcairnioideae. Triploids, tetraploids and potential hexaploids were identified. The genera show significant differences in holoploid genome size and base composition throughout the entire family. GC content is weakly positively correlated with genome size. Significant intraspecific genome size variation has been observed, including polyploidization, but no endopolyploidy and no variation in dioecious species. Within the subfamily Bromelioideae, the observed genome size between the early diverging lineages and the core Bromelioideae supports this division. The differences are due to a higher proportion of polyploids in the early diverging lineages and a significant higher 6 GC content in the core Bromelioideae. Both groups differ in their life strategies and occupy principally different habitats with corresponding morphological adaptations. Hence, the early diverging lineages are predominantly terrestrial and xeromorphic. In contrast, the prevailing epiphytic core Bromelioideae are characterised by a tank habit and mostly adapted to more humid environments. Across the family and the subfamily Bromelioideae in particular, significant genome size differences between the different life forms have been observed, but no correlation to biomes within Brazil. Tillandsia usneoides is the most widely distributed species of the family Bromeliaceae. It ranges from the southeastern United States to Argentina and Chile. Tillandsia usneoides grows epiphytic and is dispersed by seeds as well as by fragments of the plant. Within the species striking morphological differences can be observed as far as size characters are concerned. Morphotypes have shown to be stable in cultivation while growing under the same conditions. In order to investigate possible reasons for the variation the relative genome size of 75 specimens covering the whole distribution range was measured and combined with morphological, distribution and climatic data. Significant variation in the relative genome size corresponded to the morphological differences and reflected the north-south distribution gradient. Genome size and morphotypes showed a positive correlation, as well as with the mean temperature of the driest and coldest quarter and the minimal temperature of the coldest month.

Bromeliad-eating weevils.

Article

Full-text available

Jan 1999

J.Howard Frank

On the bionomics of bromeliad-inhabiting mosquitoes. I. Some factors influencing oviposition by Wyeomyia vanduzeei.

Article

Full-text available

Jan 1976

Collembola asociados a Tillandsia (Bromeliacea) en el Derrame Lávico del Chichinautzin, Morelos, México

Article

Full-text available

Jan 1981
SOUTHWEST ENTOMOL

José G. Palacios-Vargas

Collembola associated to Tillandsia Bromeliaceae in Morelos, is presented

Bionomics of the Bromeliad-Inhabiting Mosquito Wyeomyia vanduzeei and Its Nursery Plant Tillandsia utriculata

Article

Full-text available

Dec 1981

Aquatic stages of the mosquito Wyeomyia vanduzeei Dyar & Knab inhabit leaf axils of the epiphytic bromeliad Tillandsia utriculata L. in southern Florida. Rough-barked trees provide a substrate for T. utriculata and a habitat for W. vanduzeei. The volumetric capacity of T. utriculata leaf axils can be estimated from length of longest leaf (volumetric capacity in ml = 0.003251 × leaf length in cm2.7799). In each plant, older, larger axils provide most of the volumetric capacity, but the outermost axils lose their ability to retain water. Water entering bromeliad axils is throughfall, i.e. rainfall reduced in volume but enriched with nutrients as it penetrates the tree canopy (throughfall mm = --0.2715 + (0.8825 × rainfall mm) in a high marsh study area). Organic debris from the tree canopy (estimated annual mean 2.4 g/m2/day in a high marsh study area), trapped and decomposed in the bromeliad axils, also provides nutrients for aquatic organisms and for the bromeliad. The perennial bromeliad provides a stable habitat for mosquito larvae. The number of mosquito eggs laid in axils is influenced by the size, water content, senescence and flower production of the bromeliad; as many are laid in the smaller, inner axils as in the larger, outer axils. At 27°C, most floating eggs hatched within 48 and 96 h. Eggs stranded out of water suffered loss in numbers and in viability, with combined loss at 10 days of 49%, at 20 days of 77%, and at 30 days of 95% /// En el sur de la Florida (EUA), los estadios acuáticos del mosquito Wyeomyia vanduzeei Dyar & Knab habitan en las axilas de las hojas de la bromelia epifítica Tillandsia utriculata L. Arboles con la corteza rugosa proveen un substrato para T. utriculata y un habitat para W. vanduzeei. La capacidad volumétrica de las axilas de las hojas de T. utriculata puede ser estimada del largo de la hoja mas larga (capacidad volumétrica en ml = 0.003251 × largo de la hoja en cm2.7799). En cada planta, las axilas mas viejas y mayores proveen la mayoría de su capacidad volumétrica, pero las axilas exteriores pierden la capacidad de retener agua. El agua que entra en las axilas es la caída pasante, es decir, lluvia reducida en volumen, pero enriquecida con nutrientes al penetrar la bóveda del follage (caida pasante en mm = --0.2715 + (0.8825 × mm de lluvia) en una zona de estudios en la parte mas elevada de una marisma.) Escombro orgánico de la bóveda de los árboles (caida media anual 2.4 g/m2/dia en una zona de estudios en la parte elevada de una marisma), atrapado y descompuesto en las axilas de las bromelias también provee nutrientes a los organismos acuáticos y a las bromelias. La bromelia perenne provee un habitat estable para larvas de mosquitos. El número de huevos de mosquitos depositado en las axilas es influenciado por el tamaño, contenido de agua, senectud, y producción de flores de la bromelia; la misma cantidad de huevos es depositada en las mas pequeñas axilas interiores que en las mayores axilas exteriores. A 27°C, la mayoría de los huevos flotantes eclosionan dentro de un periodo de 48-96 h. Huevos varados fuera del agua sufrieron pérdidas en números y en viabilidad, con pérdidas combinadas en 10 dias de 49%, en 20 dias de 77%, y en 30 dias de 95%.

Florida's Native Bromeliads

Article

May 2019

Bromeliads are members of the pineapple family Bromeliaceae. They are perennial herbs that lack woody stems and typically grow on other plants or substrates. Bromeliads are not mosses as some of their common names suggest. They are flowering plants, although their blossoms can be very small. This document is CIR 1466, one of a series of the Wildlife Ecology and Conservation Department, UF/IFAS Extension. Original publication date September 2004. CIR 1466/UW205: Florida's Native Bromeliads (ufl.edu)

Studies of neotropical ant-plants and their ants

Article